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 floristically 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 predstavili 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 endemičnih 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, vegetacija.
1. INTRODUCTION
the development of different types of grasslands
along wide elevational and latitudinal gradients,
forming syntaxa at various ranks — Seslerietum
korabiensis Micevski 1994, Seslerietum wettsteinii
Horvat 1937, Seslerietum juncifoliae Horvat 1930,
Seslerion juncifoliae Horvat 1930, Seslerion rigidae
Zólyómi 1939, Seslerietalia juncifoliae Horvat 1930,
The genus Sesleria Scop. (Poaceae, Pooideae,
Seslerieae) is one of the most important and interesting grass genera with its’ centre of diversity
and distribution on the Balkan Peninsula. The
species of this genus play a very important role in
1 Institute of Botany and Botanical Garden Jevremovac, Faculty of Biology, University of Belgrade, Takovska 43,
11000 Belgrade, Serbia, ekabas@bio.bg.ac.rs*, dlakusic@bio.bg.ac.rs, nkuzmanovic@bio.bg.ac.rs, sneza@bio.bg.ac.rs
2 Department of Botany, Faculty of Science, University of Zagreb, Marulićev trg 20/II, 10000 Zagreb, Croatia, antun.
alegro@biol.pmf.hr
3 Faculty of Mathematics, Natural Sciences and Information Technologies, University of Primorska, Glagoljaška 8,
6000 Koper, Slovenia, bostjan.surina@prirodoslovni.com
4 Institute for Research and Development of Sustainable Ecosystems, Jagodno 100a, 10415 Novo Cice, Velika Gorica,
Croatia, vsegota@ires.hr
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Hacquetia 13/1 • 2014, 57–77
etc. However, their most important role is forming the grasslands on base-rich soils of alpine and
subalpine belts of temperate European mountain
ranges (class Elyno-Seslerietea Br.-Bl. 1948). The
domination by Sesleria species of the high mountain calcareous plant communities in Europe is
discussed by Petriccione (1995) in his overview of
this vegetation type.
Sesleria juncifolia agg. represents an amphiAdriatic group of taxa exhibiting a typical disjunct range including the western & central Balkans and the Apennine Peninsula, where neither
their systematic relationships nor the syntaxonomical scheme are entirely clear. Based on relevant taxonomic papers (Deyl 1980, Strgar 1981,
Alegro 2007, Di Pietro 2007), the following species are considered to belong to S. juncifolia agg.:
S. apennina Ujhelyi, S. calabrica (Deyl) Di Pietro, S. kalnikensis Jávorka, S. juncifolia Suffren, S.
interrupta Vis., S. ujhelyii Strgar and S. albanica
Ujhelyi. Since both names, S. juncifolia Suffren
1802 and S. tenuifolia Shrader 1806, were validly
published, we use the earlier legitimate name S.
juncifolia in this paper, as it has a priority according to Art 11.4 of ICN (McNeill et al. 2012). The
detailed discussion regarding the nomenclatural
issues in Sesleria juncifolia complex is provided in
Di Pietro et al. (2013).
For the territory of Serbia, Lakušić & Sabovljević (2005) proposed a classification scheme
in which 31 associations and eight subassociations dominanted by different Sesleria species
were included, belonging to five classes, seven
orders and 11 alliances. However, this classification was not based on serious numerical analyses, thus an objective circumscription and classification are still missing. The most investigated
grasslands dominated by Sesleria spp. belong
to the class Elyno-Seslerietea, order Seslerietalia
juncifoliae (=tenuifoliae) Horvat 1930, alliances
Seslerion rigidae Zólyómi 1939 (dry grasslands
on calcareous bedrock) and Seslerion rigidae-latifoliae D. Lakušić 1996 prov. (dry grasslands on
serpentine rocks). Furthermore, seven associations belong to the order Onobrychido-Seslerietalia
Horvat 1949, alliances Edraiantho-Seslerion Horvat 1949, Onobrychido-Festucion Horvat 1949 and
Seslerio-Festucion R. Jovanović 1955. A significant
number of communities belong to the class of alpine pastures on siliceous rocks, Caricetea curvulae Br.-Bl. 1948, order Seslerietalia comosae Simon
1957, alliance Seslerion comosae Horvat 1935. Only
one association was recorded for the class Asple-
nietea trichomanis Br.-Bl. 1934 corr. Oberd. 1977.
Finally, some communities dominated by Sesleria
spp. belong to the Festuco-Brometea Br.-Bl. & Tx.
ex Soó 1947. The dominant Sesleria species that
build up these syntaxa are S. filifolia Hoppe in
eastern Serbia and S. serbica (Adam.) Ujhelyi and
S. juncifolia in western Serbia.
Our research only included the stands dominated by Sesleria juncifolia agg. described as associations Seslerio-Edraianthetum jugoslavici Petković
et al. 1990 (Petković et al. 1990) from Mt. Mokra
Gora, Seslerietum tenuifoliae S. Vukojičić & D.
Lakušić 1990 prov. (Stanić 1990) from Mt.
Mučanj and Carici laevis-Helianthemetum alpestre
Horvat 1930 seslerietosum tenuifoliae Rajevski 1990
from Mt. Šarplanina (Rajevski 1990) in Serbia.
MATERIALS AND METHODS
Data sampling. In order to describe and resolve
the syntaxonomy of the Serbian grasslands dominated by Sesleria juncifolia agg., we processed 139
relevés belonging to 13 syntaxa dominated by S.
juncifolia agg., distributed throughout the territory of Serbia, Bosnia and Herzegovina, Montenegro, Croatia, Slovenia and northeastern Italy.
The majority of the relevés were taken from literature sources. The main criterion for the selection
of the syntaxa to be included in the analyses was
that S. juncifolia appears as both the dominant
and nominal species in the name of a syntaxa,
either at association or subassociation level (Table 1). In addition to the literature data, personal
unpublished data previously gathered on Mt.
Mučanj in Serbia, Mt. Durmitor in Montenegro
and areas of Mts. Velebit (Croatia) and Snežnik
(Slovenia; Table 1, Figure 1) were also included in
the analyses. All the relevés were sampled according to Braun-Blanquet (1964) method. The plot
size of our own relevés was 25 m2, corresponding to the standard for grasslands proposed by
Chytrý & Otýpková (2003). Plot sizes of relevés
from the literature varied, the exact sizes for each
association are given in Table 1.
Data analysis. After transforming BraunBlanquet cover-abundance values into a ninedegree ordinal scale (van der Maarel 1979) the
relevés were subjected to Detrended Correspondence Analysis (DCA) in order to detect the basic structure of the floristic composition. Finally,
the complete set was classified using Bray-Curtis
similarity and group average clustering. These
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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 (McCune & Mefford 2011) and FLORA softwares
(Karadžić et al. 1998).
In this paper, we used the concept of diagnostic and dominant species proposed by Chytrý
et al. (2002), Chytrý & Tichý (2003) and Tichý
& Chytrý (2006). Using the statistical measures
of fidelity, we quantified concentrations of species occurrences in groups of classified sites in
order to determine diagnostic species (Chytrý et
al. 2002). The size of the site groups in the data
set was standardised (virtually equalized), while
the relative frequencies of species occurrence
within and outside of these groups were kept
constant (Tichý and Chytrý 2006) to calculate
the Φ-values as a measure of fidelity independent of the number of available relevés. To assess
statistical significance of concentration of species
in vegetatation types, we performed the Monte
Carlo significance test of observed maximum indicator value for the species with 4999 permutations. PcOrd 6.0 software (McCune & Mefford
2011) was used for the calculation of Φ-values.
