HACQUETIA DOI: 10.2478/v10028-009-0004-0 A REVIEW OF POTENTILLO TERNATAE - NARDION STRICTAE ALLIANCE Nikolay I. VELEV1 & Iva I. APOSTOLOVA1 Abstract A classification of the Nardus stricta dominated communities in the Balkan Range is presented. Two associations are identified: Nardetum strictae and Campanulo alpinae - Nardetum strictae nom. nov. The alliance Potentillo ternatae - Nardion strictae is typified and some comments on its distribution range and syntaxonomical affiliation to the higher units are presented. A synopsis is included of all available associations referred to Potentillo ternatae - Nardion strictae. Key words: grasslands, nomen novum, nomenclature, numerical methods, syntaxonomy. Izvleček V članku je predstavljena klasifikacija združb z dominantno vrsto Nardus stricta na Balkanu. Prikazali smo dve asociaciji: Nardetum strictae and Campanulo alpinae - Nardetum strictae nom. nov. Zvezo Potentillo ternatae - Nardion strictae smo tipizirali in predstavili njeno razširjenost in sintaksonomsko uvrstitev v višje enote. Prikazan je sinopsis vseh opisanih asociacij, ki jih uvrščamo v zvezo Potentillo ternatae - Nardion strictae. Ključne besede: travniki, nomen novum, nomenklatura, numerične metode, sintaksonomija. INTRODUCTION The Nardus-rich communities are wide spread in the Bulgarian mountains. They are almost entirely of secondary origin and cover approximately 50000 ha (Velchev et al. 1989). A significant part of them is presented in the Balkan Range as a result of hundreds of years of extensive stock breeding. The syntaxonomical diversity of the mat-grass communities in the Bulgarian part of the Balkan Range has been studied so far following the dominant approach (see Bondev 1966; Ganchev & Kochev 1969; Meshinev et al. 2000). The same vegetation type, following the Braun-Blanquet approach, was described for Mt. Pirin (Simon 1958), Mt. Rila (Roussakova 2000) and reported for the Balkan Range, the Rhodopes and Mt. Sredna Gora (Meshinev et al. 2005). The successional changes of Nardus stricta communities in the Central Balkan Range were commented by Velev & Apostolova (2008). The authors refer communities distributed below the timberline to Nardion strictae Br.-Bl. 1926, and those occupying higher elevations (above the timberline) to Potentillo ternatae - Nardion strictae Simon 1958. Similarly, for the Serbian territory, Kojić et al. (1998, 2004) refer the colline mat-grass pastures to Nardion strictae while the mat-grass pastures in the high mountains are considered as part of Potentillo ternatae - Nardion strictae, Jasion orbiculatae Lakušić 1966 and Poion violaceae Horvat 1937. According to Krahulec (1985) Potentillo ter-natae - Nardion strictae should be considered as a Carpathian-Balkanic alliance, characteristic for the vegetation belt above the timberline. On the other hand, the associations recognized so far as being affiliated to this alliance represent an extremely broad variety of ecological and floristic peculiarities. i Department of Phytocoenology and Ecology, Institute of Botany, Bulgarian Academy of Sciences, Acad. G. Bonchev St., bl. 23, 1113 Sofia, Bulgaria, e-mail: nvelev@bio.bas.bg; iva@bio.bas.bg. 1 Hacqüetia The review of the literature revealed significant differences in authors' concepts regarding relation of the alliance to the higher units (orders and classes). The wealth of information gave us impetus to gather all available in the literature associations and to present the current overview on the Potentillo ternatae - Nardion strictae alliance. The aims of the present study are (i) to identify the syntaxonomy of Nardus stricta communities in the Balkan Range, and (ii) to review the Potentillo ternatae - Nardion strictae alliance. 2. MATERIAL AND METHODS A data set of 336 relevés containing mat-grass were analysed. They are recorded for the Balkan Range (Figure 1) following the dominant approach (Bondev 1966, Ganchev & Kochev 1969, Meshinev et al. 2000) within an altitudinal range between 1400 and 2200 m. Jp ROMANIA --' S çv M V \ R W \ A Varna• » j! V V Black Sea F.Y.R. \ O.M. .J___ i C "" '"-'"v ...... GREECE TURKEY Figure 1: Map of the sampled localities. Slika 1: Zemljevid proučevanega območja. Each relevé contains a full list of species excluding the mosses in some descriptions. The sample plots are 100 m2. The original quantitative estimations are transformed to the old Braun-Blanquet scale following Meshinev & Apostolova (2002). The relevés are stored in the TURBOVEG data base (Hennekens & Schaminèe 2001). They are exported into JUICE software (Tichy 2002) for further analysis. Subsequently unsupervised classification (TWINSPAN - Hill 1979) and supervised classification using the Cocktail method (Bruelheide 1995, 2000) are applied. Two sociological groups (cf. Bruelheide 2000; Chytry 2007) were identified within the Cocktail method: (1) setting as an initial species Nardus stricta and rep- resenting class Calluno - Ulicetea, and (2) setting as an initial species Juncus trifidus and representing class Juncetea trifidi. Each group includes five species. The classification of the relevés is based on the presence of at