Numerical Evaluation of Grasslands Dominated by Sesleria Juncifolia Agg

HACQUETIA 13/1 • 2014, 57–77 DOI: 10.2478/hacq-2014-0012

Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

Eva KABAŠ1,*, Snežana VUKOJIČIĆ1, Antun ALEGRO2, Boštjan SURINA3, Nevena KUZMANOVIĆ1, Vedran ŠEGOTA4 & Dmitar LAKUŠIĆ1

Abstract Phytosociological and numerical analyses of grasslands dominated by Sesleria juncifolia s.l. in Serbia were performed in order to resolve their syntaxonomy and nomenclature. Twelve relevés were sampled on Mt. Mučanj (western Serbia), which were then compared with similar relevés from other parts of the Balkan Peninsula by means of numerical analyses. The relevés were classified using cluster analysis, while the ordination was conducted using Detrended Correspondence Analysis (DCA). The results suggest the occurrence of two floristi- cally well defined Dinaric associations in Serbia: Seslerio juncifoliae-Edraianthetum graminifolii ass. nova from Mt. Mokra Gora (Oxytropidion urumovii, Elyno-Seslerietea) and Diantho petraeae-Seslerietum juncifoliae ass. nova (Chrysopogono-Saturejion, Festuco-Brometea) from Mt. Mučanj. Key words: Balkan Peninsula, classification, ordination, Seslerietum juncifoliae s.l., syntaxonomy, vegetation.

Izvleček Naredili smo fitocenološko in numerično analizo travišč v katerih prevladuje vrsta Sesleria juncifolia s.l. in pred- stavili sintaksonomske in nomenklaturne rešitve. Dvanajst vegetacijskih popisov smo naredili na gori Mučanj (zahodna Srbija) in jih z numeričnimi metodami primerjali s podobnimi popisi z drugih delov Balkanskega polotoka. Popise smo klasificirali s klastrsko metodo, za ordinacijo smo uporabili korespondenčno analizo z odstranjenim trendom (DCA). Rezultati kažejo na obstoj dveh floristično dobro utemeljenih dinarskih en- demičnih asociacij v Srbiji: Seslerio juncifoliae-Edraianthetum graminifolii ass. nova z Mokre Gore (Oxytropidion urumovii, Elyno-Seslerietea) in Diantho petraeae-Seslerietum juncifoliae ass. nova (Chrysopogono-Saturejion, Festuco- -Brometea) z gore Mučanj. Ključne besede: Balkanski polotok, klasifikacija, ordinacija, Seslerietum juncifoliae s.l., sintaksonomija, vege- tacija.

1. INTRODUCTION the development of different types of grasslands along wide elevational and latitudinal gradients, The genus Sesleria Scop. ( Poaceae, Pooideae, forming syntaxa at various ranks — Seslerietum Seslerieae) is one of the most important and in- korabiensis Micevski 1994, Seslerietum wettsteinii teresting grass genera with its’ centre of diversity Horvat 1937, Seslerietum juncifoliae Horvat 1930, and distribution on the Balkan Peninsula. The Seslerion juncifoliae Horvat 1930, Seslerion rigidae species of this genus play a very important role in Zólyómi 1939, Seslerietalia juncifoliae Horvat 1930,

1 Institute of Botany and Botanical Garden Jevremovac, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia, [email protected]*, [email protected], [email protected], [email protected] 2 Department of Botany, Faculty of Science, University of Zagreb, Marulićev trg 20/II, 10000 Zagreb, Croatia, antun. [email protected] 3 Faculty of Mathematics, Natural Sciences and Information Technologies, University of Primorska, Glagoljaška 8, 6000 Koper, Slovenia, [email protected] 4 Institute for Research and Development of Sustainable Ecosystems, Jagodno 100a, 10415 Novo Cice, Velika Gorica, Croatia, [email protected]

57 Hacquetia 13/1 • 2014, 57–77 etc. However, their most important role is form- nietea trichomanis Br.-Bl. 1934 corr. Oberd. 1977. ing the grasslands on base-rich soils of alpine and Finally, some communities dominated by Sesleria subalpine belts of temperate European mountain spp. belong to the Festuco-Brometea Br.-Bl. & Tx. ranges (class Elyno-Seslerietea Br.-Bl. 1948). The ex Soó 1947. The dominant Sesleria species that domination by Sesleria species of the high moun- build up these syntaxa are S. filifolia Hoppe in tain calcareous plant communities in Europe is eastern Serbia and S. serbica (Adam.) Ujhelyi and discussed by Petriccione (1995) in his overview of S. juncifolia in western Serbia. this vegetation type. Our research only included the stands domi- Sesleria juncifolia agg. represents an amphi- nated by Sesleria juncifolia agg. described as asso- Adriatic group of taxa exhibiting a typical dis- ciations Seslerio-Edraianthetum jugoslavici Petkov ić junct range including the western & central Bal- et al. 1990 (Petković et al. 1990) from Mt. Mokra kans and the Apennine Peninsula, where neither Gora, Seslerietum tenuifoliae S. Vukojičić & D. their systematic relationships nor the syntaxo- Lakušić 1990 prov. (Stanić 1990) from Mt. nomical scheme are entirely clear. Based on rel- Mučanj and Carici laevis-Helianthemetum alpestre evant taxonomic papers (Deyl 1980, Strgar 1981, Horvat 1930 seslerietosum tenuifoliae Rajevski 1990 Alegro 2007, Di Pietro 2007), the following spe- from Mt. Šarplanina (Rajevski 1990) in Serbia. cies are considered to belong to S. juncifolia agg.: S. apennina Ujhelyi, S. calabrica (Deyl) Di Piet- ro, S. kalnikensis Jávorka, S. juncifolia Suffren, S. MATERIALS AND METHODS interrupta Vis., S. ujhelyii Strgar and S. albanica Ujhelyi. Since both names, S. juncifolia Suffren Data sampling. In order to describe and resolve 1802 and S. tenuifolia Shrader 1806, were validly the syntaxonomy of the Serbian grasslands domi- published, we use the earlier legitimate name S. nated by Sesleria juncifolia agg., we processed 139 juncifolia in this paper, as it has a priority accord- relevés belonging to 13 syntaxa dominated by S. ing to Art 11.4 of ICN (McNeill et al. 2012). The juncifolia agg., distributed throughout the terri - detailed discussion regarding the nomenclatural tory of Serbia, Bosnia and Herzegovina, Monte- issues in Sesleria juncifolia complex is provided in negro, Croatia, Slovenia and northeastern Italy. Di Pietro et al. (2013). The majority of the relevés were taken from litera- For the territory of Serbia, Lakušić & Sabov ture sources. The main criterion for the selection ljević (2005) proposed a classification scheme of the syntaxa to be included in the analyses was in which 31 associations and eight subassocia- that S. juncifolia appears as both the dominant tions dominanted by different Sesleria species and nominal species in the name of a syntaxa, were included, belonging to five classes, seven either at association or subassociation level (Ta- orders and 11 alliances. However, this classifica- ble 1). In addition to the literature data, personal tion was not based on serious numerical analy- unpublished data previously gathered on Mt. ses, thus an objective circumscription and clas- Mučanj in Serbia, Mt. Durmitor in Montenegro sification are still missing. The most investigated and areas of Mts. Velebit (Croatia) and Snežnik grasslands dominated by Sesleria spp. belong (Slovenia; Table 1, Figure 1) were also included in to the class Elyno-Seslerietea, order Seslerietalia the analyses. All the relevés were sampled accord- juncifoliae (=tenuifoliae) Horvat 1930, alliances ing to Braun-Blanquet (1964) method. The plot Seslerion rigidae Zólyómi 1939 (dry grasslands size of our own relevés was 25 m2, correspond- on calcareous bedrock) and Seslerion rigidae-lati ing to the standard for grasslands proposed by foliae D. Lakušić 1996 prov. (dry grasslands on Chytrý & Otýpková (2003). Plot sizes of relevés serpentine rocks). Furthermore, seven associa- from the literature varied, the exact sizes for each tions belong to the order Onobrychido-Seslerietalia association are given in Table 1. Horvat 1949, alliances Edraiantho-Seslerion Hor- Data analysis. After transforming Braun- vat 1949, Onobrychido-Festucion Horvat 1949 and Blanquet cover-abundance values into a nine- Seslerio-Festucion R. Jovanović 1955. A significant degree ordinal scale (van der Maarel 1979) the number of communities belong to the class of al- relevés were subjected to Detrended Correspond- pine pastures on siliceous rocks, Caricetea curvu- ence Analysis (DCA) in order to detect the ba- lae Br.‑Bl. 1948, order Seslerietalia comosae Simon sic structure of the floristic composition. Finally, 1957, alliance Seslerion comosae Horvat 1935. Only the complete set was classified using Bray-Curtis one association was recorded for the class Asple- similarity and group average clustering. These