In order to determine the dominant species, we
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Hacquetia 13/1 • 2014, 57–77
calculated the coverage index (Ic) according to
Lausi et al. (1982). Species with Φ-values higher
than 0.50 were considered diagnostic. Species
with cover ≥ 50% in a minimum of 5% of the relevés for any association were accepted as dominant. Species recorded in a minimum of 60% of
the relevés for any association were considered
constant. Nomenclature of plant taxa follows
the Flora Europaea Database (Tutin et al. 2001),
except for critical taxa with unresolved relationships, which were included as species complexes
(aggregates). The names of all the syntaxa follow
Rodwell et al. (2002) with a few exceptions according to Lakušić & Sabovljević (2005). All the
underlying plot data used in the paper are stored
in Vegetation Database of Grassland Vegetation
in Serbia (Aćić et al. 2012; GIVD number: EURS-002; relevé numbers: 7000–7138).
from the Liburnian karst. The remaining two
groups represented the relevés from northeastern
Italy, Slovenia and Croatia (Group II), and the
relevés from Serbia, Bosnia and Herzegovina and
Montenegro (Group III). It can also be seen that
the relevés from Mt. Mučanj (western Serbia) and
the ones representing the Seslerio-Edraianthetum
jugoslavici (Mt. Mokra Gora) were well differentiated within this last group, while the relevés representing the Carici laevis-Helianthemetum alpestre
seslerietosum tenuifoliae from Mt. Šarplanina overlapped with the rest of the relevés in this group.
classification - Results of the cluster analysis performed on the complete data set were very
much in accordance with the ordination results,
in that they showed that two main groups of
stands (clusters) could be differentiated. Cluster I (Figure 3) corresponded to the stands of
the associations from northeastern Italy, Slovenia and Croatia. This cluster corresponded completely with Groups I and II in Figure 2, so the
only difference is related to relevés of Liburnian
heaths Rhododendro hirsuti-Juniperetum alpinae
seslerietosum tenuifoliae, which were differentiated
as a single group in DCA. On the other hand,
Cluster II (Figure 3) represented the relevés from
Serbia, Bosnia and Herzegovina and Montenegro (Group III in Figure 2). Within this cluster,
RESULTS
Ordination - A DCA conducted on the complete
data set showed three rather discrete groups of
relevés (Figure 2), well separated along first two
canonical axes. The most distinct (Group I) was
the one representing relevés of the Rhododendro
hirsuti-Juniperetum alpinae seslerietosum tenuifoliae
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.
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Information Remaining (%)
Cluster analysis
Distance (Objective function)
Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
Rhododendro hirsuti-Juniperetum alpinae
Carici humilis-Seslerietum juncifoliae
Seslerietum juncifoliae
Carici humilis-Centaureetum rupestris
Bromo-Seslerietum interruptae
Genisto sericeae-Seslerietum juncifoliae
Edraiantho graminifoliae-Sesler. juncifoliae
Diantho petrae-Seslerietum juncifoliae
Seslerietum juncifoliae
Carici laevis-Helianthemetum alpestre
Carici laevis-Seslerietum juncifolia
Seslerio-Gentianetum dinaricae
Seslerietum juncifoliae
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.
Syntaxonomical treatment
it was again noticeable that Serbian associations
represented well separated and discrete units.
On the basis of the ordination and classification analyses, we established the existence of two
well defined syntaxa with the dominance of Sesleria juncifolia in Serbia: ass. Seslerio-Edraianthetum
jugoslavici from Mt. Mokra Gora, and a new association from Mt. Mučanj: Diantho petraeae-Seslerietum juncifoliae. Relevés of the Carici laevis-Helianthemetum alpestre seslerietosum tenuifoliae from
Mt. Šarplanina overlapped with communities
from the Cincar, Vlašić, Bjelasica and Durmitor
mountains in the ordination, while in the cluster
graph analysis they formed a joint cluster together with the community from Mt. Cincar and only
a few relevés from the community from Mt. Durmitor. A synoptic table comparing all community
types is presented in the Table 3 in order to show
the differences between the individual clusters
obtained by numerical classification.
Ass. Diantho petraeae-Seslerietum juncifoliae
vukojičić & D. lakušić ass. nova hoc loco (Holotypus Table 2, rel. 5 hoc loco)
Original: Seslerietum tenuifoliae Stanić & D. Lakušić 1990 prov., nom. ined. (Art. 1, ICPN)
Note: The Seslerietum tenuifoliae Stanić & D. Lakušić 1990 prov. was not effectively published
(Def. III, Art. 1, ICPN; Weber et al. 2000), since it
was only recorded and preliminarily described in
a Diploma thesis (Stanić 1990). Therefore, we describe it here as a new association, in accordance
with the International Code of Phytosociological
Nomenclature (ICPN; Weber et al. 2000).
Dominant species: Sesleria juncifolia
Diagnostic species: Campanula rotundifolia,
Chamaecytisus ciliatus, Chamaespartium sagittale,
Cotoneaster integerrimus, Dianthus petraeus subsp.
petraeus, Draba lasiocarpa, Festuca panciciana, Ga61
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Hacquetia 13/1 • 2014, 57–77
lium corrudifolium, Helianthemum nummularium
subsp. nummularium, Ornithogalum collinum, Pedicularis heterodonta, Potentilla cinerea, Sanguisorba minor subsp. minor
constant species: Carex kitaibeliana, Edraianthus 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 inclination of about 35° on average (Table 2). The
dominant species Sesleria juncifolia with its dense
tussocks formed stands up to 40 cm high, covering 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-Carpathian or Balkan-Apennine subendemic species,
such as Cerastium decalvans, Daphne blagayana, Dianthus petraeus subsp. petraeus, Draba lasiocarpa,
Edraianthus graminifolius, Festuca panciciana, Laserpitium siler subsp. garganicum, Minuartia bosniaca, Pedicularis heterodonta and Sesleria juncifolia
were recorded in this association. Also three glacial relicts (Arabis alpina, Poa alpina, and Saxifraga
paniculata) occuring within the stands may point
to the glacial-refugial character of the association.
However, the occurrence of some differential taxa
from forests, such as Daphne mezereum, Fagus sylvatica, and Poa nemoralis, could point to the fact
that the montane beech forests of Fagetalia sylvaticae Pawłowski 1928 prevailed in the recent past,
or might even represent a natural vegetation type
on sites nowadays covered by the stands of the
Diantho petraeae-Seslerietum juncifoliae.
Figure 4: Summer aspect of the association Diantho petraeaeSeslerietum juncifoliae Vukojičić & D. Lakušić ass. nova on Mt.
Mučanj (photo: N. Kuzmanović).
Slika 4: Poletni aspekt asociacije Diantho petraeae-Seslerietum
juncifoliae Vukojičić & D. Lakušić ass. nova na gori Mučanj
(foto: N. Kuzmanović).
May, June and August, while the driest are February and March. Mean monthly temperatures
below 0 °C were noted during November, December, January, February and March, confining
the growing season to between April and September. Considering the relatively low elevation and
the mountain temperate climate, montane beech
forests represent the potential vegetation of the
investigated area. However, since the major parts
of the forests were completely degraded, artificial
forest stands of black pine (Pinus nigra), fir (Abies
alba) and spruce (Picea abies) dominate the landscape (Stanić 1990).
ecology and synchorology of the association
Stands of Diantho petraeae-Seslerietum juncifoliae
are so far known only from the area of Mt. Mučanj
in southwestern Serbia. The direction of the
mountain is NW-SE, while its maximum height
is 1534 m a.s.l. Mt. Mučanj is seperated from the
neighboring mountains by river valleys, pointing
to the fact that fluvial erosion was the factor shaping the relief, earlier formed by tectonic movements. The dominant geological substrate in this
area is limestone. The soils are shallow and very
skeletal. The climate type is temperate-continental, but in its modified mountain variant (Stanić
1990). The annual mean temperature is 2.9 °C,
and the annual precipitation is 944 mm. January is
the coldest month with a temperature of –8,5 °C,
while July is the warmest with a temperature of
+12.3 °C (Stanić 1990). The wettest months are
Ass. Seslerio juncifoliae-edraianthetum graminifolii B. Petković et al. ex Kabaš et al. ass. nov.
hoc loco
validated name: Seslerio-Edraianthetum jugoslavici Petković et al. 1990 nom. inval. (Art. 5, ICPN)
Holotypus: Petković et al. (1990: Table 2, rel. 2)
Note: The association Seslerio-Edraianthetum
jugoslavicii B. Petković et al. 1990 was not validly
published, because the holotype relevé was not
assigned (Art. 5, ICPN; Weber et al. 2000). Also,
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the name Edraianthus jugoslavicus Lakušić was
not validly published (Art. 39.1 of ICN, McNeill
et al. 2012), since it was not accompanied by a
Latin description or diagnosis. Accordingly, we
used the accepted and validly published name
Edraianthus graminifolius (L.) A. DC. (Euro+Med
2010) when naming the association.
