COBISS 1.01 AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION PREZRTA SUBASOCIACIJA V SEKUNDARNI SMREKOVI ZDRUŽBI Mitja ZUPANČIČ 1 & Vinko ŽAGAR 2 ABSTRACT UDC:582.47(497.4) An overlooked sub-association in secondary spruce asso- ciation A new sub-association of secondary spruce forest Aposeri- do-Piceetum Zupančič 1999 galietosum rotundifolii subass. nova is described. Key words: phytocoenology, ecology, Picea abies, Slo- venia IZVLEČEK UDK: 582.47(497.4) Prezrta subasociacija v sekundarni smrekovi združbi Opisana je nova subasociacija sekundarnega smrekovega gozda Aposerido-Piceetum Zupančič 1999 galietosum rotundi- folii subass. nova. Ključne besede: fitocenologija, ekologija, Picea abies, Slo- venija INTRODUCTION D u r i n g a r e v i e w o f t h e m a t e r i a l o n s p r u c e p h y t o c o e n o s e s , we found that we had overlooked five phytocoenological relevés on rather problematic spruce phytocoenoses. These relevés had not actually been overlooked but we had been unable to decide in which spruce phytocoeno- sis to place them, so we did not present them in the mon- ograph on spruce forests of Slovenia (Zup nčič 1999). We did not rush into publication of these relevés for a number of reasons, one of the main ones of which was that it concerns a secondary spruce phytocoenosis on beech habitat, which is not so important for operational forestry but interesting from a scientific point of view because of its synsystematic classification and increas - ing the biodiversity of forest vegetation. In terms of flo - ristic and vegetation research, we hesitated between two possible solutions; whether it is the new central Europe- an spruce association, not yet recognised here, Galio ro- tundifolii-Piceetum J. & M. Bartsch 1940 or a sub-associ - ation of our local secondary association Aposerido-Pi - ceetum Zupančič 1999. This paper provides an answer to this synsystematic question. The phytocoenological relevés shown in the phyto - coenological table were recorded in 1962 on the slopes between Podvolovljek and Veža – Planica in the foothills of the Savinja Alps (Savinjska dolina, Štajerska). The processing of the vegetation material has been done according to the standard Central European meth- od (Br un-Bl nquet 1964). In naming the flora, we respected Mala flora Slovenije (M rtinčič et al. 2007). For the determination of horological groups and bio - logical forms of flora, we used Poldini’s atlas (Poldini 1991). 1 Dr., Mitja ZUPANČIČ, SAZU, Novi trg 3, SI – 1000 Ljubljana 2 Vinko ŽAGAR, Bevkova c. 1, SI – 1290 Grosuplje FOLIA BIOLOGICA ET GEOLOGICA 51/2, 109–130, LJUBLJANA 2010 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION DISCUSSION The dilemma indicated in the introduction, of the syn- taxonomical classification of the spruce phytocoenosis shown in the Phytocoenological Table with five relevés, guided us in two directions. The first, superficial view of the phytocoenological relevés suggested that the relevés should be classified in the Central European spruce as - sociation Galio rotundifolii-Piceetum J. & M. Bartsch 1940. The association was described by J. & M. B rtsch (1940) in a monograph on the vegetation of Schwarz- wald, then designated Picea-Galium rotundifolium-ass., with four relevés. Oberdorfer (1957) later designated it correctly synsystematically as Galio rotundifolii-Picee- tum and presented it with twenty-four relevés in synthe - sised tables. He classified the already published relevés of J. & M. Bartsch among these relevés. The association was supposed to be distributed not only in southern Germany but also in Austria (W llnöfer in Mucin et al. 1993). What reminded us of a similarity with the as - sociation Galio rotundifolii-Piceetum? Primarily the presence of the diagnostically important species Galium rotundifolium with piceetal species and the relatively considerable presence of fagetal species in both the Bart- sch and Oberdorfer relevés and in our tables and the si - milarity in terms of the characteristic and differential species of the association. There was also similarity in the occurrence of the phytocoenosis, its syndynamics and, partially, also soil conditions. From the description of J. & M. Bartsch, it is evident that the association is secondary on habitats of beech forest, although they do not mention this particularity but say that the associati- on Galio-Piceetum sometimes eats into fagetal forest, which means that it overgrows beech habitats. There are even often whole carpets of the species Oxalis acetosella in the association, because of which they believe that it is a type of beech forest. Their finding is important that the association Galio-Piceetum approaches the vegetati- on of broad-leaved forest. The soils, similar to here, are fresh, moderately deep, light. There is a great difference in the geological base, whereby in Germany it consists of gneiss and variable sandstones ( Buntsandstein) (B rt- sch 1940), and here a carbonate base with very acid humus (pH 3.6–5.1) (Zup nčič 1999). It is similar with the description of the association by Oberdorfer ( 1957), when he says that it is rich in species of mixed fir forest on a silicate base. When he compares the vitality of beech in the association Galio- Piceetum he says that this is always better than in the association Piceo-Abietetum in western Schwarzwald. He stresses that the habitat of the association Galio-Pi - ceetum, is richer than in the western fringe mountains (of Schwarzwald). He finds that the soils are fresh, mull humus, in basins brown, skeletal. The geological base or bedrock consists of granite and gneiss. From these two descriptions and on the basis of the phytocoenological tables of J. & M. B rtsch (1940) and Oberdorfer (1957) it is clear that the association Galio- Piceetum in the area of Schwarzwald is of secondary emergence on habitats of acid beech forest. The difference between our phytocoenosis and the Schwarzwald association Galio-Piceetum at first site is in the geological base. In Schwarzwald this is non-car- bonate (silicate), and the bedrock here is carbonate (limestone and dolomite). The soils also differ, although they are sometimes similar since, because of decades (or centuries) long culture of spruce, they have changed and can be distric, deep brown, moderate podsol or there is a ra nge of soi ls, f rom rend zi na to moderately deep brow n soils, with poor absorption capacities. Floristic analysis showed that our phytocoenosis is species richer and more varied than the association Galio-Piceetum. Com- parison between them shows low similarity, the index according to Sørensen is 42 and by Jaccard only 27. The low indexes confirm the difference of the phytocoenoses, so that our phytocoenosis cannot be classified in the as - sociation Galio rotundifolii-Piceetum, although it is fair - ly close to it. We were additionally interested in the presentation of the association Galio-Piceetum in Austria (W ll - nöfer in Mucin et al. 1993). Wallnöfer selected only the species Galium rotundifolium as characteristic, which she says is poor and transgressional. She adds the distin- guishing species: Anemone nemorosa, Doronicum aus- triacum, Primula elatior and additionally some others in relation to the association Veronico latifoliae-Piceetum and dominant and constant accompanying species of the association Galio-Piceetum. She says that piceetal species predominate in the association and that it settles mon- tane and the lower subalpine zone on silicate, mineral rich and moderately acid bedrocks, in which there are deep, fresh, sometimes podzol brown soils with mull or moder humus. She concludes that the association Galio rotundifolii-Piceetum is a climactic association. We were encouraged to present the opinion of Wall- nöfer about the association Galio -Piceetum because of her classification of M. Wr ber’s (1959) association Galio rotundifolii-Abietetum M. Wraber (1955) 1959 in the association Galio rotundifolii-Piceetum J. & M. Bar- tsch 1940. She considers the fir association of M. Wraber to be a synonym for Bartsch’s spruce association. Will- ner & Gr bherr (2007) are of the same opinion as Wallnöfer. Comparison between the associations Galio- Piceetum in Schwarzwald and Galio-Abietetum in Slo- FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 110 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION venia showed a difference in the associations, which is confirmed by the index of similarity, which is 51 accord - ing to Sørensen and 34 according to Jaccard. Compari- son between our spruce association and M. Wraber’s association Galio-Abietetum shows an index of even smaller similarity: 42 by Sørensen and 25 by Jaccard. Wallnöfer’s claim, therefore, that M. Wraber’s associa - tion Galio-Abietetum is a synonym for Bartsch’s associa - tion Galio-Piceetum, does not hold. The associations are independent and syntaxonomically stable. In line with the above findings, we continued with comparisons among secondary spruce phytocoenoses known in Slovenia. We found that the spruce phytocoe - nosis is most comparable with the secondary spruce as- sociation Aposerido-Piceetum Zupančič (1978) 1999. The characteristic species of Aposeris foetida is present in four relevés, with a solid medium cover value of 