58 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

Figure 1: The map of the localities of analyzed syntaxa. Numbers on the map correspond to ordinal numbers in Table 1. Blue squares correspond to groups A and C, red dots and green triangle (stands from Mt. Šarplanina excluded from second step of analysis) correspond to group B from Figure 2. Both dots and squares correspond with UTM 10 x 10 squares. Area above 1000 m a.s.l. is shaded. Country abbreviation: IT – Italy, SLO – Slovenia, CRO – Croatia, HU – Hungary, BIH – Bosnia and Herzegovina, SR – Serbia, MNE – Montenegro, RO – Romania, BU – Bulgaria, MA – Macedonia, AL – Albania. Slika 1: Karta lokacij proučevanih sintaksonov. Številke na karti so enake kot v Tabeli 1. Modri kvadrati ustrezajo skupinama A in C, rdeči krožci in zeleni trikotniki (sestoji s Šarplanine so izvzeti iz drugega koraka v analizi) ustrezajo skupini B na Sliki 2. Krožci in kvadrati se ujemajo z UTM 10 x 10 kvadranti. Območja nad 1000 m nad morjem so osenčena. Okrajšave držav: IT – Italija, SLO – Slovenija, CRO – Hrvaška, HU – Madžarska, BIH – Bosna in Hercegovina, SR – Srbija, MNE – Črna gora, RO – Romunija, BU – Bolgarija, MA – Makedonija, AL – Albanija. analyses were processed using PcOrd 6.0 (Mc- the relative frequencies of species occurrence Cune & Mefford 2011) and FLORA softwares within and outside of these groups were kept (Karadžić et al. 1998). constant (Tichý and Chytrý 2006) to calculate In this paper, we used the concept of diag- the Φ-values as a measure of fidelity independ- nostic and dominant species proposed by Chytrý ent of the number of available relevés. To assess et al. (2002), Chytrý & Tichý (2003) and Tichý statistical significance of concentration of species & Chytrý (2006). Using the statistical measures in vegetatation types, we performed the Monte of fidelity, we quantified concentrations of spe- Carlo significance test of observed maximum in- cies occurrences in groups of classified sites in dicator value for the species with 4999 permuta- order to determine diagnostic species (Chytrý et tions. PcOrd 6.0 software (McCune & Mefford al. 2002). The size of the site groups in the data 2011) was used for the calculation of Φ-values. set was standardised (virtually equalized), while In order to determine the dominant species, we

59 Hacquetia 13/1 • 2014, 57–77 calculated the coverage index (Ic) according to from the Liburnian karst. The remaining two Lausi et al. (1982). Species with Φ-values higher groups represented the relevés from northeastern than 0.50 were considered diagnostic. Species Italy, Slovenia and Croatia (Group II), and the with cover ≥ 50% in a minimum of 5% of the rel- relevés from Serbia, Bosnia and Herzegovina and evés for any association were accepted as domi- Montenegro (Group III). It can also be seen that nant. Species recorded in a minimum of 60% of the relevés from Mt. Mučanj (western Serbia) and the relevés for any association were considered the ones representing the Seslerio-Edraianthetum constant. Nomenclature of plant taxa follows jugoslavici (Mt. Mokra Gora) were well differenti- the Flora Europaea Database (Tutin et al. 2001), ated within this last group, while the relevés rep- except for critical taxa with unresolved relation- resenting the Carici laevis-Helianthemetum alpestre ships, which were included as species complexes seslerietosum tenuifoliae from Mt. Šarplanina over- (aggregates). The names of all the syntaxa follow lapped with the rest of the relevés in this group. Rodwell et al. (2002) with a few exceptions ac- Classification - Results of the cluster analy - cording to Lakušić & Sabovljević (2005). All the sis performed on the complete data set were very underlying plot data used in the paper are stored much in accordance with the ordination results, in Vegetation Database of Grassland Vegetation in that they showed that two main groups of in Serbia (Aćić et al. 2012; GIVD number: EU- stands (clusters) could be differentiated. Clus- RS-002; relevé numbers: 7000–7138). ter I (Figure 3) corresponded to the stands of the associations from northeastern Italy, Slove- nia and Croatia. This cluster corresponded com- RESULTS pletely with Groups I and II in Figure 2, so the only difference is related to relevés of Liburnian Ordination - A DCA conducted on the complete heaths Rhododendro hirsuti-Juniperetum alpinae data set showed three rather discrete groups of seslerietosum tenuifoliae, which were differentiated relevés (Figure 2), well separated along first two as a single group in DCA. On the other hand, canonical axes. The most distinct (Group I) was Cluster II (Figure 3) represented the relevés from the one representing relevés of the Rhododendro Serbia, Bosnia and Herzegovina and Montene- hirsuti-Juniperetum alpinae seslerietosum tenuifoliae gro (Group III in Figure 2). Within this cluster,

Figure 2: Detrended Correspondence Analysis (DCA) of stands of 13 grasslands dominated by Sesleria juncifolia from the Balkan Peninsula. Slika 2: Korespondenčna analiza z odstranjenim trendom (DCA) sestojev 13 traviščnih združb v katerih prevladuje Sesleria juncifolia z Balkana.

60 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

Cluster analysis 0 2.6E+01 25 1.9E+01 50 1.3E+01 Information Remaining (%) Distance (Objective function) 75 6.5E+00 0 100 1346527891111777344444455555111111111111111111111111144441111667777991191999191911111111111122222222223933333383366678778888885555656666889 0000000000123456925679801423111113131313331222222222201343333890312230040567080900000145678912345607890112563497856770891456235678091234780 000000000000000000000000000024356092817345104152637890000678900000000050 7 0002 0 3 0 41689000000000000000000000000000000000000000000000000000000 1 2 343 562 798 13 13 3 10 11 12 13

Cluster I Cluster II 1) 113 Rhododendro hirsuti-Juniperetum alpinae, Slovenia and Croatia 17) 1418 Edraiantho graminifoliae-Sesler. juncifoliae, Mt. Mokra gora 2) 6876 Carici humilis-Seslerietum juncifoliae, Mt. Velebit 18) 1930 Diantho petrae-Seslerietum juncifoliae, Mt. Muèanj 3) 3954 Seslerietum juncifoliae, Mt. Snenik 19) 3138 Seslerietum juncifoliae, Mt. Cincar 4) 111135 Carici humilis-Centaureetum rupestris, Slovenia and Italy 10) 6567 Carici laevis-Helianthemetum alpestre, Mt. ar planina 5) 136139 Bromo-Seslerietum interruptae, Mt. Biokovo 11) 7786 Carici laevis-Seslerietum juncifolia, Mt. Bjelasica 6) 092110 Genisto sericeae-Seslerietum juncifoliae, Slovenia and Italy 12) 5564 Seslerio-Gentianetum dinaricae, Mt. Vlaiæ 13) 8791 Seslerietum juncifoliae, Mt. Durmitor

Figure 3: Cluster Analysis of stands of 13 grasslands dominated by Sesleria juncifolia from the Balkan Peninsula. Slika 3: Klastrska analiza sestojev 13 traviščnih združb v katerih prevladuje Sesleria juncifolia z Balkana. it was again noticeable that Serbian associations Syntaxonomical treatment represented well separated and discrete units. On the basis of the ordination and classifica- Ass. Diantho petraeae-Seslerietum juncifoliae tion analyses, we established the existence of two Vukojičić & D. Lakušić ass. nova hoc loco (Holo well defined syntaxa with the dominance of Sesle- typus Table 2, rel. 5 hoc loco) ria juncifolia in Serbia: ass. Seslerio-Edraianthetum Original: Seslerietum tenuifoliae Stanić & D. La jugoslavici from Mt. Mokra Gora, and a new asso- kušić 1990 prov., nom. ined. (Art. 1, ICPN) ciation from Mt. Mučanj: Diantho petraeae-Sesleri- Note: The Seslerietum tenuifoliae Stanić & D. La etum juncifoliae. Relevés of the Carici laevis-Heli- kušić 1990 prov. was not effectively published anthemetum alpestre seslerietosum tenuifoliae from (Def. III, Art. 1, ICPN; Weber et al. 2000), since it Mt. Šarplanina overlapped with communities was only recorded and preliminarily described in from the Cincar, Vlašić, Bjelasica and Durmitor a Diploma thesis (Stanić 1990). Therefore, we de- mountains in the ordination, while in the cluster scribe it here as a new association, in accordance graph analysis they formed a joint cluster togeth- with the International Code of Phytosociological er with the community from Mt. Cincar and only Nomenclature (ICPN; Weber et al. 2000). a few relevés from the community from Mt. Dur- Dominant species: Sesleria juncifolia mitor. A synoptic table comparing all community Diagnostic species: Campanula rotundifolia, types is presented in the Table 3 in order to show Chamaecytisus ciliatus, Chamaespartium sagittale, the differences between the individual clusters Cotoneaster integerrimus, Dianthus petraeus subsp. obtained by numerical classification. petraeus, Draba lasiocarpa, Festuca panciciana, Ga-

