HACQUETIA 6/2 • 2007, 195-208 DOI: 10.2478/v10028-007-0011-y SUBALPINE BEECH FOREST WITH HAIRY ALPENROSE (POLYSTICHO LONCHITIS-FAGETUM RHODODENDRETOSUM HIRSUTI subass. nova) on mt. snežnik (liburnian karst, dinaric mts) Boštjan SURINA* & Marash RAKAJ** Abstract Subalpine beech stands with Hairy Alpenrose (Rhododendron hirsutum) were phytosociologically studied on Mt. Snežnik (Dinaric Mts). They thrived on stony and steep slopes of northern exposure. Comparisons with other subalpine Beech stands (Polysticho lonchitis-Fagetum s. lat.), dinaric Fir-Beech stands with Hairy Alpenrose (Omphalodo-Fagetum s. lat. rhododendretosum hirsuti), and prealpine fir-beech stands with Hairy Alpenrose (Homogyno sylvestris-Fagetum s. lat. rhododendretosum hirsuti), stands of Hairy Alpenrose and Beech (Rhododendro hirsuti-Fagetum s. lat.), as well as Austrian subalpine beech stands (Saxifrago rotundifoliae-Fagetum s. lat.) showed their unique floristical composition due to ecological conditions, and thus distinct syntaxonomical position within the association Polysticho-Fagetum. Therefore, a new subassociation Polysticho-Fagetum rhododendretosum hirsuti subass. nova was described, and - as differential species for the subassociation - Rhododendron hirsutum, Rubus saxatilis, Rosapendulina, and Clematis alpina were chosen. Key words: Dinaric Mts, Mt. Snežnik, Polysticho-Fagetum, subalpine beech forest, Rhododendron hirsutum, vegetation Izvleček Prispevek podaja fitocenološko oznako subalpinskega bukovega gozda (Polysticho lonchitis-Fagetum) z dlakavim slečem (Rhododendron hirsutum) na Snežniku. Obravnavani sestoji uspevajo na kamnitih in/ali skalnatih ter strmih pobočjih severnih ekspozicij. Primerjave z ostalimi subalpinskimi bukovimi ter (jelovo-)bukovimi sestoji z dlakavim slečem ( Omphalodo-Fagetum rhododendretosum hirsuti, Homogyno sylvestris rhododendretosum hirsuti, Rhododendro-Fagetum) ter subalpinskimi bukovji iz Avstrije (Saxifrago rotundifolii-Fagetum) so pokazale njihov poseben sintaksonomski položaj v okviru asociacije Polysticho-Fagetum. Zato smo te sestoje uvrstili v novo suba-sociacijo Polysticho-Fagetum rhododendretosum hirsuti subass. nova, za razlikovalnice pa izbrali vrste Rhododendron hirsutum, Rubus saxatilis, Rosa pendulina in Clematis alpina. Ključne besede: Dinaridi, Snežnik, Polysticho-Fagetum, subalpinski bukov gozd, Rhododendron hirsutum, vegetacija INTRODUCTION Subalpine beech forests in Dinaric Mts: a brief synsystem-atic overview Horvat (1938) classified beech forests in NW Dinaric Mts into three subassociations, namely: (a) montane beech forests Fagetum sylvaticae croaticum australe montanum, (b) montane and altimontane Dinaric fir-beech forests Fagetum sylvaticae croaticum australe abietetosum, and (c) subalpine beech for- ests Fagetum sylvaticae croaticum australe subalpinum. Subsequent studies clearly showed that Horvat's classification of beech forests from the alliance Aremonio-Fagion (= Fagion illyricum) was too broad; consequently, beech forest subassociations were ranked as associations and several new floristically and ecologically more or less well defined associations were described (e.g. Tregubov 1957, Košir 1979, Marinček 1983, 1996, Dakskobler 1997, Marinček 1998). * University of Vienna, Faculty of Life Sciences, Department of Biogeography, Rennweg 14, A-1030 Vienna, Austria Natural History Musem Rijeka, Lorenzov prolaz 1, 51000 Rijeka, Croatia. Bostjan.Surina@univie.ac.at ** Universiteti i Shkodrës »Luigj Gurakuqi«, Departimenti Biologji-Kimisë, Shkodër, Albania 195 Hacquetia 6/2 • 2007, 183-193 After the nomenclature revision of the alliance Aremonio-Fagion (Marinček et al. 1993, but see also Poldini & Nardini 1993), the subalpine beech association (Fagetum sylvaticae croaticum australe subalpi-num Horvat 1938) was split into two new associations: Ranunculo platanifolii-Fagetum (altimontane belt) and Polysticho lonchitis-Fagetum (extrazonally on exposed and/or northern slopes in altimontane belt, but commonly (zonally) in subalpine belt up to the forest line). There were attempts to further phytogeographically differentiate association Polys-ticho-Fagetum into geographical variants: (a) Anemone trifolia (Poldini & Nardini 1993) for NE Italy and w Slovenia, (b) salix waldsteiniana (Marinček 1996) for the south-eastern foothills of the Julian Alps (namely Mts Blegoš and Ratitovec), and (c) allium victorialis (Marinček 1996) for the Dinaric Mts, but according to our analyses they seemed to be of low synsystematic significance. Subalpine beech forests on Mt. Snežnik Tomažič was one of the first who studied subalpine beech stands on Mt. Snežnik but unfortunately the results (i.e. phytosociological tables) were never published (e.g. Zupančič 2001). Nevertheless, together with colleagues he successfully conducted extensive vegetation mappings throughout the study area and, as a result, detailed vegetation maps were provided (Tomažič & Tregubov 1958, 1959). He classified subalpine beech