least three of these species. We used the logical operator AND (Bruelheide 1997) to compose formal definitions for the two associations for the Balkan Range. All available literature is referred to in order to elaborate a full list of associations of Potentillo ternatae - Nardion strictae alliance. The syntaxonomical interpretation follows the floristic and ecological principles (Braun-Blan-quet 1965; Westhoff & der Maarel 1973; Mueller-Dombois & Ellenberg 1974). The syntaxonomical nomenclature of the revisions is in accordance with the International Code of Phytosociological Nomenclature (Weber et al. 2000) (in the text ICPN). The nomenclature of the species follows Delipavlov et al. (2003), Kozhuharov (1992), Natcheva & Ganeva (2005) and Wirth (1995), while the floristic elements are used after Oberdorfer (1994) and Assyov & Petrova (2006). 3. RESULTS AND DISCUSSION The unsupervised classification of the relevés reveals that the mat-grass communities in the Balkan Range are divided into two distinct groups in relation to altitude and referring to different alliances. A part of the studied relevés is located within the subalpine belt and another part within the alpine belt. Although there are common species presented, the groups are well separated by a number of diagnostic species. The same groups are established as a result of the implementation of supervised classification and the Cocktail method (Bruelheide 1995, 2000) as well. Two associations are clearly identified: Nardetum strictae and Campanulo alpinae - Nardetum strictae (Table 1). The remaining relevés are spread among other cluster groups and are not considered in the subsequent analysis. Association Nardetum strictae Grebenščikov 1950 Sociological group Nardus stricta: Nardus stricta, Festuca rubra, Agrostis capillaris, Hieracium hop-peanum, Verbascum longifolium. Formal definition for the Balkan Range: Group 2 NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance Nardus stricta AND Nardus stricta cover > 25 %. The identified Nardetum strictae association of the Balkan Range is developed at an altitudi-nal range of 1400 - 2080 m. It occupies formerly intensive pasturelands, which are currently extensively used. The communities are distributed over all the range of the studied area (Fig. 1). They are localized on ridges or slightly inclined slopes, creating conditions for preservation of the relatively well developed soil layer. The rocks are silicate. The association was described by Grebenščikov for Mt. Stara Planina (Serbia) (Kojić et al. 1995). It is confirmed for the same locality also by Mišić et al. (1978) and Kojić et al. (1998). Janković (1982) and Kojić et al. (1995, 1998) stated that Nardus stricta dominated pastures (recognized as Nardetum strictae sensu lato) are a complex of many associations with different ecological characteristics. According to Kojić et al. (1998) the complex Nardetum strictae sensu lato is divided into 18 associations, one of which is ass. Narde-tum strictae Grebenščikov 1950. This association is treated as a separate syntaxon also by Kojić et al. (2004) and Lakušić et al. (2005). Nevertheless, this association is accepted as having a broad ecological amplitude (Mišić et al. 1978, Kojić et al. 1995) but never interpreted as a synonym of the complex Nardetum strictae sensu lato. Kojić et al. (1995) mentioned that many authors misjudge the described association for the Stara Planina association. According to ICPN rules such associations are pseudonyms and in this case should be considered as Nardetum strictae auct. non Grebenščikov 1950. Association Nardetum strictae Greb. 1950 ecologically is very close to that described for Romania by Simon (1966) as association Violo declinatae - Nardetum strictae. The difference between these associations is that Viola declinata Waldst. & Kit. is missing in Nardetum strictae. This species does not occur in Serbia and Bulgaria (see Diklić 1972; Delipavlov 1979; Assyov & Petrova 2006) where the taxonomically related species Viola dacica Borb. (= V. declinata Waldst. & Kit. ssp. bulgarica Form.) was recorded. According to Grigoriu & Alda (2004) the Violo declinatae - Nardetum strictae association unifies the mat-grass communities in Eastern Europe. Such a statement could be proved by the fact that numerous relevés related to the Violo declinatae - Nardetum strictae association (Simon 1966; Mihäilescu 2001; Grigoriu & Alda 2004), including its synonyms (e.g. Nardetum alpigenum austro - carpaticum Borza 1959 and Nardo - Vaccinietum Resm. 1970) do not contain V. declinata even in their original records. Association Campanulo alpinae - Nardetum strictae (Simon 1958) nom. nov. hoc loco Nomenclatural type: Simon (1958: Table 8, rel. 3), lectotypus hoc loco designatus Homotypical synonym: Nardetum alpinum moesia-cum Simon 1958 nomen illegitimum (Art. 34a). Sociological group Juncus trifidus: Juncus trifi-dus, Campanula alpina, Festuca airoides, Agrostis rupestris, Sesleria comosa. Formal definition for the Balkan Range: Group Juncus trifidus AND Nardus stricta cover > 5 %. The association was described by Simon for Mt. Pirin (Simon 1958: 