Successional Stages in the Development of Forest Vegetation in Cirques of Two Valleys in the Julian Alps (Nw Slovenia)

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

http://dx.doi.org/10.3986/fbg0052

ABSTRACT IZVLEČEK Successional stages in the development of forest vegeta- Sukcesijske stopnje v razvoju gozdne vegetacije v krnicah tion in cirques of two valleys in the Julian Alps (NW Slo- dveh alpskih dolin v Julijskih Alpah (severozahodna venia) Slovenija) We investigated different types of vegetation in cirques Raziskali smo različne oblike vegetacije v krnicah dveh of two valleys in the Julian Alps, Zadnjica and Kot, which dolin v Julijskih Alpah, Zadnjici in Kotu, ki nastajajo na develop on slope screes and are heavily influenced by ava- pobočnem grušču in nanje odločilno vplivajo vsakoletni lanches that occur here every year. Our phytosociological snežni plazovi. S fitocenološko analizo smo lahko prepozna- analysis identified the following shrub-forest developmental li naslednje grmiščno-gozdne razvojne stopnje: Saliceteum stages: Saliceteum eleagno-purpureae hieracietosum porrifolii eleagno-purpureae hieracietosum porrifolii var. Achnatherum var. Achnatherum calamagrostis, Homogyno sylvestris-Sali- calamagrostis, Homogyno sylvestris-Salicetum glabrae, Poly- cetum glabrae, Polysticho lonchitis-Rhamnetum fallacis, Rho- sticho lonchitis-Rhamnetum fallacis, Rhododendro hirsuti- dodendro hirsuti-Laburnetum alpini, Rhodothamno-Larice- -Laburnetum alpini, Rhodothamno-Laricetum deciduae ane- tum deciduae anemonetosum trifoliae and Rhododendro hir- monetosum trifoliae in Rhododendro hirsuti-Fagetum sylvati- suti-Fagetum sylvaticae var. Sorbus chamaemespilus. On un- cae var. Sorbus chamaemespilus. Na nesprijetih rečno- consolidated fluvio-glacial deposits in the Kot Valley we ledeniško nanosih v dolini Kot pa smo ugotovili naslednje determined the following successional stages: Salicetum ele- sukcesijske stopnje: Salicetum eleagno-purpurae hieracieto- agno-purpurae hieracietosum porrifolii var. Pinus mugo, sum porrifolii var. Pinus mugo, Amelanchiero ovalis-Pinetum Amelanchiero ovalis-Pinetum mugo laricetosum and Rhodo- mugo laricetosum in Rhododendro hirsuti-Fagetum sylvaticae dendro hirsuti-Fagetum sylvaticae petasitetosum paradoxi. petasitetosum paradoxi. Key words: phytosociology, succession, glacial valley, Ključne besede: fitocenologija, sukcesija, ledeniška doli- Julian Alps, Natura 2000, Triglav National Park, Slovenia na, 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, [email protected]

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 develop- of vegetation on slope screes and alluvial fans in their ment of forest vegetation in Alpine valleys in Slovenia cirques, where annual avalanche activity is obviously were studied in particular by M. Zupančič and collab- one the main ecological factors. The human impact on orators (Zupančič & Žagar 2009, Zupančič, Skuma- the natural processes can be seen only in Zadnjica (the vec & Mihorič 2017) as well as several geographers vicinity of a marked mountain trail) and is insignifi- (Lovrenčak 2002, Blatnik & Repe 2013, Geršič et cant in the gable end of the Kot Valley (above the Ko- al. 2014). On our sporadic visits to the mountains tarica falls). We made a phytosociological inventory of through the Alpine valleys of Zadnjica in Trenta and these, spatially relatively well-delimited shrub and for- Kot in the Upper Sava Valley (both in the Triglav Na- est communities and tried to classify them into a syn- tional Park) our attention was drawn to different types 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

Seliškar, Vreš & A. Seliškar 2003). The relevés were 2.1 Ecological description of the study area arranged in two tables. Those on slope screes in the cirques of Zadnjica and Kot (above the Kotarica falls) Both cirques, in Zadnjica and Kot, have a predomi- and the relevés on gravelly plains in the upper part of nantly shady aspect (northeast and partly northwest); Kot (Klin) were analysed separately. They were mutu- parent material is unconsolidated moraine (till) and ally compared by means of hierarchical classification, above all slope screes (Jurkovšek 1987 a, b). The ele- using the (unweighted) average linkage – UPGMA vation of both cirques is similar, spanning 1000 to method with Wishart’s similarity ratio. For this pur- 1300 m. There are no significant differences in the pre- pose, we transformed combined cover-abundance val- cipitation volume either, with the average exceeding ues into ordinal values 1– 9 (van der Maarel 1979). 2000 mm (B. Zupančič 1998), a large portion of which We used SYN-TAX 2000 (Podani 2001) software falls as snow. The mean annual temperature is similar package. The nomenclatural source for the names of too, ranging between 4 and 6 °C (Cegnar 1998), with vascular plants is Martinčič & al. (2007), except for the Zadnjica Valley still showing some sub-Mediterra- Sesleria caerulea (Aeschimann et al. 2004b). Martin- nean influence. In recent decades the area has seen a čič (2003, 2011) is the nomenclatural source for the distinctly downward trend in precipitation and higher names of mosses and Suppan et al. (2000) for the average temperatures (Dolinar 2018). Both cirques names of lichens. The nomenclatural source for the have a characteristically cold local climate and over names of syntaxa are Šilc & Čarni (2012), except for the winter they receive snow avalanches from their the name of the class Querco-Fagetea Braun-Blanquet rock walls or steps (Volk Bahun, Zorn & Pavšek et Vlieger in Vlieger 1937. 2018). The soil is therefore very shallow, partly colluvial and undeveloped, lithosol and moder rendzina.

3 RESULTS AND DISCUSSION

3.1 Phytosociological analysis of the shrub-for- Slovene Alps than previously thought (see also Bla- est vegetation sequence on slope screes in the tnik & Repe 2013). In recent years it was spotted, but avalanche area in the gable end of the Zadnjica not yet inventoried, also in the Krma Valley. The dom- and Kot valleys inant shrub layer species in one of the Rhamnus fallax stands in the gable of Kot (relevé 2 in Table 1) is Salix Table 1 comprises 22 relevés of stands on slope glabra, and this stand could be classified also into the screes that grouped by floristic similarity as indicated association Homogyno sylvestris-Salicetum glabrae in Figure 2. (comp. Dakskobler & Surina 2017). Similarly, larch In Table 1 we arranged the relevés by taking into dominated the highest stand layers only in the gable account the dominant species of the highest stand end of Kot, on one location on fully consolidated grav- layer, which means that our classification does not el. Its initial stand (relevé 6 in Table 1) can be classified fully follow the results of hierarchical classification. into the association Rhodothamno-Laricetum. Al- The most unique form, the community of red and grey though Anemone trifolia was not found in the recorded willow (Salicetum eleagno-purpureae hieracietosum por- stand, based on the whole floristic composition it rifolii var. Achnatherum calamagrostis) with Achnathe- should be classified into the subassociation anemone- rum calamagrostis, Athamanta cretensis and Petasites tosum trifoliae, which is characterised by abundant Se- paradoxus dominating the herb layer, was found only sleria caerulea in the herb layer (comp. Dakskobler, along a small torrential brook running from under the Seliškar & Rozman 2018: 259) waterfall in the Kanjavec rock face down the slope scree Most relevés on the left side of the dendrogram in until it sinks (relevé 1in Table 1). Figure 2, i.e. relevés in the central part of Table 1, char- Open Rhamnus fallax shrub stands were found acterise pioneer shrub communities with Laburnum only in the gable end of Kot, on rather coarse and not alpinum dominating the highest stand layer. Labur- entirely consolidated gravel – relevés 3–5 in Table 1. In num alpinum is a south-European montane species terms of species composition they obviously belong to distributed across most of the Alps (except the North- the association Polysticho lonchitis-Rhamnetum fallacis, eastern and partly Central Alps), in the south and cen- which was described a few years ago (Dakskobler, tral Jura, the north and central Apennines, the Car- Franz & Rozman 2013) and is more frequent in the pathians and the Dinaric Alps (Aeschimann et al.