stands into the association AIUo victorialis-Fagetum Tomažič nom. nud., and mapped them throughout the southern part of the Snežnik plateau. Afterwards, Tregubov (1957) provided three relevés (from Kujavič, drče and NE slopes of the summit) and classified them within the association Fagetum subalpinum (sensu Horvat 1938). only in 1996 Marinček published 29 relevés of subalpine beech stands from the plateaus of Trnovski gozd and Snežnik (Nw dinaric Mts), classifying them into the geographical variant Polysticho-Fagetum var. geogr. allium victorialis, he recognised three subassociations: -polystichetosum (with variants Gymnocarpium dryopteris and adoxa moschatellina), -adenostyletosum alliariae, and -hacquetietosum. Unfortunately, from the phytosociological table it is hardly possible to distinguish localities of relevés between the plateaus of Trnovski gozd and Snežnik. one of the most distinctive characteristics of subalpine beech stands (beside the floristic composition) is their unique physiognomy. Specially on the upper limit of their altitudinal range (forest line), distorted, twisted stems and curved branches of beech trees make stands appear picturesque. due to harsh environmental conditions (strong Bora, low winter temperature and heavy snowfalls) beech stands rarely exceed 3-5 m in height and 10-30 cm in trunk diameter. Subalpine beech stands of Polysticho-Fagetum are fairly widespread throughout the plateau, but they rarely (only above 1300-1400 m a.s.l. and mostly on the southern part of the Snežnik plateau) extend over larger areas. They usually thrive on mountain ridges, summits and northern slopes of mountains between 1300-1500 (1600) m a.s.l., forming a distinct forest line in direct contact with stands of Mountain pine (Hyperico grisebachii-Pinetum mugo var. geogr. arabis scopoliana), as well as Dinaric subalpine tussock grasslands (alliance Seslerion juncifo-liae). Probably, as a consequence of pasture activites in 19th century (according to Smerdel 1989), the upper altitudinal limit of stands (forest line!) might have been significantly lowered (e.g. Kindler 1957, Pogačnik & Prosen 1998). Subalpine beech stands may well occur also on lower altitudes of the plateau in freezing ravines as a result of extreme ecological conditions due to temperature inversion (Surina & Vreš 2004). On the other hand, the transition area on the lower altitudinal limit of stands might extend (mostly on northerly exposed and colder slopes) well into the altitudinal range of stands of other syntaxa (namely Ranunculo-Fagetum var. geogr. Cala-mintha grandiflora) and/or Omphalodo-Fagetum var. geogr. Calamintha grandiflora), and thus making sufficient delimitation of stands of different syntaxa (specialy Ranunculo-Fagetum) hardly possible. During our research on flora and vegetation of the Liburnian karst we noticed on several occasions physiognomically, floristically and ecologically distinct subalpine beech stands with predominating Hairy Alpenrose (rhododendron hirsutum) in shrub and/or herb layer thriving on steep, rocky and most frequently northerly exposed slopes. The aim of the present study was thus to determine their ecological and syntaxonomical position in comparison to other similar (fir-)beech stands with Hairy Alpenrose within the alliance aremonio-Fagion. Study area The Snežnik plateau is a high-karst region in the northwesternmost part of the Liburnian karst (NW Dinaric Mts, SW Slovenia). The highest mountain of the plateau is Mt. Snežnik (1796 m), which creates a 196 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) distinctive orographic barrier between the northern and the southern part of the plateau. On average, the higher (and also with more diversified relief) southern part of the plateau received considerably higher amounts of precipitation (Gomance, 937 m, 3143 mm) than its northern part (Leskova dolina, 806 m, 2166 mm), while the entire region is among the wettest in Slovenia (Manohin 1957, Zupančič 1995). The precipitation regime of the southern part is strongy influenced by the Mediterranean. on account of the higher altitude of the plateau, karst relief, windness and abundant rainfall the climate is relatively cold. The mean temperature of Gomance was 6.7 °C (Mekinda-Majaron 1995), and the vegetation season lasts from May till September. The geological bedrock consists of Jurassic and Cretaceous limestones, dolomitized limestones and their breccias (Pleničar 1956, Pavlovec & Pleničar 2000). At the time of the last glacial period the larger part of the study area was located above the perpetual snow line (Šifrer 1959). For detailed microclimatic and general vegetation descriptions of the study area compare also Zupančič (1970, 1980), Wraber (1997) and Surina & Vreš (2004). METHODS Between 2002 and 2005 we made 11 relevés of subalpine beech stands on Mt. Snežnik applying the sigmatistic method (Braun-Blanquet 1964, Dier-schke 1994). when arranging the relevés into a phy-tosociological table