Numerical analysis of the order Scorzoneretalia villosae
Massimo Terzi*
Abstract: The Scorzoneretalia villosae (= Scorzonero-Chrysopogonetalia) order was originally defined to describe
the sub-Mediterranean grasslands of western Croatia, but its distribution range has since been extended
to Albania, Bosnia and Herzegovina, Bulgaria, Italy, Kosovo, Montenegro, Slovenia and Serbia. Major syntaxonomic
revisions of the order have been performed at the regional or national scale and show inconsistencies
both for the syntaxonomic schemes and relative diagnostic taxa. This situation presents some difficulties in
comparing or transferring findings between regions or when using syntaxonomic information for practical
purposes. To tackle these inconsistencies, nearly nine hundred relevés already assigned to the Scorzoneretalia
villosae were classified with the goal of establishing syntaxonomic relationships among associations. Diagnostic
taxa of the main clusters of relevés were identified via Indicator Species Analysis (ISA). Results of statistical
analyses were then interpreted from a syntaxonomic standpoint. Associations were ordinated (non-metric multidimensional
scaling) on the basis of taxa frequencies in order to visualise their floristic and chorological relationships.
Based on the results, associations were grouped within four alliances: Scorzonerion villosae, Chrysopogono-Saturejion
subspicatae, Saturejion subspicatae and Centaureion dichroanthae were assigned to two new
suborders, the meso-xerophytic Scorzonerenalia villosae and the xerophytic Koelerienalia splendentis, classified
within the Scorzoneretalia villosae of the Festuco-Brometea. The order covers the western part of the Balkan
Peninsula and, in the northern part of its range, the southeastern portion of the pre-Alpine sectors. Floristic
similarities between the Scorzonerenalia villosae and other eastern meso-xerophytic syntaxa of the class have
been highlighted. I review the nomenclature for the order, describing or validating two suborders and four associations
for the first time, and lectotypifying three syntaxa .
Keywords: Balkan Peninsula; dry grassland; Festuco-Brometea; phytosociological nomenclature; ScorzoneroChrysopogonetalia;
syntaxonomy.
Nomenclature: Taxonomic nomenclature follows the Euro+Med Plantbase (http://ww2.bgbm.org/EuroPlusMed/,
accessed 1 June 2014) and subordinately Conti et al. (2005) and the Flora Croatica Database (FDC;
http://hirc.botanic.hr/fcd/Search.aspx). Syntaxonomical nomenclature refers to Terzi (2011) if not indicated
otherwise.
Abbreviations: ICPN = International Code of Phytosociological Nomenclature (Weber et al. 2000); IndSp =
Indicator Species; IndVal = Indicator Value index (Dufrêne & Legendre 1997); ISA = Indicator Species Analysis;
NMDS = Nonmetric Multi-Dimensional Scaling (Kruskal 1964; Mather 1976).
Submitted: 12 June 2014; revised version submitted: 11 November 2014; accepted: 12 November 2014
Co-ordinating Editor: Erwin Bergmeier
© 2015 Gebrüder Borntraeger, 70176 Stuttgart, Germany
DOI: 10.1127/phyto/2015/0009
www.borntraeger-cramer.de
0340-269X/2015/0009 $ 9.90
Phytocoenologia Vol. 45 (2015), Issue 1–2, 11–32 Research Paper
Published online June 2015
*Author’s address: Institute of Bioscience and Bioresources (IBBR), National Research Council of Italy (CNR), Via Amendola 165/a,
70126 Bari, Italy; massimo.terzi@ibbr.cnr.it.
Introduction
The sub-Mediterranean dry grasslands of the Adriatic
western Balkan Peninsula were united by Horvatić and
Horvat (Horvatić 1958, 1963; Horvat 1962) within the
Scorzonero-Chrysopogonetalia and assigned to the
Brachypodio-Chrysopogonetea. According to these authors,
the order would substitute the continental inland
vegetation of the Festuco-Brometea towards the Adriatic
coastal strip and give way to the eu-Mediterranean vegetation
of the Cymbopogono-Brachypodietalia at lower
altitude, near the sea. The concept of a high-rank syntaxon
with the intermediate role between the eu-Mediterranean
and continental vegetation was subsequently
extended to the whole of Southern Europe by Barbero &
Loisel (1972) who included the Scorzonero-Chrysopo-
eschweizerbart_xxx
12 Massimo Terzi
gonetalia into the European class Brachypodio-Brometea
nom. illeg. (Art. 29c of the International Code of Phytosociological
Nomenclature, ICPN, Weber et al. 2000).
In the following years, the biogeographic context of
the Scorzonero-Chrysopogonetalia was interpreted differently
and the order was classified within the FestucoBrometea
and the Thero-Brachypodietea or it was divided
into two new orders, Scorzoneretalia villosae and
Koelerietalia splendentis, which were classified in the
aforementioned classes (e.g., Horvatić 1973; Blečić &
Lakušić 1976; Royer 1991). In the following text, reference
is made to the Scorzoneretalia villosae instead of the
Scorzonero-Chrysopogonetalia for reasons explained be-
low.
The distribution range of syntaxa of the Scorzoneretalia
villosae was at first limited to the western side of Croatia
(Horvat 1931; Horvatić 1934, 1949), and it was subsequently
extended to other countries: Italy (Ferlan &
Giacomini 1957), Slovenia (Tomazić 1941), Bosnia and
Herzegovina (Kovačević 1959), Serbia (Nikolić & Diklić
1966), Montenegro (Černjavski et al. 1949), Macedonia
(Jovanović et al. 1986), Albania (Dring et al. 2004) and
Bulgaria (Tzonev 2009).
Many syntaxonomic revisions of the order have been
performed at national or regional scales, the most influential
being those of Horvatić (1958, 1963, 1975), Horvat
(1962), Wendelberger (1965), Barbero & Loisel (1972),
Horvat et al. (1974), Blečić & Lakušić (1976), Jovanović
et al. (1986), Poldini (1989), Royer (1991), Feoli Chiapella
& Poldini (1993), Poldini & Kaligarič (1997), Redžić
(1999), Dring et al. (2004), Antonić et al. (2005) and
Trinajstić (2008). Nonetheless, there has not as yet been
no revision that takes into consideration all the associations
across the entire distribution range of the order. Papers
dealing with the Scorzoneretalia villosae very often
refer to different syntaxonomic arrangements (see Terzi
2011) and indicate different character and differential
species of high-rank syntaxa. Moreover, in many cases,
new syntaxonomic schemes were proposed without any
indication of the data set from which they were obtained
nor the statistical methods utilised. This causes difficulties
in comparing or transferring results from one region
to another or when using syntaxonomic information for
practical purposes, such as in the context of biodiversity
conservation.
In addition, inconsistencies also occur in syntaxa
names, and many contradictory or incorrect names are,
in fact, used in scientific and technical literature. To solve
this problem, I reviewed the nomenclature for the order
in a previous publication (Terzi 2011). In this study, I
have additionally found other papers rarely quoted in
scientific literature (e.g., Černjavski et al. 1949; Kovačević
1959; Tomić-Stanković 1970), where some pivotal names
were validly published for the first time. For example, the
order name Scorzoneretalia villosae was validly published
for the first time by Kovačević (1959).
The syntaxonomic and nomenclatural inconsistencies
highlighted above are mainly due to the lack of an overall
and supranational revision of phytosociological literature
with modern statistical methods. The aim of this study
was twofold: (1) to establish the syntaxonomic relationships
among associations along the entire distribution
range of the Scorzoneretalia villosae, and (2) to identify
diagnostic species of high rank syntaxa using statistical
methods.
Data and Methods
Data set
The data set consisted of 936 relevés already assigned to
the Scorzoneretalia villosae and recorded in Italy (398
relevés), Slovenia (115), Croatia (338), Bosnia and Herzegovina
(42), Serbia (18), Montenegro (22) and Bulgaria
(3) (Fig. 1 and Supplement S1). Incomplete relevés such
as those from Albania quoted in Buzo (1991) were not
taken into consideration.
Some taxa were combined under the same tag if their
differentiation in tables was uncertain or not possible;
however, they were treated separately when discussing
their diagnostic roles if some of them were traditionally
dealt with as diagnostic species. I considered Koeleria
splendens C. Presl also for the western Balkans even
though some authors questioned its presence in that area
(Brullo et al. 2009; see also Quintanar & Castroviejo
2013). Records of Koeleria lobata in the northeast of Italy
(Feoli Chiapella & Poldini 1993) were assigned to K.
splendens. In fact, according to Brullo et al. (2009), K. lobata
should be excluded from the flora of Italy. Taxa
quoted only at the genus level were omitted from the data
set as well as bryophytes and lichens. Abundance-dominance
values of the Braun-Blanquet scale were converted
into the ordinal scale as proposed by van der Maarel
(1979).
Plot size
Information regarding plot sizes was missing in 78 relevés.
For the relevés reported by Poldini (1989) and Feoli-Chiapella
& Poldini (1993), I considered an average
plot size of 85 m
2
, because each relevé was recorded on
an area ranging 70−100 m
2
(Poldini, personal communication).
For the entire data set, plot size varied from 4 to
2000 m
2
. However in 90% of cases, it ranged between 10
and 150 m
2
with a median of 85 m
2
and a mean of 90 m
2
.
These values are clearly higher than those proposed
for herbaceous vegetation by Chytrý & Otýpková
(2003).
Even if the plot size could affect classification and
ordination results (Dengler et al. 2009), the knowledge of
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 13
such effects is insufficient (Otýpková & Chytrý 2006).
Relevés with an unusually large plot size (> 200 m
2
) and
the small-sized ones (< 10 m
2
) were deleted from the data
set. However, the type-relevés of associations described
in literature were always kept even if their plot size
exceeded the aforementioned thresholds.
Resampling and outliers
In order to reduce redundancy of information, relevés
representing similar ecological conditions, as reflected by
their floristic composition, were excluded from the matrix.
Once the Sørensen distance was calculated between
each pair of relevés on the basis of presence/absence of
species data, starting from the lowest distance value up to
the arbitrary threshold of 0.3 (see also Westhoff & van
der Maarel 1980), one relevé for each pair was randomly
deleted (De Cáceres et al. 2008); if the selection involved
a nomenclatural type, it was always included.
Because outlier relevés can influence multivariate analysis
(McCune & Grace 2002), they were identified using
the outlier analysis in PC-ORD 6.11 (McCune & Mefford
2011). Once the Sørensen distance matrix on presence/absence
data was calculated, relevés that were more
than two standard deviation units away from the mean
were removed. Type relevés of the following associations
turned out to be outliers and they were removed together
with the remaining relevés of the associations: Lino-Gypsophiletum
glomerati, Potentillo-Achilleetum clypeolatae,
Poo-Saturejetum subspicatae, Physospermo-Saturejetum
montanae, Saturejo-Festucetum dalmaticae and Artemisio-Salvietum
officinalis. After resampling and outlier
analysis, the data matrix consisted of 649 relevés.