400 and sociability (1-2) and a high level of presence (V = 80%). A considerable representation of fagetal species is charac - teristic of the secondary association Aposerido-Piceetum on habitats of primary basiphilous beech association. The presence is also important of characteristic species of the Illyrian alliance of beech forests Aremonio-Fagion , among which are distinguishing species for a geographic variant of the secondary spruce association Aposerido- Piceetum var. geogr. Helleborus niger Zupančič 1999. In our case, it is an original habitat of pre-Alpine fir-beech forest Homogyno sylvestris-Fagetum, in which spruce have advanced. Part of the spruce is planted, part has settled spontaneously. Because of the many years of impact of the fall of spruce needles, exhaustion of the upper soil horizons (Oh, A) and the activity of the spruce root system, the soils have worsened over the course of time, above all become more acid and enabled the settle- ment of acidophilic piceetal flora. The carbonate bed - rock has enabled the survival of neutral and moderately basiphilous fagetal flora. This secondary spruce phyto - coenosis thus contains a mosaic of intermixed piceetal and fagetal flora in fairly equal ratio in terms of surface coverage, and in terms of number of species a prevalence of fagetal species. Of fagetal species, eight species are present from the Illyrian alliance of beech forests Are- monio-Fagion, three of which are distinguishing species for the geographic variant: Helleborus niger subsp. niger, Cyclamen purpurascens and Cardamine trifolia with the piceetal species Homogyne sylvestris, and eleven South- easteuropean-Illyrian species are represented in our spruce phytocoenosis. In relation to the above floristic similarities, which are clearly shown in the phytocoenological table, we cal- culated an index of similarity according to Sørensen (σ) s and Jaccard (σ j ) between our spruce phytocoenosis and a) the Bavarian association Galio-Piceetum, for which σ = 42 and σ j = 27, and b) the association Aposerido-Pi- s ceetum Zupančič 1999 (Zup nčič 1999, Table 11), for which σ = 59 and σ j = 42. s Floristic analysis, together with the index of simi - larity of the phytocoenoses, indicates a relatedness of our secondary spruce phytocoenosis with the associa- tion Aposerido-Piceetum; although not very convinc- ingly, satisfactorily. However, the index of similarity with the association Galio-Piceetum is smaller and does not achieve optimality. The difference is in the floristic content of the phytocoenoses, above all the number of fagetal species - there are fewer than half of these in the association Galio-Piceetum - and partly also in species of the classes Betulo-Adenostyletea (=Mulgedio-Aco- nitetea), Tr i f o l i o - G e ra n i e t e a and Asplenietea trichoma- nis, which are all more or less basiphilous. In terms of coverage, acidophilic and basiphilous species are equally divided in the association Galio-Piceetum. Floristic and, consequently, vegetation analysis show that our secondary spruce phytocoenosis can be classified in the association Aposerido-Piceetum, but as a new sub-association, which we have named after the species Galium rotundifolium, so Aposerido-Piceetum galietosum rotundifolii subass. nova or Aposerido-Picee- tum var. geogr. Helleborus niger galietosum rotundifolii subass. nova. The sub-association A.-P. galietosum shows a partial link or relatedness with the association Galio- Piceetum, as the indexes of similarity of the phyto - coenoses according to Sørensen and Jaccard confirm. FLORISTIC AND SOCIOLOGICAL COMPOSITION OF THE SUB-ASSOCIATION In the presentation of the association Aposerido-Picee- tum (Zup nčič 1999), because of the extensiveness of the monograph on spruce phytocoenoses, we left out some of the elements of the sociological composition of the association, and we are taking this opportunity to complete them. It is primarily a matter of supplemen - ting the characteristic combinations of species, stratifi - cation, life form spectrum, phytocoenological and horo - logical groups of the association. Before that, we must additionally describe the choice of differential species of the sub-association. The floristic table, with five relevés, shows the sub- association Aposerido-Piceetum galietosum rotundifolii subass. nova with 153 plant species. There are 123 flow- FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 111 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION ering plants, 8 ferns, 21 mosses and 1 fungus. Relevés 5 and 1 have the lowest number of floristic species (61, 62), relevés 4 and 2 have the most (88, 81), and relevé 3 has 71 species. The relevés are sufficiently homogeneous. The ecological conditions of the sub-association do not differ essentially from those previously described for the association Aposerido-Piceetum . Mean annual temperature is between 4 and 8 o C with annual precipi- tation of 1750 mm and more. Soils are brown (carbon- ate), acid to moderately acid with pH KCl below 4 in Oh and around 6 in horizons A and B (Zup nčič 1999). The distinguishing species for the sub-association Aposerido-Piceetum galietosum rotundifolii are Galium rotundifolium and Goodyera repens. Their ecological and vegetation designations are summarised on the basis of Oberdorfer (1979) and our own observations. The species Galium rotundifolium L. appears in fir, spruce and mixed fir-spruce and, more rarely, in acid beech forests. It is found here and there in dwarf pine. It grows on fresh, nutrient and base moderately rich, but lime poor, moderately acid soils with moder humus. It is classified among shade-loving species. Synsystemati - cally, it is described as a species of the sub-alliance Abi- eti-Piceenion or Galio-Abietion. Phytogeographically it belongs among Subatlantic-Submediterrannean or Eu- roasian species. The species Goodyera repens (L.) R. Br. grows in fir, spruce and acid Austrian pine forests, rarely in scrub pine. It inhabits moderately dry to fresh habitats where the soils are more or less base-rich (with acidic moder humus). It is a semi-shade species, which tends to appear among acidophilic mosses in the acidic needle-fall of fir, spruce or Austrian pine. For the most part, it is classi- fied as a characteristic species of the order Va c c inio -P i- ceetalia. Phytogeographically, this is a Northeastconti- nental-Circumpolar or Circumboreal species. The characteristic combination of species gives the following picture: CHARACTERISTIC SPECIES Aposeris foetida* DISTINGUISHING SPECIES OF THE GEOGRAPHIC VARIANT Helleborus niger subsp. niger* Cyclamen purpurascens* Cardamine trifolia* DISTINGUISHING SPECIES OF THE SUB-ASSOCIATION Galium rotundifolium VACCINIO-PICEETEA s. lat. Galium rotundifolium Hylocomium splendens Melampyrum sylvaticum Rhytidiadelphus triquetrus Mnim punctatum Hypnum cupressiforme Picea abies Aposeris foetida* Hieracium sylvaticum Maianthemum bifolium Vaccinium myrtillus Gentiana asclepiadea Larix decidua Peltigera leucophlebia Polytrichum formosum Luzula pilosa Atrichum undulatum Oxalis acetosella Luzula luzuloides QUERCO-FAGETEA s. lat. Helleborus niger subsp. niger* Epipactis helleborine Cyclamen purpurascens* Lamiastrum flavidum Aremonia agrimonioides* Stachys sylvatica Cardamine trifolia* Viola reichenbachiana Knautia drymeia subsp. drymeia* Euphorbia amygdaloides Fagus sylvatica Melica nutans Sanicula europaea Pteridium aquilinum Mycelis muralis Veronica officinalis Mercurialis perennis Digitalis grandiflora Acer pseudoplatanus Campanula witasekiana* Daphne mezereum Hypericum montanum Eurhynchium zettersledtii Ctenidium molluscum Salvia glutinosa Carex digitata FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 112 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION ERICO-PINETEA s. lat. Carex alba Buphthalmum salicifolium Polygala chamaebuxus Calamagrostis varia Rubus saxatilis BETULO-ADENOSTYLETEA s. lat. (=MULGEDIO-ACONITETEA s. lat.) Senecio fuchsii TRIFOLIO-GERANIETEA s. lat. Cruciata glabra MOLINIO-ARRHENATHERETEA s. lat. Ajuga reptans ASPLENIETEA TRICHOMANIS s. lat. Asplenium trichomanes OTHER SPECIES Fragaria vesca Cladonia pyxidata * Southeasteuropean-Illyrian species As the table shows, the chosen characteristic species include the characteristic species of the association Apo- seris foetida, three distinguishing species of the geo - graphic variant Helleborus niger subsp. niger, Cyclamen purpurascens and Cardamine trifolia and the distin- guishing species of the sub-association Galium rotundi- folium, which confirms our decision on membership of the sub-association Aposerido-Piceetum (var. geogr. Hel- leborus niger). Species are balanced between the classes Va c c inio-Piceetea s. lat. and Querco-Fagetea s. lat., which signifies confirmation of our finding that it is a second - ary spruce phytocoenosis on original beech (fir-beech) habitats. The association Aposerido-Piceetum must cer- tainly be placed in the class Va c c inio -P ic e e te a. The mod - est position of species of the classes Erico-Pinetea s. lat., Betulo-Adenostyletea s. lat. (=Mulgedio-Aconitetea s. lat.), Molinio-Arrhenatheretea s. lat. and »other species«, of which there are a total of twelve, additionally shows the secondary nature and partial degradation