and comparing them with the stands of similar syntaxa we used the hierarchical classification (complete linkage method (farthest neighbour) - FNC, incremental sum of squares -ISSq, and unweighted average linkage method - UPGMA), and principal coordinates analysis (PCoA) within the SYN-TAX programme package (Podani 2001). For this purpose we transformed cover values with van der Maarel's numerical (1-9) scale (van der Maarel 1979). The measures of dissimilarity were a complement of the »similarity ratio« coefficient and »Euclidian distance«. On the final arrangement of relevés (Tab. 1) we combined the results obtained by numerical methods with the arrangement based on diagnostic species. The phytosociological groups were generally in agreement with numerous authors but formed on our own criteria. The nomenclature source for the names of vascular plants was the Mala flora Slovenije (Martinčič et al. 1999), while names for mosses and lichens were in agreement with the Annotated check-list of the mosses of Slovenia (Martinčič 2003), and the Catalogue of the lichen-ised and lichenicolous fungi of Slovenia (Suppan et al. 2000). Only the most common taxa of mosses and lichens were determined and therefore excluded from the numerical analyses. For the names of syntaxa we followed Marinček et al. (1993) with some minor annotations, but the complete list is given in an appendix. Subalpine beech stands with Hairy Alpenrose from Mt. Snežnik were compared with some similar beech communities in the Northeastern and Southeastern Alps, and in the Dinaric Mts: 1. Polysticho lonchitis-Fagetum s. lat.: Italy: Friuli (Poldini & Nardini 1993); Slovenia: Mts Blegoš and Ratitovec (Marinček 1980), Mt. Snežnik (Tregubov 1957), and Snežnik and Trnovski gozd plateaus (Marinček 1996); Croatia: Velebit Mts (Horvat 1938); 2. Omphalodo-Fagetum var. geogr. Saxifraga cuneifo-lia rhododendretosum hirsuti: Slovenia, Trnovski gozd plateau (Dakskobler et al. 2000); 3. Homogyno sylvestris-Fagetum var. geogr. Anemone trifolia rhododendretosum hirsuti, and Homogyno sylvestris-Fagetum var. geogr. Luzula nivea rhododendretosum hirsuti; Slovenia, Julian Alps (Dakskobler 2002, 2004); 4. Rhododendro hirsuti-Fagetum var. geogr. anemone trifolia & var. geogr. Phyteuma columnae: Slovenia, NW Dinaric Mts and Julian Alps (Dakskobler 2003). 5. Saxifrago rotundifoliae-Fagetum Zukrigl 1989 s. lat.: Austria, NE and SE Alps (Willner 2007). The synoptic table is available within electronic appendix. RESULTS AND DISCUSSION Floristic composition and physiognomic aspect of stands In the tree layer Fagus sylvatica4-5 completely prevailed, reaching up to 10-15 m in height and 10-30 cm of trunk diameter. Basal parts of beech-trunks, specially on slopes, were typically curved due to pressure of the heavy and long-lasting snow cover. Sor-bus aria+ occurred frequently in the studied stands, albeit with rather low coverage value. Less frequent and with low cover values were Acer pseudoplatanus+, Abies alba+, Picea abies+, and Sorbus aucuparia+. Among the shrub species (occurring in shrub and/or herb layer) Rhododendron hirsutum1-4 totaly 197 Hacquetia 6/2 • 2007, 195-208 dominated in stands and together with Fagus sylvatica (in tree layer), Lonicera alpigena+~2, Rosa pen-dulina+~3, Daphne mezereum+-1, Salix appendiculata+~l, and Rubus saxatilis+~% most significantly contributed to the physiognomic aspect of stands. Herb layer was well developed. Clematis alpina,1-2, Anemone nemorosa1-2, Valeriana tripteris+~2, Homogyne sylvestris1-, Adenostyles glabra,+-2, Phyteuma ovatum+~2, and Calamagrostis varia,1-2 dominated with rather high coverage values, whereas Polystichum lonchi-tis+~l, Cardamine enneaphyllos+~l, Euphorbia carniolica+, Cardamine trifolia+~l, Festuca altissima+~l, Mercurialis perennis+~l, Polygonatum verticillatum+~l, Gentiana as-clepiadea+~l, and Solidago virgaurea+ occurred quite frequently but with smaller coverage values. Moss layer covered 5-20 % of the relevé area and the most frequent species were Ctenidium mol-luscum1-2, Tortella tortuosa+~l, and Schistidium apocar-pum+-1. The complete floristic composition of the studied stands is given in the phytosociological table (Tab. 1). Characteristic and differential species of the association Polysticho lonchitis-Fagetum are fully represented, with Polystichum lonchitis being the most (100 %) and Ribes alpinum (36 %) the least frequent. The number of seed plants per relevé varied between 35 and 56, with the median of 47, and the coefficient of variation of 16.14 %, indicating a quite uniform floristical composition of stands. Ecology of stands Most frequently stands thrived on northerly exposed slopes (Tab. 1). Slopes were moderately steep to very steep with inclination varying between 15 ° and 50 ° (Me=35 %). Stoniness made up a large proportion of relevé areas (10-70 %, Me=40 %). In comparison to other (fir-)beech and subalpine beech stands in the studied area, we noticed considerable delay in phenophases in