Cluster analysis
Relevés were classified and, for the main clusters of relevés,
IndSp were identified by ISA (Dufrêne & Legendre
1997). Cluster analysis and ISA were based on relevés
(instead of synoptic tables) to achieve a higher accuracy
of results. All the statistical analyses described below
were performed by the software PC-ORD 6.11 (McCune
& Mefford 2011).
Two agglomerative clustering methods, which were
previously suggested as good choices for dealing with
community data, were used: the flexible beta method, β =
–0.25, with Bray and Curtis distance measure with and
without standardisation by relevé totals (i.e., Sørensen
and Relative Sørensen distance measures); and Ward’s
method on a chord distance matrix (Ward 1963, Legendre
& Gallagher 2001, McCune & Grace 2002). Species values
(y) were transformed (y’) as follows: y’ = y
x
, with x
ranging from 1 (untransformed data) to 0.5 (lowered species
value), 0.25 (near presence/absence) and 0 (presence/
absence). In total, 12 dendrogram solutions were ob-
tained.
Since this paper focuses on high rank syntaxa, I considered
only the first 15 partitioning levels for every dendrogram,
and each ISA was performed among relevant
clusters of relevés in order to identify the partition level
with the highest number of significant (p < 0.01) IndSp as
a result of a Monte Carlo test with 5000 permutations.
Fig. 1. Localities of relevés (square symbols). The shaded area corresponds to the distribution area of the Centaureion dichroanthae
and Hypochoeridenion maculatae. The Scorzonerenion villosae, Chrysopogono-Saturejion subspicatae and Saturejion
subspicatae lie in the remaining part of the distribution range of the Scorzoneretalia villosae (full squares), with the
Chrysopogono-Saturejion subspicatae mostly at lower altitude. Empty squares indicate localities of relevés that should be
excluded from the Scorzoneretalia villosae. Star symbols refer to the Danthonio-Brachypodion described by Boșcaiu (1972).
Circles indicate the capital cities in the area.
eschweizerbart_xxx
14 Massimo Terzi
The ISA was performed on presence/absence data of taxa
present in at least three relevés (Tichý & Chytrý 2006:
813). Of the 12 dendrograms, the one containing the
node with the largest number of IndSp was chosen as the
best-clustering solution. The number of IndSp has been
suggested as a criterion to compare dendrograms (Aho et
al. 2008, Tichý et al. 2010). The largest number of IndSp
also identified the most informative node, where the selected
dendrogram was pruned (McCune & Grace 2002).
To increase interpretability of results – starting from the
first level with all the relevés in only one cluster and descending
the dichotomic hierarchy of the dendrogram –
each significant IndSp was assigned to the cluster of relevés
where its IndVal first reached the maximum value
(Dufrêne & Legendre 1997). In this way, each significant
IndSp was associated with only one cluster of relevés.
The main clusters of relevés were interpreted in syntaxonomic
terms. For each of them, I selected a set of diagnostic
taxa. These were chosen among the significant
IndSp associated with that cluster or with its further subdivisions
on the basis of information reported in scientific
literature (Supplement S2) and personal judgement.
That is, among taxa already judged as character or differential
taxa, I selected the ones whose diagnostic roles
were coherent with results of the ISA. Although each IndSp
can be considered a diagnostic species, the subjective
selection was needed because the small size of the data set
did not allow overall evaluation of the sociological roles
of all plant taxa (Terzi et al. 2010). The terms ingressive
and transgressive species were used according to the definitions
given by Poldini & Sburlino (2005).
Ordination
For each association, the diagnosis containing the nomenclatural-type
relevé or subordinately the original diagnosis
were selected (Supplement S1). Association diagnoses
with fewer than four relevés were ignored and,
when available, another diagnosis for those associations
was used. Given the importance of the Scorzonero-Danthonietum
calycinae for the nomenclature of the order
and its wide distribution range, more diagnoses were
considered (see below). Some associations from western
Romania, previously assigned to the Danthonio-Brachypodion
and classified within the same Balkan order
(Brachypodio-Chrysopogonetalia nom. inval.) together
with alliances of the Scorzoneretalia villosae (Boșcaiu
1972), were also added for comparative purposes.
In the association x taxa matrix, taxa variables were
represented by percentage frequencies in the phytosociological
tables. For the associations described only by synoptic
tables, constancy classes were substituted with the
central values of corresponding frequency ranges. Before
ordination, outlier analysis in PC-ORD (McCune &
Mefford 2011) was used to identify and remove outlier
associations (Artemisio-Salvietum officinalis, Poo-Saturejetum
subspicatae, Genisto-Caricetum mucronatae
[from Horvat et al. 1974] and Leontodonto-Seslerietum
calcariae). Nonmetric multidimensional scaling ordination
(NMDS; Kruskal 1964; Mather 1976) was performed
with the chord distance in the PC-ORD slow and thorough
autopilot mode (McCune & Mefford 2011).
Life-form and chorological spectra
For each association, life-form and chorological spectra
were calculated, weighted by species frequencies. Life
forms and chorotypes were selected from works of Poldini
(1991) and, to a lesser extent, Pignatti (1982),
Horvatić et al. (1968), Assyov et al. (2006) and Lakušić &
Redžić (1991). The following chorotypes were used: Alpine;
Arctic-Alpine; Atlantic; Atlantic-Mediterranean;
Circumboreal; Eurasian; European; Southeastern European,
including Balkan species; Eurosiberian species; Illyrian
species, composed of south- and north-Illyrian
species sensu Poldini (1991) and endemics of the eastern
and north Adriatic Region; Steno-Mediterranean; EuryMediterranean;
Southern European orophyte, including
Montane-Mediterranean sensu Pignatti (1982); Paleotemperate;
Pontic species, including Mediterranean-Pontic
species sensu Pignatti (1982); and widespread plants. Joint
plots with an r
2
cut-off of 0.30 were used to show the
strengths of the relationships between life-forms and
chorotypes and ordination scores.
Phytosociological nomenclature
Decisions about nomenclature were taken following the
3rd edition of the ICPN (Weber et al. 2000), in addition
to the suggestions of Willner et al. (2011). Complete syntaxon
names are listed with specific epithets and names of
authors in Appendix 1.
Results and discussion
Ward’s clustering
The clustering solution obtained using Ward’s method on
square-root-transformed data contained the node with
the largest number of IndSp followed by those obtained
with fourth-root transformed data and the flexible-beta
method with relative Sørensen distance on presence/absence
data (491, 489 and 486 IndSp, respectively). Once
the three dendrograms were pruned, they showed
roughly similar groups of associations. Main differences
existed in the positions of relevés of the Ononido-Brometum
condensati, Teucrio-Chrysopogonetum grylli, Leontodonto-Seslerietum
calcariae and in some other asso-
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 15
ciations described by Trinajstić & Šugar (1972), Trinajstić
(1977, 1999, 2005) and Lakušić & Redžić (1991). These
differences have been considered in the interpretation of
results. The dendrogram obtained with Ward’s method
on square-root data was selected as the best solution and
will be addressed in the following sections.
The dendrogram (Fig. 2) was cut at the seventh partition
level resulting in eight clusters of relevés. Each association
was assigned to a cluster on the basis of the
nomenclatural-type positions. Associations not yet typified
were assigned to clusters containing the vast majority
of their relevés.
Cluster 6H: Avenulo-Brometum erecti, Chamaecytiso-Chrysopogonetum
grylli, Gladiolo-Molinietum arundinaceae,
Onobrychido arenariae-Brometum erecti,
Schoeno-Chrysopogonetum grylli, Gentianello-Brometum
erecti and the relevés assigned to the AvenuloBrometum
by Lasen (1995) that, according to Pignatti &
Pignatti (2014), would be ascribed to the Thlaspio-Trifolietum
pratensis. This cluster can be considered representative
of the Hypochoeridenion maculatae.
Cluster 6V1: Scorzonero-Danthonietum calycinae
(relevés from Poldini 1989; Poldini & Kaligarič 1997).
Cluster 6V2: Scorzonero-Danthonietum calycinae
(other relevés assigned to this association), GlobularioChrysopogonetum
grylli, Festuco-Armerietum canescentis,
Scorzonero-Hypochoeridetum maculatae nom. nud.
(only one relevé), Carici-Scabiosetum leucophyllae,
Globulario-Scabiosetum leucophyllae, Stipo-Genistetum
dalmaticae, Ononido-Brometum condensati, TeucrioChrysopogonetum
grylli, Artemisio-Rutetum graveolentis,
Thymo-Teucrietum chamaedryos, Achilleo-Dorycnietum
herbacei. Cluster 6V1 and part of cluster 6V2 are representative
of Scorzonerenion villosae; some associations
of this group must be classified within other alliances or
orders.
Cluster 5C1: Asphodelo-Chrysopogonetum grylli,
Bromo-Chrysopogonetum grylli, Koelerio-Festucetum illyricae,
Bromo-Festucetum lapidosae, Stipo-Salvietum officinalis,
Narcisso-Asphodeletum microcarpi, Festucetum
illyrico-valesiacae, Koelerio-Brachypodietum retusi and
Salvio-Seslerietum juncifoliae.
Cluster 5C2: Dichanthio-Cleistogenetum serotinae,
Salvio-Euphorbietum fragiferae, Seseli-Artemisietum albae,
Centaureo-Chrysopogonetum grylli and LactucoBothriochloetum
ischaemum. Cluster 5C represents the
Chrysopogono-Saturejion subspicatae.
Cluster 5S1: Centaureo-Caricetum humilis (relevés
from Trinajstić 1987; Trinajstić & Pavletić 1990; Trinajstić
et al. 1993; and few others from Poldini 1989), AstragaloSeslerietum
robustae, Seslerio-Juniperetum sibiricae,
Bromo-Seslerietum interruptae, Stipo-Caricetum humilis,
Minuartio-Genistetum pulchellae, Genisto-Globularietum
bellidifoliae, Genisto-Seslerietum tenuifoliae, Genisto-Seslerietum
kalnikensis, and Seslerio-Caricetum hu-
milis.
Cluster 5S2: Centaureo-Caricetum humilis (other relevés
assigned to this association), Saturejo-Caricetum humilis.
The cluster 5S is representative of the Saturejion
subspicatae.
Cluster 5D: Bupleuro-Brometum condensati, Centaureo-Globularietum
cordifoliae, Leontodonto-Seslerietum
calcariae, Bromo-Stipetum eriocaulis, Saturejo-Brometum
condensati and the relevés assigned to Genisto-Caricetum
mucronatae by Poldini (1989). The cluster represents
the Centaureion dichroanthae.