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 103 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

2004a, Brus 1999, 2005). It occurs on calcareous, mixed tum, Saxifrago petraeae-Tilietum, Seslerio albicantis- and silicate bedrock, in the colline, montane and partly -Ostryetum carpinifoliae, Fraxino orni-Ostryetum car- the subalpine belt, mainly on beech forest sites. As a pinifoliae, Rhododendro hirsuti-Ostryetum carpinifoli- shade-tolerant species it frequently occurs in the lower ae, Rhodothamno-Pinetum mugo, Amelanchiero ovalis- tree and shrub layer, and establishes itself as a pioneer -Pinetum mugo, but it occurs also in several other for- in windthrow and landslide areas, and on erosion est and shrub communities (source: FloVegSi database, slopes. Figure 3 shows its distribution in Slovenia. T. Seliškar, Vreš & A. Seliškar, 2003). The highest it Laburnum alpinum is the most frequent in the was recorded in the association Rhodothamno-Pine- stands of associations Arunco-Fagetum, Ostryo-Fage- tum mugo at 1560 m a.s.l. (pasture Stržiškarska Plani- tum, Seslerio autumnalis-Fagetum, Lamio orvalae-Fage- na under Mt. Poljanski Vrh and Mt. Hohkovbl), in the tum, Ranunculo platanifolii-Fagetum, Anemono trifoli- association Polysticho lonchitis-Fagetum at 1570 m ae-Fagetum, Homogyno sylvestris-Fagetum, Saxifrago (Planinica under Loška Stena rock wall), in the subas- cuneifolii-Fagetum, Rhododendro hirsuti-Fagetum, sociation Anemono-Fagetum laricetosum at 1615 m Polysticho lonchitis-Fagetum, Luzulo-Fagetum sylvati- (Pod Pečmi above the Tamar Valley, under Srednja cae, Laburno alpini-Piceetum, Rhodothamno-Larice- Ponca), in the association Rhodothamno-Laricetum at

Kot-Zadnjica

0,85 0,8 0,75 0,7 0,65 0,6 0,55 0,5 0,45 0,4

Dissimilarity 0,35 0,3 0,25 0,2 0,15 0,1 0,05 0 RL Sep RhF RhF RhF RhF PlRf PlRf PlRf HsSa RhLa RhLa RhLa RhLa RhLa RhLa RhLa RhLa RhLa RhLa RhLa ChLa

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

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; litt.). In the Apennines, a successional stage Geranio under the peak Kumlehova Glava above the Suha nodosi-Laburnetum alpini Castelli, Biondi & Ballelli Pišnica brook). These are the highest elevations of its 2001 and its subassociation sorbetosum ariae Biondi, occurrence known to us in the Julian Alps. Brus (1999: Pinzi et Gubellini 2004 were described on the sites of 206, 2005: 241) reports that on southern (the warmest, beech forests from the association Cardamino kitaibe- sunny) slopes this species occurs up to 1900 m a.s.l., lii-Fagetum sylvaticae (Castelli, Biondi & Ballelli which almost certainly does not refer to the Julian 2001, Biondi, Pinzi & Gubellini 2004). Alps, but to the Alps in general, (Brus, in litt.). As a Our relevés are distinctly different from the shrub pioneer it establishes itself on windthrow areas, in par- communities in the Jura and the Apennines. The spe- ticular in stands of associations Anemono trifoliae-Fa- cies shared with the stands of the association Sorbo getum, Ranunculo platanifolii-Fagetum and Rhododen- mougeotii-Laburnetum alpini from the Jura are Labur- dro hirsuti-Fagetum. In the northern part of the Di- num alpinum, Sorbus mougeotii, S. aria, S. chamaeme- naric Alps we described a successional stage in natural spilus, Salix appendiculata, Fagus sylvatica, Lonicera gaps of montane beech and fir-beech forest as the as- xylosteum, Mercurialis perennis, Acer pseudoplatanus, sociation Laburno alpini-Rhamnetum fallacis Polygonatum verticillatum, Fragaria vesca, Rubus idae- (Dakskobler, Franz & Rozman 2013). In the Jura, us, Lonicera alpigena and Aconitum lycoctonum s. lat. the association Sorbo mougeotii-Laburnetum alpini The species shared with the stands of the subassocia- Géhu & Géhu-Franck ex Foucault 2012 is described tion Gerenio nodosi-Laburnetum alpini sorbetosoum and designated as the edge shrub community on the ariae from the Apennines are Laburnum alpinum, Sor- sites of thermo-basophilous beech forests (Richard bus aria, Rhamnus fallax, Mercurialis perennis, Fagus 1968, Géhu & Géhu-Franck 1987, Biondi, Pinzi & sylvatica, Solidago virgaurea, Hepatica nobilis, Carex Gubellini 2004, Foucault 2012, Theurillat 2019, in digitata, Ostrya carpinifolia, Fragaria vesca, Thalic-

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 105 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

trum aquilegiifolium, Euonymus latifolia, Galium on the debris cone under Mt. Zadnjiški Ozebnik. It is album and Hypericum montanum. Despite the listed characterised by character scree and stony grassland species that they have in common, the studied Labur- species such as Achnatherum calamagrostis, Ade- num alpinum community from the Julian Alps differs nostyles glabra, Calamagrostis varia and Carex humilis. from its communities from the Jura and the Apen- This stand is tentatively classified into the provisional nines mainly with the presence of distinctly frigophi- association Carici humilis-Laburnetum alpini nom. lous species characteristic for subalpine beech, larch, prov. and is syndynamically related to the stands of the spruce and dwarf pine communities, such as Rhodo- association Anemono trifoliae-Fagetum. dendron hirsutum, Polystichum lonchitis, Pinus mugo, The tree layer in the forest stands located on the Clematis alpina, Homogyne sylvestris, Picea abies and edge or at the bottom of the studied glacial cirques, in Larix decidua. Most of the stands with dominating La- areas that are less exposed to avalanches (relevés 19–22 burnum alpinum in the highest layer are therefore clas- in Table 1), is dominated by beech. The entire species sified into the new association Rhododendro hirsuti- composition indicates a community on extreme sites, -Laburnetum alpini. Its nomenclatural type, holotypus, very similar to beech stands on the upper forest line, is relevé 16 in Table 1. The diagnostic species of the classified into the association Polysticho lonchitis-Fage- new association are Laburnum alpinum, Rhododen- tum. Also possible is classification into the association dron hirsutum, Polystichum lonchitis, Pinus mugo, Rhododendro hirsuti-Fagetum, as these stands com- Gymnocarpium robertianum, Clematis alpina, Homo- prise almost all of the diagnostic species of this asso- gyne sylvestris, Larix decidua, Sorbus chamaemespilus ciation that otherwise characterises beech sites in the and Salix appendiculata and its phytogeographical dif- submontane and montane belt up to some 1200 m a.s.l. ferential species are Rhodothamnus chamaecistus, Given the elevation of our relevés (under 1200 m) the Cyclamen purpurascens and Anemone trifolia. The latter association is considered more appropriate, but listed species characterise cold, stony sites where beech to demonstrate the similarity with the stands of the can still establish itself in favourable conditions (ab- first, Polysticho lonchitis-Fagetum, it is described as a sence of avalanches), as manifested by the entire spe- special altitudinal variant with Sorbus chamaemespilus cies composition. The new association is thus classified (Rhododendro hirsuti-Fagetum var. Sorbus chamaeme- into the alliance Aremonio-Fagion, order Fagetalia syl- spilus). In the Julian Alps, this species otherwise char- vaticae and class Querco-Fagetea (Carpino-Fagetea), acterises communities of the altimontane and subal- even though it could also be classified into the classes pine belts, and in the (lower) montane belt it is usually Erico-Pinetea or Roso pendulinae-Pinetea mugo. limited to frost hollows. In the stands of the associa- There are several differences between the stands in tion Rhododendro hirsuti-Fagetum in the Julian Alps it the Zadnjica and Kot valleys. The stands in Kot on was reported on only a few occasions (see Dakskobler moister and coarser gravel are characterised in partic- 2003). ular by a considerable admixture of sycamore maple From the relevés in Table 1 we can infer the likely (Acer pseudoplatanus), so they are described as a new sequence in the development of shrub and forest vege- subassociation aceretosum pseudoplatani (its nomen- tation on slope screes in avalanche areas in cirques of clatural type is the same as the nomenclatural type of glacial valleys in the Julian Alps: the new association, relevé 16 in Table 1). The stands in Polysticho lonchitis-Rhamnetum fallacis (Homo- the gable end of the Zadnjica Valley are mainly situat- gyno sylvestris-Salicetum glabrae) – Rhododendro hir- ed on finer gravel, on slightly drier and warmer sites, suti-Laburnetum alpini (Amelanchiero-Pinetum mugo) and are differentiated by Ostrya carpinifolia and Carex – Rhododendro hirsuti-Fagetum sylvaticae (Rhodotha- alba. The nomenclatural type, holotypus, of the new mno-Laricetum). 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 3.2 Phytosociological analysis of the sequence completely dominated by dwarf pine (Pinus mugo). Al- in shrub-forest vegetation on fluvio-glacial de- though this stand could also be classified into the as- posits in the Kot Valley (Klin). sociation Amelanchiero-Pinetum mugo, it was included in the association Rhododendro-Laburnetum alpini, In Table 2 we arranged six relevés made on the because in terms of entire species composition it gravelly plain Klin in the upper part of the Kot Valley. grouped with other relevés of the latter association. They were mutually compared using hierarchical clas- However, the same cannot apply to relevé 7 in Table 1, sification, which produced the dendrogram in Figure which characterises an open Laburnum alpinum shrub 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 inventoried several typical stands and contact commu- upper part of the Kot Valley was studied in detail by nities of Pinus mugo and Salix eleagnos. Only six rele- Zupančič, Skumavec & Mihorič (2017). They deter- vés, however, are not sufficient to allow us to infer the mined and presented with a phytosociological table right sequence in the development of shrub-forest veg- the sequence of the following plant communities: etation. Our goal is therefore not to correct or supple- Petasitetum paradoxi – Petasiti-Salicetum purpu- ment the findings of the much more detailed recent reo-albae – Calamagrostio variae-Pinetum mugo – analysis, but merely to classify our relevés into a syn- Dryado-Piceetum – Rhodothamno-Pinetum mugo – taxonomic system. Rhododendro-Fagetum – Anemono-Fagetum lycopodie- Relevé 1 in Table 1 could be classified also into the tosum. association Salicetum eleagno-purpureae. Although a In this area, i.e. on the gravelly plain under the Ko- comparison with the relevés of this community from tarica falls, we were interested mainly in beech stands the foothills of the Julian Alps, in the Nadiža Valley with hairy alpenrose (Rhododendro hirsuti-Fagetum), (Šilc & Čušin 2000) does indicate some distinct differ- as the findings by Zupančič et al. (ibid.) were new to ences, also with the variant with Petasites paradoxus. the forest vegetation in this part of the Julian Alps. We Oriolo & Poldini (2002) classify slightly similar