179-180, Table 8). The originally recorded relevés of the association are included in Table 1 (relevés 52 - 57, and is typified by relevé 54). The confirmation of this syntaxon 50 years after its establishment reveals its particular character and does not provide reasons for it to be a nomen dubium (Art. 37); followng the ICPN rules a nomen novum should be given. According to ICPN the new name is typified by the typus of the rejected one. We selected the typus following the recommendations of ICPN where both Campanula alpina and Nardus stricta are presented. Nardus stricta dominates (projection coverage > 25 %) and remains in the name (Art. 3k, 10b, 29b). The species Campanula alpina is well presented in the relevés under this association simultaneously from Mt. Pirin and Mt. Balkan (see Table 1). This species indicates the alpine character of the association and is mentioned as diagnostic for alliance Seslerion comosae by Horvat, Paw-lowski & Walas (1938), Simon (1958) and Horvat et al. (1974). According to Cheshmedzhiev (2003) in Bulgaria only Campanula alpina Jacq. ssp. or-belica (Pančič) Urum. is presented within an alti-tudinal range from 1700 to 2900 m a.s.l. In the studied area the association is distributed occasionally within the alpine belt between 1950 and 2200 m altitiude. The communities are localized mainly on ridges or highly inclined slopes where the soils are slightly eroded. The rocks are silicate. Considering the floristic composition of the association (see Table 1) we definitely support Simon's (1958) opinion that this type of vegetation is derived from Agrostidi 3 Hacqüetia - Seslerietum comosae Horvat, Pawlowski & Walas 1938, or from Seslerietalia comosae meadows in the way of pasture digression. Simon (1958) includes the association in Po-tentillo ternatae - Nardion strictae (Nardetalia, Nardo-Callunetea). At the same time, Simon (1958 in Table 8) points out diagnostic species for Ses-lerion comosae, Poion violaceae and Seslerietalia comosae. The main difference between Seslerion comosae and Potentillo ternatae - Nardion strictae after the cited reference is high abundance of Nar-dus stricta. The species Potentilla ternata is pointed out in this source as diagnostic also for Seslerieta-lia comosae. We do not support the decision taken by the author regarding association affiliation to higher syntaxa, because of lacking in diagnostic species both for the order and the class (see Table 1, rel. 52-57). Following the rules of flo-ristic resemblance (Westhoff & van der Maarel 1973), and taking into account the presence of several diagnostic species (Table 1), we refer the Campanulo alpinae - Nardetum strictae association to Seslerion comosae alliance (Seslerietalia comosae, Juncetea trifidi). The dominant position of Nardus stricta in the communities is not sufficient reason for relating an association to Nardetalia strictae and Calluno Ulicetea. In maintainance of such an opinion many authors (Bo§caiu 1971; Coldea 1990; Krahulec et al. 1996; Roussakova 2000; Devillers & Devillers-Terschuren 2001; Rodwell et al. 2002; Chytry & Tichy 2003; Koči 2007, etc.) refer the mat-grass communities which contain Arctic-Alpine species to the Juncetea trifidi class. Alliance Potentillo ternatae - Nardion strictae Simon 1958 Nomenclatural type: Violo declinatae - Nardetum strictae Simon 1966, [Simon (1966): p. 260, Table 2], neotypus hoc loco designatus. This alliance is described by Simon (1958) from Mt. Pirin (Bulgaria). It is established by a single association - Nardetum alpinum moesiacum Simon 1958, which shares the same diagnostic species. According to Def. VIII (ICPN) this association should be considered as holotypus, being the only one described for the alliance. In this particular case it is not applicable because the association belongs to Seslerion comosae as was already pointed out (see Table 1). As a neotypus we suggest Violo declinatae - Nardetum strictae association described from Mt. Bihar (Romania). Simon (1966) did not point out its affiliation to higher units when he published the association. Later on, many Romanian authors (Coldea 1987; Ivan et al. 1993; Sanda et al. 1997; Mihäilescu 2001; Pop et al. 2002; Niculescu 2004; Grigoriu & Alda 2004; Sarbu et al. 2004, etc.) included the association within Potentillo ternatae - Nardion strictae. Simon (1958) refers Potentillo ternatae - Nardion strictae to Nardo - Callunetea, and many other researchers follow this concept, as for example Horvat et al. (1974), Dihoru (1975), Mišić et al. (1978), Pop et al. (1988, 2002), Sanda et al. (1997), Kojić et al. (1997, 1998, 2004), Mihäilescu (2001), Grigoriu & Alda (2004), Niculescu (2004), Sarbu et al. (2004). Some authors, however, refer it to Juncetea trifidi (cf. Bo§caiu 1971; Coldea 1990; Roussakova 2000; Devillers & Devillers-Tersch-uren 2001; Rodwell et al. 2002; Chifu et al. 2006). The problem with the syntaxonomic position of the alliance comes after its distribution close to the alpine belt and thus containing numerous alpine species, such as Agrosris rupestris All., Alo-pecurus gerardii Vill., Dianthus microlepis Boiss, Euphrasia minima Jacq. ex DC., Festuca riloensis (Hack. ex Hajek) Markgr.