Indicator Species Analysis
The results of the ISA (Supplement S2) showed that IndSp
of the first clustering level (all the relevés belonging
to the same cluster, 3Z in Fig. 2) included taxa of the Festuco-Brometea
and others already considered diagnostic
for the Scorzoneretalia villosae, such as Salvia pratensis
Fig. 2. Ward’s clustering (chord distance) of relevés based on
square-root-transformed species data. Cluster 4Z = suborder
Scorzonerenalia villosae and alliance Scorzonerion villosae;
cluster 4K = suborder Koelerienalia splendentis; cluster
5C = Chrysopogono-Saturejion subspicatae; cluster 5S =
Saturejion subspicatae; cluster 5D = Centaureion dichroanthae;
6H = Hypochoeridenion maculatae; 6V = Scorzonerenion
villosae.
eschweizerbart_xxx
16 Massimo Terzi
subsp. bertolonii (together with S. pratensis subsp. pratensis
in the data set), Plantago holosteum, Festuca illyrica
(with F. stricta subsp. sulcata in the dataset), Stachys officinalis,
Scabiosa triandra, Veronica barrelieri, Medicago
prostrata, Noccaea praecox, and, transgressive from the
class, Eryngium amethystinum, Dorycnium pentaphyllum
subsp. germanicum, Inula hirta, Linum tenuifolium,
and Chrysopogon gryllus.
With the first dendrogram partition, the cluster 4Z resulted
as floristically well defined due to the many IndSp
and the high IndVal. Some of the IndSp have been considered
character species of meso-xerophytic syntaxa of
the Scorzoneretalia villosae (Hypochaeris maculata, Dorycnium
pentaphyllum subsp. herbaceum, Danthonia alpina),
or are transgressive-differential from the FestucoBrometea
(e.g., Ononis spinosa, Filipendula vulgaris,
Plantago media, Euphorbia verrucosa) or ingressive-differential
from other classes (e.g., Buphthalmum salicifolium,
Agrostis capillaris, Trifolium rubens). There are also
many species that are widespread in mesophilous alliances
of the Festuco-Brometea and ingressive from the
Molinio-Arrhenatheretea. IndSp of further subdivisions
of cluster 4Z are numerous, indicative of meso-xerophytic
communities and some of relatively acidic soil
conditions (e.g., Genista germanica, Danthonia decumbens,
Festuca filiformis), but few can be listed as character
taxa (e.g., Scorzonera villosa subsp. villosa, Centaurea
jacea subsp. weldeniana, Knautia illyrica). For this reason,
only one alliance (Scorzonerion villosae), divided
into two sub-alliances (Hypochoeridenion maculatatae
and Scorzonerenion villosae), was considered. It is interesting
to note that some IndSp of clusters 4Z and 6H
were included among the character species of the Brachypodietalia
pinnati (Supplement S2 columns 15 and 16).
Very few IndSp, without high IndVal and transgressive
from the class, were associated with cluster 6V2. In effect,
this cluster includes some associations that must be excluded
from the alliance. The Artemisio-Rutetum graveolentis,
Thymo-Teucrietum chamaedryos and AchilleoDorycnietum
herbacei were originally classified within
the Saturejion montanae (presumably instead of the Saturejenion
montanae) and the Saturejion subspicatae even
if they contained few species of the Scorzoneretalia villosae.
The Carici-Scabiosetum leucophyllae was originally
classified within the Saturejion subspicatae but then
moved to the Xerobromion erecti (Redžić et al. 1984,
2013; Redžić 1999). There are important floristic differences
with other central European associations of the Xerobromion
erecti and Redžić (1999) proposed a Balkan
suballiance Fumano-Scabiosenion leucophyllae. In fact,
the relationships between the Scorzoneretalia villosae and
Balkan associations classified within the Xerobromion,
and the Brometalia erecti / Artemisio-Brometalia erecti
(Mucina et al. 2009) in general, remain to be clarified; the
Carici-Scabiosetum leucophyllae and Globulario-Scabiosetum
leucophyllae share important species with the Scorzoneretalia
villosae, where they are provisionally arranged.
Two other associations of this group, the Stipo-
GenistetumdalmaticaeandtheTeucrio-Chrysopogonetum
grylli, have been provisionally moved to the Saturejion
subspicate and the Centaureion dichroanthae, respectively,
due to the presence of a few but important diagnostic
species of these alliances, such as Centaurea rupestris,
Satureja subspicata, and Cytisus purpureus (see
Redžić et al. 1984; Sburlino et al. 2008).
On the other side of the dendrogram, IndSp of cluster
4K grouped taxa already regarded as diagnostic to more
xerothermic syntaxa of the Scorzoneretalia villosae (i.e.,
Koelerietalia splendentis in the syntaxonomic scheme of
Horvatić 1973). IndSp of cluster 5C (Chrysopogono-Saturejion
subspicatae) or of its further subdivisions were
composed of many Mediterranean and Illyrian taxa.
Some can be considered diagnostic of the alliance. Others
are ingressive from the Mediterranean Thero-Brachypodietea,
while others are common to southern xerothermic
syntaxa of the Festuco-Brometea (e.g., Artemisia
alba, Melica ciliata, Petrorhagia saxifraga, Bothriochloa
ischaemum, Asperula aristata). Cluster 5C2 includes associations
of the northern part of the alliance range (Karst
area surrounding Trieste and Gorizia and the Island of
Krk), which is an area without proper summer drought
and with cooler winters than in Dalmatia (Hršak 2003;
Poldini 2009). Nonetheless, due to edaphic and mesoclimatic
conditions (Ferlan & Giacomini 1957; Poldini
1989, 2009), associations of this cluster show high percentage
of Mediterranean chorotypes (Supplement S3)
and can be assigned to the Chrysopogono-Saturejion subspicatae
owing to the presence of important diagnostic
taxa of this alliance (Table 1).
The IndSp of cluster 5C+5D were shared by the Saturejion
subspicatae and the Centaureion dichroanthae
(e.g. Genista sericea, Inula ensifolia, Scorzonera austriaca,
Plantago argentea). The Centaureion dichroanthae (cluster
5D) was differentiated by ingressive taxa from the
Elyno-Seslerietea and de-Alpine taxa, such as Globularia
cordifolia, Hieracium porrifolium, Polygala nicaeensis
subsp. carniolica and Carduus defloratus. Despite some
floristic similarities with the Saturejion subspicatae, it
was considered an autonomous alliance, vicariant in the
pre-Alpine region (shaded area in Figure 1), for both the
numerous IndSp and Alpine taxa. A very recent study by
Pignatti & Pignatti (2014) extended the distribution area
of the Centaureion dichroanthae up to the Dolomites,
thereby integrating new associations with relevés not included
in my data set. Nevertheless, based on my results
but different from their findings, the Bromo-Stipetum
eriocaulis and the Avenulo-Brometum erecti sensu Lasen
1995 non Feoli Chiapella et Poldini 1993 were classified
as originally proposed by Lasen (1995). The Centaureion
dichroanthae (cluster 5D) also included the relevés assigned
to the Genisto-Caricetum mucronatae by Poldini
(1989). The presence of many species of the Centaureion
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 17
dichroanthae within this association had already been
noted (Poldini 1989). Unfortunately, the original diagnosis
of the Genisto-Caricetum consisted only of a synoptic
table (Horvat et al 1974), and there were no other relevés
from its original area (Gorski Kotar, Croatia). In effect,
Euphorbia triflora subsp. triflora turned out to be among
IndSp of the Centaureion dichroanthae, even though it is
at the edge of its range in the Trnovski Gozd (Slovenia).
Regardless, on the basis of the diagnosis given by Poldini
(1989), the Genisto-Caricetum mucronatae can be classified
within the Centaureion dichroanthae.
The IndSp belonging to the clusters derived from further
divisions of cluster 5S were assigned to the Saturejion
subspicatae (Table 1). Numerous IndSp (but with
low IndVal), many of which are thermophilous taxa,
were associated with cluster 5S1, which represents a transition
towards the more thermo-xeric Chrysopogono-Saturejion
subspicatae.
NMDS
The three-axis solution of the NMDS ordination attained
a minimum stress of 11.7, indicating a fairly usable and
good ordination (McCune & Grace 2002). The proportion
of variance represented by the three axes were 36.9%,
25.4% and 15.5%, respectively, thus explaining 77.8% of
total variation. For simplification purposes, I presented
only the first two dimensions (Fig. 3). The relationships
between ordination scores and chorological spectra (Fig.
3, Supplement S3) showed a gradient along axis 1 from
the chorotypes Steno-Mediterranean (r
2
= 0.40) and Illyrian
(r
2
= 0.60) to Eurosiberian (r
2
= 0.77), Eurasian (r
2
=0.76)
and European (r
2
= 0.59); along axis 2, a second gradient
from Paleotemperate (r
2
= 0.36) and Eury-Mediterranean
(r
2
= 0.51) to Orophyte (r
2
= 0.66) was present. Moreover,
there were positive correlations of therophytes (r
2
= 0.45)
with axis 2 and hemicryptophytes (r
2
= 0.62) with axis 1
and a negative correlation of chamaephytes (r
2
= 0.56)
with axis 1.
Three main groups of associations, roughly corresponding
to clusters 4Z, 5C and 5S+5D, occupied different
parts of the diagram (Fig. 3). Associations of the clusFig.
3. Nonmetric multidimensional scaling ordination diagram (axes 1 and 2). Empty squares indicate Chrysopogono-Saturejion
subspicatae, full circles Saturejion subspicatae; empty circles Centaureion dichroanthae; triangles Scorzonerion villosae;
full squares Danthonio-Brachypodion. Abbreviations: eua = Eurasian; eur = European; esi = Eurosiberian species; ill = Illyrian
species; mds = Steno-Mediterranean; mde = Eury-Mediterranean; oro = Southern European orophyte; pal = Paleotemperate;
h = hemicryptophyte; t = therophyte; ch = chamaephyte. Syntaxa abbreviations in Appendix 1.
eschweizerbart_xxx
18 Massimo Terzi
ter 4Z were on the right side of the ordination space, together
with associations of the Danthonio-Brachypodion.
This group, composed of meso-xerophytic associations,
showed the lowest percentage of Illyrian and Mediterranean
species and high percentages of Eurasian, European
and Eurosiberian species (Fig. 3; Supplement S3). As expected,
orophilous species were well-represented within
the associations of the Saturejion subspicatae and the
Centaureion dichroanthae, which are partly overlapping
in the upper part of the diagram. The Centaureion dichroanthae
is differentiated by higher percentages of Alpine
species (Supplement S3). Associations of cluster 5S1
are located in the upper-left side of the diagram with
some of its associations close to the Chrysopogono-Saturejion
subspicatae, as noted above. These associations
showed the highest percentage of Illyrian species; Mediterranean
chorotypes were more prevalent within the
Chrysopogono-Saturejion subspicatae.