Diagnostic species: Acinos arvensis, Alchemilla
plicatula, Alyssum montanum, Anthyllis vulneraria
subsp. pulchella, Asplenium trichomanes-ramosum,
Helianthemum nummularium subsp. grandiflorum,
Juniperus sabina, Minuartia verna, Polygonum viviparum, Scabiosa ochroleuca, Silene pusilla, Polygala
supina subsp. supina
ated group (Group II, Figure 2) corresponds to
the stands from northwestern Italy, Slovenia and
Croatia, representing grasslands classified mostly within the class Festuco-Brometea. Finally, the
third well defined and separated group (Group
III, Figure 2; Cluster II, Figure 3) includes Serbian, Bosnian and Herzegovinian and Montenegrian grassland stands, classified mostly within
the vegetation of the high mountain rocky calcareous grasslands of the class Elyno-Seslerietea.
Furthermore, both DCA and cluster analysis also
showed that both Serbian associations are floristically well differentiated from the rest of the
similar syntaxa, i.e. the association Seslerio juncifoliae-Edraianthetum graminifolii from Mt. Mokra
Gora, along with the stands of the associations
from the Bjelasica, Durmitor, Cincar and Vlašić
mountains belonging to the class Elyno-Seslerietea. Corroboration of this statement can be seen
in the floristic composition, which is determined
by the position of these stands at high elevations,
usually above the upper forest line.
The newly described association from Mt.
Mučanj, on the other hand, significantly differs
from the subalpine and alpine rocky grasslands
of the class Elyno-Seslerietea in its floristic properties. These stands host 43 taxa (e.g., Briza media,
Bromus erectus, Euphorbia myrsinites, Globularia
cordifolia, Hieracium pilosella, Hypericum perforatum, Juniperus communis, Leucanthemum vulgare,
Phleum pratense, Plantago lanceolata, Plantago media, Poa badensis, Potentilla cinerea, Sanguisorba
minor, Teucrium chamaedrys, Thymus pulegioides,
Trifolium campestre, Trifolium montanum etc.)
which, within the eastern and southeastern Dinaric Alps, prefer calcareous grasslands developed within deciduous broadleaved forests. Additionally, the Φ-indices of most of these taxa are
highly statistically supported, hence we find their
classification within the class Festuco-Brometea to
be fully justified. The fact that only 12 taxa typical for subalpine and alpine grasslands (Acinos
alpinus, Arabis alpina, Carex kitaibeliana, Daphne
alpina, Edraianthus graminifolius, Helianthemum
canum, Hieracium villosum, Pedicularis heterodonta, Poa alpina, Rosa pendulina, Sesleria juncifolia and Thlaspi kovatsii) occur in stands from
Mt. Mučanj further supports our classification
scheme. To that end, Φ-indices of the majority of
the subalpine and alpine grassland taxa from the
class Elyno-Seslerietea do not reflect statistically
significant fidelity or the diagnostic value for the
association in general.
Syntaxonomical scheme
Festuco-Brometea Br.-Bl. & Tx. ex Soó 1947
Scorzonero-Chrysopogonetalia Horvat & Horvatić 1958
Chrysopogono-Saturejion Horvat & Horvatić
1934
Diantho petraeae-Seslerietum juncifoliae
Vukojičić & D. Lakušić 2014
Elyno-Seslerietea Br.-Bl. 1948
Crepidetalia dinaricae Lakušić 1966
Oxytropidion urumovii Lakušić 1964
Seslerio juncifoliae-Edraianthetum graminifolii Petković et al. ex Kabaš et al. 2014
DISCUSSION
Our numerical analyses showed that the analyzed
stands of different syntaxa dominated by Sesleria
juncifolia agg. in the mountains of the central and
western part of the Balkan Peninsula are very heterogenous and probably have different origins.
Their relationships in a broader, amphi-Adriatic
context are a subject of the ongoing research, preliminarily presented in Di Pietro et al. (2013).
Our results showed that the first clearly distinct group (Group I, Figure 2) represents the
stands of the Rhododendro hirsuti-Juniperetum alpinae seslerietosum tenuifoliae. These results support the opinion of Surina (2013) that despite
the domination of S. juncifolia in these stands,
the syntaxon should actually be classified within the heath vegetation of Erico-Pinetea Horvat
1959, and not within the grassland communities
of Elyno-Seslerietea. The second well differenti63
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Confirmation of the opinion that the new
community from Mt. Mučanj belongs to the class
of Festuco-Brometea should also be sought in its
coenotic surroundings and its origin. The potential natural vegetation of the highest part of Mt.
Mučanj is deciduous broadleaved forest of the
Fagetalia sylvaticae order (Jovanović et al. 1986).
Therefore, at an altitude of 1500 m a.s.l., this
isolated mountain does not provide conditions
for the development of the potential vegetation
of (sub)alpine grasslands of the Elyno-Seslerietea
class.
While there is no doubt the new association
from Mt. Mučanj belongs to the class FestucoBrometea, and not Elyno-Seslerietea, as indicated
in Lakušić & Sabovljević (2005), its assignment
to a lower syntaxa is not completely clear. Considering their geographic position and floristic
composition, these stands show transitional characteristics connecting them to calcareous karstic
grasslands of the Illyric-Dinaric region with
Chrysopogono-Saturejion Horvat & Horvatić 1934,
meso-xerophytic swards in sub-oceanic regions
of western Europe with Bromion erecti Koch 1926,
and meso-xerophytic swards in sub-continental
regions of central and eastern Europe with CirsioBrachypodion pinnati Hadač & Klika 1944. The
most important species connecting this association with the Chrysopogono-Saturejion are: Asperula
cynanchica, Dianthus petraeus, Galium corrudifolium, Globularia cordifolia, Hieracium villosum, Laserpitium siler, Leontodon crispus, Sesleria juncifolia,
and Teucrium chamaedrys. The species relating it
with the associations of the alliance Bromion erecti
are the following: Plantago media, Sanguisorba minor, Scabiosa columbaria, considered as diagnostic
for the Bromion erecti (Dengler 2003, Jarolímek &
Šibík 2008, Chytrý 2010), while it also hosts diagnostic species of the Cirsio-Brachypodion pinnati
(Dengler 2003, Jarolímek & Šibík 2008, Chytrý
2010) such as Asperula cynanchica, Brachypodium
pinnatum, Bromus erectus, Plantago media, Sanguisorba minor, Trifolium montanum. Therefore,
considering the number of common species and
their diagnostic character as well as their geographic position within the continental Dinarides, the authors think the position of the new association Diantho petraeae-Seslerietum juncifoliae is
within the alliance Chrysopogono-Saturejion.
The presence of 11 chasmophytic taxa in the
stand of the newly described association is also
significant, when the syntaxonomic position
and the origin of the association are considered.
Specific chasmophytic taxa are present in rocky
grasslands due to the proximity of chasmophytic
stands in the investigated area (Stanić & Lakušić
1993) and the fact that steep rocky grasslands
share many features with rock crevices or even
screes, generating ecological conditions suitable
for both grassland and chasmophytic taxa, e.g.