subalpine stands with predominating Rhododendron hirsutum, most probably due to unfavourable ecological conditions. Dolomi-tized limestones and limestone prevailed. Synsystematic position of studied stands Firstly, in order to accurately determine the synsystematic position of the studied stands, we performed comparisons with stands of the subalpine beech association Polysticho-Fagetum in Slovenia and Croatia (Fig. 1). D issimilarity Figure 1: Dendrogram of the subalpine beech stands (Polysticho-Fagetum s. lat.) in the Julian Alps and Dinaric Mts (complete linkage, similarity ratio); 1-3: Mt. Snežnik (Tregubov 1957), 4-33: Snežnik and Trnovski gozd plateaus (Marinček 1996), 34-44: Mt. Snežnik (Surina, this work), 45-49: NE Italy (Poldini & Nardini 1993), 50-59: Mts Blegoš and Ratitovec (Marinček 1981). Slika 1: Dendrogram subalpinskih bukovih sestojev (Polysti-cho-Fagetum s. lat.) v Julijskih Alpah in Dinaridih (complete linkage, similarity ratio); 1-3: Snežnik (Tregubov 1957), 4-33: Snežnik in Trnovski gozd (Marinček 1996), 34-44: Snežnik (Surina, to delo), 45-49: SV Italija (Poldini & Nardini 1993), 50-59: Blegoš in Ratitovec (Marinček 1981). Stands from the Snežnik plateau with predominating Rhododendron hirsutum nested in a separate cluster, supporting their distinct synsystematic position, and were -somewhat surprisingly- most similar to stands from Mt. Blegoš. Some relevés from the phytosociological table of Marinček (1996; rel. 1-5; -polystichetosum, var. Gymnocarpium dryopteris) joined the cluster with the studied stands and therefore were treated separately in subsequent analyses. 198 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) Figure 2: Two-dimensional scatter-diagram of subalpine (fir-) beech stands with Hairy Alpenrose (Rhododendron hirsutum) in the Julian Alps, NE and SE Alps in Austria, and Dinaric Mts (PCoA, Euclidian distance); Polysticho-Fagetum s. lat.: 1 - NE Italy (Poldini & Nardini 1993), 2 - Mts. Blegoš and Ratitovec (Marinček 1981), 3 - Velebit Mts (Horvat 1938), 4 - Trnovski gozd and Snežnik plateaus (Marinček 1996, rel. 6-29), 5 - Trnovski gozd and Snežnik plateus (Marinček 1996, rel. 1-5), 6 - Mt. Snežnik (Surina, this work); Omphalodo-Fagetum var. geogr. Saxifraga cuneifolia rhododendretosum: 7 - Trnovski gozd plateau (Dakskobler et al. 2000); 8-11 - Rhododendro-Fagetum var. geogr. Anemone trifolia & var. geogr. Phyteuma columnae (Dakskobler 2003, Tabs. 8, 5, 6 & 7); Homogyno-Fagetum rhododendretosum: 12-13 - Trnovski gozd plateau and the Julian Alps (Dakskobler 2002, 2004); Saxifrago rotundifoliae-Fagetum s. lat. 14-23 (SE and NE austrian Alps): 14 & 19 - Saxifrago-Fagetum calamagrostietosum variae, 15 & 20 - Saxifrago-Fagetum typicum, 16-21 - Saxifrago-Fagetum adenostyletosum al-liariae, 17-22 - Saxifrago-Fagetum petasitetosum, 18-23 - Saxifrago-Fagetum stellarietosum nemorum (Willner & Grabherr 2007, Tab. 26). Slika 2: Dvorazsežni ordinacijski diagram subalpinskih (jelovo-) bukovih z dlakavim slečem (Rhododendron hirsutum) v Julijskih Alpah in Dinaridih (PCoA, Euclidian distances); Polysticho-Fagetum s. lat.: 1 - SV Italija (Poldini & Nardini 1993), 2 - Blegoš in Ratitovec (Marinček 1981), 3 - Velebit (Horvat 1938), 4 - Trnovski gozd in Snežnik (Marinček 1996, rel. 6-29), 5 - Trnovski gozd in Snežnik (Marinček 1996, rel. 1-5), 6 - Snežnik (Surina, to delo); Omphalodo-Fagetum var. geogr. Saxifraga cuneifolia rhododendretosum: 7 - Trnovski gozd in Julijske Alpe (Dakskobler et al. 2000); Rhododendro-Fagetum var. geogr. Anemone trifolia & var. geogr. Phyteuma columnae: 8-11 - (Dakskobler 2003, Tabs. 8, 5, 6 & 7); Homogyno-Fagetum rhododendretosum: 12-13 - Julijske Alpe (Dakskobler 2002, 2004); Saxifrago rotundifoliae-Fagetum s. lat. 14-23 (SE and NE austrian Alps): 14 & 19 - Saxifrago-Fagetum calamagrostietosum variae, 15 & 20 - Saxifrago-Fagetum typicum, 16-21 - Saxi-frago-Fagetum adenostyletosum alliariae, 17-22 - Saxifrago-Fagetum petasitetosum, 18-23 - Saxifrago-Fagetum stellarietosum nemorum (Willner & Grabherr 2007, Tab. 26). Secondly, comparisons with other (fir-)beech stands with Rhododendron hirsutum in Slovenia and Austria (Saxifrago-Fagetum) were conducted (Figs. 2 & 3, Tab. 2). Again, the results showed the rather unique syntaxonomic position of the studied stands. Results from the Principal Coordinates Analysis (Fig. 2) generally indicated 6 distinct groups of syntaxa: group A is composed mainly of stands of Saxifrago-Fagetum s. lat., but (somewhat surprisingly) also with stands of Polysticho-Fagetum s. lat. from NE Italy (1) and Velebit Mts - Croatia (3); group B presents stands of Saxifrago-Fagetum ca-lamagrostietosum; group C is composed of fir-beech stands Homogyno-Fagetum s. lat. from Trnovski gozd plateau and Julian Alps, while group D of stands of Rhododendro-Fagetum. Groups E and F are composed of stands of Polysticho-Fagetum, where group 199 Hacquetia 6/2 • 2007, 183-193 rVl ^^ 1 2 4 3 5 6 7 14 15 19 20 16 17 18 21 22 23 8 10 11 9 12 13 rhod cal