Based on the data, the floristic autonomy of the Festucion
illyricae seems questionable, because this alliance
lacks character species, while its associations (FestucoLinetum
flavi and Festucetum illyrico-valesiacae) share
taxa with both the Scorzonerion villosae (Centaurea jacea
subsp. weldeniana, Plantago media, Dorycnium pentaphyllum
subsp. herbaceum, Knautia illyrica, Prunella
laciniata) and the Chrysopogono-Saturejion subspicatae
(Koeleria splendens, Ornithogalum orthophyllum subsp.
kochii, Bupleurum veronense, Potentilla tommasiniana).
These associations, together with the Festuco-Poetum
bulbosae, Ononido-Brometum condensati and Euphorbio-Chrysopogonetum
grylli, are also intermediate between
these two alliances from a chorological standpoint.
The ratio of the sum of Eurasian, European and Eurosiberian
species and the sum of Mediterranean species is
generally equal to or less than 1 in the ChrysopogonoSaturejion
subspicatae and near to or more than 2 in the
Scorzonerion villosae. However, a precise comparative
analysis is difficult when dealing only with synoptic tables,
which often lack some of the rarer taxa (see synoptic
tables in Horvatić 1963 and Horvat et al. 1974). For this
reason, the classification of these associations has to be
considered as provisional, and new data are required.
Syntaxonomy, ecology and distribution
The numerical analysis of the Scorzoneretalia villosae differentiated
two main groups of associations, corresponding
to meso-xerophytic meadows (Scorzonerion villosae)
on the one hand and sub-Mediterranean thermo-xerophytic
grasslands (Chrysopogono-Saturejion subspicatae)
and pre-Alpine/ Dinaric Mediterranean-montane pastures
on the other (Centaureion dichroanthae and Saturejion
subspicatae).
Pastures and stony grasslands on shallow calcareous
soils are classified within the Chrysopogono-Saturejion
subspicatae, Saturejion subspicatae and Centaureion dichroanthae.
The Chrysopogono-Saturejion subspicatae
occurs along the eastern and northern Adriatic coast; at
higher altitude, the alliance gives way to the Saturejion
subspicatae (Dinarides) and the Centaureion dichroanthae
(southeastern part of pre-Alps: shaded area in Fig. 1).
The Scorzonerion villosae was originally conceived to
include the Illyrian-Dinaric dry grasslands on deep and
partly decalcified terra rossa soil and on variably acidic
soils (Horvatić 1949, 1958, 1963). Feoli-Chiapella & Poldini
(1993) unified the Scorzonerion villosae with the Hypochoeridion
maculatae and extended the alliance distribution
area up to the pre-Alpine region in north-eastern
Italy and Slovenia (shaded area in Fig. 1). This resultant
alliance was floristically well differentiated, with few
character species and many differential ones shared with
other meso-xerophytic alliances of the Festuco-Brometea
that are often shared with mesotrophic meadows belonging
to the Molinio-Arrhenatheretea. Similar findings
have been reported by Pipenbaher et al. (2011), specifically
for the Slovenian karst area. Associations of the
Scorzonerion villosae showed a prevalence of hemicryptophytes
and high percentages of European, Eurasian
and Eurosiberian taxa.
On the other hand, Mediterranean and Illyrian taxa
are more concentrated in the other alliances. The Chrysopogono-Saturejion
subspicatae has a higher percentage of
Mediterranean taxa with many ingressive taxa from the
Thero-Brachypodietea s.l., while orophilous (and Mediterranean-montane)
and Alpine taxa penetrate into the
Saturejion subspicatae and Centaureion dichroanthae.
These differences were translated into syntaxonomic
terms with two suborders of the Scorzoneretalia villosae,
Scorzonerenalia villosae (including Scorzonerion villosae)
and Koelerienalia splendentis (including the other three
alliances).
It is interesting to note that already Horvat and
Horvatić (in Horvatić 1958: 69) proposed separating the
associations of the Scorzonerion villosae from those of
the Chrysopogono-Saturejion subspicatae (part of which
was later moved to Saturejion subspicatae) at the order
level. Some years later, Horvatić (1973, 1975) proposed a
new syntaxonomic scheme with the Saturejion subspicatae
and Scorzonerion villosae being assigned to the Scorzoneretalia
villosae and the Chrysopogono-Saturejion
subspicatae being assigned to the Koelerietalia splenden-
tis.
The latter has been reduced here to the rank of suborder
and the alliances Centaureion dichroanthae and Saturejion
subspicatae have been assigned to it. With increasing
altitude (or distance from the sea), the Mediterranean
influence decreases and mesophilous species of the Scorzonerion
villosae may occurr in communities of the Koelerienalia
splendentis. However, the overall floristic and
chorological patterns support the arrangement of alliances
in the two suborders as described above.
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 19
Both suborders are placed within the the Scorzoneretalia
villosae and the Festuco-Brometea. This scheme refers
to a classical interpretation of the class, with geographically
defined orders and with the Scorzoneretalia
villosae lying between the western suboceanic Brometalia
erecti and the eastern sub-continental Festucetalia
valesiacae (Royer 1991; see also Dengler et al. 2012).
Braun-Blanquet (1974) refuted the floristic autonomy of
Scorzoneretalia villosae and considered only one alliance,
Chrysopogono-Saturejion subspicatae, within the Brometalia
erecti. For many other authors, the floristic autonomy
of Scorzoneretalia villosae is due to a set of subMediterranean
and Illyrian species (Horvat 1962; Horvat
et al. 1974; Barbero & Loisel 1972; Royer 1991). Most
Illyrian taxa are concentrated within the xerophytic
Koelerienalia splendentis, which is, consequently, better
differentiated than the Scorzonerenalia villosae. The latter
has fewer Illyrian and sub-Mediterranean taxa but
shares many taxa with neighbouring alliances, especially
with Balkan associations of the Bromion erecti.
The transition between the Bromion erecti and the
Scorzonerion villosae is abrupt in some areas but gradual
in others; it has been related to decreasing inland temperature
(Horvat et al. 1974; Gaži-Baskova & Šegulja
1978; Redžić 1999). Pipenbaher et al. (2013) directly
compared plant traits and floristic compositions of the
Slovenian records of the associations Scorzonero-Danthonietum
calycinae and Onobrychido-Brometum erecti
T. Müller 1966, concluding that two orders are justified
for these vegetation types. However, while in Slovenia
the boundary between the Scorzonerion villosae and the
Bromion erecti is clearly related to the bioclimatic transition
from Mediterranean to more continental types, in
the western Balkan Peninsula the floristic relationships
between the two alliances are more complex (Kovačević
1959; Ritter-Studnička 1972; Ilijanić et al. 1972; Trinajstić
et al. 1981; Lakušić et al. 1982; Royer 1991). Over time,
Mediterranean plants must have migrated inland (i.e.
Lathyrus latifolius, Eryngium amethystinum, Scabiosa
triandra), across the Dinaric Alps penetrating into Balkan
communities of the Brometalia erecti (Horvat et al.
1974; Royer 1991). Ilijanić et al. (1972) highlighted the
similarities between the Globulario-Chrysopogonetum
grylli, recorded in a karst area west of Karlovac (Croatia)
and originally classified within the Brometalia erecti, and
the Scorzonero-Danthonietum calycinae, with taxa such
as Danthonia alpina, Chrysopogon gryllus, Filipendula
vulgaris, Trifolium montanum, Plantago media, Dorycnium
pentaphyllum subsp. herbaceum, Scabiosa triandra,
Globularia bisnagarica, Brachypodium pinnatum and
Prunella laciniata, among others. Many of these taxa can
be considered diagnostic of the Scorzoneretalia villosae
or its subordinated syntaxa.
The nomenclatural reconsideration of the order leads
to the conclusion that the type relevé of the ScorzoneroDanthonietum
calycinae, and consequently of the Scorzonerion
villosae, was recorded in the area of Bosanski
Petrovac, in Bosnia (Kovačević 1959), and brings the
Scorzonerenalia villosae nearer to other meso-xerophytic
alliances of the Balkans. For example, plant communities
from the same area which had been assigned to the
Bromo-Plantaginetum mediae (Kovačević 1959), are very
similar to the Scorzonero-Danthonietum calycinae.
Boșcaiu (1972) reduced the Brachypodio-Chrysopogonetea
to the order level (contravening ICPN Art. 27,
Note 1; Weber et al. 2000) and proposed to extend its distribution
range as far as western Romania, including the
Danthonio-Brachypodion. The new order would constitute
a Balkan vicariant of the western Brometalia erecti
(Boșcaiu 1972). Some authors (Sanda et al. 2008) followed
this approach. It is now clear that the similarities pointed
out by Boșcaiu (1972) regarded only the suborder Scorzonerenalia
villosae even though, among the diagnostic
species of the Danthonio-Brachypodion and Brachypodio-Chrysopogonetalia
(Boșcaiu 1972), only Danthonia
alpina and Chrysopogon gryllus had a significant IndVal
in my analysis (Fig. 3, Supplement S2). The DanthonioBrachypodion
recently has been considered part of the
Cirsio-Brachypodion pinnati, within the Brachypodietalia
pinnati (Dengler et al. 2012), following a proposal to
group all meso-xerophytic syntaxa of the Festuco-Brometea
within only one order. The name of the meso-xerophytic
order would be Brometalia erecti, a name proposed
to be rejected as nomen ambiguum and substituted
with Brachypodietalia pinnati as the next younger valid
name (Dengler et al. 2003). Results of ISA (Supplement
S2: columns 15 and 16) showed that some diagnostic species
of Scorzonerenalia villosae coincide with those identified
for the Brachypodietalia pinnati. Floristic and
chorological similarities of the Scorzonerenalia villosae
with Balkan associations of the Bromion erecti and further
eastwards with the Danthonio-Brachypodion might
suggest to group all these communities within only one
order. Consequently, the Scorzonerenalia villosae (and
thus the order name), or at least its more mesophilous
associations, could be moved to the Brachypodietalia
pinnati while the remaining xerophytic alliances would
be assigned to the Illyrian xerophytic order Koelerietalia
splendentis.
However, the most appropriate model for the FestucoBrometea
should be determined by means of a large-scale
comparative study. No such revision has yet been carried
out, with the exception of the thorough works performed
by Royer (1991) and, for the Balkan Peninsula, by Horvat
et al. (1974). Both these studies support the classical
concept of the class with geographically defined orders.
Therefore, the traditional approach is followed here, considering
two suborders within the Scorzoneretalia villo-
sae.
The distribution range of the Scorzoneretalia villosae
(Fig. 1) covers the western part of the Balkan Peninsula,
omitting the associations described by Tzonev (2009) in
eschweizerbart_xxx
20 Massimo Terzi
Bulgaria and the Artemisio-Salvietum officinalis in Serbia.
The latter contains many taxa indicative of a different ecological
context (Linaria concolor, Koeleria simonkaii, Euphorbia
taurinensis, Achillea clypeolata, Erysimum crepidifolium,
Achillea crithmifolia, Dianthus pelviformis, Thymus
praecox subsp. jankae), and diagnostic taxa of the
Scorzoneretalia villosae were lacking almost entirely.