Asplenium ceterach, Asplenium ruta-muraria, Asplenium trichomanes, Campanula rotundifolia, Erysimum sylvestris, Hieracium humile, Hieracium pannosum, Laserpitium siler, Leontopodium alpinum,
Saxifraga tridactylites, or Silene saxifraga. This circumstance was observed and recently discussed
by Surina & Martinčič (2012). Nevertheless, low
coverage and fidelity indices of chasmophytes in
the studied stands do not justify their classification within the class Asplenietea trichomanis.
Finaly, the syntaxonomical position of the
Carici laevis-Helianthemetum alpestre seslerietosum
tenuifoliae from Mt. Šarplanina remains unclear,
given that in the DCA analysis its stands are
overlapping with the stands of associations from
Bosnian and Montenegrin mountains. A possible
reason for this is the fact that this subassociation
was described on the basis of only three relevés,
which is not representative enough for an objective understanding of its syntaxonomical position. While all of the other investigated syntaxa
are found within the Dinaric floristic province,
the stands of this subassociation belong to the
Scardo-Pindic floristic province. Accordingly,
the comparison of these three relevés with the
rest of the 136 dinaric relevés would not reflect
the real relationships of these syntaxa within the
Balkan Peninsula.
ACKNOWLEDGEMENTS
The authors are grateful to the Serbian Ministry
of Science and Technological Development (Project No. 173030 Biodiversity of the plant life of
Serbia and Balkan Peninsula – Assessment, sustainable use and conservation, 2011-2014) for financial support. The authors also wish to thank
Jozef Šibík, Kiril Vassilev and Jürgen Dengler for
their valuable comments and suggestions, which
have significantly improved this manuscript. Finally, we thank Laura Sutcliffe for the linguistic
editing of the manuscript and EDGG for making
this possible through an IAVS grant.
64
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Co-ordinating editor: Jürgen Dengler
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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
1.
Rhododendro hirsuti-Juniperetum
alpinae Horvat ex Horvat et al. 1974
subas. seslerietosum tenuifoliae
Surina 2013
33T VL53
Slovenia
(Liburnian
karst), Croatia
(Liburnian karst)
Surina
2013
30
13
2.
Seslerio-Edraianthetum jugoslavicii
Petković et al. 1990
34T DN44
Serbia (Mt.
Mokra Gora)
Petković
et al. 1990
25–100
5
3.
Seslerietum tenuifoliae Stanić &
D. Lakušić 1990 prov.
34T DP22
Serbia (Mt.
Mučanj)
Vukojičić &
Lakušić, D.
unpubl.
20–400
12
Seslerietum juncifoliae
Horvat 1930
33T XJ66
Bosnia and
Herzegovina
(Mt. Cincar)
Lakušić et al.
1984
20–100
24
6.
Seslerio-Gentianetum dinaricae
Lakušić et al. 1982
33T YK00, YK01
Bosnia and
Herzegovina
(Mt. Vlašić)
Lakušić et al.
1982
10–100
10
7.
Carici laevis-Helianthemetum
alpestre Horvat 1930 subas.
seslerietosum tenuifoliae
Rajevski 1990
34T EM07
Serbia (Mt. Šarplanina)
Rajevski
1990
-
3
8.
Carici humilis-Seslerietum
juncifoliae
33T VK95
Croatia (northern
part of Mt.
Velebit)
Alegro
& Segota
2009, 2010,
manuscript
50
9
9.
Carici laevis-Seslerietum tenuifoliae
(Lakušić 1966) Redžić 2011
(=Seslerietum juncifoliae
montenegrinum Lakušić 1964)
34T CN94
Montenegro
(Mt. Bjelasica)
Lakušić
1966
100–200
10
10.
Seslerietum juncifoliae s.l.
34T DN46, DN47,
DN48
Montenegro (Mt.
Durmitor)
Lakušić, D.,
unpubl.
20–100
5
11.
Genisto sericeae-Seslerietum
juncifoliae Poldini 1980
33T UL96, VL06,
VL13, VL14, VL15,
VL23, VL26, VL27,
VL44
Slovenia
(Liburnian
karst), Italy
(Karst)
Poldini 1989,
Kaligarič
1997
70–80
19
12.
Carici humilis-Centaureetum
rupestris Horvat 1931 seslerietosum
juncifoliae Horvat 1962
33T VL13, VL15,
VL23, VL24, VL26,
VL27, VL44, VL54
Slovenia
(Liburnian
karst), Italy
(Karst)
Poldini 1989,
Kaligarič
1997, Surina
unpubl.
100
25
13.
Bromo-Seslerietum interruptae
Trinajstić 1965
33T XH69
Croatia
(Mt. Biokovo)
Trinajstić
1987
-
4
4, 5.
67
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Plot size Number
(m²)
of relevés
Hacquetia 13/1 • 2014, 57–77
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).
No. Relevé
Dominant taxa
Sesleria juncifolia agg.
Constant taxa
Globularia cordifolia
Edraianthus graminifolius
Saxifraga paniculata
Carex kitaibeliana
Diagnostic taxa (*also constant taxa)
Dianthus petraeus subsp. petraeus*
Chamaecytisus ciliatus*
Festuca panciciana*
Pedicularis heterodonta*
Potentilla cinerea*
Ornithogalum collinum*
Cotoneaster integerrimus*
Campanula rotundifolia*
Sanguisorba minor subsp. minor
Helianthemum nummularium subsp.
nummularium
Draba lasiocarpa
Chamaespartium sagittale
Galium corrudifolium
Other taxa
Bromus erectus
Minuartia verna
Leontodon crispus subsp. crispus
Trifolium montanum
Arenaria serpyllifolia subsp. serpyllifolia
Trifolium alpestre
Euphrasia stricta
Asplenium ceterach
Juniperus communis
Poa alpina
Hieracium pilosella subsp. pilosella
Saxifraga tridactylites
Gymnadenia conopsea
Rhamnus saxatilis subsp. saxatilis
Stachys recta
Sedum acre
Euphorbia myrsinites
Rosa pendulina
Asplenium ruta-muraria subsp. ruta-muraria
Verbascum sp.
Cover index according to
Lausi et al. (1982) (Ic,%)
Date
1450 1400 1450 1450 1450 1450 1450 1450 1450 1350 1350 1300
–
–
W NW NW NW W
NE NW
E
E
S
0
0
10
10
10
30
70
70
70
50
65
70
60
60
80
80
80
50
80
70
60
60
80
25
25
25 100 20
30
75
24
80
30 400 50
30
VII VII VII VII VII VII VII VII VII VII VII VII
1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989 1989
1
2
3
4
5*
6
7
8
9
10
11
12
Constancy (%)
Altitude (m a.s.l.)
Exposition
Slope (°)
Total cover (%)
Plot size (m²)
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
3.4
+
.
1.2
3.3
1.1
.
1.2
1.3
1.2
1.1
.
1.3
1.2
1.2
1.2
1.2
1.1
1.2
1.2
1.3
1.1
1.2
1.2
1.2
1.2
.
.
1.1
1.2
1.1
1.2
3.4
1.2
.
1.2
1.3
1.3
1.2
.
.
.
1.2
.
1.2
1.1
1.2
.
92
92
67
67
33
42
22
22
1.2
1.2
1.1
+
1.2
1.1
1.2
.
+
1.3
1.3
.
1.1
.
.
1.2
.
.
1.3
1.2
1.2
1.1
1.2
.
+
.
1.1
1.3
1.2
1.2
1.1
1.3
1.1
+
.
1.1
1.3
1.2
1.2
1.1
1.2
1.1
1.1
.
+
1.2
1.2
1.2
1.1
1.2
1.1
1.2
.
.
1.3
1.3
1.2
1.1
1.3
1.1
.
1.2
1.1
1.3
1.3
1.2
1.1
1.3
+
+
1.2
1.1
1.3
1.2
.
1.1
1.3
+
.
1.1
.
1.4
1.3
1.2
+
1.2
1.1
1.3
1.2
1.2
1.3
1.2
2.2
.