typ cal typ —H-1-1-1 h P-F O-Frh Sr-F Rh-F H-Frh Figure 3: Dendrogram of (fir-)beech syntaxa with Hairy Alpenrose (Rhododendron hirsutum) in the Julian Alps, Austrian NE and SE Alps, and Dinaric Mts (MISSQ, similarity ratio; numbers correspond with Fig. 2). Slika 3: Dendrogram subalpinskih bukovih gozdov, sub-alpinskih bukovih gozdov z dlakavim slečem, bukovih gozdov z dlakavim slečem in jelovo-bukovih sestojev z dlakavim slečem v Julijskih Alpah, SZ in JV avstrijskih Alpah in Dinaridih (MISSQ, similarity ratio; številke se ujemajo s številkami v sl. 2). F represents somewhat typical stands, and group E stands of Polysticho-Fagetum (5 & 6) and Ompha-lodo-Fagetum var. geogr. saxifraga cuneifolia (7) with predominating Rhododendron hirsutum in herb or shrub layer. Cluster analysis (Fig. 3) showed slightly different but still similar results. Syntaxa were split into three groups: the first cluster (P-F, O-Frh, and Sr-F cal & typ) was rather homogeneous and composed mostly from subalpine beech stands (Polysticho-Fagetum s. lat. - 1-6, saxifrago-Fagetum calamagrostietosum - 14 & 19 and typicum - 15 & 20) and dinaric fir-beech forest with rhododendron hirsutum (omphalodo-Fagetum var. geor. saxifraga cuneifolia rhododendretosum hirsuti - 7); the second cluster (Sr-F) comprehended syntaxa entirely of saxifrago-Fagetum - 16-23, while the third cluster (Rh-F and H-Frh) gathered syntaxa of rhododen-dro-Fagetum (8-11) and Homogyno-Fagetum (12-13). The studied subalpine beech stands with rhododendron hirsutum from Mt. Snežnik nested well within subalpine beech stands (Polysticho lonchitis-Fagetum s. lat.), but in a separate cluster (rhod). Not quite unexpectedly, fir-beech stands with rhododendron hirsutum from the Trnovski gozd plateau (O-Frh) were clustered together with our studied stands (and not separately or even together with pre-alpine fir-beech stands with rhododendron hirsutum - Table 2: Phytosociological groups (in %) in (fir-)beech syntaxa with Hairy Alpenrose (Rhododendron hirsutum) in the Julian Alps and Dinaric Mts (successive numbers from 1-13 correspond to Figs. 2 & 3). Tabela 2: Fitocenološke skupine (v %) v (jelovo-) bukovih sintaksonih z dlakavim slečem (Rhododendron hirsutum) v Julijskih Alpah in Dinaridih (številke od 1 do 13 se ujemajo s sl. 2 in 3). Syntaxon 1 2 3 4 5 6 7 8 9 10 11 12 13 Number of relevés 5 10 15 25 5 11 15 11 23 26 18 25 19 Number of seed plants 64 102 100 150 64 96 104 119 169 172 127 114 151 Vaccinio-Piceetea 30 24 17 19 33 26 28 27 15 19 17 26 23 Fagetalia sylvaticae 25 23 24 17 16 16 13 13 14 14 15 9 7 Mulgedio-Aconitetea 16 19 15 19 11 13 14 10 6 7 9 9 9 Erico-Pinetea 6 7 8 9 8 9 9 10 11 9 12 14 13 Aremonio-Fagion 6 9 11 10 8 9 6 7 8 6 7 4 6 Asplenietea trichomanis 3 2 3 6 8 11 10 8 10 11 10 10 Other species 8 13 7 11 5 3 4 6 7 9 5 4 5 Querco-Fagetea 2 2 13 7 6 5 7 3 13 5 6 4 3 Elyno-Seslerietea 2 1 1 3 3 4 3 7 4 8 5 8 10 Thlaspietea rotundifolii 3 2 3 2 3 5 5 6 3 5 5 5 8 Quercetalia pubescentis 1 1 2 1 1 2 8 4 5 3 2 Quercetalia roboris-petraeae 3 2 2 2 2 Festuco-Brometea 2 2 2 3 2 200 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) H-Frh) since they thrive on their upper elevational limit, in close contact with subalpine beech stands and in very similar ecological conditions (compare also Dakskobler et al. 2000). The reason why stands of pre-alpine fir-beech forests with R.. hirsutum (Ho-mogyno-Fagetum s. lat. rhododendretosum) clustered so separately from other subalpine beech stands might be explained by their close proximity to the Julian Alps and thus by the large proportion of SE-Alpine taxa. Comparison of phytosociological groups in (fir-)beech syntaxa with Hairy Alpenrose (Rhododendron hirsutum) in the Julian Alps and Dinaric Mts showed several distinctions and significantly explained the cluster topology: species of the Erico-Pinetea, aspknietea and Elyno-Seslerietea were most frequently represented in Rhododendro-Fagetum and Homogyno-Fagetum rhododendretosum. Furthermore, species of the Festuco-Brometea and the Quercetalia roboris-petraeae were present only in stands of Homogyno-Fagetum and Rhododendro-Fage-tum, while species of the Quercetalia pubescentis were most frequent in stands of Rhododendro-Fagetum. These stands, although physiognomically resembling Polysticho-Fagetum rhododendretosum, are characterised by the presence of thermophilous taxa and floristic richness, indicating more moderate ecological conditions as a consequence of thriving at lower altitudes. Typically among beech forests, species of the Vaccinio-Piceetea are most frequent in fir-beech forests (Omphalodo-Fagetum s. lat., Homog-yno-Fagetum s. lat.). Generaly, tall herbs (Mulgedio-aconitetea) were more frequent in subalpine beech stands (Polysticho lonchitis-Fagetum). In terms of phytosociological groups, subalpine beech stands with Hairy Alpenrose differ from other subalpine beech stands in hosting a smaller number of the Fagetalia sylvaticae (aremonio-Fagion) and the Mulgedio-aconitetea