61 Hacquetia 13/1 • 2014, 57–77 lium corrudifolium, Helianthemum nummularium subsp. nummularium, Ornithogalum collinum, Pe- dicularis heterodonta, Potentilla cinerea, Sanguisor- ba minor subsp. minor Constant species: Carex kitaibeliana, Edraian- thus graminifolius, Globularia cordifolia, Saxifraga paniculata Diagnosis: Rocky calcareous grasslands at elevations between 1300 and 1450 m a.s.l., on the NW and W (rarely E) exposed slopes, with an in- clination of about 35° on average (Table 2). The dominant species Sesleria juncifolia with its dense tussocks formed stands up to 40 cm high, cover- ing 25–80% (average 65%) of the plots (Table 2). The average number of species per plot of 16 m² was 26. Some Balkan endemic and Balkan-Car- pathian or Balkan-Apennine subendemic species, such as Cerastium decalvans, Daphne blagayana, Di- anthus petraeus subsp. petraeus, Draba lasiocarpa, Edraianthus graminifolius, Festuca panciciana, La- serpitium siler subsp. garganicum, Minuartia bosniaca, Pedicularis heterodonta and Sesleria juncifolia were recorded in this association. Also three gla- Figure 4: Summer aspect of the association Diantho petraeae- cial relicts (Arabis alpina, Poa alpina, and Saxifraga Seslerietum juncifoliae Vukojičić & D. Lakušić ass. nova on Mt. paniculata) occuring within the stands may point Mučanj (photo: N. Kuzmanović). to the glacial-refugial character of the association. Slika 4: Poletni aspekt asociacije Diantho petraeae-Seslerietum However, the occurrence of some differential taxa juncifoliae Vukojičić & D. Lakušić ass. nova na gori Mučanj (foto: N. Kuzmanović). from forests, such as Daphne mezereum, Fagus sylvatica, and Poa nemoralis, could point to the fact that the montane beech forests of Fagetalia sylvati- May, June and August, while the driest are Feb- cae Pawłowski 1928 prevailed in the recent past, ruary and March. Mean monthly temperatures or might even represent a natural vegetation type below 0 °C were noted during November, De- on sites nowadays covered by the stands of the cember, January, February and March, confining Diantho petraeae-Seslerietum juncifoliae. the growing season to between April and Septem- ber. Considering the relatively low elevation and Ecology and synchorology of the association the mountain temperate climate, montane beech Stands of Diantho petraeae-Seslerietum juncifoliae forests represent the potential vegetation of the are so far known only from the area of Mt. Mučanj investigated area. However, since the major parts in southwestern Serbia. The direction of the of the forests were completely degraded, artificial mountain is NW-SE, while its maximum height forest stands of black pine (Pinus nigra), fir (Abies is 1534 m a.s.l. Mt. Mučanj is seperated from the alba) and spruce (Picea abies) dominate the land- neighboring mountains by river valleys, pointing scape (Stanić 1990). to the fact that fluvial erosion was the factor shap- ing the relief, earlier formed by tectonic move- Ass. Seslerio juncifoliae-Edraianthetum gramini- ments. The dominant geological substrate in this folii B. Petković et al. ex Kabaš et al. ass. nov. area is limestone. The soils are shallow and very hoc loco skeletal. The climate type is temperate-continen- Validated name: Seslerio-Edraianthetum jugosla- tal, but in its modified mountain variant (Stanić vici Petković et al. 1990 nom. inval. (Art. 5, ICPN) 1990). The annual mean temperature is 2.9 °C, Holotypus: Petković et al. (1990: Table 2, rel. 2) and the annual precipitation is 944 mm. January is Note: The association Seslerio-Edraianthetum the coldest month with a temperature of –8,5 °C, jugoslavicii B. Petković et al. 1990 was not validly while July is the warmest with a temperature of published, because the holotype relevé was not +12.3 °C (Stanić 1990). The wettest months are assigned (Art. 5, ICPN; Weber et al. 2000). Also,

62 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia the name Edraianthus jugoslavicus Lakušić was ated group (Group II, Figure 2) corresponds to not validly published (Art. 39.1 of ICN, McNeill the stands from northwestern Italy, Slovenia and et al. 2012), since it was not accompanied by a Croatia, representing grasslands classified most- Latin description or diagnosis. Accordingly, we ly within the class Festuco-Brometea. Finally, the used the accepted and validly published name third well defined and separated group (Group Edraianthus graminifolius (L.) A. DC. (Euro+Med III, Figure 2; Cluster II, Figure 3) includes Ser- 2010) when naming the association. bian, Bosnian and Herzegovinian and Montene- Diagnostic species: Acinos arvensis, Alchemilla grian grassland stands, classified mostly within plicatula, Alyssum montanum, Anthyllis vulneraria the vegetation of the high mountain rocky cal- subsp. pulchella, Asplenium trichomanes-ramosum, careous grasslands of the class Elyno-Seslerietea. Helianthemum nummularium subsp. grandiflorum, Furthermore, both DCA and cluster analysis also Juniperus sabina, Minuartia verna, Polygonum vivi- showed that both Serbian associations are flo- parum, Scabiosa ochroleuca, Silene pusilla, Polygala ristically well differentiated from the rest of the supina subsp. supina similar syntaxa, i.e. the association Seslerio juncifoliae-Edraianthetum graminifolii from Mt. Mokra Gora, along with the stands of the associations Syntaxonomical scheme from the Bjelasica, Durmitor, Cincar and Vlašić mountains belonging to the class Elyno-Sesleriet- Festuco-Brometea Br.-Bl. & Tx. ex Soó 1947 ea. Corroboration of this statement can be seen Scorzonero-Chrysopogonetalia Horvat & Horva in the floristic composition, which is determined tić 1958 by the position of these stands at high elevations, Chrysopogono-Saturejion Horvat & Horvatić usually above the upper forest line. 1934 The newly described association from Mt. Diantho petraeae-Seslerietum juncifoliae Mučanj, on the other hand, significantly differs Vukojičić & D. Lakušić 2014 from the subalpine and alpine rocky grasslands of the class Elyno-Seslerietea in its floristic proper- Elyno-Seslerietea Br.-Bl. 1948 ties. These stands host 43 taxa (e.g., Briza media, Crepidetalia dinaricae Lakušić 1966 Bromus erectus, Euphorbia myrsinites, Globularia Oxytropidion urumovii Lakušić 1964 cordifolia, Hieracium pilosella, Hypericum perfora- Seslerio juncifoliae-Edraianthetum grami- tum, Juniperus communis, Leucanthemum vulgare, nifolii Petković et al. ex Kabaš et al. 2014 Phleum pratense, Plantago lanceolata, Plantago media, Poa badensis, Potentilla cinerea, Sanguisorba minor, Teucrium chamaedrys, Thymus pulegioides, Discussion Trifolium campestre, Trifolium montanum etc.) which, within the eastern and southeastern Di- Our numerical analyses showed that the analyzed naric Alps, prefer calcareous grasslands devel- stands of different syntaxa dominated by Sesleria oped within deciduous broadleaved forests. Ad- juncifolia agg. in the mountains of the central and ditionally, the Φ-indices of most of these taxa are western part of the Balkan Peninsula are very het- highly statistically supported, hence we find their erogenous and probably have different origins. classification within the class Festuco-Brometea to Their relationships in a broader, amphi-Adriatic be fully justified. The fact that only 12 taxa typi- context are a subject of the ongoing research, pre- cal for subalpine and alpine grasslands (Acinos liminarily presented in Di Pietro et al. (2013). alpinus, Arabis alpina, Carex kitaibeliana, Daphne Our results showed that the first clearly dis- alpina, Edraianthus graminifolius, Helianthemum tinct group (Group I, Figure 2) represents the canum, Hieracium villosum, Pedicularis hetero- stands of the Rhododendro hirsuti-Juniperetum al- donta, Poa alpina, Rosa pendulina, Sesleria junci- pinae seslerietosum tenuifoliae. These results sup- folia and Thlaspi kovatsii) occur in stands from port the opinion of Surina (2013) that despite Mt. Mučanj further supports our classification the domination of S. juncifolia in these stands, scheme. To that end, Φ-indices of the majority of the syntaxon should actually be classified with- the subalpine and alpine grassland taxa from the in the heath vegetation of Erico-Pinetea Horvat class Elyno-Seslerietea do not reflect statistically 1959, and not within the grassland communities significant fidelity or the diagnostic value for the of Elyno-Seslerietea. The second well differenti- association in general.