of the habitat. The stratification of the sub-association Aposerido- Piceetum galietosum rotundifolii is simple and divides into upper stand layers of majority spruce and minority beech, with here and there the presence of trees of Larix decidua, Prunus avium and Fraxinus ornus. Other tree species, Acer pseudoplatanus, A. campestre, Fraxinus ex- celsior, Ostrya carpinifolia, Sorbus aria and Juglans regia appear only in the shrub layer and, with other shrubs, cover from 0 to 60 % of the surface. The herb layer is not particularly richly developed and, as has already been said, is balanced between piceetal and fagetal species. The moss layer is for the most part modest, but here and there also strongly represented. The plant life spectrum of the phytocoenosis shows the more or less calm living conditions, which predomi- nate in the moderate zone. The increasing presence of phanerophytes indicates warmer conditions, although it Clinopodium vulgare Euphorbia cyparissias is not always the case, since they are for the most part shrub growth (13.7 %) and, with the slightly increased share of chaemophytes, indicate a shift towards slightly more unfavourable conditions than prevail in the moder - ate zone. These living conditions are more favourable for secondary spruce phytocoenoses than for primary spruce phytocoenoses. The life form spectrum indicates living conditions of the habitat between original beech (fir- beech) phytocoenoses and the partial influence of sec - ondary spruce phytocoenoses. Spruce worsens the soil through its needle-fall, shallow roots and constant shad- PLANT LIFE FORMS SPECTRUM No % Total % PHANEROPHYTA 31 20,2 P. caespitosa 11 7,2 P. scaposa 10 6,5 Nano P. 8 5,2 P. scandentia (P. lian.) 2 1,3 CHAMAEPHYTA 27 17,7 Ch. suffrutescentia 2 1,3 Ch. fruticosa 2 1,3 Ch. reptantia 1 0,7 Ch. succulenta. 1 0,7 Ch. briophyta 16 10,4 Ch. lichenosa 5 3,3 HEMICRYPTOPHYTA 65 42,4 H. scaposa 33 21,5 H. caespitosa 15 9,8 H. rosulata 11 7,2 H. reptantia 4 2,6 H. biennis 2 1,3 GEOPHYTA 27 17,7 G. rhizomata 22 14,4 G. bulbosa 4 2,6 G. parasitica 1 0,7 THEROPHYTA 1 0 , 7 T. scaposa 1 0,7 UNIDENTIFIED SPECIES 2 1,3 2 1 , 3 Tot a l 153 100 153 100 Table 1: Plant life forms spectrum FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 113 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION ing of the habitat. There is ongoing acidification of the soil through needle-fall, which poorly decomposes, with unilateral exploitation of nutrients of the upper horizons. Analysis of the horological groups shows that more than two thirds of the geoelements are cool loving (70.5%). We included mosses and lichens within the cool group (vascular flora 56.8% and moss-lichen flora 13.7%). There are most European, Circumboreal and Euroasiatic geoelements, closely followed by Paleotem - perate and Eurosiberian geoelements. All this confirms that the secondary spruce phytocoenosis thrives in somewhat harsher living conditions, as the life form spectrum also indicates. GEOELEMENTS No % Mediterranean-montane 15 9,8 Circum-Boreal 23 15,0 European 28 18,3 Eurasiatic 17 11,1 Paleotemperate 10 6,5 Eurosiberian 9 5,9 Northeast-Illyrian 7 4,7 South-Illyrian 2 1,3 Southeast-European 2 1,3 Euro-Mediterranean 2 1,3 Pontic 3 2,0 Mediterranean-Pontic 3 2,0 Mediterranean-Atlantic 2 1,3 Cosmopolitan 6 3,9 Cultivated species 1 0,6 Mosses and lichens 21 13,7 Unidentified species 2 1,3 Tot a l 153 10 0 , 0 0 Table 2: Horological groups Table 3 shows the ratio among phytocoenological groups. It is to be expected that fagetal species of the original beech (fir-beech) phytocoenoses are best repre - sented (45.8 %). In view of the carbonate bedrock there are most basiphilous species (54.3 %) from the classes Querco-Fagetea s. lat., Betulo-Adenostyletea s. lat. (=Mulgedio-Aconitetea s. lat.), Tr i f o l i o - G e ra n i e t e a s. lat. and Asplenietea trichomanis s. lat. Then follow piceetal species (24.8 %), which have become established after long ‘sprucification’ of the habitat and are acidophilic. We classif y as acidophilic species, in addition to the class Va c c inio -P ic e e te a s. lat., also Erico-Pinetea s. lat. and moss and lichen flora (34.6 %). There are fewer neutral species (11.1 %) from the class Molinio-Arrhenatheretea s. lat. and »other species«. The picture of the phytocoe - nological groups confirms the secondary nature of the spruce phytocoenosis. CLASS No % VACCINIO-PICEETEA 38 24,8 QUERCO-FAGETEA 70 45,8 ERICO-PINETEA 7 4 , 6 BETULO-ADENOSTYLETEA (MULGEDIO-ACONITETEA) 5 3 , 3 TRIFOLIO-GERANIETEA 4 2 , 6 MOLINIO-ARRHENATHERETEA 6 3 , 9 ASPLENIETEA TRICHOMANIS 4 2 , 6 OTHER SPECIES 11 7,2 MOSES AND LICHENES 8 5 , 2 Tot a l 153 10 0 , 0 Table 3: Phytocoenological groups CONCLUSION In accordance with the floristic and vegetation analysis of the secondary spruce phytocoenosis, we found that we can place it in the association Aposerido-Piceetum Zupančič 1999 as an independent sub-association Apo- serido-Piceetum galietosum rotundifolii subass. nova. The holotype of the sub-association is relevé 2 from the phytocoenological table 1. The sub-association Aposerido-Piceetum galietosum rotundifolii, in terms of its floristic composition, is close to the Central European Galio rotundifolii-Piceetum J. & M. Bartsch 1940. There is partial (lesser) similarity or (distant) relatedness between them. It must also be taken into account that the associations are located in differ - ent floral provinces. The association Aposerido-Picee- tum is in the Illyrian floral province, as is already indi - cated by the designation of the association according to Southeasteuropean-Illyrian species and the characteris- tic species of the association, Aposeris foetida. In addi- tion to the characteristic species, other Southeasteuro- pean-Illyrian species appear in the association: Helle- borus niger subsp. niger, Cyclamen purpurascens, Anem- one trifolia, Homogyne sylvestris, Cardamine trifolia; which defined the association for us as a geographic variant of the Illyrian floral province Aposerido-Picee- tum var. geogr. Helleborus niger Zupančič 1999. The as - sociation Galio rotundifolii-Piceetum J. & M. Bartsch 1940 is native to the Central European floral province. After several generations of spruce forest on a beech habitat, basiphilous Southeasteuropean-Illyrian and fagetal species will gradually disappear, especially those that are strictly bound to carbonate bedrock. At that time, the secondary spruce phytocoenosis would very similar to those from the Central European province, e.g., the association Galio-Piceetum. FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 114 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Finally, we would draw attention to the comparison between the associations Galio rotundifolii-Piceetum J. & M. Bartsch 1940 and Galio rotundifolii-Abietetum M. Wraber (1955) 1959. Despite the claims of Central Euro - pean phytocoenologists (Mucin , Gr bherr & W ll- nöfer 1993, Willner & Gr bherr 2007), that it is the same association in both cases, thus the originally de- scribed Galio-Piceetum ( th e n am e o f th e a s s o c i a ti o n Galio-Abietetum is claimed to be a synonym), this is not the case. Comparative analysis according to Sørensen and Jaccard clearly showed that they are two different associations, i.e., secondary spruce and primary fir phy - tocoenoses. The finding of a new forest phytocoenosis is a fur - ther contribution to the great biodiversity of forest veg - etation in Slovenia. It is caused by the phytogeographic position, the diversity of the geological base, mezocli - matic and microclimatic conditions and, not least, the orographic articulation of Slovenia, as well as other, less well-known biotic and abiotic factors. POVZETEK Uvod Pri pregledu gradiva o smrekovih fitocenozah smo ugo - tovili, da smo prezrli pet fitocenoloških popisov o neko - liko problematični smrekovi fitocenozi. Pravzaprav ti popisi niso bili prezrti, temveč se nismo mogli odločiti, v katero smrekovo fitocenozo bi jih uvrstili, zato jih v monografiji o smrekovih gozdovih Slovenije (Zup nčič 1999) nismo predstavili. Z objavo teh popisov nismo hi- teli iz več razlogov. Med glavnimi razlogi je bilo dejstvo, da gre za sekundarno smrekovo fitocenozo na bukovem rastišču, ki za operativno gozdarstvo ni tako pomemb - na, zanimiva pa je z znanstvenega vidika, in sicer zaradi njene sinsistematske uvrstitve in povečanja biodiverzi- tete gozdne vegetacije. Glede na floristične in vegetacij - ske raziskave smo omahovali med dvema mogočima rešitvama, in sicer, ali gre za novo, pri nas še neuvelja- vljeno srednjeevropsko smrekovo asociacijo Galio ro- tundifolii-Piceetum J. & M. Bartsch 1940 ali za subasoci - acijo naše znane sekundarne asociacije Aposerido-Picee- tum Zupančič 1999. Odgovor na to sinsistematsko vpra - šanje je v tej razpravi. Fitocenološki popisi, prikazani v Fitocenološki ta- beli, so bili posneti v letu 1962 na pobočju med Podvolo - vljekom in Vežo - Planico v podnožju Savinjskih Alp (Savinjska dolina, Štajerska). Obdelava vegetacijskega gradiva je urejena po stan- dardni srednjeevropski metodi (Br un-Bl nquet 1964). Pri poimenovanju flore smo upoštevali Malo floro Slovenije (M rtinčič et al. 2007). Za določitev ho- roloških skupin in bioloških oblik flore smo