-Dannb., Juncus trifidus L., Scleranthus neglectus Rochel ex Baumg., Sesle-ria comosa Velen. etc (see Table 1). Particularly in Bulgaria the alpine belt is situated between 2400 - 2925m and very rarely, depending on the local climatic peculiarities, could start from 2000 m altitude and even lower (Velchev 1984, 2002; Bondev 1991). Simon (1958) describes the alliance at the range of 2100 - 2400 m a.s.l. and the relevés contain species from different altitudinal belts. The species composition causes ambiguous interpretation of the alliance by different authors. Sarbu et al. (2004) report Potentillo ternatae - Nardion strictae at 1450 - 1900 m altitude for Romania. They describe it as „acidic oligo-trophic subalpine pastures". In Ukraine it occupies altitudes of 1200 - 2061 m (Malinovsky & Kricsfalusy 2000; Prots et al. 2008). Rodwell et al. (2002) determine Potentillo ternatae - Nardion strictae as „ Mat-grass swards of the alpine belt in the mountains of the eastern Balkans". The information included in Table 2 reveals relatively broad altitudinal range (500-2400 m) of the described associations within the Potentillo ternatae - Nardion strictae alliance. In fact, associations from different vegetation belts are related to the same alliance and thus combine ecologically different habitats. This contradicts to the Braun-Blanquet statement that the alliance unifies „the most closely related associations" (Braun-Blan-quet 1965: 363). 4 NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance The following 22 associations are reffered to Potentillo ternatae - Nardion strictae: In Romania: Nardetum strictae subalpinum Domin 1931, Krajina 1933; Campanulo serratae - Festucetum ovinae (Morariu 1942) Coldea 1987; Festucetum ovinae Morariu 1943; Nardo - Callune-tum vulgaris Šmarda 1953, Hrynnc. 1959, Csürös 1964; Poëtum mediae Csürös et al. 1956; Scorzonero roseae - Festucetum nigricantis (Pu§c. et al. 1956) Coldea 1987; Nardetum alpigenum austro - car-paticum Borza 1959; Festuco - Nardetum strictae montanum Csürös & Resm. 1960; Potentillo - Festucetum ovinae Resm. (1965) 1969, 1970; Agrosteto - Callunetum vulgaris Resm. & Csürös 1966; Violo declinatae - Nardetum strictae Simon 1966; Campanulo abietunae - Nardo - Festucetum commutatae Bo§caiu 1971; Antennario - Festucetum sudeticae Dihoru 1975; Carici - Nardetum strictae (Resm. 1984) Resm. & Pop 1986; Hieracio pilosellae - Nardetum strictae Pop et al. 1988 (Resmeritä & Csürös 1966; Bo§caiu 1971; Dihoru 1975; Coldea 1987, 1990; Coldea & Pop 1988; Pop et al. 1988; Ivan et al. 1993; Sanda et al. 1997; Mihäilescu 2001; Pop et al. 2002; Niculescu 2004; Grigoriu & Alda 2004; Chifu et al. 2006). In former Yugoslavia: Nardetum strictae Greb. 1950; Trifolio pallescens - Nardetum strictae Blečić & Tatić 1964; Hygronardetum strictae Pu§c.-Soroc. 1965; Succiso - Nardetum strictae Stanković-Tomić 1969; Festuco halleri - Nardetum strictae Rajev. 1974; Festuco fallaci - Nardetum strictae Rajev. 1974; Helianthemo - Nardetum strictae Rajev. 1974; Festuco nigrescenti - Nardetum strictae Rexh. & Ranđ. 1980; Festuco - Nardetum strictae subalpinum Rexh. & Ranđ. 1980; Carici oederi - Nardetum strictae Petk. 1981; Ophioslosso - Nardetum strictae Gajić 1989 (Horvat et al. 1974; Mišić et al. 1978; Kojić et al. 1998, 2004). In Bulgaria: Nardetum strictae Greb. 1950; Carici - Festucetum microphyllae Roussakova 2000 (Roussakova 2000; Velev & Apostolova 2008). In Ukraine: Soldanello - Nardetum Malinovsky & Kricsfalusy 2000 (Malinovsky & Kricsfalusy 2000). According to some authors, several of the associations listed above are referred to other alliances, as for example: Nardetum strictae Greb. 1950, Hygronardetum strictae Pu§c.-Soroc. 1965 to the Nardion strictae alliance (Kojić et al. 1998; Puzović et al. 2006; Obratov-Petkovic et al. 2007); Nardetum strictae Greb. 1950 to the Poion viola-ceae alliance (Kojić et al. 2004); Antennario - Festucetum sudeticae Dihoru 1975 to the Caricion curvulae Br.-Bl. 1925 alliance (San-da 1997); Nardo - Callunetum vulgaris Šmarda 1953, Hrynnc. 1959, Csürös 1964 to the Genistion Böch. 1943 alliance (Sanda 1997); Violo declinatae - Nardetum strictae Simon 1966 to the Nardion strictae alliance (Chifu et al. 2006). In the literature as diagnostic for Potentillo ternatae - Nardion strictae are listed the following species: Agrostis rupestris All., Alchemillaflabellata Buser, A. glaucescens Wallr., A. monticola Opiz, A. xanthochlora Rothm., Antennaria dioica (L.) Gaert-ner, Anthoxanthum alpinum Ä.Löve & D.Löve, Bellardiochloa violacea (Bell.) Chiov., Botrychium lunaria (L.) Sw., Bruckenthalia spiculifolia Rchb., Campanula excisa Presl., C. kladniana (Schur) Witasek, C. patula L. ssp. abietina (Griseb.) So-monkai, C. rotundifolia L.,C. serrata (Kit.) Hen-drych, Carex bulgarica (Domin) Lazare, Carlina acaulis L., Centaurea nervosa Willd., Cerastum fontanum Baumg., Crocus neapolitanus Mordant & Loisel., Diphasium alpinum (L.) Rothm., Euphrasia minima Lam. & DC., Festuca airoides Lam., F. nigrescens Lam., F. ovina L., Gentiana acaulis L., Gentianella lutescens (Velen.) J. Holub, Geum