species, but a higher number of asplenietea trichomanis species, the latter due to the higher proportion of stoniness. Interestingly, certain subalpine beech stands from Austria (Saxifrago-Fagetum calamagrostietosum and typicum) clustered together with stands of Polysticho-Fagetum from the SE Alps and Dinaric Mts, while other, more mesophilic stands, formed a distinct cluster (Figs. 2 & 3). According to Willner (2002, 2007), subalpine (Polysticho lonchitis-Fagetum) and altimontane (Ra-nunculo platanifolii-Fagetum) beech stands from the SE Alps and Dinaric Mts from the alliance aremo-nio-Fagion show no significant floristical differences from the Central-European subalpine and alti-montane beech stands (alliance Fagion sylvaticae). Therefore, according to the nomenclature code (Weber et al. 2000) and principle of priority, they should be treated as Saxifrago rotundifoliae-Fagetum Zukrigl 1989. However - our analysis, although not comprehensive enough - does not fully support this statement. Only thorough and extensive analysis covering Central European and SE European beech stands could solve the problem of the syn-taxonomic position of alliances (Fagion sylvaticae vs. aremonio-Fagion). Nevertheless, we classified subalpine beech stands with Hairy Alpenrose into a new subassociation Polysticho lonchitis-Fagetum rhododendretosum hirsuti, and as differential species for the subassociation we chose Rhododendron hirsutum, Rubus sax-atilis, Rosa pendulina, and Clematis alpina (Tab. 1). As inferred from Figs. 1-3, relevés 1-5 from the phytosociological table of Polysticho-Fagetum var. geogr. allium victorialis (Marinček 1996, cluster Ib - 5) were included into the new subassociation -rhododendretosum hirsuti, since the differential species (Polystichum lonchitis for the subassociation and (Gymnocarpium dryopteris, Solidago virgaurea, Vaccinium vitis-idaea for the variant) of the previously described subassociation and variant (-polysticheto-sum var. Gymnocarpium robertianum) were not most accurately chosen, and do not fully indicate specific ecological conditions of the selective stands. As inferred from our synoptic table and numerical analyses (but also field diagnostics) those relevés do not represent a "typicum" but rather distinct stands with a particular floristic combination, frequency and coverage of differential species, indicating unfavourable ecological conditions. While classification of the studied stands within subalpine beech stands was not questionable, delimitation of stands of syntaxa Polysticho-Fagetum s. lat. and Ranunculo platanifolii-Fagetum s. lat., specially in terms of good characteristic and differential species, is less conspicuous and in many cases (according to our personal observations and while comparing available phytosociological data, partly also following judgments of Horvat (1938) and Tomažič (1958, 1959) - see the introduction) the distinction between stands of the two syntaxa was based till now most frequently on their physiognomy rather than floristic principles, since appropriate characteristic and differential species are hard to define. Therefore only a thorough synoptic approach taking into account all (fir-)beech stands of the Illyrian floral province would properly challenge the current syntaxonomy. Although much beyond the scope of the present treatise, we sug- 201 Hacquetia 6/2 • 2007, 183-193 gest reconsidering the present synsystematics of subalpine-altimontane beech stands of the alliance Aremonio-Fagion and possibly trying to define them with the help of altitudinal variants. Additionally, and as already pointed out, the geographical delimitation of subalpine beech syntaxa in terms of different geographical variants proved to be of no significant synsystematic value (e.g. figs. 1-3). We classified the studied stands as follows: Querco-Fagetea Br.-Bl. et Vlieg. 1937 Fagetalia sylvaticae Pawl. 1928 Aremonio-Fagion (Horvat 1938) Borhidi in Török, Podani et Borhidi 1989 Polysticho lonchitis-Fagetum (Horvat 1938) Marinček in Poldini & Nardini 1993 rhododendretosum hirsuti Surina subass. nova Nomenclatural type for the subassociation Polys-ticho lonchitis-Fagetum rhododendretosum hirsuti subass. nova: relevé no. 4 in table 1, holotypus hoc loco. Generally, stands of the Polysticho-Fagetum rhododendretosum on the Snežnik plateau are neither frequent nor do they extend over larger areas. Although subalpine beech stands are quite widespread throughout the plateau, we found stands with prevailing Rhododendron hirsutum in the undergrowth only on extreme sites, namely: western and north-western slopes of the dolina of Stanišče, northerly exposed slopes between Lom and Kapetanova bajta (above the dolinas of Grda draga and Pekel), in the area of forest reserves of Planinc-Za-trep and Zdrocle, on ridges above dolinas of Grdobe and Medvedje doline, in the vicinity of Herbade above the dolina of Ilovca, and on the ridge west from Mt. Zaknovec. As a rule, stands of subalpine beech