North- and westward, the order Scorzoneretalia villosae
follows the south-eastern pre-Alpine sectors (Feoli
Chiapelli & Poldini 1993; Lasen 1995; Pignatti & Pignatti
2014) where it came in connection with the Festucetalia
valesiacae. Its syntaxonomic relationships with the
Diplachnion, especially with associations descibed by
Meyer (1976) along the southern pre-Alpine lakes, remain
to be solved. Regarding the Italian Peninsula, some
authors extended the order range to xerophytic and
meso-xerophytic basophilous grasslands of the Apennines
(Bonin 1978; Biondi & Galdenzi 2012). However,
preliminary results of a comparative study among xerophytic
grasslands of the amphi-Adriatic area (Terzi & Di
Pietro 2013) highlighted important floristic differences
between the Balkan and peninsular Italian sides of the
Adriatic Sea, notably Illyrian/south-eastern European
taxa on the Balkan side (e.g., Chrysopogon gryllus, Festuca
illyrica, Festuca valesiaca, Genista spp., Scorzonera
villosa subsp. villosa, Stipa eriocaulis) and other taxa,
mostly Italian endemics, on the other side (e.g., Festuca
circummediterranea, Phleum ambiguum, Cerastium suffruticosum,
Crepis lacera, Scorzonera villosa subsp. columnae,
Thymus spinulosus). As some of these taxa are
often dominant, a differentiation at the order level was
proposed (Terzi & Di Pietro 2013). This idea has been
strengthened by the recent description of some new endemic
orders for peninsular Italian grasslands (Ubaldi
2011; Biondi et al. 2014).
Syntaxonomic scheme and phytosociological
nomenclature
Based on these results, the syntaxonomic scheme below
is proposed together with a set of diagnostic taxa. The
classification of associations described by few relevés or
by synoptic tables is only provisional, and more in-depth
studies are required to determine their correct syntaxonomic
positions. Moreover, associations quoted in Pignatti
& Pignatti (2014) but not included in the analysis,
were added to the scheme for completeness but marked
by an asterisk. Numbers within square brackets refer to
the nomenclatural notes in Appendix 2.
1F. Class: Festuco-Brometea Br.-Bl. & Tx. ex Klika &
Hadač 1944
3Z. Order: Scorzoneretalia villosae Kovačević 1959 [1]
Nomenclatural synonym: Scorzonero villosae-Chrysopogonetalia
grylli Horvatić & Horvat in Horvatić
1963; Scorzoneretalia villosae Horvatić 1973. Holotypus:
Scorzonerion villosae Horvatić ex Kovačević
1959. Diagnostic taxa: Anthyllis vulneraria subsp.
polyphylla; Centaurea jacea subsp. gaudinii; Chrysopogon
gryllus; Cyanus triumfettii; Dorycnium pentaphyllum
subsp. germanicum; Eryngium amethystinum;
Euphorbia nicaeensis; Festuca illyrica; Festuca
valesiaca; Leucanthemum platylepis; Medicago
prostrata; Noccaea praecox; Plantago argentea; Plantago
holosteum; Potentilla heptaphylla subsp. australis;
Salvia pratensis subsp. bertolonii; Scabiosa triandra;
Seseli montanum subsp. tommasinii; Stachys officinalis
subsp. serotina; Veronica barrelieri.
4K. Suborder: Koelerienalia splendentis (Horvatić 1973)
stat. nov. hoc loco (Horvatić 1973: 7)
Holotypus: Chrysopogono grylli-Koelerion splendentis
Horvatić 1973 (that is a syntaxonomic synonym of
Chrysopogono grylli-Saturejion subspicatae Horvat &
Horvatić ex Černjavski, Grebenščikov & Pavlović
1949). Diagnostic taxa: Dianthus sylvestris subsp. tergestinus;
Euphrasia illyrica; Genista holopetala; Genista
sericea; Genista sylvestris subsp. sylvestris; Globularia
cordifolia; Hyacinthella dalmatica; Inula ensifolia;
Koeleria splendens; Linum narbonense; Ornithogalum
orthophyllum subsp. kochii; Potentilla heptaphylla
subsp. australis; Potentilla tommasiniana; Satureja
montana subsp. variegata; Scorzonera austriaca; Seseli
kochii; Stachys recta subsp. subcrenata; Stipa eriocaulis;
Teucrium montanum; Trinia glauca.
5C. Alliance: Chrysopogono grylli-Saturejion subspicatae
Horvat & Horvatić ex Černjavski, Grebenščikov &
Pavlović 1949 [5]
Syntaxonomic Synonym: Chrysopogono grylli-Koelerion
splendentis Horvatić 1973. Lectotypus: Bromo
erecti-Chrysopogonetum grylli Horvatić 1934. Diagnostic
taxa: Achillea nobilis; Astragalus muelleri; Bupleurum
veronense; Carduus nutans subsp. micropterus;
Centaurea cristata; Centaurea spinosociliata;
Centaurea tommasinii; Dianthus ciliatus; Euphorbia
fragifera; Festuca lapidosa; Genista sylvestris subsp.
dalmatica; Onobrychis arenaria subsp. tommasinii;
Onosma echioides subsp. dalmatica; Salvia officinalis;
Satureja montana subsp. variegata.
Associations: ac – Bromo erecti-Chrysopogonetum grylli
Horvatić 1934 [6]; bf – Bromo erecti-Festucetum lapidosae
Trinajstić 1992; cc – Centaureo cristatae-Chrysopogonetum
grylli Ferlan & Giacomini 1957 [7]; dc – Dichanthio
ischaemum-Cleistogenetum serotinae Horvatić
ex Trinajstić in Poldini 1975 nom. mut. propos.; fi – Festucetum
illyrico-valesiacae Horvat in Horvat et al. 1974
corr. Trinajstić 2000; fl – Festuco-Linetum flavi RitterStudnička
1972; ha – Helichryso italici-Armerietum
dalmaticae Horvatić 1963; kb – Koelerio macranthaeBrachypodietum
retusi ass. nov. hoc loco [9]; kf –
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 21
Table 1. Abridged synoptic table of the Scorzoneretalia villosae, from the level of suballiance to the level of class. Diagnostic
taxa are selected from IndSp associated to the clusters in Figure 2, corresponding to the following syntaxa: 5C = Chrysopogono-Saturejion
subspicatae; 5S = Saturejion subspicatae; 5D = Centaureion dichroanthae; 6H = Hypochoeridenion maculatae;
6V = Scorzonerenion villosae. The percentage frequencies of taxa are given in columns. For each taxon, the IndVal and
the associated cluster are indicated in the last two columns. The list of IndSp is reported in the Supplement S2.
5C 5S 5D 6H 6V IndVal Cluster
Number of relevés 159 198 51 98 142
Alliance: Chrysopogono grylli-Saturejion subspicatae
Satureja montana subsp. variegata 42 15 21 1 4 75 5C2
Bupleurum veronense 40 4 . . 4 35 5C
Salvia officinalis 31 8 . . . 27 5C1
Onosma echioides subsp. dalmatica 18 4 . . 7 13 5C
Euphorbia fragifera 15 4 . . 1 19 5C2
Centaurea cristata 15 6 . . . 16 5C2
Carduus nutans subsp. micropterus 14 5 . . 2 23 5C1
Genista sylvestris subsp. dalmatica 11 5 . . 4 13 5C1
Centaurea spinosociliata 9 3 . . . 14 5C1
Centaurea tommasinii 9 1 . . 3 13 5C1
Astragalus muelleri 7 2 . . . 10 5C1
Festuca lapidosa 4 . . . . 8 5C1
Achillea nobilis 4 1 . . 2 6 5C1
Onobrychis arenaria subsp. tommasinii 3 . . . . 7 5C2
Alliance: Saturejion subspicatae
Centaurea rupestris 9 51 8 . 8 32 5S
Sesleria tenuifolia subsp. tenuifolia 3 42 . . 1 38 5S1
Satureja subspicata . 42 2 . 1 39 5S
Anthyllis montana subsp. jacquinii . 29 4 . 1 25 5S
Jurinea mollis 3 20 . . . 16 5S
Muscari botryoides 5 19 . 2 4 11 5S
Edraianthus tenuifolius 5 16 . . 2 29 5S1
Gentiana verna subsp. tergestina . 14 4 . 4 11 5S2
Pulsatilla montana . 13 . 1 2 13 5S2
Globularia meridionalis . 11 . . . 26 5S1
Crepis chondrilloides 1 11 . . . 9 5S
Bunium alpinum subsp. montanum 2 10 . . 1 23 5S1
Narcissus poeticus subsp. radiiflorus . 9 . 5 1 7 5S
Astragalus croaticus . 7 . . . 19 5S1
Iris pallida subsp. illyrica 3 6 . . . 4 5S1
Klasea radiata subsp. cetinjensis . 4 . . . 10 5S1
Alliance: Centaureion dichroanthae
Sesleria caerulea . . 87 16 . 80 5D
Cytisus purpureus . 3 54 16 3 45 5D
Stachys recta subsp. labiosa 1 . 52 7 1 48 5D
Hieracium porrifolium . . 44 . . 44 5D
Carex mucronata . 1 38 1 . 37 5D
Lomelosia graminifolia subsp. graminifolia . 1 35 3 . 33 5D
Polygala nicaeensis subsp. carniolica . . 31 2 . 30 5D
Leontodon incanus . 2 31 1 . 29 5D
eschweizerbart_xxx
22 Massimo Terzi
5C 5S 5D 6H 6V IndVal Cluster
Number of relevés 159 198 51 98 142
Euphorbia triflora subsp. kerneri . . 25 5 . 23 5D
Cytisus pseudoprocumbens . 7 23 7 4 15 5D
Carduus defloratus . . 21 3 . 20 5D
Gypsophila repens . . 19 3 . 18 5D
Centaurea dichroantha . . 19 6 . 17 5D
Moltkia suffruticosa . . 19 . . 19 5D
Calamagrostis varia . 1 19 3 . 17 5D
Allium ericetorum 1 3 17 4 . 12 5D
Gentiana clusii . 2 15 6 . 12 5D
Potentilla pusilla 1 . 12 1 2 8 5D
Erysimum sylvestre . 4 10 . . 7 5D
Euphrasia cuspidata . . 8 . . 8 5D
Suborder: Koelerienalia splendentis
Teucrium montanum 38 63 71 20 8 43 4K
Koeleria splendens 60 50 15 4 17 40 4K
Globularia cordifolia 5 36 79 11 3 48 5D
Stipa eriocaulis 24 30 48 2 4 27 4K
Scorzonera austriaca 6 32 37 5 4 25 5D+5S
Seseli kochii 13 12 42 3 1 21 5D
Potentilla heptaphylla subsp. australis 16 21 25 5 6 15 4K
Genista sericea 7 36 19 5 . 26 5D+5S
Inula ensifolia 2 21 33 . 1 21 5D+5S
Dianthus sylvestris subsp. tergestinus 26 24 . 1 5 19 4K
Genista sylvestris subsp. sylvestris 13 32 4 . 2 20 4K
Trinia glauca 1 17 19 4 4 14 5D+5S
Potentilla tommasiniana 9 24 2 . 1 14 4K
Stachys recta subsp. subcrenata 15 19 . . 1 14 4K
Linum narbonense . 11 13 2 1 11 5D+5S
Ornithogalum orthophyllum subsp. kochii 7 5 . . 2 8 5C1
Euphrasia illyrica 2 5 2 . . 3 4K
Hyacinthella dalmatica 3 . . . . 5 5C1
Suballiance: Hypochoeridenion maculatae
Centaurea scabiosa subsp. fritschii 2 1 15 47 4 31 6H
Cirsium pannonicum . 5 2 41 13 28 6H
Knautia ressmannii . . 15 40 . 29 6H
Rhinanthus alectorolophus subsp. freynii 1 1 2 35 10 24 6H
Prunella grandiflora . 9 10 31 4 18 6H
Carex montana . 4 8 27 4 17 6H
Achillea roseoalba . . . 18 . 18 6H
Linum viscosum . 1 6 14 1 10 6H
Alliance: Scorzonerion villosae and
Suballiance: Scorzonerenion villosae
Filipendula vulgaris 1 29 2 57 43 38 4Z
Plantago media 5 18 4 45 50 39 4Z
Table 1. cont.