1.2
1.1
1.3
1.2
1.2
1.2
1.2
1.2
.
1.2
.
.
1.2
+
100
92
92
83
83
75
75
75
58
33
30
30
30
26
23
22
22
19
.
1.1
.
.
1.1
1.1
1.1
1.1
.
.
1.2
.
50
16
+
1.2
.
.
+
.
.
1.1
1.2
.
.
.
.
.
1.2
.
.
1.2
1.2
.
1.3
+
.
.
.
+
.
1.1
1.2
1.3
1.2
1.2
1.3
+
1.2
.
50
50
50
16
15
14
1.1
1.1
1.1
1.2
1.2
1.1
1.1
.
.
1.1
.
.
+
.
.
.
.
.
.
.
.
.
+
1.1
.
.
.
.
+
.
.
.
+
.
.
.
.
1.1
.
.
1.2
1.2
+
1.1
.
+
1.1
.
.
1.1
.
.
.
.
.
.
.
.
.
.
1.2
1.2
.
.
1.2
+
1.1
.
.
1.2
1.2
1.1
.
.
.
+
.
.
.
.
1.1
1.2
.
1.1
.
.
+
.
.
+
.
1.1
+
.
.
.
.
1.1
.
.
1.1
.
.
1.1
.
.
.
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Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
No. Relevé
Acinos alpinus
Teucrium chamaedrys
Sedum album
Erysimum sylvestre subsp. sylvestre
Leontopodium alpinum subsp. alpinum
Arabis hirsuta
Phleum pratense subsp. pratense
Thlaspi kovatsii
Aethionema saxatile subsp. saxatile
Daphne mezereum
Thymus pulegioides
Vicia incana
Cerastium decalvans
Minuartia bosniaca
Briza media subsp. media
Scabiosa columbaria subsp. columbaria
Hieracium villosum
Poa nemoralis
Rhamnus alpinus subsp. fallax
Epipactis atrorubens
Thymus glabrescens
Hieracium pannosum
Arabis glabra
Cerastium brachypetalum subsp.
brachypetalum
Arabis alpina
Leucanthemum vulgare
Plantago lanceolata
Plantago media
Heracleum sp.
Fagus sylvatica
Valeriana montana
Myosotis arvensis subsp. arvensis
Erigeron annuus
Sorbus aria
Hypericum perforatum
Poa badensis
Asperula aristata subsp. scabra
Polygala supina subsp. supina
Brachypodium pinnatum subsp. pinnatum
Asplenium trichomanes subsp. trichomanes
Lotus corniculatus
Fragaria vesca
Allium lavum subsp. lavum
Laserpitium siler subsp. siler
Vincetoxicum hirundinaria
Daphne blagayana
Trifolium campestre
Silene saxifraga
Asperula cynanchica
Helianthemum canum subsp. canum
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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
69
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12 C. (%) Cov.
1.2
25
7
1.2
25
7
.
25
6
.
17
7
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17
6
.
17
6
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17
6
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17
6
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6
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+
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3
Seslerietum juncifoliae,
Durmitor
Genisto sericeae-Seslerietum
juncifoliae, Slovenia & Italia
Carici humilis-Centaureetum
rupestris, Slovenija & Italia
Bromo-Seslerietum interruptae,
Biokovo
Group with Max Φ-value observed
Max Φ-values
Overall constancy
B
C
100
100
100
89
100
100
12 0.6 13
.
.
33
33
30
.
40
.
32
16
60
56
.
.
13 0.38 7
5 0.33 7
.
67
.
100
67
.
67
.
.
67
.
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11
.
22
.
22
.
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11
11
11
11
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30
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40
.
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20
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40
20
40
.
20
60
40
20
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26
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16
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20
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68
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12
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1
6
3
4
2
3
6
2
12
4
2
5
2
4
5
92
60
100
100
100
100
100
.
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40
58
8
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75
30
30
85
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100
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80
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80
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60
25
8
100
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83
17
58
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31
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75
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20
100
40
10
70
.
30
40
.
50
30
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10
Carici laevis-Helianthemetum
alpestre ses. juncifoliae, Sara
3
Seslerio-Gentianaetum dinaricae,
Vlašić
2
Carici humilis-Seslerietum
juncifoliae, Velebit
A
Seslerietum juncifoliae, Snežnik
13
Seslerietum juncifoliae, Cincar
12
Diantho petraeae-Seslerietum juncifoliae, Mučanj
11
Seslerio juncifoliae-Edraianthetum
graminifolii, Mokra gora
10
Rhododendro hirsuti-Juniperetum
alpinae, Slovenia & Hrvatska
Dominant taxa
Sesleria juncifolia agg.
Constant taxa
Bromus erectus agg.
Lotus corniculatus L.
Diagnostic taxa
Rosa pendulina L.
Helianthemum canum (L.) Baumg. ssp. canum
Dianthus petraeus Waldst. & Kit. ssp. petraeus
Helianthemum oelandicum (L.) DC. ssp. alpestre (Jacq.) Breistr.
Helianthemum nummularium (L.) Miller ssp. grandilorum (Scop.) Schinz & Thell.
Potentilla cinerea Chaix ex Vill.
Cerastium decalvans Schloss. & Vuk.
Minuartia verna agg.
Dorycnium pentaphyllum Scop. ssp. germanicum (Gremli) Gams
Globularia meridionalis (Podp.) O. Schwarz
Polygonum viviparum L.
Coronilla vaginalis Lam.
Silene pusilla Waldst. & Kit.
9
1
6
7
8
Constancy (%)
0.57
0.81
0.75
0.71
0.68
0.62
0.52
0.51
0.51
0.5
0.72
0.68
0.62
5
4
4
4
4
4
4
4
4
4
3
3
3
Hacquetia 13/1 • 2014, 57–77
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Taxa
Carici laevis-Seslerietum
tenuifoliae, Bjelasica
Table 3: 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
columns. The last three columns refer to the max Φ-values of the species, the number of the syntaxa where it was observed and the overall constancy. Species are
sorted in descending order of constancy and Φ-values. The companion species with less than 10% overall constancy are not shown.
Tabela 3: Sinoptična tabela vseh proučevanih združb. V stolpcih so podane stalnosti pojavljanj vrst v odstotkih v 13 analiziranih sintaksonih. V zadnjih treh
stolpcih so predstavljene maksimalna Φ vrednost posamezne vrste, število sintaksonov v katerih se pojavlja in celotna stalnost. Vrste so razvrščene padajoče
glede na stalnost in Φ vrednosti. Spremljevalne vrste s stalnostjo pod 10% niso prikazane.
1
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54
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31
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54
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100
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100
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85
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77
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2
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40
40
20
40
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100
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40
40
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3
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92
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50
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92
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50
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50
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8
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13
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6
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63
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44
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56
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6
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80
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60
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60
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10
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40
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10
7
67
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33
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100
67
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100
67
100
67
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8
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67
33
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33
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11
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67
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44
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11
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9
20
20
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90
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.
70
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70
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60
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50
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40
10
60
.
20
20
40
.
20
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.
20
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.
20
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20
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20
40
.
20
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20
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11
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16
42
16
.
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.
21
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5
21
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12
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28
8
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72
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.
36
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52
4
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12
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64
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13
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75
50
.
100
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.
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50
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25
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A
4
3
3
4
4
1
3
3
13
13
1
13
1
3
2
9
1
12
7
7
7
9
12
3
13
1
6
9
3
3
5
9
12
5
4
1
2
2
9
B
0.6
0.59
0.58
0.56
0.54
0.53
0.53
0.53
0.53
0.51
0.5
1
0.95
0.95
0.88
0.85
0.78
0.78
0.77
0.75
0.72
0.72
0.71
0.69
0.69
0.68
0.65
0.65
0.63
0.63
0.63
0.63
0.59
0.58
0.57
0.56
0.55
0.55
0.55
C
3
3
3
3
3
3
3
3
3
3
3
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
Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
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Oxytropis dinarica (Murb.) Wettst.