montanum L., Helictotrichon versicolor (Vill.) Pilger., Hieracium alpinum L., H. aurantiacum L., H. hop-peanum Schultes, H. lactucella Wallr., Homogyne alpina (L.) Cass., Hypericum maculatum Crantz, Hypochaeris uniflora Vill., Leontodon rilaensis Hayek, Ligusticum mutellina (L.) Crantz, Luzula sudetica (Willd.) DC., Nardusstricta L., Phleum alpinum L., Plantago atrata Hoppe, P. gentianoides Sibth. & Sm., Poa alpina L., P. media Schur, Po-tentilla aurea L., P. erecta (L.) Rausch., P. ternata C. Koch., Primula elatior (L.) Hill., Pseudorchis albida (L.) A. & D. Löve, Scorzonerapurpurea L. ssp. rosea (Waldst. & Kit.) Nym., Solidago virgaurea L. ssp alpestris (W. & K.) Gaud., Thesium alpinum L., Thlaspi dacicum Heuffel, Thymus alpestris Tausch, T. pulegioides L., T. vandasii Velen., Veronica alpina L., V. officinalis L., Viola canina L., V. dacica Borbas and V. declinata Waldst. & Kit. (Simon 1958; Bo§caiu 1971; Dihoru 1975; Coldea 1987, 1990; Coldea & Pop 1988; Sanda et al. 1988, 1997; Ivan et al. 1993; Roussakova 2000; Malinovsky & Kric-sfalusy 2000; Mihäilescu 2001; Niculescu 2004; Grigoriu & Alda 2004; Sarbu et al. 2004; Petrik et al. 2005; Chifu et al. 2006). Some species, as for example Homogyne alpina, Festuca airoides, Nardus stricta, Poa media, Po- 5 Hacqüetia tentilla ternata, are pointed out also as diagnostic for Seslerietalia comosae or Juncetea trifidi (Simon 1958; Mucina et al. 1990; Roussakova 2000; Chy-try & Tichy 2003; Koči 2007). Horvat et al. (1937) point out Potentilla ternata as diagnostic for the alpine vegetation included in Caricetalia curvulae Br.-Bl. 1926. At the same time, species as Veronica officinalis, Cerastumfontanum, Thymus pulegioides, Hieracium hoppeanum and Potentilla erecta have broad distribution and practically are not considered characteristic for the sub-alpine and alpine vegetation. The proximity of alpine vegetation much influences Potentillo ternatae - Nardion strictae. Floristic-geographical analysis of the diagnostic species shows that 49 % have Alpine and Arctic-Alpine distribution. Another 18 % of the species represent the sub-Atlantic floristic element. Car-pathian-Balkanic flora is presented by 14 % of diagnostic species and could be an argument for distinction of the alliance. The list of diagnostic species reveals significant resemblance between Potentillo ternatae - Nardion strictae and Nardion strictae. Mention should also be made of Nardo strictae - Agrostion tenuis Sillinger 1933 which is presented in the Krkonoše and the West Carpathian Mts (Krahulec 1990; Krahulec et al. 1996). The species composition of the relevés from the Krkonoše Mts is similar to our data. It could be suggested that one large alliance (Nardion ?) actually includes geographically differentiated suballiances like Potentillo ternatae - Nardion strictae and Nardo strictae - Agrostion tenuis. But large data set analysis is needed to prove this hypothesis. 4. CONCLUSIONS The mat-grass communities in Bulgaria, as in many European countries, are most often a sequence of long-time intensive grazing and trampling. The pastures are distributed in the mountains, mostly above the timberline and occur in different altitudinal vegetation belts. In the alpine vegetation, the communities characterized by significant Nardus stricta abundance are related to Juncetea trifidi class, which is supported by the presence of many preserved diagnostic species (e.g. Juncus trifidus, Veronica bellidioides, Festuca airoides, Campanula alpina, Agrostis rup-estris). At lower altitudes, in the subalpine belt, the grazing has been more intensive. These ter- ritories are covered by huge mountain pastures, quite uniform due to the strong dominance of the mat-grass. In the Balkan Range these communities are referred to Potentillo ternatae - Nardion strictae. We typified the Potentillo ternatae - Nardi-on strictae alliance by Violo declinatae - Nardetum strictae association. The Potentillo ternatae - Nardion strictae vegetation includes many character species (e.g. Potentilla erecta, Agrostis capillaris, Hypericum maculatum, Festuca rubra) for Calluno - Ulicetea and should be subordinated to this class. The high abundance of Nardus stricta in a vegetation type does not necessarily refer this vegetation to Calluno - Ulicetea. The mat-grass pastures developed on places of former Juncetea trifidi syntaxa remain syntaxonomically in the same class. It might be predicted that a decrease in the grazing and trampling intensity, which is ongoing currently at many places in the high mountains, will reduce the mat-grass abundance and will result in demutation to the primary vegetation communities. 5. SYNTAXONOMICAL SYNOPSIS: Class Juncetea trifidi Hadač in Klika & Hadač 1944 [Syn.: Caricetea curvulae Br.-Bl. 1948)] Order Seslerietalia comosae (Simon 1958) Lakušić 1966 Alliance Seslerion comosae Horvat 1935 em. Laku-šić 1966 Ass. Campanulo alpinae - Nardetum strictae (Simon 1958) nom. nov. Class Calluno-Ulicetea Br.