Kot-Klin

0,75 0,7 0,65 0,6 0,55 0,5 0,45 0,4 0,35 Dissimilarity 0,3 0,25 0,2 0,15 0,1 0,05 0 Sep AoPm AoPm RhFpp RhFpp RhFpp

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

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 107 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

stands into the montane altitudinal form of the subas- with certain ecological characteristics. The stands of sociation typicum. There is also a certain similarity this association dominate on steep to precipitous shady with relevés of the subassociation Salicetum eleagno- slopes, whereas ours were found on levelled terrain -purpureae hieracietosum porrifolii described by Čušin and gravel deposits. Based on the above, they can be & Šilc (2006) on gravel bars of the Soča River near described and typified as a new subassociation Rhodo- Bovec. The species that differentiate our relevé against dendro hirsuti-Fagetum sylvaticae petasitetosum para- the relevés published by Šilc & Čušin (2000) and Ori- doxi subass. nov. The nomenclatural type, holotypus, is olo & Poldini (2002) are in particular Pinus mugo, relevé 6 in Table 2. The differential species of the new Hieracium porrifolium, Rhododendron hirsutum, Rha- subassociation are Petasites paradoxus and Brachypo- mnus fallax, Salix waldsteiniana, Dryas octopetala, Pri- dium rupestre. mula clusiana and many species of the classes Elyno-Se- slerietea and Thlaspietea rotundifolii. Some of the diagnostic species shared with the stands of the subassocia- 3.3 An overview of syntax described or dis- tion -hieracietosum porrifolii are Hieracium porrifolium, cussed in the article: H. piloselloides, Carduus crassifolius, Biscutella laevigata, Campanula cespitosa, Silene vulgaris subsp. glareosa, Salicetea purpureae Moor 1958 Achnatherum calamagrostis, Rumex scutatus, Sesleria Salicetalia purpureae Moor 1958 caerulea and Dryas octopetala. Our relevé at least slight- Salicion eleagno-daphnoidis (Moor 1958) Grass 1993 ly resembles also the relevé of the syntaxon Petasiti-Sali- Salicetum eleagno-purpureae Sillinger 1933 cetum purpureo-albae (Zupančič, Skumavec & Miho- -hieracietosum porrifolii Čušin et Šilc 2006 rič 2017, relevé 2 in the phytosociological table), but var. Pinus mugo var. nov. with a significant difference – the dominant species var. Achnatherum calamagrostis var. nov. there is Salix eleagnos and not Salix alba – which is a Querco-Fagetea Br.-Bl. & Vlieg. 1937 result of different site conditions (M. Zupančič, in litt.). Rhamnetali fallacis Fukarek 1969 The above comparisons allow us to classify our relevé Seslerio calcariae-Rhamnion fallacis Dakskobler, into the new variant Salicetum eleagno-purpureae hiera- Franz et Rozman 2013 cietosum porrifolii var. Pinus mugo var. nov. The differ- Polysticho lonchitis-Rhamnetum fallacis Dakskob- ential species of the new variant are Pinus mugo, Rhodo- ler, Franz et Rozman 2013 dendron hirsutum, Salix waldsteiniana and Potentilla Fagetalia sylvaticae Pawł. in Pawł. & al. 1928 clusiana. Aremonio-Fagion (Ht. 1938) Borhidi in Török, Po- Relevés 2 and 3 in Table 2 can be classified into the dani & Borhidi 1989 association Amelanchiero ovalis-Pinetum mugo, as it Rhododendro hirsuti-Laburnetum alpini ass. nov. comprises several its diagnostic species, but not Fraxi- hoc loco nus ornus and Ostrya carpinifolia. Another option -ostryetosum carpinifoliae subass. nov. hoc loco would be classification into the association Rhodotha- -aceretosum pseudoplatani subass. nov. hoc loco mno-Pinetum mugo. Given the elevation of occurrence Carici humilis-Laburnetum alpini nom. prov. we prefer (like in the case of similar associations Rho- Rhododendro hirsuti-Fagetum sylvaticae Accetto ex dodendro hirsuti-Fagetum / Polysticho lonchitis-Fage- Dakskobler 1998 tum) the dwarf pine community of lower altitudes -petasitetosum paradoxi subass. nov. hoc loco (Amelanchiero-Pinetum mugo), indicating its similari- Vaccinio-Piceetea Br.-Bl. in Br.-Bl., Sissingh et Vlieger ty with the stands of the association Rhodothamno-Pi- 1939 netum mugo with the provisional name of the subas- Piceetalia excelsae Pawłowski in Pawłowski et al. 1928 sociation, laricetosum. Erico-Pinion mugo Leibundgut 1948 Stands in relevés 4–6 can be classified into the as- Rhodothamno-Laricetum deciduae Willner et Zuk sociation Rhododendro hirsuti-Fagetum sylvaticae, as rigl 1999 they comprise most of its diagnostic species. However, -anemonetosum trifoliae Dakskobler 2006 these stands could also be classified into the associa- Erico-Pinetea Horvat 1959 tion Polysticho lonchitis-Fagetum. The species support- Erico-Pinetalia Horvat 1959 ing this classification is Sorbus chamaemepsilus, so Fraxino orni-Pinion nigrae-sylvestris Zupančič 2007 these stands can also be treated within the already Helleboro nigri-Pinenion (Horvat 1959) Zupančič mentioned altitudinal variant. On the other hand, the 2007 studied stands differ from other previously described Amelanchiero-Pinetum mugo Minghetti in Pe- forms of the association Rhododendro hirsuti-Fagetum 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. Ericion carneae Rübel ex Grabherr, Greimler et Rhododendro hirsuti-Ericetea carneae Schubert et al. Mucina 1993 2001 Homogyno sylvestris-Salicetum glabrae Dakskobler Rhododendro hirsuti-Ericetalia carneae Grabherr, et Surina 2017 Greim ler et Mucina 1993