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Confirmation of the opinion that the new and the origin of the association are considered. community from Mt. Mučanj belongs to the class Specific chasmophytic taxa are present in rocky of Festuco-Brometea should also be sought in its grasslands due to the proximity of chasmophytic coenotic surroundings and its origin. The poten- stands in the investigated area (Stanić & Lakušić tial natural vegetation of the highest part of Mt. 1993) and the fact that steep rocky grasslands Mučanj is deciduous broadleaved forest of the share many features with rock crevices or even Fagetalia sylvaticae order (Jovanović et al. 1986). screes, generating ecological conditions suitable Therefore, at an altitude of 1500 m a.s.l., this for both grassland and chasmophytic taxa, e.g. isolated mountain does not provide conditions Asplenium ceterach, Asplenium ruta-muraria, Asple- for the development of the potential vegetation nium trichomanes, Campanula rotundifolia, Erysi- of (sub)alpine grasslands of the Elyno-Seslerietea mum sylvestris, Hieracium humile, Hieracium pan- class. nosum, Laserpitium siler, Leontopodium alpinum, While there is no doubt the new association Saxifraga tridactylites, or Silene saxifraga. This cir- from Mt. Mučanj belongs to the class Festuco- cumstance was observed and recently discussed Brometea, and not Elyno-Seslerietea, as indicated by Surina & Martinčič (2012). Nevertheless, low in Lakušić & Sabovljević (2005), its assignment coverage and fidelity indices of chasmophytes in to a lower syntaxa is not completely clear. Con- the studied stands do not justify their classifica- sidering their geographic position and floristic tion within the class Asplenietea trichomanis. composition, these stands show transitional char- Finaly, the syntaxonomical position of the acteristics connecting them to calcareous karstic Carici laevis-Helianthemetum alpestre seslerietosum grasslands of the Illyric-Dinaric region with tenuifoliae from Mt. Šarplanina remains unclear, Chrysopogono-Saturejion Horvat & Horvatić 1934, given that in the DCA analysis its stands are meso-xerophytic swards in sub-oceanic regions overlapping with the stands of associations from of western Europe with Bromion erecti Koch 1926, Bosnian and Montenegrin mountains. A possible and meso-xerophytic swards in sub-continental reason for this is the fact that this subassociation regions of central and eastern Europe with Cirsio- was described on the basis of only three relevés, Brachypodion pinnati Hadač & Klika 1944. The which is not representative enough for an objec- most important species connecting this associa- tive understanding of its syntaxonomical position with the Chrysopogono-Saturejion are: Asperula tion. While all of the other investigated syntaxa cynanchica, Dianthus petraeus, Galium corrudifo- are found within the Dinaric floristic province, lium, Globularia cordifolia, Hieracium villosum, La- the stands of this subassociation belong to the serpitium siler, Leontodon crispus, Sesleria juncifolia, Scardo-Pindic floristic province. Accordingly, and Teucrium chamaedrys. The species relating it the comparison of these three relevés with the with the associations of the alliance Bromion erecti rest of the 136 dinaric relevés would not reflect are the following: Plantago media, Sanguisorba mi- the real relationships of these syntaxa within the nor, Scabiosa columbaria, considered as diagnostic Balkan Peninsula. for the Bromion erecti (Dengler 2003, Jarolímek & Šibík 2008, Chytrý 2010), while it also hosts diagnostic species of the Cirsio-Brachypodion pinnati Acknowledgements (Dengler 2003, Jarolímek & Šibík 2008, Chytrý 2010) such as Asperula cynanchica, Brachypodium The authors are grateful to the Serbian Ministry pinnatum, Bromus erectus, Plantago media, San- of Science and Technological Development (Pro- guisorba minor, Trifolium montanum. Therefore, ject No. 173030 Biodiversity of the plant life of considering the number of common species and Serbia and Balkan Peninsula – Assessment, sus- their diagnostic character as well as their geo- tainable use and conservation, 2011‑2014) for fi- graphic position within the continental Dinar- nancial support. The authors also wish to thank ides, the authors think the position of the new as- Jozef Šibík, Kiril Vassilev and Jürgen Dengler for sociation Diantho petraeae-Seslerietum juncifoliae is their valuable comments and suggestions, which within the alliance Chrysopogono-Saturejion. have significantly improved this manuscript. Fi- The presence of 11 chasmophytic taxa in the nally, we thank Laura Sutcliffe for the linguistic stand of the newly described association is also editing of the manuscript and EDGG for making significant, when the syntaxonomic position this possible through an IAVS grant.

64 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

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66 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

Table 1: Analysed syntaxa dominated by Sesleria juncifola agg. from the Balkan Peninsula used in the analysis. Tabela 1: Proučevani sintaksoni v katerih prevladuje Sesleria juncifola agg. z Balkanskega polotoka uporabljeni v analizi.

No. Syntaxon UTM Locality Reference Plot size Number (m²) of relevés 1. Rhododendro hirsuti-Juniperetum 33T VL53 Slovenia Surina 30 13 alpinae Horvat ex Horvat et al. 1974 (Liburnian 2013 subas. seslerietosum tenuifoliae karst), Croatia Surina 2013 (Liburnian karst) 2. Seslerio-Edraianthetum jugoslavicii 34T DN44 Serbia (Mt. Petković 25–100 5 Petković et al. 1990 Mokra Gora) et al. 1990 3. Seslerietum tenuifoliae Stanić & 34T DP22 Serbia (Mt. Vukojičić & 20–400 12 D. Lakušić 1990 prov. Mučanj) Lakušić, D. unpubl. 4, 5. Seslerietum juncifoliae 33T XJ66 Bosnia and Lakušić et al. 20–100 24 Horvat 1930 Herzegovina 1984 (Mt. Cincar) 6. Seslerio-Gentianetum dinaricae 33T YK00, YK01 Bosnia and Lakušić et al. 10–100 10 Lakušić et al. 1982 Herzegovina 1982 (Mt. Vlašić) 7. Carici laevis-Helianthemetum 34T EM07 Serbia (Mt. Šar- Rajevski - 3 alpestre Horvat 1930 subas. planina) 1990 seslerietosum tenuifoliae Rajevski 1990 8. Carici humilis-Seslerietum 33T VK95 Croatia (northern Alegro 50 9 juncifoliae part of Mt. & Segota Velebit) 2009, 2010, manuscript 9. Carici laevis-Seslerietum tenuifoliae 34T CN94 Montenegro Lakušić 100–200 10 (Lakušić 1966) Redžić 2011 (Mt. Bjelasica) 1966 (=Seslerietum juncifoliae montenegrinum Lakušić 1964) 10. Seslerietum juncifoliae s.l. 34T DN46, DN47, Montenegro (Mt. Lakušić, D., 20–100 5 DN48 Durmitor) unpubl. 11. Genisto sericeae-Seslerietum 33T UL96, VL06, Slovenia Poldini 1989, 70–80 19 juncifoliae Poldini 1980 VL13, VL14, VL15, (Liburnian Kaligarič VL23, VL26, VL27, karst), Italy 1997 VL44 (Karst) 12. Carici humilis-Centaureetum 33T VL13, VL15, Slovenia Poldini 1989, 100 25 rupestris Horvat 1931 seslerietosum VL23, VL24, VL26, (Liburnian Kaligarič juncifoliae Horvat 1962 VL27, VL44, VL54 karst), Italy 1997, Surina (Karst) unpubl. 13. Bromo-Seslerietum interruptae 33T XH69 Croatia Trinajstić - 4 Trinajstić 1965 (Mt. Biokovo) 1987

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Table 2: Analytical table of the association Diantho petraeae-Seslerietum juncifoliae ass. nova. from Mt. Mučanj in Serbia. Species are sorted in descending order of constancy and cover values. (* = holotypus). Tabela 2: Analitična tabela asociacije Diantho petraeae-Seslerietum juncifoliae ass. nova. z gore Mučanj v Srbiji. Vrste so razvrščene padajoče glede na stalnost in pokrovnost (* = holotip).