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 23
5C 5S 5D 6H 6V IndVal Cluster
Number of relevés 159 198 51 98 142
Ononis spinosa 4 4 2 27 42 32 4Z
Euphorbia verrucosa . 14 4 44 21 25 4Z
Trifolium rubens 1 5 2 30 28 26 4Z
Hypochaeris maculata 1 8 6 35 19 21 4Z
Scorzonera villosa subsp. villosa 11 30 . 7 46 30 6V2
Knautia illyrica 1 30 8 8 45 39 6V2
Centaurea jacea subsp. weldeniana 4 14 . 1 45 44 6V2
Danthonia alpina . 6 . 12 30 20 4Z
Dorycnium pentaphyllum subsp. herbaceum 8 8 6 12 27 16 4Z
Ferulago campestris 3 6 2 13 20 25 6V2
Prunella laciniata 4 2 . 4 29 19 6V
Lathyrus latifolius . 2 . 1 29 33 6V2
Tragopogon tommasinii . 3 . . 7 11 6V2
Order: Scorzoneretalia villosae
Salvia pratensis 32 35 15 54 55 40 3Z
Eryngium amethystinum 69 46 25 11 32 42 3Z
Chrysopogon gryllus 48 10 21 37 62 36 3Z
Festuca illyrica + F. stricta subsp. sulcata 32 41 17 56 31 37 3Z
Plantago holosteum 38 48 21 15 30 34 3Z
Anthyllis vulneraria subsp. polyphylla 2 23 56 34 13 30 5D
Stachys officinalis 4 35 10 45 27 25 3Z
Scabiosa triandra 18 19 8 34 32 23 3Z
Centaurea jacea subsp. gaudinii 1 9 44 37 2 27 5D
Dorycnium pentaphyllum subsp. germanicum 19 47 2 3 20 24 3Z
Festuca valesiaca 43 17 2 4 15 51 5C1
Veronica barrelieri 8 16 12 9 11 12 3Z
Medicago prostata 23 14 2 1 14 13 3Z
Noccaea praecox 9 7 17 11 4 8 3Z
Plantago argentea . 31 19 18 6 21 5D+5S
Cyanus triumfettii . 23 23 16 8 16 5D+5S
Leucanthemum platylepis 1 29 . 5 27 23 5S2
Euphorbia nicaeensis 4 26 . . 10 21 5S2
Seseli montanum subsp. tommasinii 6 6 . . 1 7 5C1
Class: Festuco-Brometea
Bromopsis erecta s.l. + B. condensata 79 86 81 87 82 83 3Z
Lotus corniculatus 26 52 46 85 66 53 3Z
Sanguisorba minor 52 43 40 39 48 45 3Z
Carex humilis 17 84 75 24 13 58 5D+5S
Helianthemum nummularium subsp. obscurum 21 33 58 43 39 35 3Z
Brachypodium pinnatum + B. rupestre 11 17 23 87 51 54 4Z
Thymus longicaulis s.l. 58 34 42 11 37 38 3Z
Galium lucidum 33 47 63 8 30 35 3Z
Teucrium chamaedrys 26 33 60 24 32 32 3Z
Galium verum 4 7 31 78 39 47 4Z
Table 1. cont.
eschweizerbart_xxx
24 Massimo Terzi
5C 5S 5D 6H 6V IndVal Cluster
Number of relevés 159 198 51 98 142
Briza media 1 14 6 81 44 52 4Z
Euphorbia cyparissias 28 28 33 30 23 27 3Z
Asperula cynanchica 24 19 33 44 15 24 3Z
Hippocrepis comosa 25 29 25 31 23 27 3Z
Globularia bisnagarica 9 25 38 24 17 20 3Z
Anthericum ramosum 7 34 35 29 9 21 3Z
Carex flacca 1 6 12 53 30 35 4Z
Leontodon crispus 23 27 23 8 16 21 3Z
Carex caryophyllea 8 11 13 37 22 21 4Z
Linum tenuifolium 20 14 38 2 12 15 3Z
Asperula purpurea 19 14 29 5 9 25 5C2
Sedum sexangulare 48 8 6 5 8 37 5C
Pilosella officinarum 20 8 8 19 18 18 5C1
Carlina acaulis 2 14 21 26 6 12 3Z
Petrorhagia saxifraga 42 8 6 2 10 31 5C
Campanula glomerata 1 6 8 45 4 31 6H
Pimpinella saxifraga + P. alpina 2 3 12 39 6 24 6H
Allium lusitanicum 9 18 23 2 1 14 4K
Bothriochloa ischaemum 35 3 6 3 6 52 5C2
Asperula aristata 18 18 8 1 6 16 5C1
Medicago falcata 1 6 8 10 15 10 4Z
Centaurea jacea 3 1 4 22 4 13 6H
Koeleria pyramidata . 5 21 72 27 42 6H
Trifolium montanum . 18 4 61 32 37 4Z
Fumana procumbens + F. ericoides 30 15 44 . 6 21 4K
Linum catharticum . 8 21 43 14 21 6H
Koeleria macrantha 12 9 21 . 10 10 3Z
Artemisia alba 19 7 23 . 2 30 5C2
Thymus pulegioides 1 5 . 31 13 19 6H
Dianthus sylvestris . 5 37 1 1 32 5D
Melica ciliata 23 12 4 . 2 16 5C
Medicago lupulina 4 1 . 19 15 15 4Z
Trifolium campestre 11 5 . 5 14 12 5C1
Ranunculus bulbosus 1 2 . 16 11 12 4Z
Echinops ritro 4 12 4 . 4 11 5S1
Anthyllis vulneraria 26 34 . . 13 21 4K
Polygala nicaeensis 9 35 . . 24 19 5S
Teucrium capitatum subsp. capitatum 36 6 . . 6 55 5C1
Polygala comosa . . 13 21 6 11 6H
Galium corrudifolium 19 18 . . 2 26 5C1
Satureja montana 13 20 . . 3 18 5S1
Carex hallerana 19 8 . . 4 17 5C1
Kengia serotina 18 . 8 . 1 35 5C2
Paronychia kapela 7 12 . . 1 25 5S1
Table 1. cont.
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 25
5C 5S 5D 6H 6V IndVal Cluster
Number of relevés 159 198 51 98 142
Alyssum montanum 14 3 2 . . 13 5C2
Poa bulbosa 8 2 . . 4 12 5C1
Armeria canescens 1 6 . . 2 13 5S1
Thymus praecox . . 17 3 . 16 5D
Argyrolobium zanonii 9 . . . 1 12 5C2
Thymus striatus . 5 . . 1 14 5S1
Lactuca viminea 6 . . . . 12 5C2
Onobrychis alba subsp. laconica . 4 . . . 12 5S1
Table 1. cont.
Koelerio splendentis-Festucetum illyricae Horvatić 1963
corr. Trinajstić 1992; na – Narcisso tazettae-Asphodeletum
microcarpi Šegulja 1969; se – Salvio officinalis-Euphorbietum
fragiferae Lausi & Poldini 1962; ss – Salvio
officinalis-Seslerietum juncifoliae Trinajstić 1977 (transition
to Saturejion subspicatae); sd – Saturejo montanaeDichanthietum
ischaemum Horvat in Horvat et al. 1974
nom. corr.; so – Stipo bromoidis-Salvietum officinalis
Horvatić 1963.
5S. Alliance: Saturejion subspicatae Tomić-Stanković
1970 [3]
Lectotypus: Centaureo rupestris-Caricetum humilis
Horvat 1931. Diagnostic taxa: Anthyllis montana
subsp. jacquinii; Astragalus croaticus; Bunium alpinum
subsp. montanum; Centaurea rupestris; Crepis chondrilloides;
Edraianthus tenuifolius; Gentiana verna
subsp. tergestina; Globularia meridionalis; Iris pallida
subsp. illyrica; Jurinea mollis; Klasea radiata subsp. cetinjensis;
Muscari botryoides; Narcissus poeticus subsp.
radiiflorus; Pulsatilla montana; Satureja subspicata;
Sesleria tenuifolia subsp. tenuifolia.
Associations: as – Astragalo croatici-Seslerietum robustae
Trinajstić ex Terzi 2011; bs – Bromo erecti-Seslerietum interruptae
Trinajstić ex Terzi 2011; cr – Centaureo rupestris-Caricetum
humilis Horvat 1931 [4]; gk – Genisto
sericeae-Seslerietum kalnikensis Dakskobler & Peljhan ex
Terzi ass. nov. hoc loco [8]; gs – Genisto sericeae-Seslerietum
tenuifoliae Poldini 1980; gg – Genisto-Globularietum
bellidifoliae Tomić-Stanković 1970; mg – Minuartio
capillaceae-Genistetum pulchellae Šegulja & Bedalov ex
Terzi 2011; pc – Pediculari acaulis-Caricetum humilis
Horvat in Horvat et al. 1974; sm – Saturejo subspicataeCaricetum
humilis Trinajstić 1999; su – Saturejo subspicatae-Edraianthetum
tenuifolii Horvat in Horvat et al.