-Bl. & Tüxen ex Klika & Hadač 1944 [Syn.: Nardo - Callunetea Preising 1949] Order Nardetalia strictae Preising 1949 Alliance Potentillo ternatae - Nardion strictae Simon 1958 Ass. Nardetum strictae subalpinum Domin 1931, Krajina 1933 nom. illeg. (Art. 34a) [Syn.: Nardetum strictae montanum Sillinger 1933] Ass. Nardetum strictae Greb. 1950 Ass. Poëtum mediae Csürös et al. 1956 Ass. Nardo - Callunetum vulgaris Šmarda 1953, Hrynnc. 1959, Csürös 1964 [Syn.: Vaccinio - Callunetum vulgaris Bük. 1942; Agrosteto -Callunetum vulgaris Resm. & Csürös 1966; Arnica montana - Calluna vulgaris Ghi§a et al. 1970; Festuceto rubrae - Callunetum vulgaris Resm. 1970] 6 NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance Ass. Trifoliopallescens - Nardetum strictae Blečić & Tatić 1964 Ass. Hygronardetum strictae Pu§c.-Soroc. 1965 nom. illeg. (Art. 34a) Ass. Violo declinatae - Nardetum strictae Simon 1966 [Syn.: Nardetum strictae alpinum Borza 1934 (Art. 34a) p. p.; Nardetum strictae alpinum Ghi§a 1940 (Art. 34a) p. p.; Nardetum strictae alpinum Csürös 1955 (Art. 34a) p. p.; Nardetum strictae alpinum Pu§caru et al. 1956 (Art. 34a); Nardetum alpigenum austro - carpaticum Borza 1959 (Art. 34a); Nardetum strictae subalpinum Buia et al. 1962 (Art. 34a); Nardetum strictae alpinum Buia et al. 1962 (Art. 34a); Nardetum strictae alpinum Buia 1963 (Art. 34a) p. p.; Nardetum strictae montanum Resmeritä & Csürös 1963 nom. illeg. (Art. 34a); Agrosteto-Festucetum rubrae montanum nardosum Resm. & Csürös 1966 nom. illeg. (Art. 34a); Nardetum strictae alpinum Beldie 1967 (Art. 34a) p. p.; Nardo - Vaccinietum Resm. 1970; Nardo-Fes-tucetum tenuifoliae Buiculescu 1971; Festuco rubrae - Agrostetum capillaris Horvat 1951 var. Nardus stricta Danciu 1974; Nardo-Fes-tucetum tenuifoliae Buiculescu 1971; Festuco rubrae - Agrostetum capillaris Horvat 1951 subass. nardetosum strictae Pop 1976] Ass. Succiso - Nardetum strictae Stanković-Tomić 1969 Ass. Festuco fallaci - Nardetum strictae Rajev. 1974 Ass. Festuco halleri - Nardetum strictae Rajev. 1974 Ass. Helianthemo - Nardetum strictae Rajev. 1974 Ass. Antennario - Festucetum sudeticae Dihoru 1975 [Syn.: Festucetum supinae Domin 1933; Festucetum supinae subalpinum Sillinger 1933 p. p.; Festuca supina - Nardus stricta §erbänescu 1939; Festuca supina Pu§caru et al. 1959; Festuca supina - Deschampsia flexuosa - Vaccinium myrtillus Pu§caru et al. 1959; Festuca supina - Festuca rubra fal-lax Pu§caru et al. 1959; Festucetum supinae Vicol et al. 1971; Potentillo chrysocraspedae -Festucetum airoidis Bo§caiu 1971; Potentillo ternatae - Festucetum supinae Bo§caiu 1971] Ass. Festuco nigrescenti - Nardetum strictae Rexh. & Rand. 1980 Ass. Festuco - Nardetum strictae subalpinum Rexh. & Rand. 1980 nom. illeg. (Art. 34a) Ass. Carici oederi - Nardetum strictae Petk. 1981 Ass. Carici - Nardetum strictae (Resm. 1984) Resm. & Pop 1986 [Syn.: Hygronardetum strictae Borza 1934, Hygronardetum strictae Pu§c. et al. 1956; Hygronardetum subalpinum Resm. & Csürös (1960) 1963; Hygronardetum strictae alpinum Buia et al. 1962, Hygronardetum strictae alpinum Resm. & Csürös 1963; Hygronardetum montanum Resm. & Csürös 1963, Hygronardetum strictae montanum Buia 1963] Ass. Campanulo serratae - Festucetum ovinae (Morariu 1943) Coldea 1987 [Syn.: Festucetum ovinae Morariu 1943; Festuceto ovinae - Potentilletum aureae Resm. 1965; Potentillo - Festucetum ovinae Resm. 1965, 1986; Potentillo aureae - Festucetum ovinae Resm. 1969, 1970, Resm. & Pop 1984; Violo declinatae - Nardetum strictae Simon 1966 subass. festucetosum ovinae (Morariu 1943) Chifu et al. 2006] Ass. Scorzonero roseae - Festucetum nigricantis (Pu§c. et al. 1956) Coldea 1987 [Syn.: Festucetum rubraefallax Pu§c. et al. 1956; Nardo - Festucetum rubrae fallax Pu§c. et al. 1959; Festuca supina - Festuca rubra Pu§c. et al. 1959; Festucetum rubrae montanum Csürös & Resm. 1960; Festucetum rubrae subalpinum siliciculum Csürös & Resm. 1960; Festuco - Nardetum strictae montanum Csürös & Resm. 1960; Festuco rubrae - Nardetum Csürös & Resm. 1960; Festuco - Alchemil-letum vulgaris Csürös & Resm. 1960; Campanulo abietinae - Nardo - Festucetum com-mutatae Bo§caiu 1971 nom. nud. (Art. 2a)] Ass. Hieracio polosellae - Nardetum strictae Pop et al. 1988 pro syn., nom. nud. (Art. 2b, 3a, Recomm. 46f) [Syn.: Xeronardetum Soo 1931; Xeronardetum montanum Resm. & Csürös 1963] Ass. Ophioslosso - Nardetum strictae Gajić 1989 Ass. Carici - Festucetum microphyllae Roussa-kova 2000 Ass. Soldanello - Nardetum Malinovsky & Krics-falusy 2000 ACKNOWLEDGEMENTS We thank the two anonymous reviewers for the helpful comments and critical remarks on a previous version of the manuscript. 7 Hacqüetia 6. REFERENCES Assyov, B. & Petrova, A. (eds): 2006. Conspectus of the Bulgarian Vascular Flora. Distribution Maps and Floristic Elements. Ed. 3. BBF, Sofia, 453 pp. Beldie, A. 1967: Flora and vegetation of Bucegi Mountain. Acad. Press, Bucharest, 578 pp. (in Romanian). Bondev, I. 1966: Die Hochgebirgs-Pflanzendecke des Berkovischen und Čiprovschen Gebirges. Izv. Bot. Inst. (Sofia), 16: 79-169 (in Bulgarian). Bondev, I. 1991: The Vegetation of Bulgaria. Map 1: 600000 with explanatory text. St. Kliment Ohridski Univ. Press, Sofia, 184 pp. (in Bulgarian). Borza, A. 1959: Flora and vegetation of Sebe§uliu valley. Acad. Press, Bucharest, 326 pp. (in Romanian). Bo§caiu, N. 1971: Flora and Vegetation of Tarcu, Godeanu and Cerna Mountains. Acad. Press, Bucharest, 494 pp. (in Romanian). 