4 CONCLUSIONS

On slope screes and gravel deposits in the gable end of Kot valleys are characterised by their similarity with two glacial valleys in the Julian Alps, in Zadnjica and stands of the association Polysticho lonchitis-Fagetum Kot (Triglav National Park), we phytosociologically (and are therefore treated as a special altitudinal vari- described several shrub and forest communities whose ant with Sorbus chamaemespilus). Its form on fluvio- development is affected not only by the size, consolida- glacial gravel deposits that can be treated as a new sub- tion and moisture level of the gravel, but most deci- association petasitetosum paradoxi was found only on sively by avalanches that occur there every year. Only levelled terrain in the Kot Valley. Our findings con- two shrub and one forest community were document- firm and supplement the findings of Zupančič, Sku- ed with a sufficient number of relevés. Stands of asso- mavec & Mihorič (2017) on the exceptionally diverse ciations Polysticho lonchitis-Rhamnetum fallacis and and fascinating shrub-forest vegetation in the gable Amelanchiero ovalis-Pinetum mugo are new to the Kot end of the Kot Valley and its substantial biotic value. Valley. We described a new association Rhododendro Most of this vegetation belongs to Natura 2000 habitat hirsuti-Laburnetum alpini, which belongs to the group types: 91K0 Illyrian Fagus sylvatica forests (Aremonio- of shrub-forest communities of the montane-subalpine -Fagion), 4070 Pinus mugo and Rhododendrum hirsu- belt (Rhododendro hirsuti-Sorbetum aucupariae, Rho- tum shrubs (Mugo-Rhododendretum hirsuti) and 9420 dodendro hirsuti-Salicetum appendiculatae, Rhododen- Alpine Larix decidua forests. It comprises several pro- dro hirsuti-Betuletum carpaticae, Alno viridis-Sorbe- tected (Cypripedium calceolus – also a Natura 2000 tum aucupariae, Calamagrostio arundinaceae-Sorbe- species, Gymnadenia odoratissima, G. conopsea, Epi- tum ariae) – comp. Dakskobler, Kutnar & Rozman pactis helleborine, Dianthus sternbergii, Gentiana clu- (2016) that are a more or less long-term stages on po- sii, Lycopodium annotinum), endemic (Cerastium sub- tential beech sites and their occurrence could, although triflorum, Papaver alpinum subsp. ernesti-mayeri = P. not necessarily, be associated with human impact or ernesti-mayeri) and rare species (Sorbus mougeotii). past activities. In our case, the stands of the new asso- The study area is situated in the inner Triglav National ciation are a stage in development towards beech forest Park area, so human impact on the development of on extreme sites in the montane belt classified into the vegetation here (other than increased mountaineering association Rhododendro hirsuti-Fagetum. The stands in recent years and human-induced climate changes) is of this association in the gable ends of Zadnjica and insignificant.

5 POVZETEK

V zatrepu dveh ledeniških dolin v Julijskih Alpah, v linah smo našli sestoje subasociacije Salicetum elea- Zadnjici in Kotu (Triglavski narodni park), smo na po- gno-purpureae hieracietosum porrifolii, ki smo jo do bočnih gruščih in prodnatih nanosih fitocenološko zdaj poznali le v Zgornjem Posočju (Čušin & Šilc opisali več grmiščnih in gozdnih združb, na katerih 2006). Kot novo smo opisali asociacijo Rhododendro razvoj poleg velikosti in ustaljenosti ter vlažnosti gru- hirsuti-Laburnetum alpini, ki sodi v skupino grmišč- šča najbolj odločilno vplivajo predvsem vsakoletni no-gozdnih združb montansko-subalpinskega pasu snežni plazovi. Z zadostnim številom popisov smo v (Rhododendro hirsuti-Sorbetum aucupariae, Rhodo- obeh dolinah dokumentirali le dve grmiščni in eno dendro hirsuti-Salicetum appendiculatae, Rhododendro gozdno združbo. Sestoji asociacij Polysticho lonchitis- hirsuti-Betuletum carpaticae, Alno viridis-Sorbetum -Rhamnetum fallacis in Amelanchiero ovalis-Pinetum aucupariae, Calamagrostio arundinaceae-Sorbetum mugo so novost v dolini Kota. V obeh raziskovanih do- ariae) – prim. Dakskobler, Kutnar & Rozman

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 109 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

(2016), ki so bolj ali manj dolgotrajen stadij na potenci- stju v zatrepu doline Kot in o njegovi veliki biotski alno bukovih rastiščih in je njihov nastanek lahko, ne vrednosti. Večinsko sodi v evropsko varstveno po- pa nujno, povezan tudi s človekovimi vplivi oz. njego- membne habitatne tipe: 91K0 Ilirski bukovi gozdovi vo preteklo dejavnostjo. V našem primeru so sestoji (Aremonio-Fagion), 4070 *Ruševje z dlakavim slečem nove asociacije razvojna stopnja proti bukovemu gozdu (Mugo-Rhododendretum hirsuti) in 9420 Alpski mace- na skrajnih rastiščih v montanskem pasu, ki jih uvr- snovi gozdovi. V njem raste več zavarovanih vrst ščamo v asociacijo Rhododendro hirsuti-Fagetum. Za (Cypripedium calceolus – tudi Natura 2000 vrsta, sestoje te asociacije v zatrepih dolin Zadnjice in Kota je Gymnadenia odoratissima, G. conopsea, Epipactis hel- značilna podobnost s sestoji asociacije Polysticho lon- leborine, Dianthus sternbergii, Gentiana clusii, Lycopo- chitis-Fagetum (zato jih obravnavamo kot posebno vi- dium annotinum), endemitov (Cerastium subtriflorum, šinsko varianto z vrsto Sorbus chamaemespilus). Samo Papaver alpinum subsp. ernesti-mayeri = P. ernesti-ma- v dolini Kot pa smo doslej našli njeno obliko na rečno- yeri) in redkih vrst (Sorbus mougeotii). Ker raziskano -ledeniških prodnatih nanosih, na uravnavi, ki jo območje leži v strožje varovanem delu Triglavskega na- lahko vrednotimo kot novo subasociacijo petasiteto- rodnega parka, so človekovi vplivi na tukajšnji razvoj sum paradoxi. Naša spoznanja potrjujejo in dopolnju- vegetacije (z izjemo povečanega obiska planincev v za- jejo ugotovitve Zupančiča in sodelavcev (2017) o izre- dnjih letih in od človeka povzročenih podnebnih spre- dno raznolikem in zanimivem grmiščno-gozdnem ra- memb) majhni.

ACKNOWLEDGEMENTS

Academician Dr. Mitja Zupančič peer-reviewed my Jura. Brane Anderle, Andrej Seliškar and Branko Vreš paper and offered some useful corrections and improve- are co-authors of Figure 3. Iztok Sajko prepared Figure 1 ments. Prof. dr. Andraž Čarni kindly helped with litera- for print. I acknowledge the financial support from the ture references. Sincere thanks to Prof. Dr. Jean-Paul Slovenian Research Agency (research core funding No. Theurillat for his valuable information regarding the P1-0236). English translation by Andreja Šalamon valid name of the Laburnum alpinum community in the 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. Gubellini, 2004: Vegetazione e paesaggio vegetale del Massiccio del Monte Cucco (Appen- nino centrale – Dorsale Umbro-Marchigiana). Fitosociologia 41 (2) suppl. 1: 3–81. Blatnik, M. & B. Repe, 2013: Klasifikacija izbranih melišč glede na vegetacijske značilnosti. Geografski vestnik (Ljubljana) 85 (2): 9–24. Braun-Blanquet, J., 1964: Pflanzensoziologie. Grundzüge der Vegetationskunde. 3. Auf., Springer Verlag, Wien– New York. Brus, R., 1999: Alpski nagnoj (Laburnum alpinum (Mill.) Bercht. & J. Presl.): In: M. Kotar & R. Brus: Naše drevesne vrste. Slovenska matica, Ljubljana, pp. 204–208. Brus, R., 2005: Dendrologija za gozdarje. Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za gozdarstvo in obnovljive gozdne vire, Ljubljana. Castelli, M., E. Biondi & S. Ballelli, 2001: La vegetazione erbacea, arbustiva e preforestale del piano montano dell’ Appennino piemontese (Valli Borbera e Curone – Italia). Fitosociologia 38 (1): 125–151. Cegnar, T., 1998: Temperatura zraka. In: J. Fridl, D. Kladnik M. Orožen Adamič, D. Perko (eds.): Geografski atlas Slovenije. Država v prostoru in času. Državna založba Slovenije, Ljubljana, pp. 100–101. Čušin, B. & U. Šilc, 2006: Vegetation development on gravel sites of the Soča river between the towns of Bovec and Tolmin. Sauteria (Salzburg) 14: 279−292.