uporabili Poldinijev atlas (Poldini 1991). Razprava V uvodu nakazana dilema sintaksonomskega uvrščanja smrekove fitocenoze, kot jo prikazuje Fitocenološka ta - bela s petimi popisi, nas je vodila v dve smeri. Prvič, po- vršen pregled fitocenoloških popisov nam je narekoval, da popise uvrstimo v srednjeevropsko smrekovo asocia- cijo Galio rotundifolii-Piceetum J. & M. Bartsch 1940. Asociacijo sta popisala zakonca B rtsch (1940) v mo- nografiji o vegetaciji Schwarzwalda, tedaj pod imenom Picea-Galium rotundifolium-ass. s štirimi popisi. Kasne- je jo je Oberdorfer (1957) sinsistematsko pravilno po - imenoval Galio rotundifolii-Piceetum in jo predstavil s štiriindvajsetimi popisi v sintezni tabeli. Med te popise je uvrstil že objavljene popise zakoncev Bartsch. Asocia- cija naj ne bi bila razširjena le v južni Nemčiji, temveč tudi v Avstriji (W llnöfer v Mucin et al. 1993). Kaj nas je spominjalo na podobnost z asociacijo Galio rotun- difolii-Piceetum? Predvsem zastopanost diagnostično pomembne vrste Galium rotundifolium s piceetalnimi vrstami in razmeroma precejšnja navzočnost fagetalnih vrst tako v Bartschevi in Oberdorfejevi kot v naši tabeli ter podobnosti z značilnimi oziroma diferencialnimi vr - stami asociacije. Podobnost pa je bila tudi pri nastanku fitocenoze, njeni sindinamiki in deloma v talnih razme- rah. Iz opisa zakoncev Bartsch je razvidno, da je asocia- cija sekundarna na rastišču bukovih gozdov, čeprav tega posebej ne navajata, govorita pa, da se asociacija Galio- Piceetum včasih zajeda v fagetalne gozdove, kar pomeni, da porašča bukova rastišča. Večkrat so v asociaciji cele preproge vrste Oxalis acetosella, za katero menita, da je vrsta bukovih gozdov. Pomembna je njuna ugotovitev, da se asociacija Galio-Piceetum približuje vegetaciji li- stnatih gozdov. Tla so, podobno kot pri nas, sveža, zmer- no globoka, rahla. V elika razlika je v geološki podlagi, kjer so v Nemčiji gnajsi in pisani peščenjaki ( Buntsan- dstein) (B rtsch 1940), pri nas pa karbonatna podlaga z zelo kislim humusnim horizontom (pH 3,6–5,1) (Zu- p nčič 1999). Podobno je z opisom združbe Oberdorferj (1957), ko ta pravi, da je bogata z vrstami mešanega jelo- vega gozda na silikatni podlagi. Ko primerja vitalnost FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 115 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION bukve v asociaciji Galio-Piceetum pove, da je ta vedno boljša kot v asociaciji Piceo-Abietetum v zahodnem Sch- warzwaldu. Poudarja, da je rastišče asociacije Galio-Pi- ceetum v primerjavi z zahodnim obrobjem gorovja (Sch- warzwalda) bogatejše. Ugotavlja, da so tla sveža, sprste- ninasta, v kotanjah rjava, skeletna. Geološko podlago sestavljata granit in gnajs. Iz teh dveh opisov in na osnovi fitocenoloških tabel zakoncev B rtsch (1940) in Oberdorferj (1957) se jasno vidi, da je asociacija Galio-Piceetum na območju Schwarzwalda sekundarno nastala na rastišču kislih bu- kovih gozdov. Razlika med našo fitocenozo in schwarzwaldsko asociacijo Galio-Piceetum je na prvi pogled v geološki podlagi. V Schwarzwaldu je nekarbonatna ( silikatna ), matična podlaga pri nas pa je karbonatna (apnenčasta in dolomitna). Tudi tla se razlikujejo, vendar so včasih po - dobna, saj so se zaradi večdesetletne (stoletne) kulture smreke spremenila in so lahko distrična, globoka rjava, zmerno opodzoljena oziroma imamo serijo tal od rend- zin do srednje globokih rjavih tal s slabo adsorpcijsko sposobnostjo. Floristična analiza pa je pokazala, da je naša fitocenoza vrstno bogatejša in raznovrstnejša od asociacije Galio-Piceetum. Primerjava med njima kaže majhno podobnost fitocenoz, po Sørensenu je indeks 42 in po Jaccardu le 27. Nizka indeksa potrjujeta različnost fitocenoz, tako da naše fitocenoze ne moremo uvrstiti v asociacijo Galio rotundifolii-Piceetum, čeprav se ji neko- liko približuje. Zanimala nas je še predstavitev asociacije Galio-Pi- ceetum v Avstriji (W llnöfer v Mucin et al. 1993). Wallnöferjeva je izbrala za značilnico le vrsto Galium rotundifolium, za katero pravi, da je slaba in transgresij- ska. Doda pa razlikovalnice: Anemone nemorosa, Doro- nicum austriacum, Primula elatior in še druge glede na asociacijo Veronico latifoliae-Piceetum ter dominantne in stalne spremljevalke asociacije Galio-Piceetum. Pravi, da v asociaciji prevladujejo piceetalne vrste in da združ- ba naseljuje montanski in spodnji subalpski pas na sili- katni, mineralno bogati in zmerno kisli geološki podla- gi, kjer so globoka, sveža, včasih podzoljena rjava tla s sprsteninastim ali prhninastim humusom. Ugotavlja, da je asociacija Galio rotundifolii-Piceetum klimaksna združba. Da predstavimo mnenje Wallnöferjeve o asociaciji Galio-Piceetum, nas je spodbudila njena uvrstitev M. Wr berjeve (1959) asociacije Galio rotundifolii-Abie- tetum M. Wraber (1955) 1959 v asociacijo Galio rotun- difolii-Piceetum J. & M. Bartsch 1940. Jelovo združbo M. Wraberja ima za sinonim Bartscheve smrekove aso- ciacije. Enakega mnenja kot Wallnöferjeva sta Will- ner & Gr bherr (2007). Primerjava med asociacija- ma Galio-Piceetum v Schwarzwaldu in Galio-Abiete- tum v Sloveniji je pokazala različnost asociacij, kar po- trjujeta indeksa podobnosti, in sicer po Sørensenu 51 in po Jaccardu 34. Primerjava med našo smrekovo fitoce- nozo in M. Wraberjevo jelovo asociacijo Galio-Abiete- tum kaže in d eks še man jše pod o b n osti , in s i ce r po Sørensenu 42 in po Jaccardu 25. Torej trditev Wallnö - ferjeve, da gre pri M. Wraberjevi asociaciji Galio-Abie- tetum za s in o nim Bartsch ev e asociaci j e Galio-Picee- tum, ne drži. Asociaciji sta samostojni in sintaksonom- sko stabilni. Skladno z gornjimi ugotovitvami smo nadaljevali s primerjavami med pri nas znanimi sekundarnimi smre- kovimi fitocenozami. Ugotovili smo, da je smrekova fi - tocenoza najbolj primerljiva s sekundarno smrekovo asociacijo Aposerido-Piceetum Z u p a n č i č ( 1 9 7 8 ) 1 9 9 9 . Značilnica asociacije Aposeris foetida je prisotna v štirih popisih s solidno srednjo pokrovno vrednostjo 400 in sociabilnostjo (1–2) ter visoko stopnjo navzočnosti (pre - zenca V = 80 %). Za sekundarno asociacijo Aposerido- Piceetum na rastišču primarne bazifilne bukove združ- be j e zn ačiln a p r ece j š n j a zast o p an o st fag etalnih vrst. Pomembna je tudi prisotnost značilnic ilirske zveze bu- kovih gozdov Aremonio-Fagion, med katerimi so razli- kovalnice za geografsko varianto sekundarne smrekove asociacije Aposerido-Piceetum var. geogr. Helleborus niger Zupančič 1999. V našem primeru gre za prvotno rastišče predalp- skega jelovo-bukovega gozda Homogyno sylvestris-Fage- tum, na katerem so pospeševali smreko. Del smreke je sajen, del se je naselil spontano. Zaradi dolgoletnega vpliva opada smrekovih iglic, izčrpavanja zgornjih hori - zontov tal (Oh, A) in delovanja smrekovega koreninske- ga sistema so se tla sčasoma slabšala, predvsem zakiso- vala in omogočala naselitev acidofilne piceetalne flore. Karbonatna matična podlaga pa je omogočala preživetje nevtralni in zmerno bazični fagetalni flori. Tako imamo v tej sekundarni smrekovi fitocenozi mozaično preple - tanje piceetalne in fagetalne flore v precej enakomernem sorazmerju glede zastiranja površine, po številu vrst pa prednjačijo fagetalne vrste. Od fagetalnih vrst je iz ilir - ske zveze bukovih gozdov Aremonio-Fagion prisotnih osem vrst, od tega tri, ki so razlikovalnice za geografsko varianto: Helleborus niger subsp. niger, Cyclamen purpu- rascens in Cardamine trifolia s piceetalno vrsto Homo- gyne sylvestris, sicer pa je v naši smrekovi fitocenozi za - stopanih enajst jugovzhodnoevropsko-ilirskih vrst. Glede na gornje floristične podobnosti, ki jih nazor- no kaže Fitocenološka tabela, smo izračunali indekse podobnosti po Sørensenu (σ ) in Jaccardu (σ j ) med našo s smrekovo fitocenozo in a) bavarsko asociacijo Galio-Pi- ceetum, kjer je σ = 42 in σ j = 27, in b) asociacijo Aposeri- s do-Piceetum Zupančič 1999 (Zup nčič 1999, tabela 11), kjer je σ = 59 in σ j = 42. s FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 116 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Floristična analiza z indeksom podobnosti fitoce - noz kaže na sorodnost naše sekundarne smrekove fito - cenoze z asociacijo Aposerido-Piceetum, sicer ne zelo prepričljivo, vendar zadovoljivo, indeks podobnosti z asociacijo Galio-Piceetum pa je manjši in ne dosega op- timalnosti. Razlika je v floristični vsebnosti fitocenoz, predvsem v številčnosti fagetalnih vrst; teh je več kot po - lovica manj v asociaciji Galio-Piceetum in deloma še v vrstah razredov Betulo-Adenostyletea (=Mulgedio-Aco- nitetea), Tr i f o l i o - G e ra n i e t e a in Asplenietea trichomanis, ki so vse bolj ali manj bazične. Glede zastrtosti pa si v asociaciji Galio-Piceetum acidofilne in bazifilne vrste enakomerno delijo prostor. Floristična in posledično vegetacijska analiza sta pokazali, da lahko našo sekundarno smrekovo fitoceno - zo uvrstimo v asociacijo Aposerido-Piceetum, vendar kot novo subasociacijo, ki smo jo poimenovali po vrsti Galium rotundifolium, torej Aposerido-Piceetum galieto- sum rotundifolii subass. nova oziroma Aposerido-Picee- tum var. geogr. Helleborus niger galietosum rotundifolii subass. nova. Subasociacija A.