Altitude (m a.s.l.) 1450 1400 1450 1450 1450 1450 1450 1450 1450 1350 1350 1300 Exposition – – W NW NW NW W NE NW E E S Slope (°) 0 0 10 10 10 30 70 70 70 50 65 70 Total cover (%) 60 60 80 80 80 50 80 70 60 60 80 25 Plot size (m²) 25 25 100 20 30 75 24 80 30 400 50 30 VII VII VII VII VII VII VII VII VII VII VII VII Date 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989

No. Relevé 1 2 3 4 5* 6 7 8 9 10 11 12 Constancy (%) Cover index according to Lausi et al. (1982) (Ic,%) Dominant taxa Sesleria juncifolia agg. 2.3 2.3 3.4 4.5 4.5 3.5 4.4 4.5 3.4 3.4 2.3 2.3 100 74 Constant taxa Globularia cordifolia 3.4 3.3 1.3 1.3 1.2 1.3 1.2 1.1 3.4 1.3 . 1.2 92 33 Edraianthus graminifolius + 1.1 1.2 1.2 1.1 1.1 1.2 1.2 1.2 1.3 . 1.1 92 42 Saxifraga paniculata . . 1.1 1.2 1.2 1.2 . 1.1 . 1.2 1.2 1.2 67 22 Carex kitaibeliana 1.2 1.2 . 1.2 1.2 1.2 . 1.2 1.2 . . . 67 22 Diagnostic taxa (*also constant taxa) Dianthus petraeus subsp. petraeus* 1.2 1.3 1.3 1.3 1.3 1.2 1.3 1.3 1.3 1.4 1.3 1.2 100 33 Chamaecytisus ciliatus* 1.2 1.3 1.2 1.2 1.2 1.2 1.3 1.3 1.2 1.3 1.2 1.2 92 30 Festuca panciciana* 1.1 . 1.2 1.2 1.2 1.2 1.2 1.2 . 1.2 2.2 1.2 92 30 Pedicularis heterodonta* + 1.1 1.1 1.1 1.1 1.1 1.1 1.1 1.1 + . . 83 30 Potentilla cinerea* 1.2 . 1.2 1.3 1.2 1.2 1.3 1.3 1.3 1.2 1.2 1.2 83 26 Ornithogalum collinum* 1.1 . . 1.1 1.1 1.1 1.1 + + 1.1 1.1 . 75 23 Cotoneaster integerrimus* 1.2 1.2 + + 1.1 1.2 . + . 1.3 1.3 . 75 22 Campanula rotundifolia* ...... 1.2 1.2 1.1 1.2 1.2 1.2 75 22 Sanguisorba minor subsp. minor + . 1.1 1.1 + . 1.1 1.1 . 1.2 1.2 + 58 19 Helianthemum nummularium subsp. . 1.1 . . 1.1 1.1 1.1 1.1 . . 1.2 . 50 16 nummularium Draba lasiocarpa + . . . . . 1.2 + . 1.1 1.2 + 50 16 Chamaespartium sagittale 1.2 + 1.1 . . . . . + 1.2 1.2 1.2 50 15 Galium corrudifolium . . 1.2 . 1.2 1.2 1.3 . . 1.3 1.3 . 50 14 Other taxa Bromus erectus 1.1 . 1.2 1.2 1.1 1.1 1.2 . . 1.1 . 58 19 Minuartia verna 1.1 . 1.2 1.2 1.2 . 1.2 1.2 . 1.2 1.2 . 58 18 Leontodon crispus subsp. crispus 1.1 + + . . . . . 1.2 1.3 . . 50 17 Trifolium montanum 1.2 1.1 1.1 . 1.1 1.1 ...... 42 14 Arenaria serpyllifolia subsp. serpyllifolia 1.2 . . 1.2 . . . . . 1.1 1.2 . 42 13 Trifolium alpestre 1.1 . + + . . . . . 1.2 1.2 1.1 42 12 Euphrasia stricta 1.1 . 1.1 1.1 + . . . . 1.1 . . 42 10 Asplenium ceterach ...... 1.1 1.1 1.1 33 11 Juniperus communis . + . . . 1.2 . . . 1.2 . + 33 11 Poa alpina 1.1 . 1.1 1.2 + ...... 33 10 Hieracium pilosella subsp. pilosella . . . 1.2 . 1.2 1.1 . . . . . 33 9 Saxifraga tridactylites . . . 1.1 1.1 . 1.1 . . 1.1 . . 33 9 Gymnadenia conopsea + + . . + 1.1 . + . . . . 25 8 Rhamnus saxatilis subsp. saxatilis ...... 1.3 1.3 1.2 25 8 Stachys recta ...... 1.2 1.2 1.2 25 8 Sedum acre . . . + . . . . . 1.2 1.2 + 25 8 Euphorbia myrsinites ...... 2.3 + 25 8 Rosa pendulina . 1.1 . . 1.1 + ...... 25 8 Asplenium ruta-muraria subsp. ruta-muraria . . . . . + . . . 1.1 . . 25 8 Verbascum sp...... 1.2 1.1 1.1 25 7

68 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia

No. Relevé 1 2 3 4 5* 6 7 8 9 10 11 12 C. (%) Cov. Acinos alpinus ...... 1.1 . . 1.2 . 1.2 25 7 Teucrium chamaedrys ...... 1.3 1.3 1.2 25 7 Sedum album . . . . + . . . . 1.2 1.2 . 25 6 Erysimum sylvestre subsp. sylvestre . . . . + + . . . . 1.1 . 17 7 Leontopodium alpinum subsp. alpinum . . . . . 1.2 . 1.1 . . . . 17 6 Arabis hirsuta ...... 1.1 1.2 . 17 6 Phleum pratense subsp. pratense ...... 1.1 1.3 . 17 6 Thlaspi kovatsii ...... 1.1 1.1 . . . . 17 6 Aethionema saxatile subsp. saxatile ...... 1.1 1.2 . 17 6 Daphne mezereum . . . . . 1.1 ...... 17 6 Thymus pulegioides ...... 1.1 . . . . . 17 6 Vicia incana . + ...... 1.2 . 17 6 Cerastium decalvans ...... 1.2 1.2 . 17 6 Minuartia bosniaca ...... 1.2 . + 17 5 Briza media subsp. media + . . . . 1.1 ...... 17 5 Scabiosa columbaria subsp. columbaria . . 1.1 ...... 1.1 . . 17 5 Hieracium villosum ...... 1.2 1.1 . . . 17 5 Poa nemoralis . . . . . 1.3 1.2 . . . . . 17 4 Rhamnus alpinus subsp. fallax ...... 1.3 . . 17 4 Epipactis atrorubens . . . . . + + . . . + . 8 3 Thymus glabrescens . . + . + ...... 8 3 Hieracium pannosum ...... 1.4 . . 8 3 Arabis glabra ...... 1.2 . . 8 3 Cerastium brachypetalum subsp...... 1.3 . 8 3 brachypetalum Arabis alpina ...... 1.3 . 8 3 Leucanthemum vulgare . 1.1 ...... 8 3 Plantago lanceolata . . 1.1 ...... 8 3 Plantago media 1.1 ...... 8 3 Heracleum sp...... 1.1 ...... 8 3 Fagus sylvatica . . . . . + ...... 8 3 Valeriana montana ...... + . . . . . 8 3 Myosotis arvensis subsp. arvensis ...... + . . . . 8 3 Erigeron annuus ...... + . . 8 3 Sorbus aria . . . . + ...... 8 3 Hypericum perforatum ...... + . . 8 3 Poa badensis ...... 1.1 . . 8 3 Asperula aristata subsp. scabra ...... 1.1 . . 8 3 Polygala supina subsp. supina ...... 1.2 . . 8 3 Brachypodium pinnatum subsp. pinnatum . . + ...... 8 3 Asplenium trichomanes subsp. trichomanes ...... 1.1 . 8 3 Lotus corniculatus + ...... 8 3 Fragaria vesca . . . . . + + . . . . . 8 2 Allium flavum subsp. flavum ...... 1.1 . . 8 2 Laserpitium siler subsp. siler ...... 1.3 . . 8 2 Vincetoxicum hirundinaria ...... 1.3 . 8 2 Daphne blagayana . 1.2 ...... 8 2 Trifolium campestre ...... + . . 8 2 Silene saxifraga ...... 1.3 . . 8 2 Asperula cynanchica ...... 1.1 . . 8 2 Helianthemum canum subsp. canum . . 1.1 ...... 8 2

Relevés 1, 3–9: Serbia, Mt. Mučanj, UTM grid 34T DP22, 43.544924° N, 20.038117° E Relevé 2: Serbia, Mt. Mučanj, UTM grid 34T DP22, 43.543552° N, 20.038329° E Relevés 10–12: Serbia, Mt. Mučanj, UTM grid 34T DP22, 43.542562° N, 20.038277° E

Hacquetia 13/1 • 2014, 57–77 Overall constancy Overall

7 5 7 4 4 4 4 4 4 4 4 4 3 3 3

C 13 Max Φ-values Max

0.6 0.38 0.57 0.33 0.81 0.75 0.71 0.68 0.62 0.52 0.51 0.51 0.5 0.68 0.72 0.62 Group with Max Φ-value observed Φ-value Max with Group

1 5 6 3 4 2 3 6 2 4 5 2 2 A

12 13 12

Biokovo

, ,

Bromo-Seslerietum interruptae Bromo-Seslerietum ......