110 FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

Dakskobler, I., 2003: Asociacija Rhododendro hirsuti-Fagetum Accetto ex Dakskobler 1998 v zahodni Sloveniji. Razprave 4. razreda SAZU (Ljubljana) 44–2: 5–85. Dakskobler, I., 2014: Association Amelanchiero ovalis-Pinetum mugo in northwestern Slovenia. Acta Biologica Slovenica (Ljubljana) 57 (1): 15–43. Dakskobler, I., W. R. Franz & A. Rozman, 2013: Phytosociology and ecology of Rhamnus fallax in the Southea- stern Alps and in the northern part of the Dinaric Alps. Wulfenia (Klagenfurt) 20: 101–144. Dakskobler, I., L. Kutnar, A. Rozman, 2016: Macesnovje, ruševje, zelenojelševje in druge gorske grmovne združ- be v Sloveniji. Silva Slovenica, Gozdarski inštitut Slovenije. Dakskobler, I. & B. Surina, 2017: Phytosociological analysis of montane-subalpine dwarf willow shrub communities in the Julian Alps and on the Trnovski gozd plateau (NW and W Slovenia). Hacquetia (Ljubljana) 16 (2): 213–280. Dakskobler, I., A. Seliškar & A. Rozman, 2018: Phytosociological analysis of European larch forests in the Southe- astern Alps. Hacquetia (Ljubljana)17 (2): 247–519. Dolinar, M. (ed.) 2018: Ocena podnebnih sprememb v Sloveniji do konca 21. stoletja: sintezno poročilo (Assessment of climate change in Slovenia until the end of the 21st century: synthesis report). Agencija Republike Slovenije za okolje, Ljubljana (Elaborat, 156 pp.) Foucault, B., 2012: Contribution au prodrome des végétations de France: les Betulo carpaticae-Alnetea viridis Rej- mánek in Huml, Lepš, Prach et Rejmánek 1979. J. Bot. Soc. Bot. France 60: 47–68. Géhu J. M. & J. Gehu-Franck, 1987: Groupements arbustifs et mégaphorbiaies di Haut Jura francais. Quelques aspects particuliers. Lazaroa 7: 25–35. Geršič, M., B. Repe, M. Blatnik, V. Brečko Grubar, B. Kovač, N. Pozvek, A. Seifert, 2014: Geografija in ra- stlinska sukcesija – izbrani primeri iz slovenskih pokrajin. Založba ZRC, Ljubljana. Jurkovšek, B., 1987a: Tolmač listov Beljak in Ponteba. - Osnovna geološka karta SFRJ 1:100 000. Zvezni geološki zavod, Beograd, 58 pp. Jurkovšek, B., 1987b: Osnovna geološka karta SFRJ. Beljak in Ponteba 1: 100 000. Zvezni geološki zavod, Beograd. Lovrenčak, F., 2002: Povezave med prstjo in rastlinstvom na vršajih v Planici. Geografski vestnik (Ljubljana) 74 (1): 54–63. Maarel van der, E., 1979: Transformation of cover-abundance values in phytosociology and its effects on community similarity. Vegetatio (Den Haag) 39 (2): 97–114. Martinčič, A., 2003: Seznam listnatih mahov (Bryopsida) Slovenije. Hacquetia (Ljubljana) 2 (1): 91–166. Martinčič, A., 2011: Seznam jetrenjakov (Marchanthiophyta) in rogovnjakov (Anthocerotophyta) Slovenije. Anno- tated Checklist of Slovenian Liverworts (Marchanthiophyta) and Hornworts (Anthocerotophyta). Scopolia (Lju- bljana) 72: 1–38. Martinčič, A., T. Wraber, N. Jogan, A. Podobnik, B. Turk, B. Vreš, V. Ravnik, B. Frajman, S. Strgulc Krajšek, B. Trčak, T. Bačič, M. A. Fischer, K. Eler & B. Surina, 2007: Mala flora Slovenije. Ključ za dolo- čanje praprotnic in semenk. Četrta, dopolnjena in spremenjena izdaja. Tehniška založba Slovenije, Ljubljana. Oriolo, G. &, L. Poldini, 2002: Willow gravel bank thickets (Salicion eleagni-daphnoides (Moor 1958) Grass 1993) in Friuli Venezia Giulia (NE Italy). Hacquetia 1 (2): 141–156. Podani, J., 2001: SYN-TAX 2000. Computer Programs for Data Analysis in Ecology and Systematics. User‘s Manual, Budapest. Richard J. L., 1968: Quelques groupements végétaux à la limite supérieure de la forêt dans les hautes chaines du Jura. Vegetatio 16 (1–4): 205–219. Seliškar, T., B. Vreš & A. Seliškar, 2003: FloVegSi 2.0. Računalniški program za urejanje in analizo bioloških podatkov. Biološki inštitut ZRC SAZU, Ljubljana. Šilc, U. & B. Čušin, 2000: The association Salicetum incano-purpureae Sillinger 1933 on the gravel bars of the Na- diža river (northwestern Slovenia). Gortania (Udine) 22: 91–109. Šilc, U. & A. Čarni, 2012: Conspectus of vegetation syntaxa in Slovenia. Hacquetia (Ljubljana) 11 (1): 113–164. Volk Bahun, M., M. Zorn & M. Pavšek, 2018: Snežni plazovi v Triglavskem pogorju. In: M. Zorn et al. (eds.): Triglav 240, Založba ZRC, Ljubljana, pp.147–157. Zupančič, B., 1998: Padavine. In: J. Fridl, D. Kladnik M. Orožen Adamič, D. Perko (eds.): Geografski atlas Slove- nije. Država v prostoru in času. Državna založba Slovenije, Ljubljana, pp. 98–99. Zupančič, M. & V. Žagar, 2009: The development of forest vegetation in Alpine valleys in Slovenia = Razvoj gozdne vegetacije v alpskih dolinah Slovenije. Folia biologica et geologica (Ljubljana) 50 (1): 189–211.

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 111 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 Slo- venija). 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

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 113 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

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 115

I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

273176 414528 1190 20 NE 9/19/2018 Kot 9549/3 Sc

10 51 70 10 70 35 20 20 22 Re

400 273185 414572 1190 10 SE 9/19/2018 Kot 9549/3 Sc

. 5

14 52 21 80 20 50 50 40 Re

400 273149 408395 1090 15 NW 9/21/2018 Zadnjica-Utro 9648/2 D

16 61 35 80 30 20 20 50 45 Re

400 273162 408264 1030 60+30 NW 9/21/2018 Zadnjica-Utro 9648/2 Rs

16 70 52 10 19 15 30 30 80 Re

400 273183 414701 1115 20 N 9/19/2018 Kot 9549/3 Sc

. 5

18 10 15 80 70 20 68 40 Co

200 273171 414680 1130 20+30 NW 9/19/2018 Kot 9549/3 Sc

10 10 17 10 55 15 60 80 40 Co

200 273182 414640 1140 30 SE 9/19/2018 Kot 9549/3 Rs

. 6

16 10 80 30 48 20 30 60 Re

200 273177 414558 1170 20+20 NE Sc 9/19/2018 Kot 9549/3

. 6

10 15 15 60 35 20 42 60 Re

400 273161 408208 1030 90+50 NW Sc 9/21/2018 Zadnjica-Utro 9648/2

. 5

14 10 41 20 20 25 30 60 Re

200 273147 408385 1070 NW 40+30 Sc-Mo 9/21/2018 Zadnjica-Utro 9648/2

. 8

10 15 13 60 20 48 20 80 Re

400 273148 408392 1080 W 60+40 Ta 9/21/2018 Zadnjica-Utro 9648/2

. 6

70 10 15 12 50 20 20 80 Re

400 273146 408395 1070 W 50 9/21/2018 Zadnjica-Utro Sc-Mo 9648/2

11 10 15 15 15 80 49 20 80 Re

400 273145 408335 1050 W 80+30 9/21/2018 Zadnjica-Utro Sc-Mo 9648/2

10 10 52 10 10 10 15 70 20 Re

400 273164 408127 1000 NW 20 9/21/2018 Zadnjica-Utro Sc-Mo 9648/2

. 5 9 5

70 10 15 45 20 80 Re

200 273144 NW 408199 1015 20 9/21/2018 Zadnjica-Utro Sc-Mo 9648/2

. 8 8 5

10 57 15 60 20 80 Re

400 258936 SEE 407384 1280 60 7/3/2015 dol Zadnjiški 9648/2 Sc

. . 6 7

Li 33 15 60 30 30 80 100

273186 E 414595 1150 70+40 Sc 9/19/2018 Kot 9549/3

. 6

10 10 10 52 10 30 25 80 Re

200 273184 SE 414510 1200 80 Sc 9/19/2018 Kot 9549/3

. . . . 5 5

Li 36 30 50 40 100

273180 1200 E 414482 Sc 70 9/19/2018 Kot 9549/3

. . . . 4

10 35 50 48 40

200 273178 1180 NE 414509 Sc 70 9/19/2018 Kot 9549/3

. . . . . 3

57 35 75 30

400 273181 1190 SE Sc 70 9/19/2018 Kot 9549/3 414505

. . . . 5 2

Li 35 42 40 50

200 273163 1030 NW Gr-Sc 90 9/21/2018 Zadnjica-Utro 9648/2 408256 . . . . . 1 15 49 40 40 Co 100 2

m m E2 E1 E0 m cm E3a E3b Shrub layer (Grmovna plast) Lower tree layer (Spodnja drevesna plast) Number species of (Število vrst) Relevé area (Velikost popisne ploskve) Cover in % (Zastiranje v %): Upper tree layer (Zgornja drevesna plast) Stoniness in % (Kamnitost v %) Maximum height trees of (Največja drevesna višina) Maximum diameter trees of (Največji prsni premer dreves) Moss layer (Mahovna plast) Database number relevé of (Delovna številka popisa) Date relevé of (Datum popisa) Parent material (Matična podlaga) Table 1: Successional 1: Table stages of shrub-forest vegetation on slope screes in glacial cirques of Zadnjica and Kot valleys in the JulianPreglednica Alps Sukcesijski 1: stadiji rastja na pobočnem grušču v krnicah dolin Zadnjice in Kota v Julijskih Alpah Number relevé of (Zaporedna številka popisa) Elevation in m (Nadmorska višina v m) Aspect (Lega) Slope in degrees (Nagib v stopinjah) Soil (Tla) Herb layer (Zeliščna plast) Locality (Nahajališče) (Kvadrant) Quadrant Coordinate GK Y (D-48)

116 FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 5 5 9 9 9 5 9 9 18 18 14 18 91 32 18 41 59 32 41 73 32 41 32 32 50 27 68 77 23 36 28 23 36 23 45 36 50 68 64 64 Fr. Fr.