1974; sj – Seslerio robustae-Juniperetum sibiricae Domac
1962; sc – Seslerio tenuifoliae-Caricetum humilis Horvat
1930; st – Stipo eriocaulis-Caricetum humilis Trinajstić ex
Terzi 2011; sg – Stipo pennatae-Genistetum dalmaticae
Redžić et al. ex Terzi 2011
5D. Alliance: Centaureion dichroanthae Pignatti 1952
Holotypus: Centaureo dichroanthae-Globularietum
cordifoliae Pignatti 1952. Diagnostic taxa: Allium ericetorum;
Calamagrostis varia; Carduus defloratus;
Carex mucronata; Centaurea dichroantha; Cytisus
pseudoprocumbens; Cytisus purpureus; Erysimum sylvestre;
Euphorbia triflora subsp. kerneri; Euphrasia
cuspidata; Gentiana clusii; Gypsophila repens; Hieracium
porrifolium; Leontodon incanus; Lomelosia
graminifolia subsp. graminifolia; Moltkia suffruticosa;
Polygala nicaeensis subsp. carniolica; Potentilla pusilla;
Sesleria caerulea; Stachys recta subsp. labiosa.
Associations: si – Bromo condensati-Stipetum eriocaulis
Lasen ex Terzi ass. nov. hoc loco [8]; bb – Bupleuro ranunculoides-Brometum
condensati ass. nov. hoc loco [9];
cg – Centaureo dichroanthae-Globularietum cordifoliae
Pignatti 1952; gc – Genisto-Caricetum mucronatae Horvat
in Horvat, et al. 1974 (sensu Poldini 1989); Laserpitido-Festucetum
alpestris Pedrotti 1970*; ls – Leontodonto
brumatii-Seslerietum calcariae Dakskobler et al. 2012; sb
– Saturejo variegatae-Brometum condensati Poldini &
Feoli Chiapella in Feoli Chiapella & Poldini 1993; tc –
Teucrio capitati-Chrysopogonetum grylli Sburlino et al.
2008 (transition to Chrysopogono-Saturejion subspi-
catae).
4Z. Suborder: Scorzonerenalia villosae subord. nov. hoc
loco
Holotypus: Scorzonerion villosae Horvatić ex
Kovačević 1959 (Kovačević 1959: 18). Same diagnostic
taxa of Scorzonerion villosae.
5V. Alliance: Scorzonerion villosae Horvatić ex Kovačević
1959 [1]
Holotypus: Scorzonero villosae-Danthonietum calycinae
Kovačević 1959. Diagnostic taxa: Centaurea jacea
subsp. weldeniana; Cirsium pannonicum; Danthonia
alpina; Dorycnium pentaphyllum subsp. herbaceum;
Euphorbia verrucosa; Ferulago campestris; Filipendula
vulgaris; Hypochaeris maculata; Knautia illyrica;
eschweizerbart_xxx
26 Massimo Terzi
Lathyrus latifolius; Ononis spinosa; Plantago media;
Prunella laciniata; Scorzonera villosa subsp. villosa;
Tragopogon tommasinii; Trifolium rubens.
6V. Suballiance: Scorzonerenion villosae Poldini & Feoli
Chiapella 1993
Holotypus: Scorzonero villosae-Danthonietum calycinae
Kovačević 1959. Same diagnostic taxa of Scorzonerion
villosae.
Associations: ec – Euphorbio nicaeensis-Chrysopogonetum
grylli Horvatić 1963; fp – Festuco illyricae-Poetum
bulbosae Horvat in Horvat et al. 1974 corr. Terzi 2011; fa
– Festuco-Armerietum canescentis Trinajstić & Sugar
1972; ge – Globulario elongatae-Chrysopogonetum grylli
Ilijanić et al. ex Terzi 2011; oa – Ononido antiquorumBrometum
condensati Horvatić (1934) 1963; ds – Scorzonero
villosae-Danthonietum calycinae Kovačević 1959
[1, 2]. Uncertain position: cs – Carici vernae-Scabiosetum
leucophyllae Redžić et al. ex Terzi 2011; gl – Globulario
cordifoliae-Scabiosetum leucophyllae Redžić et al. ex
Terzi 2011
6H. Suballiance: Hypochoeridenion maculatae Poldini &
Feoli Chiapella ex Terzi 2011
Holotypus: Onobrychido arenariae-Brometum erecti
Poldini & Feoli Chiapella in Feoli Chiapella & Poldini
1993. Diagnostic taxa: Achillea roseoalba; Carex montana;
Cirsium pannonicum; Centaurea scabiosa subsp.
fritschii; Knautia ressmannii; Linum viscosum; Prunella
grandiflora; Rhinanthus alectorolophus subsp. freynii.
Associations: ab – Avenulo praeustae-Brometum erecti
Poldini & Feoli Chiapella ex Terzi 2011; ch – Chamaecytiso
hirsuti-Chrysopogonetum grylli Pignatti ex Feoli
Chiapella & Poldini 1993; gb – Gentianello pilosae-Brometum
erecti Dakskobler & Završnik 2009; gm – Gladiolo
palustris-Molinietum arundinaceae Poldini & Feoli
Chiapella in Feoli Chiapella & Poldini 1993; ob – Onobrychido
arenariae-Brometum erecti Poldini & Feoli
Chiapella in Feoli Chiapella & Poldini 1993; sn – Schoeno
nigricantis-Chrysopogonetum grylli Pignatti ex Feoli
Chiapella & Poldini 1993 (transition to Centaureion dichroanthae);
ts – Thlaspio-Trifolietum pratensis Zanotto
1960*.
Conclusion
The numerical analysis of relevés and associations previously
assigned to the Scorzoneretalia villosae highlights
two main groups of associations classified within the new
meso-xerophytic Scorzonerenalia villosae and the xerophytic
Koelerienalia splendentis. The latter is well differentiated
by the presence of numerous sub-Mediterranean
and Illyrian species, while the Scorzonerenalia villosae,
except from few Illyrian and south-eastern European diagnostic
taxa, shares many taxa with mesophytic syntaxa
of the Festuco-Bometea. The results of this study clarify
the syntaxonomy and nomenclature of the Scorzoneretalia
villosae but also demonstrate the need for a syntaxonomic
revision of the Festuco-Brometea, especially of its
more mesophilous associations. Apart from the classical
model of the class, with geographically defined orders,
other proposals suggest a single order for the meso-xerophytic
plant communities of the class (Dengler et al.
2003) or a specific Balkanic order for the more mesophilous
communities (Boșcaiu 1972). These issues were beyond
the scope of this paper but it emerged that the more
mesophilous associations of the Scorzonerenalia villosae
(together with the order and suborder names) would
have to be involved in such a revision.
Acknowledgments
I would like to thank L. Poldini (University of Trieste) and
R. Di Pietro (University of Rome) for useful discussion on a
preliminary draft of this paper and three anonymous referees
and the editors, E. Bergmeier and J. Dengler, for their
helpful suggestions and comments; L. Sutcliffe is thanked
for linguistic revision of the manuscript.
References
Aho, K., Roberts, D.W. & Weaver, T. 2008. Using geometric
and non-geometric internal evaluators to compare eight vegetation
classification methods. Journal of Vegetation Science
19: 549–562.
Antonić, O., Kušan, V., Bakran-Petricioli, T., Alegro, A.,
Gottstein-Matočec, S., Peternel, H. & Tkalčec, Z. 2005.
Klasifikacija staništa Republike Hrvatske [Habitat classification
of the Republic of Croatia]. Drypis 1: 1–12.
Assyov, B., Petrova, A.S., Dimitrov, D. & Vassilev, R. 2006.
Conspectus of the Bulgarian vascular flora. Distribution
maps and floristic elements. Third edition. Bulgarian Biodiversity
Foundation, Sofia, BG.
Bajić, D. 1960. Brdske i dolinske livade i pašnjaci u slivu reka
Une i Sane [Meadows and pastures of mountains and valleys
in the basin of the rivers Una and Sana]. Radovi Poljoprivrednog
Fakulteta Univerziteta u Sarajevu 8–9: 237–268.
Barbero, M. & Loisel, R. 1972. Contribution à l’étude des pelouses
à Brome méditerranéennes et méditerranéo-montagnardes.
Anales del Instituto Botánico A. J. Cavanilles 28:
91–165.
Biondi, E. & Galdenzi, D. 2012. Phytosociological analysis of
the grasslands of Montagna dei Fiori (central Italy) and syntaxonomic
review of the class Festuco-Brometea in the Apennines.
Plant Sociology 49: 91–112.
Biondi, E., Allegrezza, M., Casavecchia, S., Galdenzi, D., Gasparri,
R., Pesaresi, S., Vagge, I. & Blasi, C. 2014. New and
validated syntaxa for the checklist of Italian vegetation. Plant
Biosystems 148: 318–332.
Blečić, V. & Lakušić, R. 1976. Prodromus biljnih zajednica
Crne Gore [Prodromus of plant communities of Montenegro].
Glasnik Republičkog Zavoda za Zaštitu Prirode i
Prirodnjačkog Muzeja u Titogradu 9: 57–98.
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 27
Bonin, G. 1978. Contribution à la connaissance de la végétation
des montagnes de l’Apennin centreméridional (tableaux et
figures). Ph.D. thesis, Univ. Droit d’Economie et de Sciences,
Aix-Marseille. Centre Regional de Documentation
Pedagogique, Marseille, FR.
Boșcaiu, N. 1972. Pajiștile xerice (Festuco-Brometea Br.-Bl. 43)
din depresiunea Caransebeș-Mehadia [Xeric grasslands (Festuco-Brometea
Br.-Bl. 43) in the depression of CaransebeșMehadia].
Acta Horti Botanici Bucurestiensis 1970–1971:
449–468.
Braun-Blanquet, J. 1974. Die höheren Gesellschaftseinheiten
der Vegetation des südeuropaïsch-westmediterranen Raums.
Station International De Géobotanique Méditerranéenne et
Alpine [communication no. 204], Montpellier, FR.
Brullo, S., Giusso del Gado, G.P. & Minissale, P. 2009. Taxonomic
revision of the Koeleria splendens C. Presl group (Poaceae)
in Italy based on morphological characters. Plant Biosystems
143: 140–161.
Buzo, K. 1991. Bimësia e kullotave dhe e livadheve natyrore të
Shqiperisë [The vegetation of natural pastures and meadows
of Albania]. Shtëpia Botuese e Librit Universitar, Tirana,
AL.
Černjavski, P., Grebenščikov, O. & Pavlović, Z. 1949. O vegetaciji
i fiori Skadarskog podrucja [Vegetation and flora of the
Skadar area].Glasnik Prirodnjačkog Muzeja Srpske Zemlje,
B, 1–2: 5–91.
Chytrý, M. & Otýpková, Z. 2003. Plot sizes used for phytosociological
sampling of European vegetation. Journal of Vegetation
Science 14: 563–570.
Conti, F., Abbate, G., Alessandrini, A. & Blasi, C. 2005. An annotated
check-list of the Italian vascular flora. Palombi Editore,
Roma, IT.
Dakskobler, I. & Peljhan, J. 2007. Viola pyrenaica Ramond ex
Dc in the northern part of the Dinaric Mountains (The Plateaus
of Trnovski Gozd and Nanos, Slovenia). Hacquetia 6:
143–169.