13 100

, Slovenija & Italia & Slovenija , rupestris

Carici humilis-Centaureetum humilis-Centaureetum Carici ......

12 60 56 20 68 12 100

, Slovenia & Italia & Slovenia , juncifoliae

Genisto sericeae-Seslerietum sericeae-Seslerietum Genisto ......

11 89 32 16 26 16

Durmitor

Seslerietum juncifoliae Seslerietum ......

40 10 40 20 40 20 60 40 20 100

, Bjelasica , tenuifoliae

Carici laevis-Seslerietum laevis-Seslerietum Carici ...... 9

30 30 40 20 100

, Velebit , juncifoliae

Carici humilis-Seslerietum humilis-Seslerietum Carici ...... 8

11 11 11 11 11 33 33 22 22 100

, Sara , juncifoliae ses. alpestre

Carici laevis-Helianthemetum laevis-Helianthemetum Carici ...... 7

67 67 67 67 100 100

Vlašić

, , dinaricae eslerio-Gentianaetum

S . . . Constancy (%) 6

30 30 20 40 10 70 30 40 50 10 30 100 100

, Snežnik , juncifoliae Seslerietum

...... 5

75 31 75 100

, Cincar , juncifoliae Seslerietum

...... 4

100

, Mučanj , foliae

Diantho petraeae-Seslerietum junci petraeae-Seslerietum Diantho ...... 3 8 8

58 25 83 17 58 100 100

, Mokra gora Mokra , graminifolii

Seslerio juncifoliae-Edraianthetum juncifoliae-Edraianthetum Seslerio ...... 2

60 40 80 60 80 100

Hrvatska & Slovenia , alpinae Rhododendro hirsuti-Juniperetum hirsuti-Juniperetum Rhododendro ...... 1 92 85 (Scop.) Schinz & Thell. ssp. grandiflorum (Scop.) Schinz & (Jacq.) Breistr. (L.) DC. ssp. alpestre Scop. ssp. germanicum (Gremli) Gams (Podp.) O. Schwarz L. Schloss. & Vuk. Schloss. & Lam. agg. agg. L . Waldst. & Kit. Waldst. Dominant taxa Sesleria juncifolia Taxa Constant taxa agg. erectus Bromus Lotus corniculatus L. Helianthemum canum (L.) Baumg. ssp. Diagnostic taxa Rosa pendulina Waldst. & Kit. ssp. petraeus Dianthus petraeus Waldst. Helianthemum oelandicum Helianthemum nummularium (L.) Miller Potentilla cinerea Chaix ex Vill. Potentilla cinerea Cerastium decalvans Minuartia verna Dorycnium pentaphyllum Globularia meridionalis Polygonum viviparum Silene pusilla Coronilla vaginalis Coronilla Synoptic table of all the analyzed communities. The percentage constancy values for each species within each of the 13 analyzed syntaxa are given in the given are syntaxa the 13 analyzed each species each of within for values constancy percentage The all communities. the of analyzed table 3: Synoptic Table Species are observed was constancy. it the overall where and the syntaxa of the species, the of number Φ -values the max to refer columns three last The columns. shown. not are constancy 10% overall species than less with companion The Φ -values. and constancy of sorted order in descending treh V zadnjih sintaksonih. v 13 analiziranih v odstotkih vrst so podane V stolpcih stalnosti pojavljanj združb. tabela vseh proučevanih 3: Sinoptična Tabela so razvrščene padajoče Vrste stalnost. in celotna se pojavlja v katerih število sintaksonov posamezne vrste, maksimalna Φ vrednost so predstavljene stolpcih pod s stalnostjo 10% niso vrste prikazane. Spremljevalne glede in Φ vrednosti. stalnost na

70 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia 3 2 3 2 3 2 3 2 3 2 2 3 2 3 2 3 2 2 3 2 3 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 C B 0.6 0.72 0.59 0.72 0.58 0.71 0.56 0.69 0.54 0.69 0.68 0.53 0.65 0.53 0.65 0.53 0.63 0.63 0.53 0.63 0.51 1 0.5 0.63 0.95 0.59 0.95 0.58 0.88 0.57 0.85 0.56 0.78 0.55 0.55 0.78 0.55 0.77 0.75 4 7 3 9 3 4 3 4 1 1 6 3 9 3 3 3 5 1 9 1 3 5 2 4 9 1 1 2 2 9 7 7 A 12 13 13 13 13 12 12 ...... 13 50 25 75 50 100 ...... 4 8 12 36 52 28 12 64 72 ...... 5 11 21 21 16 42 16 ...... 60 10 20 20 40 20 20 40 20 20 20 20 20 ...... 9 20 20 70 60 50 90 40 70 ...... 8 11 11 67 33 33 44 67 ...... 7 67 67 33 67 67 100 100 100 ...... 6 80 60 60 10 40 10 ...... 5 6 63 13 44 56 ...... 4 ...... 3 8 8 92 75 50 50 75 50 50 92 ...... 2 40 40 20 40 40 40 100 ...... 1 85 54 31 54 77 100 100 . . (Boiss.) Nyman (A. Kerner ex Borbás) Jalas (Vis.) Bornm. (Vis.) (Borbás) Martinovsky & Skalicky . ssp. mollis (L.) P. Gibbs (L.) P. Medicus L. (Desf.) Boiss. & Reuter (L.) Sprengel (Wahlenb.) Rothm (Wahlenb.) All. L. (Hackel) K. Richter L. ssp. pulchella Vill. Heuffel Vill. ssp. austriacum (Schott ex Roemer & Schultes) Hayek Vill. (Schott) Á. & D. Löve Chaix Kummer & Sendtner L. Schreber ssp. supina ssp. eriocaulis (L.) Reichenb (L.) Karsten ssp. abies Oxytropis dinarica (Murb.) Wettst. Oxytropis Festuca panciciana Thymus praecox Opiz ssp. zygiformis (H. Braun) Jalas Campanula rotundifolia Campanula rotundifolia Euphorbia nicaeensis Thymus praecox Opiz ssp. polytrichus Chamaespartium sagittale Arctostaphylos uva-ursi Arctostaphylos (L.) Ehrend. Asperula purpurea Picea abies Cyclamen purpurascens Miller Vill. ssp. grisebachii Vill. Hypericum richeri Draba lasiocarpa Rochel Jovibarba heuffelii Cotoneaster integerrimus Galium corrudifolium Vill. Stipa pennata Helianthemum nummularium (L.) Miller ssp. (L.) Moench Peucedanum oreoselinum (Dunal) Gren. & Godron Fumana procumbens Verbascum chaixii Verbascum Daphne alpina L. Salix appendiculata Asperula aristata L. Plantago argentea Chamaecytisus ciliatus Plantago lanceolata Anthyllis vulneraria Festuca bosniaca Draba aizoides L. L. Athamanta cretensis Juniperus communis ssp. alpina (Suter) Celak Polygala supina Jurinea mollis Hieracium pavichii Juniperus sabina L. (Griseb.) P. W. Ball W. (Griseb.) P. Onobrychis montana DC. ssp. scardica Dryas octopetala Bupleurum ranunculoides