4 7 1 4 8 2 5 4 9 1 5 6 2 4 2 9 2 6 9 1 7 8 5 7 8 3 7 7 2

11 11 17 14 10 16 14 15 13 15 20 Pr. Pr. 5140385

......

4 1 1 1 1 + + + + + + + + + + + + + + + + + 22 5140552

......

r 5 1 1 + + + + + + + + 21 5137177

......

r r r 4 1 1 1 1 1 1 1 2 + + + + + + + 20 5137291

......

4 1 2 1 1 1 1 1 1 1 + + + + + + + + 19 5140460

......

1 1 4 1 1 + + + + + + + + + + 18 5140443

......

2 1 1 1 3 1 1 + + + + + + + + + + + + + + + 17 5140501

......

1 2 1 2 1 4 1 1 1 2 1 + + + 16 5140389

......

r 1 2 3 + + + + + + + + + + + + 15 5137195

......

2 4 2 5 1 1 + + + + + + + + + + 14 5137308

......

r 2 3 1 4 1 4 1 + + + + + + + + + 13 5137228

......

r r r 3 1 3 4 3 1 + + + + + + + + 12 5137316

......

r 2 3 1 3 4 3 1 + + + + + 11 5137304

......

r 2 2 3 1 3 4 1 + + + + + + + 10 5137412

...... 3 9 1 4 2 1 + + + + + + + 5137390

...... r 1 4 3 1 8 1 + + + + + + + + 5136616

...... 7 4 1 1 + 5140436

...... 1 1 1 6 1 4 1 1 1 1 + + + + + 5140525

...... 2 1 5 4 1 1 + + + 5140416

...... 4 4 + + + + + + + 5140406

...... 4 1 1 1 3 1 1 1 + + + 5140471

...... 4 2 1 + + 5137338 ...... 1 4 + + + + + + + + + + m E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E2 E2 E3 E3 E3a E3a E3a E2a E2a E2a E2a E2a E2a E2a E2a E2a E3b E2b E2b E2b E2b E2b E2b E2b E2b E2b Fagus sylvatica Carex ferruginea Carex Rhamnus fallax Fagus sylvatica Aconitum lycoctonum subsp. ranunculifolium subsp. lycoctonum Aconitum Rhodothamnus chamaecistus Salix glabra Acer pseudoplatanusAcer Polystichum lonchitis Polystichum Laburnum alpinum Laburnum alpinum Rhododendron hirsutum Acer pseudoplatanusAcer Carex alba Carex Laburnum alpinum Ostrya carpinifolia Salix appendiculata Larix decidua Number relevé of (Zaporedna številka popisa) Salix eleagnos Acer pseudoplatanusAcer Ostrya carpinifolia Fagus sylvatica Homogyne sylvestrisHomogyne Larix decidua Rhamnus fallax Ostrya carpinifolia Fagus sylvatica Laburnum alpinum Pinus mugo Larix decidua Carex brachystachys Acer pseudoplatanusAcer Ostrya carpinifolia Clematis alpina Clematis Gymnocarpium robertianum Coordinate X (D-48) GK Diagnostic species of the associations and subassociations (Diagnostične vrste asociacij in subasociacij) Salix purpurea Rhamnus fallax Fagus sylvatica Pinus mugo Sorbus chamaemespilus FS FS FS FS FS FS FS FS FS SP SP ES EP EP EP EP EP AF AF BA AF TR TA BA TA TA BA AT TA VP QP QP VP QP VP VP QP VP VP MuA

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 117 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 5 5 5 5 9 5 5 9 5 9 5 9 9 5 5 9 9 5 9 Fr. 14 14 91 55 32 91 18 41 59 41 23 59 23 27 32 68 23 36 27 86 23 1 1 1 7 1 5 2 1 9 1 2 1 5 9 3 3 2 1 5 8 2 2 6 1 4 1 2 7 2 6 1 2 5 Pr. 13 12 15 19 13 20 20 ...... 1 1 22 + + + + + + + + + + ...... 1 1 1 1 1 21 + + + + + + + + + + + ...... 1 2 1 1 1 1 20 + + + + + + ...... r 1 1 1 1 1 1 1 1 19 1 + + + + ...... 1 18 1 + + + + + + + + + + + + + ...... 17 1 1 1 + + + + + ...... 16 1 1 1 + + + + + + ...... 15 1 1 + + + + + + + ...... 14 1 1 1 1 + + + + + + ...... 1 1 13 2 + + + + + + + + + + ...... 1 1 1 1 1 12 1 1 + + ...... 11 1 1 1 1 1 1 + + + + + + + ...... 1 10 1 1 1 1 1 1 1 1 + + + + + 9 ...... 1 1 1 1 1 + + + + + 8 ...... 1 1 1 2 + + + + + + + 7 ...... 1 2 1 + + + + + + 6 ...... 1 1 + + + 5 ...... 1 1 1 + + + + + + + 4 ...... 1 + + + + + 3 ...... r 1 1 1 + + + + + + + + 2 ...... 1 + + 1 ...... + E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E2a E2a E2a E3a E2a E2b E2b Number relevé of (Zaporedna številka popisa) Tilio-Acerion Fagetalia sylvaticae Fagetalia nutansMelica Galeobdolon flavidum Carex digitata Epipactis helleborine Epipactis Lonicera alpigena Actaea spicata Cardamine enneaphyllos Sorbus aria Aruncus dioicus Querco-Fagetea Hepatica nobilis Daphne mezereumDaphne Epilobium montanum Sorbus aria Dryopteris carthusiana Salvia glutinosa Helleborus niger Helleborus Euonymus latifolia Paris quadrifolia Convallaria majalis Convallaria Galium laevigatumGalium Sorbus aria Mycelis muralis Mycelis Platanthera bifolia Platanthera Knautia drymeiaKnautia Luzula nivea Geranium robertianumGeranium Euphorbia amygdaloidesEuphorbia Mercurialis perennis Quercetalia pubescenti-petraeae Quercetalia Sorbus aria (Aria edulis) Hypericum montanum Anemone trifolia Anemone Lonicera xylosteum Dryopteris filix-mas Brachypodium sylvaticum Brachypodium Polystichum aculeatum Polystichum Poa nemoralis Poa Primula veris Primula Aremonio-Fagion purpurascensCyclamen Thalictrum aquilegiifolium Potentilla erecta FS AI QF TA AF QP QR

118 FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Fr. 5 5 5 5 9 5 5 9 9 5 5 9 5 5 9 5 9 14 18 18 14 18 14 18 18 14 14 27 41 55 27 73 27 27 23 23 36 77 23 50 100 Pr. 1 1 1 3 1 2 6 1 4 1 4 2 2 9 1 4 6 1 4 3 6 2 4 1 5 3 1 8 5 6 2 3 1 3 2 5 16 11 17 12 22 22 ...... 1 3 1 1 + + + + + 21 ...... 1 + + + + + + + + 20 ...... 1 2 + + + + + + + + 19 ...... 1 1 1 1 1 1 + + + + + + + 18 ...... 2 2 + + + + + + 17 ...... 3 2 1 + + + 16 ...... 1 2 1 1 2 + + 15 ...... 1 1 1 + + + 14 ...... 3 1 + + + + + + + + + + + 13 ...... r 1 1 1 2 1 3 1 + + + + + + + + + + 12 ...... r 1 3 1 1 1 3 1 + + + + + + 11 ...... 2 1 2 1 + + + + + + + 10 ...... 1 1 1 2 2 1 1 + + + + + 9 ...... 1 + + + + + + + 8 ...... 1 + + + + + + + 7 ...... 3 6 ...... r 2 1 2 1 1 + + + 5 ...... 1 1 1 + 4 ...... r r 1 1 + + 3 ...... 1 + + + + + + + + 2 ...... 1 1 1 + + + + + 1 ...... 1 + + E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E2a E3a E2a E2a E2a E2a E2a E2b E2b E2b E2b E2b Number relevé of (Zaporedna številka popisa) Aposeris foetida Erico-Pinetea Calamagrostis varia Abies alba Abies Phegopteris connectilis Phegopteris Gentiana asclepiadea Gentiana Sorbus austriaca subsp. austriaca austriacaSorbus subsp. Abies alba Abies Luzula sylvatica Sorbus mougeotii Asperula aristata Lycopodium annotinum Lycopodium Amelanchier ovalisAmelanchier Rosa pendulina Rosa Veronica urticifolia Veronica Peucedanum rablense Peucedanum Dryopteris expansa Dryopteris Euphrasia cuspidata Lonicera nigra Betulo-Alnetea Sorbus mougeotii Calamagrostis villosa Vaccinium vitis-idaea Vaccinium Erica carnea Valeriana tripteris Valeriana Huperzia selago Huperzia Luzula pilosa Cotoneaster tomentosus Oxalis acetosella Oxalis Maianthemum bifolium Maianthemum Aquilegia nigricans Rubus saxatilis Solidago virgaurea Polygala chamaebuxusPolygala Saxifraga cuneifolia Molinia arundinaceaMolinia Gymnocarpium dryopteris Hieracium murorum Hieracium Vaccinium myrtillusVaccinium Amelanchier ovalisAmelanchier Vaccinio-Piceetea abiesPicea Rhamnus saxatilis Picea abiesPicea EP BA VP