forest (Polysticho-Fagetum s. lat.) do not have a significant role in wood exploitation, although till the middle of the 20th century these stands were (mostly due to suitable trunk diameter) abundantly used for charcoal-burning. Nevertheless, their indispensable forest protective function has been recognized for a long time. CONCLUSIONS (a) Subalpine beech stands with Hairy Alpenrose (Rhododenron hirsutum) are floristically and ecologically quite distinct from other subalpine beech stands of the association Polysticho lonchitis-Fagetum s. lat. (b) Therefore, on the basis of the synoptic table, numerical analyses, and according to the current synsystematics of subalpine beech syntaxa of the Illyrian floral province, we classified them into the new subassociation Polysticho-Fagetum rho-dodendretosum hirsuti subass. nova (alliance Aremo-nio-Fagion). Differential species of the subassociation (Rhododendron hirsutum, Rubus saxatilis, Rosa pendulina, and Clematis alpina) sufficiently illustrate the ecological peculiarities of their sites. (c) Stands thrive on ecologically unfavourable growth sites, e.g. steep, rocky and northerly exposed slopes with heavy and long-lasting snow cover, and ridges exposed to strong Bora. (d) Stands of the Polys-ticho-Fagetum rhododendretosum subass. nova on the Snežnik plateau are neither frequent nor do they extend over larger areas. Till now they were observed only on the southern part of the plateau. ACKNOWLEDGMENTS The author thanks I. Dakskobler and P. Košir for many valuable discussions and helpful suggestions that improved the manuscript. APPENDIX 1. Localities of relevés: Slovenia, Dinaric Mts, Liburnian karst, Snežnik plateau: 1-3, 9 - Stanišče, 24. 7. 2002, leg. B. Surina. 4, 5, 7, 8, 10, 11 - Kapetanova bajta, 26. 7. 2005, leg. B. Surina. 6 - Ilovca (SW from Herbade), 29. 7. 2005, leg. B. Surina. 2. List of syntaxa - Adenostyletalia G. Br.-Bl. & J. Br.-Bl. 1931 - Allio victorialis-Fagetum (Tomažič 1958) Accetto 2002 nom. inval. - Aremonio-Fagion (Horvat 1938) Borhidi in Török, Podani & Borhidi 1989 - Asplenietea trichomanis Br.-Bl. in Meier & Br.-Bl. 1934 - Elyno-Seslerietea Br.-Bl. 1948 - Erico-Pinetea Horvat 1959 - Fagetalia sylvaticae Pawl. 1928 - Fagion sylvaticae Luquet 1926 - Homogyno sylvestris-Fagetum Marinček et al. 1993 rhododendretosum hirsuti Dakskobler 2004 - Hyperico grisebachii-Pinetum mugo Zupančič et al. 2004 var. geogr. Arabis scopoliana Zupančič et al. 2004 202 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) - Mulgedio-Aconitetea Hadač & Klika in Klika & Hadač 1944 - omphalodo-Fagetum (Tregubov 1957 corr. Puncer 1980) Marinček et al. 1993 var. geogr. Saxifraga cuneifolia Surina 2002 rhododendretosum hirsuti Dakskobler et al. 2000 - Polysticho lonchitis-Fagetum (Horvat 1938) Marinček in Poldini & Nardini 1993 rhododendretosum hirsuti Surina 2007 - Quercetalia pubscentis Klika 1933 - Querco-Fagetea Br.-Bl. & Vlieg. 1937 - Ranunculo platanifolii-Fagetum Marinček et al. 1993 - rhododendro hirsuti-Fagetum Accetto ex Daksko-bler 1998 var. geogr. Anemone trifolia Dakskobler 1998 - rhododendro hirsuti-Fagetum Accetto ex Daksko-bler 1998 var. geogr. Phyteuma columnae Daksko-bler nom. prov. - saxifrago rotundifoliae-Fagetum Zukrigl 1989 s. lat. - seslerion juncifoliae Horvat 1962 - Thlaspietea rotundifolii Br.-Bl. in Br.-Bl. & Jenny 1926 - vaccinio-Piceetea Br.-Bl. 1939 emend. Zupančič (1976) 2000 REFERENCES Braun-BlanquetJ. 1964: Pflanzensoziologie. Grundzüge der Vegetationskunde. 3. Auflage. Springer, Wien - New York, 865 pp. Dakskobler, I. 1997: Geografske variante asociacije seslerio autumnalis-Fagetum (Ht.) M. Wraber ex Borhidi 1963. 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Recieved 26. 2. 2007 Revision recieved 13. 11. 2007 Accepted 19. 11. 2007 For Appendix. 1, see Hacquetia Electronic Archives; http://www.versita.com/science/lifesciences/hacquetia 204 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) Table 1: Phytosociological table of the subalpine beech stands with Hairy Alpenrose (Rhododendron hirsutum) on Mt. Snežnik. Tabela 1: Analizna tabela subalpinskih bukovih sestojev z dlakavim slečem (Rhododendron hirsutum) na Snežniku. Succesive number 1 2 3 4* 5 6 7 8 9 10 11 Altitude (m) 1310 1330 1310 1420 1420 1340 1440 1430 1340 1390 1390 Exposition N N NE NNW N N N N N N N Inclination (°) 35 35 35 30 40 30 25 15 45 40 50 Relevé area (m2) 400 400 400 400 400 400 400 400 400 400 400 Stoniness (%) 10 40 25 40 40 50 20 50 40 20 70 Cover (%) Tree layer A 90 90 95 95 90 90 80 80 80 95 80 Shrub layer B 10 10 10 10 5 5 1 5 30 40 60 Herb layer C 80 80 70 60 50 40 70 50 60 70 30 Moss layer D 5 20 5 10 5 5 5 5 20 5 10 No. of seed plants 54 49 41 56 55 45 57 47 35 40 40 Fr. % Character and differential species of the association VP Polystichum lonchitis C + + + 1 + 1 + + + + 1 11 100 MA Salix appendiculata B C + + + + + + + + 1 + 8 73 ES Carex ferruginea C + + + 1 + 5 45 VP Lonicera caerulea C + + + + 4 36 EP Ribes alpinum B C + + + + 4 2 