Festuca panciciana (Hackel) K. Richter
Campanula rotundifolia (Desf.) Boiss. & Reuter
Thymus praecox Opiz ssp. polytrichus (A. Kerner ex Borbás) Jalas
Arctostaphylos uva-ursi (L.) Sprengel
Picea abies (L.) Karsten ssp. abies
Draba lasiocarpa Rochel
Cotoneaster integerrimus Medicus
Stipa pennata ssp. eriocaulis (Borbás) Martinovsky & Skalicky
Fumana procumbens (Dunal) Gren. & Godron
Daphne alpina L.
Asperula aristata L.
Salix appendiculata Vill.
Chamaecytisus ciliatus (Wahlenb.) Rothm.
Anthyllis vulneraria L. ssp. pulchella (Vis.) Bornm.
Draba aizoides L.
Juniperus communis ssp. alpina (Suter) Celak.
Jurinea mollis (L.) Reichenb. ssp. mollis
Onobrychis montana DC. ssp. scardica (Griseb.) P. W. Ball
Dryas octopetala L.
Bupleurum ranunculoides L.
Thymus praecox Opiz ssp. zygiformis (H. Braun) Jalas
Euphorbia nicaeensis All.
Chamaespartium sagittale (L.) P. Gibbs
Asperula purpurea (L.) Ehrend.
Cyclamen purpurascens Miller
Hypericum richeri Vill. ssp. grisebachii (Boiss.) Nyman
Jovibarba heuffelii (Schott) Á. & D. Löve
Galium corrudifolium Vill.
Helianthemum nummularium (L.) Miller ssp. nummularium
Peucedanum oreoselinum (L.) Moench
Verbascum chaixii Vill. ssp. austriacum (Schott ex Roemer & Schultes) Hayek
Plantago argentea Chaix
Plantago lanceolata L.
Festuca bosniaca Kummer & Sendtner
Athamanta cretensis L.
Polygala supina Schreber ssp. supina
Juniperus sabina L.
Hieracium pavichii Heuffel
1
.
.
.
.
.
62
.
.
.
100
100
.
.
.
.
85
85
.
.
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.
.
.
69
.
.
.
.
.
.
.
.
.
.
.
2
.
.
.
.
.
.
.
40
.
.
.
100
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
60
60
.
.
.
3
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
83
.
75
58
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
4
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
75
75
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
.
.
.
.
.
38
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
6
.
.
.
.
.
.
.
50
.
.
.
.
.
.
.
.
.
.
80
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
60
.
.
7
.
.
.
.
.
.
.
.
.
.
.
.
100
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
67
67
67
67
67
67
.
.
.
.
67
8
11
11
.
.
.
.
22
.
11
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
9
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
70
70
70
.
.
.
.
.
.
.
.
.
.
60
.
10
40
40
.
.
.
.
.
.
40
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
11
.
.
16
11
5
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
12
.
.
32
16
12
.
28
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
13
.
.
.
.
.
.
.
.
.
.
.
.
.
100
100
.
.
.
.
.
.
.
.
75
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
A
10
10
12
4
13
1
5
6
10
1
1
2
7
13
13
1
1
3
6
3
3
4
4
13
9
9
9
1
7
7
7
7
7
7
2
2
6
9
7
B
0.55
0.55
0.55
0.54
0.54
0.53
0.5
0.5
0.5
1
1
1
1
1
1
0.91
0.91
0.91
0.89
0.86
0.86
0.86
0.86
0.86
0.83
0.83
0.83
0.82
0.81
0.81
0.81
0.81
0.81
0.81
0.76
0.76
0.76
0.76
0.75
C
2
2
2
2
2
2
2
2
2
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
Hacquetia 13/1 • 2014, 57–77
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Androsace villosa L.
Achillea clavennae L.
Anthyllis vulneraria L. ssp. polyphylla (DC.) Nyman
Astragalus vesicarius L. ssp. carniolicus (A.Kerner) Chater
Echinops ritro L. ssp. ruthenicus (Bieb.) Nyman
Thesium linophyllon L.
Helianthemum nummularium ssp. obscurum (Celak.) J. Holub
Ranunculus montanus agg.
Lilium bosniacum (Beck) Fritsch
Calamagrostis varia (Schrader) Host ssp. varia
Erica herbacea L.
Alchemilla plicatula Gand.
Festuca duriuscula L.
Anthyllis vulneraria L. ssp. weldeniana (Rchb.) Cullen
Edraianthus tenuifolius (Waldst. & Kit.) A. DC.
Abies alba Miller
Campanula cochlearifolia Lam.
Pedicularis heterodonta Pančić
Gentiana dinarica G. Beck
Ornithogalum collinum Guss.
Sanguisorba minor Scop. ssp. minor
Genista januensis Viv.
Iberis pruitii Tineo
Satureja montana L. ssp. illyrica Nyman
Cerastium malyi (Georgiev) Niketić
Myosotis alpestris F. W. Schmidt
Festuca varia Haenke
Gymnocarpium robertianum (Hoffm.) Newman
Achillea ageratifolia (Sibth. & Sm.) Boiss.
Campanula spatulata Sibth. & Sm. ssp. spatulata
Saxifraga sempervivum C. Koch
Sesleria korabensis (Kümmerle & Jáv.) Deyl
Poa molinerii Balbis
Silene ciliata Pourret
Alyssum montanum L.
Asplenium trichomanes-ramosum L.
Pedicularis brachyodonta Schlosser & Vuk. ssp. brachyodonta
Pedicularis verticillata L.
Allium carinatum L. ssp. pulchellum Bonnier & Layens
1
.
54
54
.
.
.
.
.
.
.
.
.
.
46
46
.
.
.
.
.
.
.
.
.
.
.
38
38
.
69
.
.
.
.
.
.
.
31
31
2
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
40
40
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
3
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
42
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
4
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
56
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
38
.
.
.
.
.
.
.
.
.
.
6
.
.
.
50
50
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
7
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
33
33
33
33
33
.
.
.
.
8
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
44
44
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
9
.
.
.
.
.
50
.
.
.
.
.
.
.
.
.
.
.
60
.
.
.
40
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
10
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
40
40
40
40
.
.
.
.
.
.
.
.
.
.
.
.
.
11
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
12
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
32
32
.
.
13
.
.
.
.
.
.
50
50
50
50
50
50
50
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
A
5
1
1
6
6
9
13
13
13
13
13
13
13
1
1
8
8
9
3
2
2
9
10
10
10
10
1
1
5
1
7
7
7
7
7
12
12
1
1
B
0.74
0.72
0.72
0.69
0.69
0.69
0.69
0.69
0.69
0.69
0.69
0.69
0.69
0.67
0.67
0.65
0.65
0.65
0.63
0.62
0.62
0.62
0.62
0.62
0.62
0.62
0.61
0.61
0.6
0.58
0.56
0.56
0.56
0.56
0.56
0.55
0.55
0.54
0.54
C
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
Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
73
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Gentiana clusii Perr. & Song.
Homogyne sylvestris Cass.
Lonicera caerulea L.
Genista pilosa L.
Saxifraga marginata Sternb. var. coriophylla (Griseb.) Engler
Alyssum scardicum Wettst.
Bunium alpinum ssp. montanum (Koch) P.W. Ball
Carlina corymbosa L.
Carex halleriana Asso
Centaurea spinosociliata ssp. cristata (Bartl.) Dostál
Euphrasia illyrica Wettst.
Ornithogalum orthophyllum Ten.
Festuca pseudovina Hackel ex Wiesb.
Aster bellidiastrum (L.) Scop.
Hieracium murorum L.
Rosa pimpinellifolia L.
Satureja subspicata Bartl. ex Vis.
Potentilla crantzii (Crantz) Beck ex Fritsch
Euphrasia stricta D. Wolff ex J. F. Lehm.