-P. galietosum kaže delno povezavo oziroma sorodnost z asociacijo Galio-Picee- tum, kar nam potrjujejo tudi indeksi podobnosti fitoce - noz po Sørensenu in Jaccardu. Floristična in sociološka sestava subasociacije Pri predstavitvi asociacije Aposerido-Piceetum (Zup n- čič 1999) smo zaradi obširnosti monografije o smreko - vih fitocenozah opustili nekatere prvine sociološke se- stave asociacije, ob tej priložnosti pa bi jih dopolnili. Gre predvsem za dopolnitev karakteristične kombinacije vrst, stratifikacije, biološkega spektra, fitocenoloških in horoloških skupin združbe. Pred tem pa moramo še opi- sati izbiro razlikovalnih vrst subasociacije. Floristična tabela s petimi popisi prikazuje subasoci - acijo Aposerido-Piceetum galietosum rotundifolii subass. nova s 153 rastlinskimi vrstami. Cvetnic je 123, 8 prapro- ti, 21 mahov in 1 gliva. Najmanjše število florističnih vrst imata popisa 5 in 1 (61, 62), največ pa popisa 4 in 2 (88, 81), popis 3 ima 71 vrst. Popisi so zadosti homogeni. Ekološke razmere subasociacije se ne razlikujejo bi- stveno od prej opisanih za asociacijo Aposerido-Picee- tum. Srednja letna temperatura se giblje med 4 in 8 0C z letno namočenostjo 1750 mm in več. Tla so rjava (kar - bonatna), kisla do zmerno kisla s pH KCl v Oh pod 4 ter v A in B horizontu okoli 6 (Zup nčič 1999). Razlikovalnici za subasociacijo Aposerido-Piceetum galietosum rotundifolii sta Galium rotundifolium in Go- odyera repens. Njune ekološke in vegetacijske oznake so povzete po Oberdorferju (1979) in po naših opaža - njih. Vrsta Galium rotundifolium L. se pojavlja v jelovih, smrekovih, mešanih jelovo-smrekovih in redkeje v ki- slih bukovih gozdovih. Tu in tam jo dobimo v ruševju. Porašča sveža, s hranili in bazami zmerno bogata, toda z apnencem revna, zmerno kisla tla s prhninastim humu- som. Uvrščamo jo v sencoljubne vrste. Sinsistematsko je opredeljena kot vrsta podzveze Abieti-Piceenion oziro- ma Galio-Abietion. Fitogeografsko spada med subatlant - sko-submediteranske oziroma evroazijske vrste. Vrsta Goodyera repens (L.) R. Br. raste v jelovih, smrekovih in kislih borovih gozdovih, redkeje v rušev- ju. Naseljuje zmerno suha do sveža rastišča, kjer so tla bolj ali manj bogata z bazami (s kislim prhninastim hu- musom). Je polsenčna vrsta, ki se rada pojavlja med ki- sloljubnimi mahovi ali na kislem opadu iglic jelke, smre- ke ali bora. Večinoma jo uvrščamo kot značilnico reda Va c c inio -P ic e e talia. Fitogeografsko je to severovzhodno kontinentalna-cirkumpolarna oziroma cirkumborealna vrsta. Karakteristična kombinacija vrst kaže naslednjo podobo: ZNAČILNICA Aposeris foetida* RAZLIKOV ALNICE GEOGRAFSKE V ARIANTE Helleborus niger subsp. niger* Cyclamen purpurascens* Cardamine trifolia* RAZLIKOV ALNICA SUBASOCIACIJE Galium rotundifolium VACCINIO-PICEETEA s. lat. Galium rotundifolium Hylocomium splendens Melampyrum sylvaticum Rhytidiadelphus triquetrus Mnim punctatum Hypnum cupressiforme Picea abies Aposeris foetida* Hieracium sylvaticum Maianthemum bifolium FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 117 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Vaccinium myrtillus Gentiana asclepiadea Larix decidua Peltigera leucophlebia Polytrichum formosum Luzula pilosa Atrichum undulatum Oxalis acetosella Luzula luzuloides QUERCO-FAGETEA s. lat. Helleborus niger subsp. niger* Epipactis helleborine Cyclamen purpurascens* Lamiastrum flavidum Aremonia agrimonioides* Stachys sylvatica Cardamine trifolia* Viola reichenbachiana Knautia drymeia subsp. drymeia* Euphorbia amygdaloides Fagus sylvatica Melica nutans Sanicula europaea Pteridium aquilinum Mycelis muralis Veronica officinalis Mercurialis perennis Digitalis grandiflora Acer pseudoplatanus Campanula witasekiana* Daphne mezereum Hypericum montanum Eurhynchium zettersledtii Ctenidium molluscum Salvia glutinosa Carex digitata ERICO-PINETEA s.lat. Carex alba Buphthalmum salicifolium Polygala chamaebuxus Calamagrostis varia Rubus saxatilis BETULO-ADENOSTYLETEA s. lat. (=MULGEDIO-ACONITETEA s. lat.) Senecio fuchsii TRIFOLIO-GERANIETEA s. lat. Cruciata glabra MOLINIO-ARRHENATHERETEA s. lat. Ajuga reptans ASPLENIETEA TRICHOMANIS s. lat. Asplenium trichomanes OSTALE VRSTE Fragaria vesca Cladonia pyxidata * Jugovzhodnoevropsko-ilirske vrste Kot kaže preglednica, so med izbranimi karakteri- stičnimi vrstami značilnica asociacije Aposeris foetida, tri razlikovalnice geografske variante Helleborus niger subsp. niger, Cyclamen purpurascens, Cardamine trifolia in raz- likovalnica subasociacije Galium rotundifolium, kar potr- juje našo odločitev o pripadnosti subasociacije asociaciji Aposerido-Piceetum (var. geogr. Helleborus niger). Urav- notežene so vrste med razredoma Va c c ini o-Piceetea s. lat. in Querco-Fagetea s. lat., kar pomeni potrditev naše ugo- tovitve, da gre za sekundarno smrekovo fitocenozo na prvotnem bukovem (jelovo-bukovem) rastišču. Vsekakor pa moramo asociacijo Aposerido-Piceetum uvrstiti v ra- zred Va c c ini o -P i c e e te a. Skromna uvrstitev vrst razredov Erico-Pinetea s. lat., Betulo-Adenostyletea s. lat. (=Mulge- dio-Aconitetea s. lat.), Molinio-Arrhenatheretea s. lat. in »ostalih« vrst, ki jih je skupno dvanajst, pa dodatno kaže na njeno sekundarnost in delno degradacijo rastišča. Clinopodium vulgare Euphorbia cyparissias Stratifikacija subasociacije Aposerido-Piceetum ga- lietosum rotundifolii je preprosta in se deli v nadstojno plast večinske smreke in manjšinske bukve s tu in tam prisotnimi drevesi Larix decidua, Prunus avium in Fraxinus ornus. Druge drevesne vrste Acer pseudoplata- nus, A. campestre, Fraxinus excelsior, Ostrya carpinifo- lia, Sorbus aria in Juglans regia so le v grmovni plasti in z drugimi grmovnicami pokrivajo od 0 do 60 % površi - ne. Zeliščna plast ni posebno bogato razvita in je, kot že rečeno, uravnotežena med piceetalnimi in fagetalnimi vrstami. Mahovna plast je večinoma skromna, tu in tam tudi močneje zastopana. Biološki spekter fitocenoze kaže na bolj ali man j umirjene življenjske razmere, ki vladajo v zmernem pasu. Nekoliko je povečan delež fanerofitov in hamefi - tov. Povečanje fanerofitov kaže na toplejše razmere, če - prav ni vedno tako, saj so večinoma grmovne rasti FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 118 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION (13,7 %) in kažejo z nekoliko povečanim deležem hame - fitov odmik v smeri nekoliko neugodnejših razmer, kot vladajo v zmernem pasu. Za sekundarno smrekovo fito - cenozo so te življenjske razmere v primerjavi s primar- nimi smrekovimi fitocenozami ugodne. Biološki spek - ter kaže na življenjske razmere rastišča med prvotno bukovo (jelovo-bukovo) fitocenozo in na delni vpliv se - kundarne smrekove fitocenoze. Smreka s svojim opa- dom iglic, plitvim koreninjenjem in stalnim zasenče- njem rastišča slabša tla. Prisotno je kontinuirano zaki - sovanje tal z opadom, ki se slabo razkraja z enostransko izrabo hranil zgornjih horizontov. BIOLOŠKI SPEKTER Št. % Skupaj % FANEROFITI-PHANEROPHYTA 31 20,2 Šopasti f. (P. caesp.) 11 7,2 Steblasti f. (P. scap.) 10 6,5 Nano f. (NP) 8 5,2 Vzpenjalni f. (P. scand.) 2 1,3 HAMEFITI-CHAMAEPHYTA 27 17,7 Polgrmičasti h. (Ch. suffr.) 2 1,3 Grmovni h. (Ch. frut.) 2 1,3 Plazeči h. (Ch. rept.) 1 0,7 Sukulentni h. (Ch. succ.) 1 0,7 Mahovni h. (Ch. brio.) 16 10,4 Lišajni h. (Ch. lich.) 5 3,3 HEMIKRIOPTOFITI - HEMICRYPTOPHYTA 65 42,4 Steblasti h. (H. scap.) 33 21,5 Šopasti h. (H. caesp.) 15 9,8 Rozetni h. (H. ros.) 11 7,2 Plazeči h. (H. rept.) 4 2,6 Dvoletni h. (H. bien.) 2 1,3 GEOFITI-GEOPHYTA 27 17,7 G. s koreniko (G. rhiz.) 22 14,4 G. z gomolji (G. bulb.) 4 2,6 G. paraziti (G. par.) 1 0,7 TEROFITI-THEROPHYTA 1 0 , 7 Steblasti t. (T. scap.) 1 0,7 NEOPREDELJENI VRSTI 2 1,3 2 1 , 3 Skupaj 153 100,0 153 100,00 Tabela 1: Biološki spekter Analiza horoloških skupin kaže, da je več kot dve tretjini geoelementov hladnih (70,5 %). K hladnim sku - pinam smo prišteli mahove in lišaje (vaskularna flora 56,8 % in mahovno-lišajna flora 13,7 %). Največ je evrop - skih, cirkumborealnih in evroazijskih geoelementov, ne zaostajajo pa niti paleotemperatni in evrosibirski geoe- lementi. Vsi ti potrjujejo, da sekundarna smrekova fito - cenoza uspeva v nekoliko ostrejših življenjskih razme- rah, kar nam kaže tudi biološki spekter. Tabela 3 kaže razmerja med fitocenološkimi sku - pinami. Pričakovano je, da so najbolj zastopane fage - talne vrste prvotne bukove (jelovo-bukove) fitocenoze (45,8 %). Glede na karbonatno geološko podlago pa je največ bazifilnih vrst (54,3 %) iz razredov Querco-Fa- Geoelementi Št. % Mediteransko-montanski 15 9,8 Cirkumborealni 23 15,0 Evropski 28 18,3 Evroazijski 17 11,1 Paleotemperatni 10 6,5 Evrosibirski 9 5,9 Severnoilirski 7 4,7 Južnoilirski 2 1,3 Jugovzhodnoevropski 2 1,3 Evromediteranski 2 1,3 Pontski 3 2,0 Mediteransko-pontski 3 2,0 Mediteransko-atlantski 2 1,3 Kozmopolitski 6 3,9 Kultivirana vrsta 1 0,6 Mahovi in lišaji 21 13,7 Nedoločeni vrsti 2 1,3 Skupaj 153 100,00 Tabela 2: Horološke skupine getea s. lat., Betulo-Adenostyletea s. lat. ( = Mulgedio- Aconitetea s. lat.), Tr ifoli o - G e ra ni e te a s. lat. in Aspleni- etea trichomanis s. lat. Nato sledijo piceetalne vrste (24,8 %), ki so se uveljavile po dolgotrajni zasmrečitvi rastišča in so acidofilne. K acidofilnim vrstam uvršča- mo poleg razreda Vaccinio -Piceetea s. lat. še Erico-Pi- netea s. lat. ter mahovno in lišajno floro (34,6 %). Manj je nevtralnih vrst ( 1 1,1 % ) iz razreda Molinio-Arrhe- natheretea