71 Hacquetia 13/1 • 2014, 57–77 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C B 0.86 0.55 0.86 0.55 0.86 0.55 0.86 0.54 0.83 0.54 0.83 0.53 0.83 0.5 0.82 0.5 0.5 1 0.81 0.81 1 0.81 0.81 1 0.81 1 0.81 1 0.76 1 0.76 0.91 0.76 0.91 0.76 0.91 0.75 0.89 0.86 3 4 4 4 9 9 1 9 5 1 6 1 7 7 1 7 7 2 7 7 7 2 2 1 6 1 9 3 7 6 3 A 10 10 12 13 13 10 13 13 ...... 13 75 100 100 ...... 12 32 16 12 28 ...... 5 11 11 16 ...... 40 40 10 40 ...... 9 70 70 70 60 ...... 8 11 11 11 22 ...... 7 67 67 67 67 67 67 67 100 ...... 6 50 60 80 ...... 5 38 ...... 4 75 75 ...... 3 83 75 58 ...... 2 40 60 60 100 ...... 1 62 69 85 85 100 100 (A.Kerner) Chater (Rchb.) Cullen (DC.) Nyman Bonnier & Layens Nyman Schlosser & Vuk. ssp. brachyodonta Vuk. Schlosser & Lam. L. (Waldst. & Kit.) A. DC. & Kit.) (Waldst. Guss. (Sibth. & Sm.) Boiss. L. Gand. L. ssp. polyphylla L. ssp. weldeniana L. L. G. Beck L. L. ssp. pulchellum L. Haenke Balbis Pourret Tineo Miller Androsace villosa Androsace Genista januensis Viv. Achillea clavennae Iberis pruitii Anthyllis vulneraria Satureja montana L. ssp. illyrica Satureja Astragalus vesicarius L. ssp. carniolicus (Georgiev) Niketić Cerastium malyi (Georgiev) L. ssp. ruthenicus (Bieb.) Nyman Echinops ritro F. W. Schmidt W. Myosotis alpestris F. Thesium linophyllon Festuca varia Helianthemum nummularium ssp. obscurum (Celak.) J. Holub (Hoffm.) Newman (Hoffm.) Gymnocarpium robertianum Ranunculus montanus agg. Achillea ageratifolia Calamagrostis varia (Schrader) Host ssp. Calamagrostis Lilium bosniacum (Beck) Fritsch Erica herbacea Campanula spatulata Sibth. & Sm. ssp. Saxifraga sempervivum C. Koch Alchemilla plicatula (Kümmerle & Jáv.) Deyl Sesleria korabensis (Kümmerle & Jáv.) Poa molinerii Festuca duriuscula Silene ciliata Anthyllis vulneraria Alyssum montanum L. Edraianthus tenuifolius Asplenium trichomanes-ramosum L. Abies alba Pedicularis brachyodonta Campanula cochlearifolia Pedicularis verticillata Pančić Pedicularis heterodonta Allium carinatum Gentiana dinarica Ornithogalum collinum Sanguisorba minor Scop. ssp.

72 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C B 0.74 0.72 0.62 0.62 0.72 0.62 0.69 0.62 0.69 0.62 0.69 0.69 0.61 0.69 0.61 0.6 0.69 0.69 0.58 0.56 0.69 0.69 0.56 0.56 0.69 0.56 0.67 0.56 0.67 0.65 0.55 0.55 0.65 0.54 0.65 0.54 0.63 0.62 0.62 5 1 9 1 6 6 9 1 1 5 1 7 7 7 7 1 7 1 8 8 1 9 1 3 2 2 A 10 10 10 10 13 13 13 13 13 13 13 12 12 ...... 13 50 50 50 50 50 50 50 ...... 12 32 32 ...... 11 ...... 10 40 40 40 40 ...... 9 40 50 60 ...... 8 44 44 ...... 7 33 33 33 33 33 ...... 6 50 50 ...... 5 56 38 ...... 4 ...... 3 42 ...... 2 40 40 ...... 1 54 54 38 38 69 46 46 31 31 (Wettst.) Hayek (Wettst.) (Griseb.) Engler

Schlosser & Vuk. ssp. grisebachii Vuk. Schlosser & (Horvatić) Vogt & Greuter Vogt (Horvatić) ssp. cristata (Bartl.) Dostál Buser L. (Halácsy) Mattf. Hackel ex Wiesb. Bartl. ex Vis. (St-Yves) Markgr.-Dannenb. (St-Yves) Schleicher ex Gaudin (Hoppe) Reichenb. (L.) Scop. Wettst. (Crantz) Beck ex Fritsch L . Boiss. & Spruner Asso (L.) Miller ssp. alpina L. Perr. & Song. Perr. L. Gentiana clusii Minuartia baldaccii Homogyne sylvestris Cass. Lonicera caerulea Trifolium noricum Wulfen Trifolium Silene parnassica Genista pilosa L. Leucanthemum illyricum Sternb. var. coriophylla Sternb. var. Saxifraga marginata Pedicularis brachyodonta Wettst. Alyssum scardicum Rubus saxatilis (Koch) P.W. Ball Bunium alpinum ssp. montanum (Koch) P.W. Carlina corymbosa L. Cirsium erisithales (Jacq.) Scop. Carex halleriana Carex (Scop.) Link Chamaecytisus purpureus Clematis alpina Centaurea spinosociliata Centaurea Euphrasia illyrica Festuca adamovicii Alchemilla flabellata Ornithogalum orthophyllum Ten. Festuca pseudovina Cerastium alpinum L. Hieracium alpicola Aster bellidiastrum Primula veris L. ssp. L. Hieracium murorum Pulsatilla montana Rosa pimpinellifolia Satureja subspicata Satureja Scabiosa triandra L. L. Solidago virgaurea Potentilla crantzii Carex digitata Carex D. Wolff ex J. F. Lehm. ex J. F. Wolff Euphrasia stricta D. Acinos arvensis (Lam.) Dandy L. Scabiosa ochroleuca

73 Hacquetia 13/1 • 2014, 57–77 5 1 1 5 5 1 1 5 5 1 1 4 1 1 4 6 1 4 6 4 6 4 6 4 6 4 6 6 6 4 4 6 4 5 5 4 4 4 C B 0.4 0.54 0.54 0.34 0.36 0.54 0.53 0.31 0.33 0.53 0.53 0.49 0.53 0.53 0.48 0.48 0.51 0.44 0.45 0.39 0.44 0.39 0.44 0.38 0.42 0.37 0.33 0.4 0.31 0.32 0.32 0.3 0.31 0.42 0.43 0.3 0.27 0.28 9 1 4 7 9 5 6 6 9 3 9 9 1 3 2 6 1 6 6 5 4 6 3 1 A 11 11 13 13 12 13 12 13 12 13 12 13 13 13 ...... 13 75 75 25 75 50 25 50 50 100 100 100 100 ...... 8 8 8 12 40 60 40 20 28 60 48 72 24 44 16 76 28 20 32 88 36 56 28 32 ...... 5 5 5 11 11 11 11 11 42 37 68 84 37 37 63 26 74 ...... 10 20 20 20 20 20 20 40 20 20 20 20 20 20 20 60 20 ...... 9 30 50 30 10 30 30 50 80 20 30 10 70 100 ...... 8 11 11 11 11 11 11 33 56 33 33 22 44 22 56 33 22 33 ...... 7 67 67 33 100 100 ...... 6 30 20 30 30 80 30 10 40 20 30 30 40 30 100 ...... 5 6 31 44 44 19 56 44 19 50 56 13 31 ...... 4 ...... 3 8 8 33 25 92 67 50 17 67 25 100 ...... 2 40 20 40 40 100 ...... 1 8 92 77 69 46 100 (A. Kerner) Hayek L. (L.) A. DC. (L.) (Bellardi) Chiov. L. L. All. ssp. adscendens (Bartl.) Dostál Miller Scop. Risso ex Koch Jacq. Latourr. C. Presl Thore Degen ex Becherer Scop. L. ssp. jacquinii agg. Wulfen (Waldst. & Kit.) Nyman ssp. simplex (Waldst. (L.) Moench L. agg. L. L. Rhododendron hirsutum L. Rhododendron Poa alpina (G. Beck) P. D. Sell ssp. dinarica (G. Beck) P. Tausch praemorsa (L.) Crepis Thalictrum aquilegiifolium Inula ensifolia Gentiana verna croaticus (Borbás) Tutin ssp. croaticus Urv. Dianthus giganteus D’ Vicia cracca Vicia Acinos alpinus Genista sylvestris Festuca rubra L. ssp. Bellardiochloa violacea Bellardiochloa Allium ericetorum Stachys germanica L. ssp. Centaurea triumfetti Centaurea Genista sericea Teucrium montanum L. Teucrium Other taxa Other Globularia cordifolia Centaurea rupestris L. Centaurea Edraianthus graminifolius Koeleria splendens Carex kitaibeliana Carex Plantago holosteum Dianthus sylvestris agg. Vill. ssp. crispus Vill. Leontodon crispus L. Phyteuma orbiculare Carlina acaulis Asperula cynanchica Galium lucidum All. Carex humilis Leysser Carex Polygala nicaeensis Hieracium villosum Trinia glauca Trinia Carex caryophyllea Carex Saxifraga paniculata Anthyllis montana Laserpitium siler L. ssp. Teucrium chamaedrys L. Teucrium