Acinos arvensis (Lam.) Dandy
Scabiosa ochroleuca L.
Minuartia baldaccii (Halácsy) Mattf.
Trifolium noricum Wulfen
Silene parnassica Boiss. & Spruner
Leucanthemum illyricum (Horvatić) Vogt & Greuter
Pedicularis brachyodonta Schlosser & Vuk. ssp. grisebachii (Wettst.) Hayek
Rubus saxatilis L.
Cirsium erisithales (Jacq.) Scop.
Chamaecytisus purpureus (Scop.) Link
Clematis alpina (L.) Miller ssp. alpina
Festuca adamovicii (St-Yves) Markgr.-Dannenb.
Alchemilla labellata Buser
Cerastium alpinum L.
Hieracium alpicola Schleicher ex Gaudin
Primula veris L. ssp. veris
Pulsatilla montana (Hoppe) Reichenb.
Scabiosa triandra L.
Solidago virgaurea L.
Carex digitata L.
1
100
.
.
.
.
.
.
.
.
2
.
.
.
.
.
.
.
.
.
3
.
.
.
.
.
.
.
.
.
4
.
.
.
.
.
.
.
.
.
5
.
.
31
.
.
.
.
.
.
6
.
.
.
30
30
.
.
.
.
7
.
.
.
.
.
.
.
.
.
8
.
.
.
.
.
.
.
.
.
9
.
30
.
.
.
30
30
30
.
10
.
.
.
.
.
.
.
.
.
11
.
.
.
.
.
.
.
.
.
12
.
.
.
.
.
.
.
.
28
13
.
.
.
.
.
.
.
.
.
A
1
4
5
6
6
9
9
9
12
B
0.54
0.54
0.54
0.53
0.53
0.53
0.53
0.53
0.51
C
1
1
1
1
1
1
1
1
1
.
.
.
.
.
77
69
.
.
.
.
.
.
.
.
.
.
92
.
.
.
.
.
8
.
.
.
46
.
.
.
100
20
.
40
.
.
.
.
.
40
40
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
100
.
92
67
.
.
.
8
.
.
.
67
33
.
.
.
25
.
.
.
.
.
50
.
17
.
.
8
25
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
56
.
.
.
50
56
.
.
.
.
.
.
.
44
44
.
19
.
44
.
19
.
.
.
13
.
31
6
.
.
80
100
30
10
.
40
20
30
.
40
30
20
.
.
.
.
.
.
.
.
.
.
30
.
30
.
.
.
.
100
100
67
.
.
.
.
33
.
.
.
67
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
33
33
.
.
.
22
11
44
56
33
11
.
.
11
33
11
.
.
56
.
.
.
.
22
11
33
22
.
11
.
.
80
100
30
.
.
.
.
.
.
70
50
.
.
.
10
50
.
.
20
.
.
.
.
.
10
.
.
20
20
20
40
20
.
20
20
20
.
20
60
20
.
.
.
20
.
.
.
20
.
20
20
20
.
.
.
.
11
37
.
.
63
.
26
5
74
11
11
.
.
5
37
42
5
68
84
37
.
11
.
.
.
.
.
.
.
20
48
.
.
8
28
76
20
88
28
56
.
.
40
40
60
.
.
60
72
24
44
16
32
.
36
8
32
8
100
100
.
.
.
.
.
.
100
.
100
.
.
.
75
75
.
.
25
75
50
25
50
.
.
50
.
.
.
3
13
2
6
11
1
12
6
13
4
13
9
9
7
13
13
3
1
11
12
13
12
13
5
6
13
6
1
3
0.48
0.45
0.44
0.44
0.42
0.4
0.33
0.31
0.3
0.43
0.42
0.4
0.36
0.34
0.33
0.31
0.49
0.48
0.44
0.39
0.39
0.38
0.37
0.32
0.32
0.31
0.3
0.28
0.27
6
6
6
6
6
6
6
6
6
5
5
5
5
5
5
5
4
4
4
4
4
4
4
4
4
4
4
4
4
Hacquetia 13/1 • 2014, 57–77
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Rhododendron hirsutum L.
Crepis praemorsa (L.) Tausch ssp. dinarica (G. Beck) P. D. Sell
Thalictrum aquilegiifolium L.
Dianthus giganteus D’ Urv. ssp. croaticus (Borbás) Tutin
Vicia cracca L.
Festuca rubra L. ssp. rubra
Bellardiochloa violacea (Bellardi) Chiov.
Stachys germanica L. ssp. germanica
Centaurea triumfetti All. ssp. adscendens (Bartl.) Dostál
Other taxa
Globularia cordifolia L.
Teucrium montanum L.
Edraianthus graminifolius (L.) A. DC.
Carex kitaibeliana Degen ex Becherer
Dianthus sylvestris agg.
Phyteuma orbiculare L.
Galium lucidum All.
Asperula cynanchica L.
Carex humilis Leysser
Trinia glauca agg.
Anthyllis montana L. ssp. jacquinii (A. Kerner) Hayek
Saxifraga paniculata Miller
Poa alpina L.
Gentiana verna agg.
Inula ensifolia L.
Genista sylvestris Scop.
Acinos alpinus (L.) Moench
Allium ericetorum Thore
Genista sericea Wulfen
Centaurea rupestris L.
Koeleria splendens C. Presl
Plantago holosteum Scop.
Leontodon crispus Vill. ssp. crispus
Carlina acaulis ssp. simplex (Waldst. & Kit.) Nyman
Hieracium villosum Jacq.
Polygala nicaeensis Risso ex Koch
Carex caryophyllea Latourr.
Laserpitium siler L. ssp. siler
Teucrium chamaedrys L.
1
38
.
.
.
23
.
.
.
.
15
.
.
.
.
62
.
.
.
.
.
.
.
.
.
.
.
.
31
23
.
.
.
.
.
.
.
.
.
.
2
.
.
.
.
20
.
.
.
.
40
.
.
.
.
.
.
.
.
.
.
.
.
.
.
20
.
.
.
.
.
.
.
.
.
.
.
.
.
.
3
.
.
.
25
25
42
.
.
.
.
.
.
.
.
.
.
17
42
.
.
.
.
.
.
8
.
.
.
.
17
.
33
42
25
33
.
8
.
.
4
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
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.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
.
6
13
25
.
38
.
.
.
.
.
88
.
.
.
.
.
.
.
.
31
.
31
38
.
6
.
.
6
.
.
.
.
.
.
.
.
.
.
6
40
.
10
.
.
.
.
.
.
.
60
.
.
40
.
.
.
30
.
30
.
.
30
.
.
.
.
.
.
20
10
.
.
.
.
40
.
.
.
7
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
33
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
33
33
.
8
33
33
33
.
.
11
22
.
33
11
22
67
.
.
11
.
.
.
.
33
.
22
.
.
.
11
.
22
.
.
.
.
.
.
.
.
.
11
.
9
.
.
.
.
.
.
.
.
.
.
70
.
.
10
.
.
.
50
.
.
.
50
.
.
20
.
.
.
.
.
.
.
10
.
.
20
.
.
40
10
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
20
.
.
.
.
.
20
.
20
20
.
.
.
.
11
.
42
.
5
5
.
26
11
5
.
.
.
11
.
.
11
11
.
5
.
42
.
.
.
.
.
11
16
.
.
21
.
.
.
.
.
.
.
.
12
4
40
32
40
.
28
36
40
52
.
.
56
28
12
24
4
4
.
44
.
60
44
8
28
.
.
36
.
12
4
8
.
.
.
.
.
.
.
.
13
.
.
.
.
.
.
25
50
.
.
.
.
50
.
.
25
.
.
25
.
.
.
.
25
.
.
25
.
.
.
.
.
.
.
.
.
.
.