s. lat. in »ostalih« vrst. Podoba fitocenolo - ških skupin nam potrjuje sekundarnost smrekove fito - cenoze. Razred št. % VACCINIO-PICEETEA 38 24,8 QUERCO-FAGETEA 70 45,8 ERICO-PINETEA 7 4,6 BETULO-ADENOSTYLETEA (MULGEDIO-ACONITETEA) 5 3 , 3 TRIFOLIO-GERANIETEA 4 2,6 MOLINIO-ARRHENATHERETEA 6 3,9 ASPLENIETEA TRICHOMANIS 4 2,6 OSTALE VRSTE 11 7,2 MAHOVI IN LIŠAJI 8 5,2 Skupaj 153 100,0 Tabela 3: Fitocenološke skupine Zaključek Skladno s floristično in vegetacijsko analizo sekundarne smrekove fitocenoze smo ugotovili, da jo lahko uvrsti- mo v asociacijo Aposerido-Piceetum Zupančič 1999 kot njeno samostojno subasociacijo Aposerido-Piceetum ga- lietosum rotundifolii subass. nova. Holotip subasociaci- je je popis 2 iz Fitocenološke tabele 1. Subasociacija Aposerido-Piceetum galietosum rotun- difolii se glede na svojo floristično sestavo približuje sre- FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 119 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION dnjeevropski asociaciji Galio rotundifolii-Piceetum J. & M. Bartsch 1940. Med njima je delna (manjša) podob- nost oziroma ( daljna) sorodnost. Upoštevati pa mora- mo, da se asociaciji nahajata v različnih flornih provin - cah. Asociacija Aposerido-Piceetum je v ilirski florni provinci, kar pove že ime asociacije po jugovzhodnoe- vropsko-ilirski vrsti in značilnici asociacije Aposeris fo- etida. Poleg značilnice se v asociaciji pojavljajo še druge jugovzhodnoevropsko-ilirske vrste: Helleborus niger subsp. niger, Cyclamen purpurascens, Anemone trifolia, Homogyne sylvestris, Cardamine trifolia; te so nam aso- ciacijo opredelile kot geografsko varianto ilirske florne province Aposerido-Piceetum var. geogr. Helleborus niger Zupančič 1999. Asociacija Galio rotundifolii-Picee- tum J. & M. Bartsch 1940 je doma v srednjeevropski florni provinci. Po več generacijah smrekovega gozda na bukovem rastišču bodo sčasoma izginile bazifilne jugovzhodnoe - vropsko-ilirske in fagetalne vrste, predvsem tiste, ki so strogo vezane na karbonatna tla. Takrat bi bila sekun - darna smrekova fitocenoza zelo podobna tistim iz sre - dnjeevropske province, npr. asociaciji Galio-Piceetum. Na koncu naj opozorimo na primerjavo med asoci- acijama Galio rotundifolii-Piceetum J. & M. Bartsch 1940 in Galio rotundifolii-Abietetum M. Wraber (1955) 1959. Kljub trditvam srednjeevropskih fitocenologov (Muci- n , Gr bherr & W llnöfer 1993, Willner & Gr b- herr 2007), da gre v obeh primerih za isto asociacijo, torej za prvo opisano Galio-Piceetum (ime asociacije Ga- lio-Abietetum naj bi bil sinonim), to ne drži. Primerjalni analizi po Sørensenu in Jaccardu sta jasno pokazali, da gre za dve različni asociaciji, to je za sekundarno smre- kovo in primarno jelovo fitocenozo. Ugotovitev nove gozdne fitocenoze je nadaljnji pri- spevek k veliki biotski različnosti (biodiverziteti) gozdne vegetacije v Sloveniji. Njen vzrok je v fitogeografskem položaju, različnosti geološke podlage, mezoklimatskih in mikroklimatskih razmerah in ne nazadnje v orograf- ski razčlenjenosti Slovenije ter še drugih, manj znanih biotskih in abiotskih dejavnikih. LITERATURA B rtsch, J. & M. B rtsch, 1940: Vegetationskunde des Schwarzwaldes . Pflanzensoziologie (Jena) 4: 1–229. Br un-Bl nquet, J., 1964: Pflanzensoziologie . Wien-New York. M rtinčič, A., T. Wr ber, N. Jog n, A. Podobnik, B. Turk & B. Vreš, 2007: Mala flora Slovenije. Ljubljana. Mucin , L., G. Gr bherr & S. W llnöfer, 1993: Die Pflanzengesselschaften Österreichs. Teil III. Wälder und Ge- büsche. Jena-Stuttgart-New York. Oberdorfer, E., 1957: Süddeutische Pflanzengesellschaften . Pflanzensoziologie (Jena) 10: 1–564. Oberdorfer, E., 1979: Pflanzensoziologische Exkursions Flora . Stuttgart. Poldini, L., 1991: Atlante corologico delle piante vascolari nel Friuli-Venezia Giulia . Udine. Willner, W. & G. Gr bherr, 2007: Die Wälder und Gebüsche Österreichs. München. (Textband und Tabellen- band). Zup nčič, M., 1999: Smrekovi gozdovi Slovenije . Dela SAZU (Ljubljana) 36: 1–212. FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 120 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION PHYTOCOENOLOGICAL TABLE (Fitocenološka tabela) 1: APOSERIDO-PICEETUM Zupančič (1978) 1999 GALIETOSUM ROTUNDIFOLII Zupančič & Žagar subass. nova Number of relevé (Zaporedna številka popisa) 1 2 3 4 5 Working No of relevé (Delovna številka popisa) 116/62 115/62 118/62 112/62 114/62 Date (Datum) 23.8.62 23.8.62 23.8.62 23.8.62 23.8.62 Altitude in m (Nadmorska višina v m) 1170 1060 1070 680 970 Aspect (Nebesna lega) S SSE SE SSE SE Slope in degrees (Nagib v stopinjah) 15–20 5–10 0–5 20 15 Sinsistematical characteristic (Sinsistematska pripadnost) Bedrock (Geološka podlaga) lime dolo lime, dolo lime, dolo lime lime, dolo Stoniness in % (Kamnitost v %) 10 0 5 70 5 Cover (Pokrovnost) %: Tree layer (drevesna pl.) I 90 80 90 80 90 Shrub layer (grmovna plast) II 0 40 10 60 5 Herb layer (zeliščna plast) III 40 80 80 60 40 Moss layer (mahovna plast) IV 10 30 10 50 70 Relevé (Velikost popisne ploskve) m2 400 400 400 400 400 Covc Planica Za tesnicami Zamorsko Zamorski vrh Location (Kraj popisov) V e ž a – Pl a n ic a S a v i n j s k a d o l i n a Province (Pokrajina) Š t a j e r s k a CHARACTERISTIC SPECIES OF THE ASSOCIATION (Značilnica za asociacijo) APOSERIDO-PICEETUM Zupančič 1999 1 2 3 4 5 Presence (Prezenca) •V P Aposeris foetida III 1.1 1.2 1.1 1.1 . 4 3 DIFERENTIAL SPECIES OF THE GEOGRAPHICAL VARIANT(Razlikovalnice geografske variante) HELLEBORUS NIGER Zupančič 1999 1 2 3 4 5 • F 1 Helleborus niger subsp. niger III + 2.2 2.2 2.1 1.2 5 • F Cyclamen purpurascens 1.1 1.1 + 1.1 1.2 5 1 • F Cardamine trifolia + 1.1 + . . 3 1 + . . . + •V P 2 Homogyne sylvestris 2 DIFERENTIAL SPECIES OF THE SUBASSOCIATION (Razlikovalnici subasociacije) GALIETOSUM ROTUNDIFOLII Zupančič & Žagar subass. nova 1 2 3 4 5 AP Galium rotundifolium III (+) + 1.2 + + 5 VP Goodyera repens . 1.1 . . 2.1 2 2 AP ABIETI-PICEENION Br.-Bl. 1939 in Br.-Bl. et al. 1939 1 2 3 4 5 Galium rotundifolium III (+) + 1.2 + + 5 I . 1.1 . + . 2 Abies alba 2 II . 1.2 . - . 1 Valeriana tripteris III . + . + . 2 Veronica urticifolia . + . . . 1 Adenostyles glabra . . +0 . . . 1 RV RHODODENDRO-VACCINIENION Br.-Bl. in Br.-Bl. & Jeny 1926 & VACCINIO-PICEENION Oberd. 1957 1 2 3 4 5 Melampyrum sylvaticum III 1.2 2.2 1.2 . + 4 Mnium punctatum IV . +.3 + + 1.5 4 Luzula luzulina III . + . . . 1 Homogyne alpina + . . . . 1 Pyrola minor . . . . + 1 Vaccinium vitis-idaea . . + . . 1 VACCINIO-PICEION Br.-Bl. (1938) 1939 & VACCINIO-PICEETALIA Br.-Bl. in Br.-Bl. 1939 emend. K.-Lund 1967 VP 1-2 1 2 3 4 5 I 5.5 3.2 5.5 4.1 5.5 5 VP 1 Picea abies II + 2.2 1.1 3.2 + 5 5 III + 1.1 + + + 5 VP Hieracium sylvaticum + 1.2 + + + 5 1 VP 2 Vaccinium myrtillus + 1.2 + . +.2 4 FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 121 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION VP 2 Larix decidua I + . 2.1 . 1.1 3 VP 2 Polytrichum formosum IV . +.2 + + . 3 VP 2 Atrichum undulatum + . + + . 3 VP 2 Goodyera repens III . 1.1 . . 2.1 2 VP 2 Monotropa hypopitys . . +.2 . + 2 •V P 2 Homogyne sylvestris + . . . + 2 VP 2 Cantharellus cibarius . +.2 . . . 1 VP 2 Orthilia secunda . +.2 . . . 1 VP 2 Rosa pendulina II . . . . + 1 VP 3 VACCINIO-PICEETEA Br.-Bl. in Br.-Bl. et al. 1939 em. Zupančič (1980) 2000 s. lat. 1 2 3 4 5 Hylocomium splendens IV + +.3 1.5 1.5 2.5 5 Rhytidiadelphus triquetrus + 1.5 2.5 1.3 +.3 5 Hypnum cupressiforme 2.5 1.2 +.3 2.5 . 4 • Aposeris foetida III 1.1 1.2 1.1 1.1 . 4 Maianthemum bifolium 1.1 1.1 1.1 + . 4 Dicranum scoparium IV . +.3 + +.3 1.3 4 Gentiana asclepiadea III + 1.1 + . + 4 Peltigera leucophlebia IV . + + + + 4 Luzula pilosa III + . 1.1 + . 3 Oxalis acetosella + . 1.1 + . 3 Luzula luzuloides + + +.2 . . 3 Grimmia pulvinata IV + + . . . 2 Solidago virgaurea III + + . . . 2 Pleurozium schreberi IV . . . . 1.5 1 EP Laserpitium peucedanoides III . . . + . 1 • F 1 AREMONIO-FAGION (Ht. 1938) Török, Podani & Borhidi 1989 1 2 3 4 5 Helleborus niger subsp. niger III + 2.2 2.2 2.1 1.2 5 Cyclamen purpurascens 1.1 1.1 + 1.1 1.2 5 Aremonia agrimonoides + + + + + 5 Cardamine trifolia + 1.1 + . . 3 Knautia drymeia subsp. drymeia . + + . + 3 Dentaria enneaphyllos + . . . . 1 Lamium orvala . + . . . 1 Rhamnus fallax II . . . + . 1 F 2 FAGETALIA SYLVATICAE Pawl. 1928 1 2 3 4 5 Fagus sylvatica I II 1.1 + + 1.1 - +.2 2.1 1.1 + + 5 5 Sanicula europaea III +.2 1.1 + 1.1 + 5 Mycelis muralis + + 1.1 1.1 + 5 Mercurialis perennis 2.2 + + + + 5 Acer pseudoplatanus II + + + + + 5 Daphne mezereum + + + + + 5 Eurhynchium zetterstedtii IV . 1.5 1.3 2.5 +.3 4 Salvia glutinosa III . 1.1 + 1.1 + 4 Epipactis helleborine + + + . + 4 Galeobdolon flavidum + + + + . 4 Stachys sylvatica + + + . + 4 Viola reichenbachiana . + + + + 4 Euphorbia amygdaloides 1.1 1.1 + . . 3 Melica nutans + + . + . 3 Brachypodium sylvaticum . . +.2 . 1.1 2 Rosa arvensis II . + . 1.1 . 2 Actaea spicata III . . . + + 2 Cephalanthera damasonium + + . . . 2 Geranium robertianum . . + + . 2 Gymnocarpium robertianum . . + + . 2 Lonicera alpigena II . + . . + 2 Paris quadrifolia III + . + . . 2 Campanula trachelium . . . 1.1 . 1 FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 122 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Asarum europaeum . . . + . 1 Fraxinus excelsior II . . . + . 1 • Galium laevigatum III . . + . . 