36 18 Differential species of the subassociation rhododendretosum hirsuti EP Rhododendron hirsutum C 1 3 3 3 2 3 3 3 4 3 3 11 100 EP Rubus saxatilis C 2 1 + 1 1 1 1 2 1 2 1 11 100 VP Clematis alpina C 1 2 2 2 2 2 2 2 2 2 2 11 100 VP Rosa pendulina B C 2 1 2 2 + 2 1 2 + 2 2 2 1 1 3 1 2 11 100 AF Aremonio-Fagion Cardamine enneaphyllos C + + + + + 1 + + + + + 11 100 Euphorbia carniolica C + + + + + + + + + + 10 91 Cardamine trifolia C 1 + + 1 + + + + 8 73 Hacquetia epipactis C + 1 + + 4 36 Cyclamen purpurascens C + 1 2 18 Vicia oroboides C + + 2 18 Rhamnus fallax B + 1 9 Omphalodes verna C + 1 9 FS Fagetalia sylvaticae A 5 5 5 5 5 5 5 5 4 5 5 Fagus sylvatica B + + 1 1 + + + 1 3 2 2 11 100 C + + + 1 + + + + + + Lonicera alpigena B C + + + + + 1 + + + 1 1 1 1 1 2 2 11 100 Dahpne mezereum B C + + + 1 1 + + + + + + + + + 1 + 10 91 Festuca altissima C + + + 1 1 1 1 + 1 9 82 Mercurialis perennis C 1 1 + + 1 + 1 + 1 9 82 Prenanthes purpurea C + + + + + + + + + 9 82 205 Hacquetia 6/2 • 2007, 183-193 Succesive number 3 4* 5 10 11 QP Quercetalia pubescentis Acer pseudoplatanus A C + + + + + + + + + + . 7 64 0 Epilobium montanum C + + + + + 5 45 Symphytum tuberosum C 1 + + + + 5 45 Ranunculus platanifolius C + + + + + .5 45 Helleborus niger C 1 + + . 3 27 Actaea spicata C + + + + . 4 36 Dryopterisfilix-mas C + + + .3 27 Geranium robertianum C + + + .3 27 Mycelis muralis C + + . 2 18 Adoxa moschatelina C + + 2 18 Lilium martagon C + + . 2 18 Aruncus dioicus C + .1 9 Galium sylvaticum C + .1 9 Lathyrus vernus C + .1 9 A Sorbus aria B C + + + + + + + + + 9 82 Querco-Fagetea Anemone nemorosa C 1 2 1 1 1 1 2 2 1 2 2 11 100 Carex digitata C + + + + + 5 45 Convalaria majalis C + + 1 + 4 36 Hepatica nobilis C + + + 3 27 Dactylorhiza maculata C + + 2 18 Poa nemoralis C + 1 9 Mulgedio-Aconitetea Veratrum album subsp. album C 1 + + + + + + + + + + 11 100 Phyteuma ovatum C 1 1 1 2 1 + + 1 1 1 10 91 Polygonatum verticillatum C + + + + + + 1 1 8 73 Pleurospermum austriacum C + + + 3 27 Athyrium filix-femina C + + 2 18 Viola biflora C + + 2 18 Aconitum ranunculifolium C + 1 9 Allium victorialis C + 1 9 Saxifraga rotundifolia C + 1 9 Cirsium waldsteinii C + 1 9 Vaccinio-Piceetea Valeriana tripteris C 1 1 1 2 1 + 1 1 1 2 2 11 100 Homogyne sylvestris C 1 1 1 2 1 1 1 1 1 1 1 11 100 Gentiana asclepiadea C 1 + + + + + + 1 + + + 11 100 Solidago virgaurea C A + + + + + + + + + + + + 10 91 Abies alba B C + 1 1 + + + + 1 + 1 + 8 73 Vaccinium myrtillus C 1 + 1 1 2 2 2 1 8 73 Maianthemum bifolium C + + + + + + 1 + 8 73 Huperzia selago C + + + + + + + 7 64 Aposeris foetida C 2 1 + + + 5 45 Veronica urticifolia C + + 1 1 + 5 45 Hieracium murorum agg. C + + + + + 5 45 1 2 6 7 8 9 + + + + + + 206 Boštjan Surina: Subalpine Beech forest with Hairy Alpenrose ... on Mt. Snežnik (Liburnian karst, Dinaric Mts) Succesive number 1 2 3 4* 5 6 7 8 9 10 11 Vaccinium vitis-idaea C + + + 2 4 36 Dryopteris dilatata C + + + + 4 36 Luzula sylvatica C A + + + + + 4 36 Picea abies B C + + + + + 4 36 Gymnocarpium dryopteris C + 1 1 3 27 Oxalis acetosella C 1 + + + 4 36 Laserpitium krapfii C + + 2 18 Thelypteris phegopteris C + + 2 18 Erico-Pinetea Calamagrostis varia C 2 1 1 1 1 2 2 1 2 1 10 91 Cirsium erisithales C + + + + + + 1 + 8 73 Aquillegia nigricans C + + + + + 5 45 Erica herbacea C + + 2 18 Juniperus sibirica C + + 2 18 Elyno-Seslerietea Aster bellidiastrum C 1 + + 1 1 + + 7 64 Erigeron glabratus C + 1 9 Heliosperma alpestre C + 1 9 Asplenietea trichomanis Asplenium viride C + + + + 1 1 + + + + + 11 100 Campanula justiniana C + + 1 + 1 + + 7 64 Carex brachystachys C 1 1 + + + 5 45 Moehringia muscosa C + + + + 4 36 Cystopteris fragilis C + + + + 4 36 Cystopteris regia C + + + 3 27 Asplenium ruta-muraria C + + + 3 27 Asplenium trichomanes C + 1 9 Festuca stenantha C + 1 9 Thlaspietea rotundifolii Adenostyles glabra C 1 2 2 1 + + 1 1 1 9 82 Campanula cochleariifolia C 1 + + + 4 36 Gymnocarpium robertianum C + + + + + 5 45 Dryopteris montana C + 1 9 Other species A + Sorbus aucuparia B C + + + + + + + 7 64 Thalictrum aquilegiifolium C + + + + 1 45 Rubus idaeus C + + 2 18 Parnassia palustris C + 1 9 Mosses and lichens Ctenidium molluscum D 2 1 1 2 2 2 2 2 2 2 2 11 100 Tortella tortuosa D 1 + 1 1 1 1 1 1 1 1 1 11 100 Schistidium apocarpum D + + + 1 1 1 + 1 + 1 10 91 Fissidens cristatus D 1 + 1 1 1 + 1 + 8 73 Cladonia pyxidata D + + + + + + + + 8 73 Dicranum scoparium D 1 + + + + + + 7 64 Plagiochila asplenioides D + + 1 1 + 1 6 55 Conocephalum conicum D + + + + 4 36 207 Hacquetia 6/2 • 2007, 195-208 Succesive number 1 2 3 4* 5 6 7 8 9 10 11 Peltigera leucophlebia D. + + + + 4 36 Hypnum cupressiforme D 1 + + . 3 27 Brachythecium sp. D. + + + 3 27 Distichium capillaceum D. . + + 1. 3 27 Pleurozium schreibersi D. . + + . + 3 27 Leptogium saturninum D. . + + 2 18 Rhytidiadelphus triquetrus D. + . + 2 18 Solorina saccata D. + . + 2 18 Bryum sp. D. + 1 9 Peltigera sp. D. + 1 9 Pohlia elongata/elongata D. . + 1 9 Polytrichum alpinum D. + 1 9 Polytrichum commune D. + 1 9 208