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Nardetum strictae N a r d e t u m s t r i c t a e Locality Balkan Mountain Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Altitude (m) 1600 1650 1650 1750 1900 1550 1550 1440 1400 1420 1540 1600 1480 1860 1930 1700 1580 1600 1500 1700 1500 1680 1850 1600 1650 1700 1600 1650 2000 Exposition Potentillo ternatae - Nardion strictae 4 5 4 5 + + . + 1 1 1 1 Nardus stricta * Bruckenthalia spiculifolia Hieracium hoppeanum Hypericum maculatum Potentilla erecta Potentilla ternata * Geum montanum Antennaria dioica Bellardiochloa violacea Thymus vandasii Alchemillaflabellata Viola dacica Luzula sudetica Festuca nigrescens Homogyne alpina * Campanula patula ssp. abietina Ligusticum mutellina Campanula rotundifolia Scorzonera purpurea ssp. rosea Carex ovalis Veronica officinalis Plantago atrata Plantago gentianoides . . . . Nardetalia strictae and Nardo - Callunetea Agrostis capillaris Festuca rubra Vaccinium myrtillus Lerchenfeldia flexuosa Anthoxanthum odoratum Vaccinium vitis-idaea Chamaespartium sagittale Luzula multiflora Rumex acetosella Hieracium pilosella Sieglingia decumbens Hypericum perforatum Ranunculus acris Luzula campestris 4433 . + + . 1 + 2 . 4444 + + + . 2 1 1 . + + . . 4453 +2++ . + + 2 . + + . + + . . . . + . . . ++++++++++ .........+ + + + + . . . . . + . + + +. + . . . . 343444 ++++++ 1 + . . + . + + . + + . . + . . . . + + + + 22 + . . . . . + . . . . . . . . 1 + . 334455 + + + 1 . . 1 + + . + + + . 2 + + . . . + + . + 1 + . . 2.+. 3 2 . . . + + + + +1 1 . + . ++++ ++++ + + + . . + . . +++++++ ++ 1 ++2 1 + 2 ++ +. + 1 1 2 1 + 1 1 1 + 1 1 1 + + + + + 1 + + + 1 + + + + + + + + + + + + + + +2222 2 2 2 1 3 + 1 + . . . + . 3 2 2 3 23 2 + 3 2 2 2 2 3 2 2 2. 3 3 + + + 1 1 + +. 2 + + 2 + + 2 2 + +. 2 2 + 1 + + + + + + + + + + + + + +. + 1 + 1 + + +. + + + + 2 + . + 1 +++ 3 2 + ++ ++ + + + + + + + + ++ + ++ + 1 +++ + + +++ + + + ++ + ++ + + + 12 NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance Tabela 1: Asociaciji Nardetum strictae Greb. 1950 in Campanula alpinae - Nardetum strictae (Simon 1958) nom. N a r d e t u m s t r i c t a e Campanulo alpinae - Nardetum strictae Balkan Mountain Mt. Pirin Mt. Balkan 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 Constancy 52 53 54 55 56 57 Constancy 58 59 60 61 62 Constancy 1620 1480 2000 1500 1900 1750 2000 1650 1620 1800 1950 1800 1850 2000 2000 2000 1950 2000 1700 2000 1850 2080 2100 2150 2300 2400 2200 2200 2050 2150 1950 2200 2150 to GO ^ W So wWtStd^ ^ ^ M w m & & & yivivi OO OO OO tB 5 3 5 3 4 3 4 3 4 3 4 3 4 5 5 5 4 5 55 4 3 V 4 3-4 3-4 1 4 2-3 V 5 3 2 2 2 V 1 + 3 + 1 2 3 + 2 1 1 + 1 1 1 .+ + IV 1 2 + + IV + + + + + + 3 + + + + + III + I + + + + + + + + + + + + III + + II 2 + + + + + + 1 + .1 + III + + + + + III 2 . +-1+-1 + + V + + II + + + 1 + + + + + II . + + + III + 1 + III + + + + + + II + + + + + V + 3 + + + + + + II 1 1 I + + + + + + + + + + 1 + + II + + 2 + IV + + + + II + + + + + + II + + + + 2 2 2 2 2 .1 II + I 2 + 1 + + I 1 2 1-2 . 1-2 IV 1 1 1 + IV + 1 + + + 2 .2 I + + + II + + 1 .+ I I + + I + I + + + I + I 2 1 1-2 2-3 + 1-2 V 2 I . + 1-2 + III + 2 2 2 + 2 + + + + 1 1 + 1 1 V . + 1 + 1 + + + 2 + 1 2 2 2 3 2 1 2 2 + V . 2 + II + 2 + 3 + 3 2 2 2 3 1 2 1 1 + 1 1 + 1 2 V . . . + . I + 1 + .1 IV + 1 + 1 3 + + + + + 1 + 1 IV . 1 + 1 1+ V + + + + + + + 1 + + III . . + . . I 2 + + III 2 + 1 + 2 2 + 2 + 1 + + III . 1 .+ II + + II . + I + + + + + + + II . .+ I + + + + + + + + + II . + I + + 1 + 1 II . + I I . + I + + + 1 I . 1 I . 13 Hacqüetia Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Seslerion comosae Campanula alpina . . . . Juncus trifidus . . . . Agrostis rupestris . . . . Dianthus microlepis . . . . Cetraria islandica . . . + Festuca airoides . . . . Veronica bellidioides . . . . Omalotheca supina . . . . Euphrasia minima . . . . Hieracium alpicola gr. . . . . Sesleria comosa . . . . Seslerietalia comosae and Juncetea trifidi Poa media . . . . Scleranthus neglectus . . . . Ranunculus montanus . . . . Alopecurus gerardii . . . . Carex kitaibeliana . . . . Crocus veluchensis . . . . Luzula italica . . . . Festuca riloensis . . . . Other species + 1 + + + + + 1 1 + ++++ 1 + + + + + + + + . . . Juniperus sibirica . + + + . + r. .+ + ++ + + + + +. .+ + 1.. Vaccinium uliginosum . . + + . . .+ + + + + 1. +. Carex caryophyllea .....+ ++ + + +. +. + + 1 + 1 +. Genista depressa s.l. .....+ +. + +. + + + + +. +2 +++ Campanula patula ssp. epigaea . . . . + + +. + ++ .+ 2. + 1 +. + + . + Verbascum longifolium .....+ +. + ++ .+ + .+ + 1 + + .2 +. + + . + Luzula luzuloides .....+ r .1 + .+ + + + 2 2. + Acinos alpinus Potentilla crantzii Trifolium repens s.l. Gentianella bulgarica Agrostis canina Arenaria biflora Bistorta major Thymus zygioides Trifolium pratense Veronica chamaedrys Thymus jankae Peucedanum oligophyllum Campanula velebitica Cruciata glabra Hieracium sparsum Koeleria eriostachya Geum coccineum Luzula pindica Omalotheca sylvatica Plantago subulata Polytrichum juniperinum Potentilla haynaldiana Veratrum lobelianum Sagina saginoides + +. . + . 