1 Poa nemoralis . + . . . 1 Polystichum aculeatum . + . . . 1 Prenanthes purpurea . + . . . 1 Pulmonaria officinalis . + . . . 1 Ranunculus lanuginosus . . + . . 1 Scrophularia nodosa . . . + . 1 Symphytum tuberosum + . . . . 1 Galium odoratum . . . +0 . . 1 RP QUERCETALIA ROBORIS-PETRAEAE R. Tx. (1931) 1937 s. lat. 2 1 2 3 4 5 Pteridium aquilinum III + + . + + 4 Veronica officinalis + . + + + 4 Polypodium vulgare . . + + . 2 Frangula alnus II . . . + . 1 Q 2 QUERCETALIA PUBESCENTIS Br.-Bl. 1931 s. lat. 1 2 3 4 5 Digitalis grandiflora III + + . 1.1 . 3 Campanula witasekiana + + . . + 3 Hypericum montanum . + + . + 3 I . . . 1.1 . 1 Fraxinus ornus 1 II . . . 2.2 . 1 Ostrya carpinifolia . . . + . 1 Sorbus aria . + . . . 1 P PRUNETALIA SPINOSAE R. Tx. 1952 s. lat. 1 2 3 4 5 Berberis vulgaris II . . . + . 1 Crataegus monogyna . . . + . 1 Juniperus communis . . . + . 1 Pyrus pyraster . . . + . 1 Viburnum lantana . . . + . 1 F QUERCO-FAGETEA Br.-Bl. & Vlieger in Vlieger 1937 3 1 2 3 4 5 Ctenidium molluscum IV 1.3 1.5 1.3 3.5 2.5 5 Carex digitata III + + + + + 5 Hepatica nobilis 1.1 + . 2.2 . 3 Corylus avellana II . + . 1.1 . 2 Lonicera xylosteum . . . + 1.1 2 Anemone nemorosa III + + . . . 2 Clematis vitalba II . . . + + 2 Isothecium myurum IV . + . + . 2 Platanthera bifolia III . + + . . 2 Acer campestre II . . . + . 1 Hedera helix . . . + . 1 Listera ovata III . . + . . 1 Prunus avium II . . . + . 1 EP ERICO-PINETEA Ht. 1959 s. lat. 3 1 2 3 4 5 Carex alba III 1.1 +.2 2.3 + 1.3 5 Polygala chamaebuxus . + + 1.2 1.2 4 Buphthalmum salicifolium + . + 1.1 . 3 Calamagrostis varia +.2 +.2 . . + 3 Rubus saxatilis II + + . . + 3 Pimpinella saxifraga III . . . + + 2 Gymnadenia odoratissima . . + . . 1 A3 BETULO-ADENOSTYLETEA Br.-Bl. & R. Tx. 1943 s. lat. (=MULGEDIO-ACONITETEA Hadač & Klika in Klika & Hadač 1944) 1 2 3 4 5 Senecio fuchsii III + + + + + 5 FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 123 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Polygonatum verticillatum 2.1 + + . . 3 Dryopteris filix-mas . + . + . 2 Athyrium filix-femina + . . . . 1 Ranunculus platanifolius . . + . . 1 TG TRIFOLIO-GERANIETEA Th. Müller 1961 s. lat. 1 2 3 4 5 Cruciata glabra III . + + + 1.1 4 Clinopodium vulgare . + . 1.1 + 3 Vincetoxicum hirundinaria . . . + + 2 Origanum vulgare . . . + . 1 MA MOLINIO-ARRHENATHERETEA Tx. 1937 s. lat. 1 2 3 4 5 Ajuga reptans III 1.1 1.1 1.1 + + 5 Dactylis glomerata (s.lat.) . . + . + 2 SC Dactylorhiza maculata . + + . . 2 Ranunculus sp. . . + . + 2 Angelica sylvestris . . + . . 1 Cirsium palustre . . + . . 1 AS ASPLENIETEA TRICHOMANIS Br.-Bl. in Meier & Br.-Bl. 1934 corr. Oberd. 1977 s. lat. 1 2 3 4 5 Asplenium trichomanes III + . + +.2 . 3 Asplenium ruta-muraria . . . + + 2 Asplenium viride . . . + + 2 Moehringia muscosa . . . +.3 . 1 O OTHER SPECIES (Ostale vrste) 1 2 3 4 5 E Fragaria vesca III . 1.1 + + + 4 FB Euphorbia cyparissias . . + + + 3 Phyteuma sp. . + . . + 2 AR Urtica dioica + . . + . 2 AR Aegopodium podagraria . . . + . 1 S Campanula cochleariifolia . . + . . 1 Juglans regia II . . . + . 1 NC Potentilla erecta III . + . . . 1 Rubus fruticosus II . . . + . 1 S Saxifraga crustata III . . . + . 1 Sorbus aucuparia II . + . . . 1 ML MOSSES AND LICHENS (Mahovi in lišaji) 1 2 3 4 5 Cladonia pyxidata IV + + . . + 3 Plagiochila asplenioides . 1.3 . + . 2 Tortella tortuosa +.2 . . + . 2 Neckera crispa 1.3 . . . . 1 Anomodon viticulosus . . . +.3 . 1 Cladonia rangiferina . + . . . 1 Cladonia squamosa + . . . . 1 Metzgeria furcata + . . . . 1 LEGEND (Legenda) Sinsistematical characteristic (Sinsistematska pripadnost) Bedrock (Geološka podlaga) EM Erico-Pinion mugi Leibundgut 1948 nom. inv. lime limestone (apnenec) SC Scheuchzerio-Caricetea fuscae (Nordh. 1936) R. Tx. 1937 dolo dolomíte (dolomit) E Epilobietea angustifolii R. Tx. & Prsg. 1950 FB Festuco-Brometea Br.-Bl. & R. Tx. 1943 AR Artemisietea Lohm., Prsg. & R. Tx. in R. Tx.1950 NC Nardo-Callunetea Preising 1949 S Seslerietea Br.-Bl. 1948 em. Oberd. 1978 • Southeast European species (Jugovzhodno evropske vrste) FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 124 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION PHYTOCOENOLOGICAL TABLE (Fitocenološka tabela) 2: GALIO ROTUNDIFOLII-PICEETUM J. et M. Bartsch 1940 s. lat. Number of anal. tab. (Številka analitične tabele) 1 2 3 4 5 6 Author of anal. table (Avtor analitične tabele) Zupančič Bartsch Oberdorfer Oberdorfer M. Wraber Zupančič Altitude (Nadmorska višina) 680-1170 850-990 750-1000 700-800 380-1230 930-1570 Aspect (Nebesna lega) S-SE N-NW all (vse) N-S Sinsistematical characteristic (Sinsistematska pripadnost) gna, gra, Bedrock (Geološka podlaga) apn gna, gra gna, gra grd, gna, ble apn, dol bsa, mor Stonines (Kamnitost) 0-70 0-10 15-45 0-40 Location, province, state (Kraj popisov, pokrajina, država) Ve ž a-Pl a n ic a Savinjska dolina Štajesko Slovenija Schwarzwald Bayern Deutschland Schwarzwald Bayern Deutschland Schwarzwald Bayern Deutschland Pohorje Štajesko Slovenija Štajesko Gorenjsko Koroško Slovenija Number of relevé (Število popisov) 5 4 12 12 38 25 CHARACTERISTIC SPECIES OF THE ASSOCIATION (Značilnica za asociacijo) APOSERIDO-PICEETUM Zupančič 1999 1 2 3 4 5 6 V 1-4 V 1 VP Aposeris foetida III . . . . 3 DIFFERENTIAL SPECIES OF THE ASSOCIATION (Razlikovalnice za asociacijo) APOSERIDO-PICEETUM Zupančič 1999 1 2 3 4 5 6 V +-2 V +-2 Helleborus niger subsp. niger III . . . . V +-1 III +-1 IV +-1 Cyclamen purpurascens . . . . I I +-2 IV +-2 Cardamine trifolia . . . VP Homogyne sylvestris III +-3 III + . . . . 2 CHARACTERISTIC SPECIES OF THE ASSOCIATION (Značilnice za asociacijo) GALIO ROTUNDIFOLII-PICEETUM J. et M. Bartsch 1940 1 2 3 4 I V 3-5 4 4-5 II V VP 1 Picea abies II III V +-3 V +-1 - - - - - - AP Galium rotundifolium V +-1 3 +-1 III (I) 5 6 V 1-4 V 3-5 V 1-2 V +-2 III +-1 III + V 1-3 . CHARACTERISTIC SPECIES OF THE ASSOCIATION (Značilnice za asociacijo) GALIO ROTUNDIFOLII-ABIETETUM M. Wraber (1955) 1959 1 2 3 4 5 6 V +-2 VP Rubus hirtus II . . . . . 3 IV +-2 VP Hieracium rotundatum III . . . . . 1 IV +-2 VP 3 Thuidium tamariscinum IV . . . . . DIFFERENTIAL SPECIES OF THE ASSOCIATION (Razlikovalnice za asociacijo) GALIO ROTUNDIFOLII-ABIETETUM M. Wraber (1955) 1959 1 2 3 4 5 6 II +-2 I . . . . II +-2 RP Castanea sativa II . . . . . 2 II +-1 III . . . . . RP 2 Betula pendula I . . . . I + . RP Carex montana III . . . . I + . 2 AP ABIETI-PICEENION Br.-Bl. in Br.-Bl. et al. 1939 1 2 3 4 5 6 V +-1 3 +-1 V 1-3 Galium rotundifolium III III (I) . III +-1 3 +-2 V 1-5 II +-1 I VI V Abies alba II II 1 - - - V +-3 . I V +-1 Valeriana tripteris III III + . . . . Veronica urticifolia II + . . . . I I I +-1 FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 125 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION II +0 2 +-1 II +-1 Adenostyles glabra . . . . . Dryopteris dilatata . III . . C i r c a e a a l p i n a . . . I . . II +-1 I + Plagiochila asplenioides var. major IV . . . . RV RHODODENDRO-VACCINIENION Br.-Bl. in Br.-Bl. et Jenny 1926 Br.-Bl. 1939 & VACCINIO-PICEENION Oberd. 1957 1 2 3 4 5 6 V +-2 2 +-1 I V +-2 IV +-3 Melampyrum sylvaticum III V V V +-1 I + I +-2 IV +-1 Rhizomnium punctatum IV . . . . Luzula luzulina III II + . . . I V +-1 Homogyne alpina II + . . . . Pyrola minor II + . . . . . IV +-1 Vaccinium vitis-idaea II + . . . I + 3 +-1 Rhytidiadelphus loreus IV . III V . . 2 +-1 I + Lonicera nigra II . 2 + IV IV I + I + Luzula sylvatica subsp. sylvatica III . 2 + I I III +-1 Listera cordata III . I . . I + Plagiothecium undulatum IV . 2 + . . . I + Lycopodium annotinum III . 1 + I . . . Moneses uniflora . . . I . III +-1 Ptilium crista-castrensis IV . . . I . . VP 1 VACCINIO-PICEION Br.-Bl. in Br.-Bl. et al. 1939 1 2 3 4 5 6 V 3-5 4 4-5 V 1-4 V 3-5 I II V V +-3 V 1-2 V +-2 Picea abies II - - - III V +-1 - - - III +-1 III + V +-1 V +-2 V +-2 Hieracium sylvaticum . I V V Thelypteris limbosperma . 2 + I . III +-1 I + Blechnum spicant . 1 + . . I + I + IV +-1 Gymnocarpium dryopteris . . III . II + I V +-2 H i e r a c i u m r o t u n d a t u m . . . . . II +-1 III +-1 Bazzania trilobata IV . . . . VP VACCINIO-PICEETALIA Br.-Bl. in Br.-Bl. et al. 1939 emend. K.-Lund 1967 2 1 2 3 4 5 6 V +-1 IV +-3 V +-3 Vaccinium myrtillus III . V IV IV +-2 Larix decidua I . . . . 3 1-4 V 1-2 II +-1 Polytrichum formosum IV IV + IV II Atrichum undulatum IV + . II . III +-1 I + Goodyera repens III III 1-2 . . . . I + Monotropa hypopitys III + . . . . . • Homogyne sylvestris III + . . . . I I I +-3 Cantharellus cibarius II + . . . I + . I I +-1 Orthilia secunda II + . I . . Rosa pendulina II II + . I . . I I + 2 1-4 V 1-4 I + Thelypteris phegopteris . 2 + II . . II + II +-1 Avenella flexuosa III . IV III Plagiothecium neglectum IV . . . . II + I +-1 Dicranum majus . . . . . VP VACCINIO-PICEETEA Br.-Bl. in Br.-Bl. et al. 1939 emend. Zupančič (1980) 2000 3 1 2 3 4 5 6 V +-2 4 +-5 Hylocomium splendens IV V . III +-2 I + V +-2 2 +-1 II +-2 I +-3 Rhytidiadelphus triquetrus III . V +-2 V +-2 III +-1 Hypnum cupressiforme . . . V 1-4 • Aposeris foetida III V 1 . . . . V +-1 3 +-1 IV +-2 Maianthemum bifolium II I II + V +-1 III +-2 IV +-1 Dicranum scoparium IV . I III V +-1 I I +-2 IV +-1 Gentiana asclepiadea III . . . I +-1 V + IV +-1 1 + V +-2 IV +-1 Peltigera leucophlebia IV . . . . Luzula pilosa III II II IV +-1 4 3-4 V +-2 V +-2 Oxalis acetosella V I V Luzula luzuloides IV + . I V V V +-3 III +-4 FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 126 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Grimmia pulvinata IV III + . . . Solidago virgaurea III III + . I I V Pleurozium schreberi II1 . . . EP Laserpitium peucedanoides II+ . . . Sphagnum sp. IV . 