25
A
6
11
12
12
1
3
12
12
12
2
4
5
13
6
5
13
3
3
12
8
12
12
5
5
9
10
5
1
1
6
11
3
3
3
3
6
7
7
9
B
0.27
0.27
0.27
0.26
0.25
0.24
0.23
0.48
0.47
0.46
0.46
0.46
0.43
0.42
0.41
0.41
0.4
0.39
0.38
0.36
0.36
0.35
0.3
0.29
0.29
0.29
0.28
0.26
0.25
0.25
0.18
0.49
0.49
0.49
0.49
0.49
0.49
0.49
0.49
C
4
4
4
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
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Biscutella laevigata L.
Anthericum ramosum L.
Hippocrepis comosa L.
Gymnadenia conopsea (L.) R. Br.
Asplenium ruta-muraria L. ssp. ruta-muraria
Trifolium montanum L.
Sanguisorba minor Scop. ssp. muricata Briq.
Eryngium amethystinum L.
Thalictrum minus L. ssp. minus
Pinus mugo Turra
Anthyllis vulneraria L. ssp. alpestris Ascherson & Graebner
Inula hirta L.
Crepis chondrilloides Jacq.
Gentiana utriculosa L.
Gentiana lutea L. ssp. symphyandra (Murb.) Hayek
Thymus longicaulis C. Presl
Erysimum sylvestre (Crantz) Scop. ssp. sylvestre
Trifolium alpestre L.
Thesium divaricatum Jan ex Mert. & Koch
Pimpinella saxifraga (L.) Loret & Barrandon
Scorzonera austriaca Willd.
Senecio doronicum (L.) L. ssp. doronicum
Euphorbia cyparissias L.
Filipendula vulgaris Moench
Asplenium trichomanes L. ssp. trichomanes
Rhinanthus aristatus Celak.
Potentilla australis Krasan
Amelanchier ovalis Medicus
Buphthalmum salicifolium L.
Briza media L. ssp. media
Stachys recta L. ssp. subcrenata (Vis.) Briq.
Saxifraga tridactylites L.
Arenaria serpyllifolia L. ssp. serpyllifolia
Sedum acre L.
Juniperus communis L.
Euphrasia dinarica (G. Beck) Murb.
Silene saxifraga L.
Dianthus integer Vis.
Armeria canescens (Host) Boiss. ssp. canescens
1
.
.
31
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.
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15
.
23
23
.
.
31
.
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2
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.
3
.
.
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.
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.
.
.
8
.
.
.
.
17
.
.
.
8
.
.
.
25
.
.
.
8
.
.
.
17
8
.
.
.
.
.
17
.
.
4
.
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5
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56
.
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6
.
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25
25
.
.
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.
19
6
.
.
10
30
30
.
30
.
.
.
.
.
.
30
30
.
.
.
.
.
.
.
20
20
20
20
20
20
10
.
.
.
.
.
10
20
.
.
.
7
.
.
.
.
33
33
.
.
33
.
.
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8
.
.
.
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.
.
.
.
.
.
11
.
.
.
.
.
22
.
.
.
.
.
.
.
11
11
.
.
.
.
.
.
.
22
.
.
.
9
.
.
.
.
.
30
30
.
.
.
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.
.
.
.
.
.
.
.
.
30
.
10
10
10
.
.
.
20
.
.
10
.
.
.
.
.
.
.
10
.
.
.
20
.
.
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.
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.
.
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.
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.
20
.
.
.
.
.
.
.
.
.
.
20
20
20
.
.
.
.
20
.
.
11
.
32
.
.
.
.
.
.
.
42
53
11
.
.
.
5
.
.
.
21
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
16
.
12
36
32
.
.
.
.
.
32
.
16
24
20
.
.
.
16
.
.
.
64
.
8
.
.
.
.
.
.
.
.
.
.
8
8
20
.
.
20
8
13
50
.
.
.
.
.
.
.
.
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25
.
.
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.
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.
A
5
12
1
6
6
9
5
5
7
11
11
12
13
6
6
12
1
1
8
12
1
3
6
6
6
6
6
6
9
10
10
10
5
5
12
8
3
12
5
B
0.47
0.47
0.46
0.46
0.46
0.46
0.44
0.44
0.44
0.44
0.43
0.43
0.41
0.4
0.4
0.39
0.38
0.38
0.37
0.37
0.36
0.35
0.35
0.35
0.35
0.35
0.35
0.35
0.35
0.35
0.35
0.35
0.34
0.34
0.34
0.32
0.3
0.28
0.27
C
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
Hacquetia 13/1 • 2014, 57–77
76
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Muscari botryoides (L.) Miller
Frangula rupestris (Scop.) Schur
Campanula scheuchzeri Vill.
Senecio papposus (Reichenb.) Less. ssp. papposus
Luzula campestris (L.) DC.
Scabiosa columbaria L. ssp. portae (A. Kerner ex Huter) Hayek
Polygala alpestris Reichenb. ssp. croatica (Chodat) Hayek
Linum narbonense L.
Asperula aristata L. ssp. scabra (J. & C. Presl) Nyman
Ruta graveolens L.
Seseli elatum ssp. gouanii (Koch) P.W. Ball
Cytisus pseudoprocumbens Markgraf
Thymus striatus Vahl
Scabiosa columbaria L. ssp. columbaria
Cotoneaster nebrodensis (Guss.) C. Koch
Scorzonera villosa Scop.
Peltaria alliacea Jacq.
Valeriana montana L.
Anthyllis vulneraria L.
Satureja subspicata ssp. liburnica Šilić
Galium anisophyllon Vill.
Stachys recta agg.
Galium verum L. ssp. verum
Veronica austriaca L. ssp. austriaca
Gentianella crispata (Vis.) J. Holub
Brachypodium pinnatum (L.) Beauv. ssp. pinnatum
Primula veris L. ssp. columnae (Ten.) Lüdi
Arabis scopoliana Boiss.
Festuca amethystina L. ssp. kummeri (G. Beck) Markgr.-Dannenb.
Rhamnus alpinus L. ssp. fallax (Boiss.) Maire & Petitmengin
Poa badensis Haenke ex Willd.
Iris reichenbachii Heuffel
Mercurialis ovata Sternb. & Hoppe
Lilium carniolicum Bernh. ex Koch
Allium senescens L. ssp. montanum (Fries) Holub
Galium mollugo L.
Vicia incana Gouan
Euphorbia fragifera Jan
Viola hirta L.
1
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8
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8
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2
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.
3
.
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.
8
8
8
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8
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.
4
.
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.
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.
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.
5
.
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13
13
.
.
.
.
6
.
6
.
6
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.
6
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10
.
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7
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8
.
.
.
.
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.
11
11
.
.
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.
.
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9
.
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10
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.
.
.
11
11
5
.
.
.
.
.
.
.
26
.
32
.
.
5
11
12
8
8
12
.
28
12
.
4
.
12
.
8
8
8
8
4
13
.
.
.
.
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.
.
.
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.
.
.
.
.
.
A
12
12
4
5
5
12
8
8
6
12
1
11
12
12
12
11
B
0.27
0.27
0.25
0.24
0.24
0.23
0.22
0.22
0.2
0.17
0.16
0.16
0.16
0.16
0.16
0.11
C
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Eva Kabaš et al.: Numerical evaluation of grasslands dominated by Sesleria juncifolia agg. in Serbia
77
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Hyssopus oficinalis L. ssp. aristatus (Godron) Briq.
Veronica spicata ssp. barrelieri (Schott ex Roemer & Schultes) Murb.
Hypochoeris maculata L.
Campanula justiniana Witasek
Vincetoxicum hirundinaria Medicus
Plantago media L.
Sorbus aria (L.) Crantz
Chamaecytisus hirsutus (L.) Link
Linum catharticum L.
Iris pallida ssp. illyrica (Tomm. ex Vis.) K. Richt.
Anemone nemorosa L.
Artemisia alba Turra
Euphorbia verrucosa L.
Hypericum perforatum L.
Saxifraga crustata Vest
Thlaspi praecox Wulfen
Nevena Kuzmanović
Eva Kabaš