1 + + +.32 . . + +11+ + + + +.+ + + . +++ . + . ..................+ ....+ .+ + + + + + 12.22321222...... + . + . . . .+2 + . 1++. .+ + + + +. . . .++........+ .1.......... ++++ 14 + + +++ ++ 1 ++ 1 + + + + + ++ + 2 + ++ + +++ + + + + + + + ++ 1 + ++ + + 1 2 + + + + + NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 C. 52 53 54 55 56 57 C. 58 59 60 61 62 C. 2 + + + + 1 11 + + + 3 + 1 1 1 + 1 + + + 2+ +1 +1 2 . .+ 1 2 2 2 1 1.21 + 22. . . . . + 2 . + . . . .+ + +.+ 1 1 + 1 +++ . . + + . . +2 + + + . . + . . . . . . . . + +.++. ++1+ + + . + . + + 2 + + 1 1+ + + + + + + . + . + + + + + . + .+ + + +. +..... + + + . . . . . . . . ......1 . . . . + + + . . + . + . . . . . . . . . . . . . ........+....... . . . + . . +.....+ .+ + ++++ .+ ++ + .+ +-1 + + V 1 2 + + 2 V I .+ + II 2 2 2 3 3 V I + . +-1+-1 1 + V 1 + II + +-1 . + + + V + + + III I + + + III + 2 II I 1 2 1 2 + V I + 1 + + + V .+ + II + + + III I +. + + III + I + 1 + III I + 2 II + + II + . II . . I . . III .+I 3 + I . . II .. I + 2 + 1 + .+ + + +11 3. .11.+1 + 1+ + .32 + 11.2.2 + . + 2 . . . . + . . . . + +-1 1 1-2+-1+-1+-1 + 1 + +-1 + + +-1 + + + +-1 .+ 1-2 +-1 IV III III III III III III II II II II I I I II II II I + + +I I . . . + + . + + IV . . + + . + . . III .I .I +I +I .I .I .I +I .I +I . I 1 ... + I +I I ++ + II I+ + + + 2 IV + + + III 1 + + III + 2 II 1 2 2 III + + + III 1 + + III 2 2 1 + IV 2 + II 1 + + III + + II + + I + I + + II 2 + 1 III + I + + II + 1 . + . . 11 + + + 3 + ++ 2 ++ + 3 + + I + + + + + + II ++ II ++ I + + + ++ + + + ++ + + + + + + I + 15 Hacqüetia Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Stellaria graminea Achillea millefolium Alchemilla vulgaris agg. Calamagrostis arundinacea Campanula sparsa Carduus carduelis Centaurea napulifera Cerastium arvense Cerastium banaticum Chamaecytisus pygmaeus Cruciata laevipes Deschampsia caespitosa Euphrasia liburnica Festuca valida Helianthemum nummularium Juncus conglomeratus Leontodon autumnalis s.l. Leontodon hispidus Lotus corniculatus Omalotheca norvegica Plantago lanceolata Poa annua Potentilla argentea Rubusidaeus Satureja montana Scleranthus perennis Senecio abrotanifolius Thlaspi kovatsii Viola tricolor Picea abies Pinus mugo Polytrichum piliferum Rhizocarpon alpicola Caloplaca sinapisperma Cladonia pyxidata Lepraria neglecta Bryum sp. Taraxacum sp. 12+ + 1 . + + + + . + + + . + +++ . . + +++++ + . . . + + + . + . . + 1+ + . + + . + . + + . + . . + + + . . . . + . + + + . + + . . + . + . . + . . + . 1 + + + + . . . + . . . . Species mentioned in the literature as diagnostic also for order Seslerietalia comosae and class Juncetea trifidi. + + + + + + + + + + + r ++ +++ + +++ + + + ++ + +++ + + + + + + + + +++ + + + + 16 NiKOLAY I. Velev & Iva I. Apostolova: A Review of Potentillo ternatae - Nardion strictae alliance 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 C. 52 53 54 55 56 57 C. 58 59 60 61 62 C. +.+ . . . + . +..... + . + . . . + . . + . ......+ + . + + .......+ . + + + . . . . . . . . . . ++ ++++ ++ II + ++ + + + + . . . . . . . + . . . .....+ 2 + + + 17 Hacqüetia Table 2: Altitudinal range of some Potentillo ternatae - Nardion strictae syntaxa. Table 2: Razpon nadmorskih višin nekaterih sintaksonov zveze Potentillo ternatae - Nardion strictae. Syntaxon Altitudinal range Country Reference Nardetum alpinum moesiacum 2100- -2400 m Bulgaria Simon 1958 Carici - Festucetum microphyllae 2170- 2200 m Bulgaria Roussakova 2000 Poa media dominated communities 2160- 2700 m Bulgaria Roussakova 2000 FFRY * Bulgaria Pavlović 1951; Mišić 1960; Mišić et al. Nardetum strictae 1000- 2080 m 1978; Petrov & Mesaroš 1988 Velev & Apostolova 2008 Hygronardetum strictae 1370- -1700 m FFRY Mišić et al. 1978 Helianthemo - Nardetum strictae 1700- -1900 m FFRY Janković 1982 Festuco fallaci - Nardetum strictae 500- -1800 m FFRY Janković 1982 Festuco halleri - Nardetum strictae 2000- 2300 m FFRY Janković 1982 Pu^caru et al. 1956; Borza 1959; Simon 1966; Beldie 1967; 2006 Violo declinatae - Nardetum strictae 570- 2200 m Romania Coldea 1987; Resmeritâ 1970; Pop et al. 2002; Grigoriu & Alda 2004 Niculescu 2004; Sarbu 2004; Sanda et al. 2006; Chifu et al. 2006 Pu^caru et al. 1956; Csürös & Resmeritâ 1960; Gergely 1969 Coldea 1987, 1990; Coldea & Pop 1988; Mihâilescu 2001 Scorzonero roseae - Festucetum nigricantis 850- -1950 m Romania Pop et al. 2002; Niculescu 2004; Sarbu 2004; Sanda et al. 2006 Chifu et al. 2006 Campanulo serratae - Festucetum ovinae 1100- 2120 m Romania Resmeritâ 1986; Coldea 1987; Pop et al. 2002 Hieracio polosellae - Nardetum strictae 700- 1250 m Romania Resmeritâ 1970; Pop et al. 2002 Puçcaru et al. 1956; Buia 1963; Beldie Carici - Nardetum strictae 1100- -1850 m Romania 1967; Resmeritâ 1970 Pop et al. 2002 Nardo - Callunetum vulgaris 800- -1600 m Romania Ghiça et al. 1970; Resmeritâ 1970; Pop et al. 2002 Poetum mediae 1700- 2020 m Romania Coldea 1990; Niculescu 2004; Sarbu 2004; Donita 2005 Antennario - Festucetum sudeticae 1550- 2150 m Romania Boçcaiu 1971 Soldanello - Nardetum 1200- 1750 m Ukraine Malinovsky & Kricsfalusy 2000 FFRY - Former Federal Republic of Yugoslavia 18