2 + . . Hieracium sp. III . 1 + . . Calamagrostis arundinacea . . III V R u b u s h i r t u s I I . . . . Thuidium tamariscinum IV . . . . Solidago virgaurea subsp. minuta I I I . . (=Solidago virgaurea subsp. alpestris) . . Leucobryum glaucum IV . . . . • F 1 AREMONIO-FAGION (Ht. 1938) Török, Podani & Borhidi in Borhidi 1989 1 2 3 4 V +-2 Helleborus niger subsp. niger III . . . V +-1 Cyclamen purpurascens . . . Aremonia agrimonoides V + . . . IV +-1 Cardamine trifolia . . . Knautia drymeia subsp. drymeia IV + . . . Dentaria enneaphyllos II + . . . Lamium orvala II + . . . Rhamnus fallax II II + . . . F 2 FAGETALIA SYLVATICAE Pawlowski 1928 1 2 3 4 V +-2 2 +-1 I Fagus sylvatica V +-1 II - IV - V - V +-2 Mercurialis perennis III . II II V +-2 Eurhynchium zetterstedtii IV . . . V +-1 Mycelis muralis III . III . V +-1 Sanicula europaea . . . V +-1 Salvia glutinosa . . . Acer pseudoplatanus II V + 2 + II I Daphne mezereum V + . I I I Epipactis helleborine III V + . I . Galeobdolon flavidum V + . (=Lamiastrum flavidum) III II Stachys sylvatica V + . . . Viola reichenbachiana V + 2 + IV III IV +-1 Euphorbia amygdaloides . . . Melica nutans IV + . I III Brachypodium sylvaticum III +-1 . . . Rosa arvensis II III+-1 . . . Actaea spicata III III + . III . Cephalanthera damasonium III + . . . Geranium robertianum III + . III III Gymnocarpium robertianum III + . . . Lonicera alpigena II III + . . . Paris quadrifolia III III + 3 + V I I Campanula trachelium II 1 . . I I Asarum europaeum II + . . . Fraxinus excelsior II II + . . . • Galium laevigatum III II + . . . Poa nemoralis II + 1 + I I I Polystichum aculeatum II + . . . Prenanthes purpurea II + 1 + IV IV Pulmonaria officinalis II + . . . Ranunculus lanuginosus II + . . . Scrophularia nodosa II + . . . Symphytum tuberosum II + . . . Galium odoratum II +0 . I I Epilobium montanum . . III III . I I +-1 II +-3 . . . II +-4 V +-2 IV +-2 I 1-2 I + 5 . . . I I +-2 I + . . . 5 IV +-2 V +-2 . III +-3 V +-1 I I + III + III +-1 . . II + . III + . . . . . . . . . . . . . . . . V +-1 . . I + . I I + III + I I + IV + I +-1 . . . I + . I + . I+ 6 V +-2 I I I +-1 I V + IV +-2 III + I I I + I + . 6 III +-1 III +- 1 III +-2 II +-1 III + I + II + II +-1 V +-1 I + IV + I + II + V +-1 III + I + . I + . . I + I I +-1 I I + . I + I + I + I I + I I + II + I + I + . I I + . II + FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 127 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION RP 2 Q 2 P F 3 Phyteuma spicatum . . II II Petasites albus . . II I Eurhynchium striatum IV . . I II Festuca altissima III . . I . Carex sylvatica . . I . Galium sylvaticum . . . II L i l i u m m a r t a g o n . . . I QUERCETALIA ROBORIS-PETRAEAE R.-Tx. (1931) 1937 s. lat. 1 2 3 4 Pteridium aquilinum III V + . . . Veronica officinalis V + . III III Polypodium vulgare III + . . . Frangula alnus II II + . . . Melampyrum pratense III . 1+ I (I) I . . . . Castanea sativa I I . . . . III . . . . B e t u l a p e n d u l a I . . . . C a r e x m o n t a n a I I I . . . . H i e r a c i u m l a c h e n a l i i . . . . QUERCETALIA PUBESCENTIS Br.-Bl. 1931 s. lat. 1 2 3 4 Digitalis grandiflora III IV +-1 . I III Campanula witasekiana IV + . . . Hypericum montanum IV + . . . Fraxinus ornus I II 1 . . . II II 2 . . . Ostrya carpinifolia II + . . . Sorbus aria II + . I I PRUNETALIA SPINOSAE R. Tx. 1952 s. lat. 1 2 3 4 Berberis vulgaris II II + . . . Crataegus monogyna II + . . . Juniperus communis II + . . . Pyrus pyraster II + . . . Viburnum lantana II + . . . QUERCO-FAGETEA Br.-Bl. & Vlieger in Vlieger 1937 1 2 3 4 Ctenidium molluscum IV V 1-2 . . . Carex digitata III V + . . II Hepatica nobilis IV +-2 . . . Corylus avellana II III +-1 . I I I V Lonicera xylosteum III +-1 . I III Anemone nemorosa III III + 3 + I . Clematis vitalba II III + . . . Isothecium myurum IV III + . . . Platanthera bifolia III III + . . . Moehringia muscosa II + . . . Acer campestre II II + . . . Hedera helix II + . . . Listera ovata III II + . . . Prunus avium I II + . . . Moehringia trinervia III . . II III Convallaria majalis . . I II I . . . . Quercus petraea II . . . . I I I . . . . C a r p i n u s b e t u l u s I . . . . Sambucus nigra II . . . . . I + . II +-4 . . . 5 I I +-2 II +-1 I I + . I + I I +-2 I I +-2 II +-1 I + I + I + 5 . . . . . . . 5 . . . . . 5 . II +-1 . I I +-1 . . . I I +-2 . . . . . I + . . I +-1 I + I + I + I+ II +-1 II + . . I + . I + 6 II +-1 III + I + I + . . . . . . 6 II + . . . . . I I + 6 I + . . . . 6 V +-2 IV +-1 I V +-1 I + I + I V +-1 . I + I + . . . I + . . . . . FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 128 3 3 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION EP A E TG S MA SC AR AS ERICO-PINETEA Ht. 1959 s. lat. 1 2 3 4 5 6 V +-2 I I I +-1 V +-1 Carex alba III . . . . Polygala chamaebuxus . . . . . IV +-1 Buphthalmum salicifolium . . . . I + I V +-1 Calamagrostis varia IV + . . . . I I +-2 Rubus saxatilis II IV + . . I . Pimpinella saxifraga III III + . . . . Gymnadenia odoratissima II + . . . . Pinus sylvestris I . . . . II+-2 BETULO-ADENOSTYLETEA Br.-Bl. & R. Tx. 1943 s. lat. (=MULGEDIO-ACONITETEA Hadač & Klika in Klika & Hadač 1944) 1 2 3 4 5 6 Senecio fuchsii III V + . IV V III +-2 II + IV +-1 IV +-1 3 + II +-1 Polygonatum verticillatum V III . Dryopteris filix-mas III + . IV III II + Athyrium filix-femina II + 2 + V III V +-2 IV + Ranunculus platanifolius II + . . . . I I + Milium effusum . 1 + III II . I + IV +-1 Rubus idaeus II . 1 + II II I + Sambucus racemosa . . III III . I + Vicia sylvatica III . . . I . . A d e n o s t y l e s a l l i a r i a e . . . I . . EPILOBIETEA ANGUSTIFOLII R. Tx. & Prsg. in R. Tx. 1950 1 2 3 4 5 6 Fragaria vesca III V +-1 . II III II + III + Gnaphalium sylvaticum . . . . I + I + Myosotis sylvatica . . . . I + I + TRIFOLIO-GERANIETEA SANGUINEI Th. Müller 1961 s. lat. 1 2 3 4 5 6 Cruciata glabra III V +-1 . . . . I I + Clinopodium vulgare V +-1 . . . . I + Vincetoxicum hirundinaria III + . . . . . Origanum vulgare II + . . . . . SESLERIETEA Br.-Bl. 1948 em. Oberd. 1978 1 2 3 4 5 6 Campanula cochleariifolia III II + . . . . I I I +-1 Saxifraga incrustata II + . . . . . Galium saxatile . I . . . 1 + MOLINIO-ARRHENATHERETEA R. Tx. 1937 s. lat. 1 2 3 4 5 6 Ajuga reptans III V +-1 2 + . III . II + Dactylis glomerata (s.lat.) III + . . . . . Dactylorhiza maculata III + . . . . I + Leucanthemum ircutianum III + . . . . . Angelica sylvestris II + . I I . I + Cirsium palustre II + . . . . . Vicia sepium . . II IV . I + Veronica chamaedrys . . II II 1 + I + Cardamine pratensis . . I III . . ARTEMISIETEA VULGARIS Lohm., Prsg. & R. Tx. in R. Tx. 1950 1 23 4 56 Urtica dioica III III + . . . . . Aegopodium podagraria II + . . . . . II +-1 Galeopsis pubescens . . . . . ASPLENIETEA TRICHOMANIS Br.-Bl. in Meier & Br.-Bl. 1934 corr. Oberd. 1977 1 2 3 4 5 6 Asplenium trichomanes III IV + . . . . I + FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 129 MITJA ZUPANČIČ & VINKO ŽAGAR: AN OVERLOOKED SUB-ASSOCIATION IN SECONDARY SPRUCE ASSOCIATION Asplenium viride III + . . . . I I I + Asplenium ruta-muraria III + . . . . I + Moehringia muscosa II + . . . . I I + O OTHER SPECIES (Ostale vrste) 1 2 3 4 5 6 Euphorbia cyparissias III IV + . . . . I + Phyteuma sp. III + . . . . . Juglans regia II II + . . . . . NC Potentilla erecta III II + . . . I + II + Rubus fruticosus II II + . . . . I + Sorbus aucuparia II + 2+-1 IV III II +-1 III + Knautia sylvatica III . . II V . . ML MOSSES AND LICHENS (Mahovi in lišaji) 1 2 3 4 5 6 Cladonia pyxidata IV IV + . . . I + III + Plagiochila asplenioides III +-1 . I . III +-1 IV +-1 Tortella tortuosa III +-1 . . . . I V +-1 Neckera crispa II 1 . . . . . Anomodon viticulosus II + . . . . . Cladonia rangiferina II + . . . . I I I + Cladonia squamosa II + . . . . . Metzgeria furcata II + . . . . I + Mnium affine . 3 +-1 . . I +-1 I + Pogonatum aloides . . . . I + . LEGEND (Legenda) Analytical tables (Analitične tabele) 1 - Aposerido-Piceetum Zupančič (1978) 1999 galietosum rotundifolii Zupančič & Žagar 2010 2 - Picea–Galium rotundifolium ass. J. et M. Bartsch 1940 3 - Galio rotundifolii-Piceetum J. et M. Bartsch 1940 typicum Oberd. 1957 4 - Galio rotundifolii-Piceetum J. et M. Bartsch 1940 calamagrostidetosum arundinaceae Oberd. 1957 5 - Galio rotundifolii-Abietetum M. Wraber (1955) 1959 6 - Aposerido-Piceetum Zupančič 1999 var. geogr. Helleborus niger (comparison selected species in column one) Sinsistematical characteristic (Sinsistematska pripadnost) EP Erico-Pinion mugi Leibundgut 1948 nom. inv. NC Nardo-Callunetea Preising 1949 SC Scheuchzerio-Caricetea fuscae (Nordh. 1936) R. Tx. 1937 • Southeast European species (Jugovzhodno evropske vrste) Bedrock (Geološka podlaga) apn limestone (apnenec) ble micashist (blestniki) bsa variable sandstones /buntsandstein/ (pisani peščenjak) gna gneiss (gnájs) gra granite (granit) grd granodiorite (granodiorit) mor moraine (morena) FOLIA BIOLOGICA ET GEOLOGICA 51/2 – 2010 130