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 119 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 5 9 5 5 5 9 9 5 9 9 5 9 9 5 9 5 9 9 5 9 14 18 18 18 14 18 32 18 14 18 18 50 77 32 32 50 36 27 23 23 Fr. 1 2 7 1 1 1 3 4 2 2 1 2 2 4 4 1 2 2 4 1 2 1 8 4 4 2 3 2 5 5 4 1 2 3 7 7 6 11 17 11 Pr...... + + + + + 22 ...... 1 1 + + + + + + 21 ...... r 1 + + + + 20 ...... r + + + + 19 ...... 1 1 1 1 + + + + + + + + 18 ...... + + + + + + + + + 17 ...... 1 + + + 16 ...... + + + + 15 ...... 1 + + 14 ...... 1 13 ...... 1 1 + + + + 12 ...... 1 + + + + + + + 11 ...... 1 2 + + + + + 10 ...... r 9 2 1 3 + + + + + + + + + ...... 8 1 2 1 1 1 3 + + + + + + ...... 7 1 2 + + + + + ...... 1 6 1 1 + + + + + ...... 5 + + + + + + ...... 1 4 1 1 + + + ...... 3 1 1 + + + + + ...... 1 2 1 + + + + + + ...... 1 1 + + + + + + E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E3a E2a E2a E3a E2a E2b E2b Sambuco-Salicion Rhamno-Prunetea capreae, Sorbus aucuparia Veratrum album Veratrum Galium lucidum Polygonatum odoratum Polygonatum Crepis paludosa Number relevé of (Zaporedna številka popisa) Pimpinella saxifraga Agropyron caninum Viola hirtaViola Stachys recta agg, Trifolio-Geranietea hirundinariaVincetoxicum Laserpitium siler Phyteuma ovatum Phyteuma Prunella grandiflora Polygonatum verticillatumPolygonatum Gentianella ciliata Juniperus communis Festuco-Brometea salicifolium Buphthalmum Rubus idaeus Rubus Carex humilis Epilobietea angustifolii Epilobietea Fragaria vesca Hypericum perforatum Senecio ovatus Linum catharticumLinum Juniperus communis Athyrium filix-femina Valeriana wallrothii (V. collina) (V. wallrothii Valeriana Sorbus aucuparia Euphorbia cyparissias Clinopodium vulgare Clinopodium Viola bifloraViola Brachypodium rupestre Brachypodium Chaerophyllum hirsutum Chaerophyllum Sorbus aucuparia Betula pendula Laserpitium latifolium Cirsium erisithalesCirsium Carlina acaulisCarlina Anthericum ramosum Sorbus aucuparia Mulgedio-Aconitetea paniculatum subsp. degenii Aconitum

FB RP RP EA TG SSC MuA

120 FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 9 5 5 5 9 5 5 5 5 5 5 5 5 5 9 5 5 9 5 5 5 5 5 14 14 14 18 14 18 95 27 68 50 36 59 32 23 27 36 23 45 Fr. 2 1 1 1 2 3 1 1 1 6 1 1 3 4 1 1 8 1 1 3 2 5 8 1 1 2 4 1 1 1 6 3 5 1 1 7 11 21 15 10 13 Pr...... 1 + + + + 22 ...... 1 1 1 + + + 21 ...... 1 2 + + + + + 20 ...... + + 19 ...... 1 2 1 2 + + + + + 18 ...... 1 2 1 + + + + 17 ...... 1 1 + + + + + + 16 ...... 1 + + + + 15 ...... 1 + + 14 ...... 1 + + 13 ...... 2 1 + + 12 ...... 1 + + + + 11 ...... 1 + 10 ...... 1 9 + + + + + + + ...... 8 + + + + + + + + + + + ...... 1 2 1 7 2 + + + ...... r 1 2 1 1 1 6 1 2 + + + + + + + ...... 2 1 1 1 5 + + + + ...... 1 1 4 + + + + + + + + ...... 1 1 3 1 + + + + + + + ...... 1 2 2 1 1 2 + + + + + + + + + ...... 1 1 + + + + + + + + + + + + E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 Trifolium pratense Trifolium Sesleria caerulea Sesleria Astrantia bavaricaAstrantia Briza media Acinos alpinusAcinos Laserpitium peucedanoides Lathyrus pratensis Valeriana montana Valeriana Phyteuma orbiculare Phyteuma Juncus monanthos Gymnadenia conopsea Achillea clavenae Festuca calva Thlaspietea rotundifolii carniolicaAstrantia Elyno-Seslerietea alopecuros Betonica Helianthemum nummularium subsp. grandiflorum subsp. nummularium Helianthemum Achillea millefolium Linum julicum Linum Euphrasia picta Teucrium montanum Teucrium Dryas octopetala Selaginella selaginoides Selaginella Carex mucronata Tofieldia calyculata Ranunculus nemorosus agg. Thymus praecox subsp. polytrichus Lotus corniculatus s. lat. (L. alpinus ?) Carex sempervirens Hippocrepis comosa Thesium alpinum Caricetalia davallianae palustrisParnassia Globularia nudicaulis Molinio-Arrhenatheretea album Galium Carduus crassifoliusCarduus Scabiosa lucida subsp. stricta Aster bellidiastrumAster Rhinanthus glacialis Rhinanthus Alchemilla alpigena Alchemilla Arabis hirsuta Arabis Adenostyles glabra Number relevé of (Zaporedna številka popisa) Arabis vochinensis Arabis

ES TR CD PaT MA

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 121 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 5 9 9 9 5 5 9 9 9 9 5 5 5 9 9 5 5 9 9 5 5 9 14 18 18 14 18 32 14 18 18 18 50 23 23 23 23 23 23 50 64 Fr. 13,6 54,5 1 7 2 3 2 2 1 1 2 4 2 2 2 1 4 1 4 1 2 5 3 2 1 5 4 1 3 2 2 4 3 5 4 5 1 5 1 5 2 11 14 11 12 Pr...... 1 1 1 + + + + 22 ...... 1 + + + + 21 ...... 1 + + + + + + + + + + + 20 ...... 1 + + + + 19 ...... 1 1 1 1 + + + + + + + + 18 ...... 1 1 1 1 + + + + + 17 ...... 1 1 + + + + + + + + 16 ...... 1 1 + + + + 15 ...... 2 1 + + 14 ...... 1 1 + 13 ...... r 2 1 1 1 + + + + 12 ...... 1 1 1 + 11 ...... 1 1 1 + + + 10 ...... 9 ...... 8 + + + + ...... 7 1 + + + + + ...... 2 1 6 1 1 1 + ...... 5 1 + ...... 1 4 1 1 + + + + + + + + + + ...... 1 3 1 1 + + + + + + + + + ...... 2 + + + + + ...... 1 1 1 3 + + + + + + + + + + + + E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E0 E0 E0 E0 E0 E0 E0 E0 E0 E0 E0 Dianthus sternbergii Paederota lutea Athamanta cretensis Asplenium fissum Schistidium apocarpum Cystopteris regia Heliosperma alpestre Asplenietea trichomanis Asplenium viride Viola pyrenaicaViola Scrophularia juratensis Asplenium ruta-muraria Conocephalum conicum Rhytidiadelphus triquetrusRhytidiadelphus Silene vulgaris glareosa subsp. Gypsophila repens Leontodon hyoseroides Trisetum argenteumTrisetum Soldanella alpina Soldanella Hylocomium splendensHylocomium Number relevé of (Zaporedna številka popisa) Moehringia muscosa Moehringia Aquilegia einseleana Aquilegia Homalothecium lutescens Tortella tortuosa Tortella Hieracium porrifoliumHieracium Hieracium piloselloides Hieracium Rumex scutatus Festuca nitida Festuca Cystopteris montana Peltigera canina Peltigera Mosses and lichens (Mahovi in lišaji) molluscumCtenidium Mnium thomsoniiMnium Kernera saxatilis Hieracium bifidum Hieracium Arabis alpina Arabis Campanula cespitosa Biscutella laevigata Petasites paradoxus Dicranum scoparium Dicranum Cerastium subtriflorum Asplenium trichomanes Achnatherum calamagrostisAchnatherum Fissidens dubius Cystopteris fragilis AT ML