74 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia 3 3 4 3 4 3 4 3 3 3 4 3 4 4 3 3 4 3 3 3 3 3 2 3 2 3 2 2 3 2 3 3 3 3 2 2 3 3 2 C B 0.36 0.35 0.27 0.3 0.27 0.29 0.27 0.28 0.29 0.29 0.26 0.26 0.24 0.25 0.25 0.48 0.23 0.25 0.46 0.47 0.18 0.46 0.49 0.46 0.49 0.43 0.49 0.49 0.42 0.49 0.4 0.41 0.41 0.39 0.49 0.49 0.38 0.36 0.49 6 5 5 5 9 1 3 1 1 6 2 4 3 5 3 3 3 6 6 3 5 3 7 7 8 9 A 11 11 12 12 12 10 12 12 12 12 13 13 12 ...... 13 25 25 50 25 50 25 25 25 ...... 4 8 4 8 4 4 12 44 40 28 32 36 40 28 12 40 36 52 56 28 12 24 44 60 ...... 5 5 5 5 11 11 11 11 11 11 42 16 26 21 42 ...... 10 20 20 20 20 ...... 9 50 20 70 10 10 20 50 40 ...... 8 11 11 11 11 11 22 33 33 33 22 22 33 22 67 33 ...... 7 33 33 33 ...... 6 40 30 10 20 10 60 40 40 30 30 ...... 5 6 6 6 31 38 13 25 38 88 31 ...... 4 ...... 3 8 8 25 25 42 17 33 42 25 33 42 17 ...... 2 20 20 40 ...... 1 38 31 23 23 15 62 canescens L. L. L. ssp. trichomanes L. L. ssp. serpyllifolia (L.) Loret & Barrandon Jacq. Moench L. ssp. alpestris Ascherson & Graebner L. Medicus L. Krasan Vis. L. L. ssp. symphyandra (Murb.) Hayek Biscutella laevigata Senecio doronicum (L.) L. ssp. doronicum Senecio doronicum Euphorbia cyparissias L. Anthericum ramosum L. Filipendula vulgaris Hippocrepis comosa L. Hippocrepis Asplenium trichomanes Gymnadenia conopsea (L.) R. Br. Amelanchier ovalis Potentilla australis Rhinanthus aristatus Celak. Asplenium ruta-muraria L. ssp. Buphthalmum salicifolium Sanguisorba minor Scop. ssp. muricata Briq. Trifolium montanum L. Trifolium Briza media L. ssp. Thalictrum minus L. ssp. Eryngium amethystinum (Vis.) Briq. (Vis.) L. ssp. subcrenata Stachys recta Anthyllis vulneraria Pinus mugo Turra Saxifraga tridactylites Inula hirta L. Arenaria serpyllifolia Arenaria Crepis chondrilloides Crepis Sedum acre L. Sedum acre Gentiana utriculosa Juniperus communis L. Euphrasia dinarica (G. Beck) Murb. Gentiana lutea Silene saxifraga Trifolium alpestre L. alpestre Trifolium Erysimum sylvestre (Crantz) Scop. ssp. sylvestre Erysimum sylvestre Thymus longicaulis C. Presl Thesium divaricatum Jan ex Mert. & Koch Armeria canescens (Host) Boiss. ssp. Dianthus integer Pimpinella saxifraga Scorzonera austriaca Willd.

75 Hacquetia 13/1 • 2014, 57–77 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 C B 0.47 0.36 0.47 0.35 0.46 0.35 0.46 0.46 0.35 0.46 0.35 0.35 0.35 0.35 0.44 0.35 0.44 0.35 0.44 0.35 0.35 0.44 0.34 0.41 0.43 0.43 0.4 0.34 0.4 0.39 0.34 0.38 0.32 0.38 0.3 0.37 0.28 0.27 0.37 5 1 3 1 6 6 6 6 9 6 6 6 6 5 9 5 7 5 6 5 6 1 8 1 3 8 5 A 11 11 12 10 10 10 13 12 12 12 12 12 ...... 50 13 25 ...... 8 8 8 8 36 12 32 32 16 20 24 16 20 20 64 ...... 5 11 11 32 42 53 16 21 ...... 10 20 20 20 20 20 20 ...... 9 10 10 30 10 30 20 10 30 ...... 8 11 11 11 22 22 ...... 7 33 33 33 ...... 6 10 20 30 30 20 20 20 30 10 20 20 30 30 10 20 ...... 5 6 56 25 25 19 ...... 4 ...... 3 8 8 8 8 25 17 17 17 ...... 2 ...... 1 31 15 23 23 31 (Chodat) Hayek Šilić (Boiss.) Maire & Petitmengin (L.) Beauv. ssp. pinnatum (L.) Beauv. (Vis.) J. Holub (Vis.) L. ssp. kummeri (G. Beck) Markgr.-Dannenb. Vill. L. ssp. liburnica Jan Bernh. ex Koch L. ssp. austriaca croatica Reichenb. ssp. croatica Sternb. & Hoppe Heuffel L. ssp. montanum (Fries) Holub Jacq. L. (Koch) P.W. Ball ssp. gouanii (Koch) P.W. Gouan L. Muscari botryoides (L.) Miller Frangula rupestris (Scop.) Schur Stachys recta agg. Stachys recta Campanula scheuchzeri Vill. Galium verum L. ssp. Senecio papposus (Reichenb.) Less. ssp. Veronica austriaca Veronica Luzula campestris (L.) DC. Scabiosa columbaria L. ssp. portae (A. Kerner ex Huter) Hayek Gentianella crispata Polygala alpestris Brachypodium pinnatum Festuca amethystina Arabis scopoliana Boiss. (Ten.) Lüdi Primula veris L. ssp. columnae (Ten.) Linum narbonense L. Rhamnus alpinus L. ssp. fallax Asperula aristata L. ssp. scabra (J. & C. Presl) Nyman Poa badensis Haenke ex Willd. Ruta graveolens Iris reichenbachii Iris reichenbachii Mercurialis ovata Mercurialis Seseli elatum Lilium carniolicum Scabiosa columbaria L. ssp. Thymus striatus Vahl Markgraf Cytisus pseudoprocumbens (Guss.) C. Koch Cotoneaster nebrodensis Allium senescens Scorzonera villosa Scop. Peltaria alliacea Galium mollugo L. Valeriana montana L. Valeriana Vicia incana Vicia Anthyllis vulneraria Euphorbia fragifera Satureja subspicata Satureja Viola hirta Viola Galium anisophyllon

76 Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 C B 0.27 0.27 0.25 0.24 0.24 0.22 0.22 0.23 0.2 0.17 0.16 0.16 0.16 0.16 0.11 0.16 4 5 5 8 8 6 1 A 11 11 12 12 12 12 12 12 12 ...... 13 . . . . 8 8 4 8 8 8 4 8 12 12 28 12 12 ...... 5 5 11 11 11 26 32 ...... 10 ...... 9 ...... 8 11 11 ...... 7 ...... 6 10 ...... 5 6 6 6 13 13 ...... 4 ...... 3 8 8 8 8 ...... 2 ...... 1 8 8 (Schott ex Roemer & Schultes) Murb. Medicus (Tomm. ex Vis.) K. Richt. Vis.) ex (Tomm. Witasek L. Vest ssp. barrelieri L. Turra Veronica spicata Veronica Hyssopus officinalis L. ssp. aristatus (Godron) Briq. Hypochoeris maculata Campanula justiniana Vincetoxicum hirundinaria Vincetoxicum Plantago media Linum catharticum L. Chamaecytisus hirsutus (L.) Link Sorbus aria (L.) Crantz Iris pallida ssp. illyrica L. Anemone nemorosa Artemisia alba Euphorbia verrucosa L. Hypericum perforatum L. Saxifraga crustata Thlaspi praecox Wulfen