122 FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 5 5 5 5 5 Fr. 1 1 1 1 1 Pr...... 22 . . . . . 21 . . . . + 20 . . . . + 19 . . . . + 18 . . . . . 17 . . . . . 16 . . . . . 15 . . . . . 14 . . . . . 13 . . . . . 12 . . . . . 11 . . . + + 10 . . . . . 9 . . . . . 8 . . . . . 7 . . . . . 6 . . . . . 5 . . . . . 4 . . . . . 3 . . . . . 2 . . . . . 1 E0 E0 E0 E0 E0 Neckera crispaNeckera Polytrichum formosum Polytrichum Mnium sp. Plagiomnium undulatumPlagiomnium Legend Legenda - PresencePr. (number relevés of in which the species is presented) - število popisov, v katerih se pojavlja vrsta FrequencyFr. in % - frekvenca v % Gr Gravel - prod Sc Scree - grušč MoraineMo (Til) - morena (til) Talus - vršajTa Rs Rockslide - podorno skalovje D Dolomite - dolomit Co Colluvial soil - koluvialna tla Li Lithosols - kamnišče RendzinaRe rendzina - QR Quercetalia roboris AI incanae Alnion PaT alpinae-Trisetetalia Poo Plagiochila porelloides Number relevé of (Zaporedna številka popisa)

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 123 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 273197 273196 273194 273191 273190 273192 (Delovna številka popisa) 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) E1 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 m 15 22 20 19 (Največja drevesna višina) 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 E1 2 . . . . . 1 17 SP Salix purpurea E2b + . . . . . 1 17 TR Petasites paradoxus E1 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 E1 + . + . . . 2 33 EP Amelanchier ovalis E2b . + . . . . 1 17 EP Amelanchier ovalis E2a . + r . r . 3 50 TR Hieracium porrifolium E1 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 E1 + + 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 E1 . . . 1 1 1 3 50 FB Brachypodium rupestre E1 . . . 1 1 + 3 50 VP Homogyne sylvestris E1 . . . + . + 2 33 EP Erico-Pinetea Calamagrostis varia E1 1 1 + 2 3 3 6 100 Erica carnea E1 + 4 4 4 4 4 6 100 Molinia arundinacea E1 1 2 4 + 1 1 6 100 Carex alba E1 . + + 1 2 1 5 83 Euphrasia cuspidata E1 + 1 1 . . . 3 50 Rubus saxatilis E1 . . . 1 1 1 3 50 Allium ericetorum E1 . + . . + . 2 33 Aquilegia nigricans E1 . + . . + . 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

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 AF 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 TA 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 FS 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 . . r + . . 2 33 Mercurialis perennis E1 . . . + . + 2 33 Prenanthes purpurea E1 . . . . + + 2 33 Galeobdolon flavidum E1 . . . . + . 1 17 Dryopteris filix-mas E1 . . . . . + 1 17 QP 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 QF Querco-Fagetea QR Potentilla erecta E1 . . 1 1 + 1 4 67 Hepatica nobilis E1 . + + 1 . . 3 50 Viola riviniana E1 . + . r . . 2 33 AI Frangula alnus E2a . . . . + . 1 17 VP 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

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 125 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS

Number of relevé (Zaporedna številka popisa) 1 2 3 4 5 6 Pr. Fr. Abies alba E2a . . . . + . 1 17 Aposeris foetida E1 . . . . + . 1 17 Solidago virgaurea E1 . . . . + . 1 17 Luzula pilosa E1 . . . . . + 1 17 BA Betulo-Alnetea Salix appendiculata E2b . + + . . . 2 33 Salix appendiculata E2a 2 . + . + . 3 50 Salix appendiculata E1 1 . . . . . 1 17 Juniperus sibirica E2a . . . . r r 2 33 Sorbus chamaemespilus E2a . . + . + + 3 50 Salix waldsteiniana E2a + . . . . . 1 17 Salix glabra E2a . + . . . . 1 17 MuA Mulgedio-Aconitetea Polygonatum verticillatum E1 . . . . + . 1 17 Athyrium filix-femina E1 . . . . . + 1 17 EA Epilobietea angustifolii Fragaria vesca E1 . . . + + + 3 50 SSC Sambuci-Salicion capreae, Rhamno-Prunetea Sorbus aucuparia E3a . + . + . + 3 50 Sorbus aucuparia E2b . . . + . . 1 17 Sorbus aucuparia E1 . . + + + + 4 67 RP Berberis vulgaris E2a . . . . . r 1 17 FB Festuco-Brometea Buphthalmum salicifolium E1 + + 1 + + . 5 83 Galium lucidum E1 + + + . + . 4 67 Carlina acaulis E1 . + + . + . 3 50 Cirsium erisithales E1 + + . . . . 2 33 Hippocrepis comosa E1 . 1 + . . . 2 33 Prunella grandiflora E1 . + 1 . . . 2 33 Gentianella ciliata E1 . 1 . . + . 2 33 Festuca rupicola E1 + . . . . . 1 17 Coronilla vaginalis E1 . + . . . . 1 17 Stachys recta s. lat. E1 . + . . . . 1 17 Gymnadenia conopsea E1 . . + . . . 1 17 TG Viola hirta E1 . . + . . . 1 17 PaT Poo alpinae-Trisetetalia, Molinio-Arrhenatheretea MA Angelica sylvestris E1 . + r . . . 2 33 Campanula scheuchzeri E1 . . . + . . 1 17 Ranunculus nemorosus E1 . . . r . . 1 17 ES Elyno-Seslerietea Laserpitium peucedanoides E1 . + 1 + 1 + 5 83 Dryas octopetala E1 1 3 2 + . . 4 67 Lotus corniculatus s. lat. (L. alpinus) E1 . + 1 + + . 4 67 Betonica alopecuros E1 . + 1 . 1 + 4 67 Aster bellidiastrum E1 + 1 2 . . . 3 50 Carex mucronata E1 + 1 + . . . 3 50 Globularia cordifolia E1 + 1 + . . . 3 50 Carex firma E1 + + 1 . . . 3 50 Senecio abrotanifolius E1 . + + + . . 3 50 Scabiosa lucida subsp. stricta E1 . + 1 . + . 3 50 Carduus crassifolius E1 + + . . . . 2 33 Sesleria caerulea E1 + . + . . . 2 33 NS Antennaria dioica E1 . + + . . . 2 33 CD Tofieldia calyculata E1 . + + . . . 2 33 Achillea clavenae E1 + . . . . . 1 17 Phyteuma orbiculare E1 + . . . . . 1 17 Globularia nudicaulis E1 . + . . . . 1 17 Thymus praecox subsp. polytrichus E1 . + . . . . 1 17 Arctostaphylos alpinus E1 . . + . . . 1 17 Gentiana clusii E1 . . + . . . 1 17 Rhinanthus glacialis E1 . . + . . . 1 17 Thesium alpinum E1 . . + . . . 1 17

126 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 3 4 5 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 + + . . . 3 50 Dianthus sternbergii E1 + + + . . . 3 50 Hieracium bifidum E1 + + + . . . 3 50 Adenostyles glabra E1 . + + + . . 3 50 Gymnocarpium robertianum E1 . . . 1 1 + 3 50 Achnatherum calamagrostis E1 + + . . . . 2 33 Biscutella laevigata E1 + + . . . . 2 33 Hieracium piloselloides E1 + + . . . . 2 33 Rumex scutatus E1 + + . . . . 2 33 Trisetum argenteum E1 + + . . . . 2 33 Valeriana montana E1 + . . 1 . . 2 33 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 E0 2 + 1 . + . 4 67 Hylocomium splendens E0 . . + 1 . + 3 50 Rhytidiadelphus triquetrus E0 . . + 1 . . 2 33 Dicranum scoparium E0 . . . + . + 2 33 Cladonia rangiferina E0 . . + . . . 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

FOLIA BIOLOGICA ET GEOLOGICA 60/1 – 2019 127