De Cáceres, M., Font, X. & Oliva, F. 2008. Assessing species
diagnostic value in large data sets: a comparison between phi
coefficient and Ochiai index. Journal of Vegetation Science
19: 779–788.
Dengler, J., Berg, C., Eisenberg, M., Isermann, M., Jansen, F.,
Koska, I., Löbel, S., Manthey, M., Päzolt, J., (…) & Wollert,
H. 2003. New descriptions and typifications of syntaxa
within the project Plant communities of Mecklenburg-Vorpommern
and their vulnerability. Part I. Feddes Repertorium
114: 587–631.
Dengler, J., Löbel, S. & Dolnik, C. 2009. Species constancy depends
on plot size – a problem for vegetation classification
and how it can be solved. Journal of Vegetation Science 20:
754–766.
Dengler, J., Becker, T., Ruprecht, E., Szabó, A., Becker, U.,
Beldean, M., Bita-Nicolae, C., Dolnik, C., Goia, I., (…) &
Uğurlu, E. 2012. Festuco-Brometea communities of the
Transylvanian Plateau (Romania) – a preliminary overview
on syntaxonomy, ecology, and biodiversity. Tuexenia 32:
319–359.
Dring, J., Hoda, P., Mersinllari, M., Mullaj, A., Pignatti, S. &
Rodwell, J. 2002. Plant communities of Albania: a preliminary
overview. Annali di Botanica (Rome) 2: 7–30.
Dufrêne, M. & Legendre, P. 1997. Species assemblages and indicator
species: the need for a flexible asymmetrical approach.
Ecological Monographs 67: 345–366.
Feoli Chiapella, L. & Poldini, L. 1993. Prati e pascoli del Friuli
(NE Italia) su substrati basici [Grasslands of Friuli (NE Italy)
on basic substrates]. Studia Geobotanica 13: 3–140.
Ferlan, L. & Giacomini, V. 1957. Appunti fitosociologici su esempi
di „pascolo carsico“: Chrysopogoneto-Centaureetum
cristatae [Phytosociological notes on karst pasture types:
Chrysopogoneto-Centaureetum cristatae]. In: Anonymous
(ed.) Convegno Friulano di Scienze Naturali (Proceedings of
the first Friulian Meeting on Natural Science) 4–5 Sept 1955,
Udine, pp. 159–183. Tip. Domenico del Bianco & figlio,
Udine, IT.
Gaži-Baskova, V. & Šegulja, N. 1978. Sindinamika vegetacije
brdskih travnjaka na podrucju Hrvatskog primorja [The
syndinamic of vegetation of mountain grasslands in the Dinaric
Region of Hrvatsko Primorje]. Godišnjak Biološkog
Instituta Univerziteta u Sarajevu 31: 37–47.
Horvat, I. 1931. Die Berg-Wiesen und Heiden in Kroatien. Acta
Botanica Croatica 6: 76–90.
Horvat, I. 1962. Vegetacija planina Zapadne Hrvatske [Mountain
vegetation of West Croatia]. Acta Biologica (Zagreb) 30:
5–179.
Horvat, I., Glavač, V. & Ellenberg, H. 1974. Vegetation Südosteuropas.
G. Fischer, Stuttgart, DE.
Horvatić, S. 1934. Flora i vegetacija otoka Paga [Flora and vegetation
of the island of Pag ]. Prirodoslovna Istraživanja 19:
116–372.
Horvatić, S. 1949. Istraživanje vegetacije u istri god [Survey of
the vegetation of Istria]. 1948. Ljetopis Jugoslavenske Akademije
Znanosti i Umjetnosti 55: 105–109.
Horvatić, S. 1958. Geographisch-typologische Gliederung der
Niederungs-Wiesen und Weiden Kroatiens. Pflanzengeographischen
Gebiete Kroatiens. Angewandte Pflanzensoziologie
(Vienna) 15: 63–73.
Horvatić, S. 1963. Vegetacijska karta otoka Paga s općim pregledom
vegetacijskih jedinica Hrvatskog primorja [Vegetation
map of the island of Pag, with a review of the vegetation
units of the coast of Croatia]. Acta Biologica (Zagreb) 4:
3–187.
Horvatić, S. 1973. Syntaxonomic analysis of the vegetation of
dry grassland and stony meadows in Eastern Adriatic coastal
Karts district based on the latest phytocoenological research.
Fragmenta Herbologica Jugoslavica 32: 1–15.
Horvatić, S. 1975. Neuer Beitrag zur Kenntnis der Syntaxonomie
der Trocken-Rasen und Steintriften Gesellschaften des
ostadriatischen Karstgebietes. In: Jordanov, D., Bondev, I.,
Kozuharov, S., Kuzmanov, B., Palamarev, E. & Velcev, V.
(eds.) Problems of Balkan flora and vegetation, pp 300–310.
Bulgarian Academy of Sciences. Sofia, BG.
Horvatić, S., Ilijanić, L. & Marković-Gospodarić, L. 1968.
Biljni pokrov okoline Senja [Vegetation in the area of Senj].
Senjski Zbornik 3: 298–323.
Hršak, V. 2003. Multivariate analysis of phyto-climatic patterns
of the Croatian part of the eastern Adriatic coast. Plant Biosystems
137: 281–292.
Ilijanić, L., Gaži, V. & Topić, J. 1972. Grasslands containing
Chrysopogon gryllus in continental regions of West Croatia.
Acta Botanica Croatica 31: 155–164.
Jovanović, B., Lakušić, R., Rizovski, .R, Trinajstić, I. &
Zupančič, M. 1986. Prodromus Phytocoenosum Jugoslaviae:
ad mappam vegetationis m 1:200 000 [Prodromus of phytocoenoses
of Yugoslavia: vegetation map at 1:200 000].
Naučno veće vegetacijske karte Jugoslavije, Bribir-Ilok, HR.
Kovačević, J. 1959. Pregled tipova travnjaka s agroekološkog
gledišta područja bivših bosanskih kotare Sanski Most,
Mrkonjić-Grad, Bosanski Petrovac I Lijevča Polja [Overview
of grassland types in the former Bosnian districts of
Sanski Most, Mrkonjic-Grad, Bosanski Petrovac and Lijevče
eschweizerbart_xxx
28 Massimo Terzi
Polje from an agroecological viewpoint]. Godišnjak
Biološkog Instituta Univerziteta u Sarajevu 1–2: 3–46.
Kruskal, J.B. 1964. Multidimensional scaling by optimizing
goodness of fit to a nonmetric hypothesis. Psychometrika 29:
1–27.
Lakušić, R., Muratspahić, D., Pavlović, I. & Redžić, S. 1982.
Vegetacija ekosistema kraških polja Hercegovine [The vegetation
of ekosystems in karst poljes of Herzegovina].
Godišnjak Biološkog Instituta Univerziteta u Sarajevu 35:
81–92.
Lakušić, R. & Redžić, S. 1991. Vegetacija refugijalno-reliktnih
ekosistema siva rijeke Une [Vegetation of the refugial-relict
ecosystems of the Una river system]. Bilten Društvo Ekologa
Bosne i Hercegovine, B, 6: 25–73.
Lasen, C. 1995. Note sintassonomiche e corologiche sui prati
aridi del massiccio del Grappa [Chorological and syntaxonomical
remarks on the xerophytic grasslands in the Grappa
Massif]. Fitosociologia 30: 181–199.
Legendre, P. & Gallagher, E.D. 2001. Ecologically meaningful
transformations for ordination of species data. Oecologia
129: 271–280.
Mather, P.M. 1976. Computational methods of multivariate
analysis in physical geography. J. Wiley & Sons, London,
UK.
McCune, B. & Grace, J.B. 2002. Analysis of ecological communities.
MjM Software Design, Gleneden Beach, Oregón, US.
McCune, B. & Mefford, M.J. 2011. PC-ORD. Multivariate
analysis of ecological data. Version 6.0. MjM Software,
Gleneden Beach, Oregon, US.
Meyer, M. 1976. Pflanzensoziologische und ökologische Untersuchungen
an insubrischen Trockenwiesen karbonathaltiger
Standorte. Veröffentlichungen des Geobotanischen Instituts
der Eidgenössischen Technischen Hochschule, Stiftung Rübel,
in Zürich 57: 1–145.
Mucina, L., Dengler, J., Bergmeier, E., Čarni, A., Dimopoulos,
P., Jahn, R. & Matevski V. 2009. New and validated highrank
syntaxa from Europe. Lazaroa 30: 267–276.
Nikolić, V. & Diklić, N. 1966. Zajednica žalfije i rudinskog
pelina, Artemisio-Salvietum officinalis (Salvia officinalisArtemisia
lobelii Grebenščikov, 1950) u Sićevačkoj klisuri
[Communities with Artemisia and Salvia, Artemisio-Salvietum
officinalis (Salvia officinalis-Artemisia lobelia
Grebenščikov, 1950) in the Sićevo Gorge]. Glasnik
Prirodnjačkog Muzeja u Beogradu, B, 21: 5–21.
Otýpková, Z. & Chytrý, M. 2006. Effects of plot size on the
ordination of vegetation samples. Journal of Vegetation Science
17: 465–472.
Pignatti, E. & Pignatti, S. 2014. Plant life of the Dolomites. Vegetation
structure and ecology. Springer, Heidelberg, DE.
Pignatti, S. 1982. Flora d’Italia [The Flora of Italy]. Ed. Agricole,
Bologna, IT.
Pipenbaher, N., Kaligarič, M., Mason, N.W.H., Škornik, S. &
Norman, W.H. 2013. Dry calcareous grasslands from two
neighboring biogeographic regions: relationship between
plant traits and rarity. Biodiversity and Conservation 22:
2207–2221.
Pipenbaher, N., Kaligarič, M. & Škornik, S. 2011. Floristic and
functional comparision of karst pastures and karst meadows
from the North Adriatic karst. Acta Carsologica 40: 515–525.
Poldini, L. 1989. La vegetazione del Carso isontino e triestino
[Vegetation of Trieste and Gorizia karst]. Ed. Lint, Trieste,
IT.
Poldini, L. 1991. Atlante corologico delle piante vascolari nel
Friuli-Venezia Giulia [Chorological atlas of vascular plants
in the region Friuli-Venezia Giulia]. Regione Autonoma
Friuli-Venezia Giulia, Udine, IT.
Poldini, L. 2009. La diversità vegetale del Carso fra Trieste e
Gorizia [Plant diversity of the Karst between Trieste and
Gorizia]. Goliardiche, Trieste, IT.
Poldini, L. & Kaligarič, M. 1997. New contribution on the typology
of the vegetation of dry grasslands (Scorzoneretalia
villosae H-ić1975) in the north Adriatic Karst. Gortania 19:
119–148.
Poldini, L. & Sburlino, G. 2005. Terminologia fitosociologica
essenziale [Essential phytosociological terminology]. Fitosociologia
42: 57–69.
Quintanar, A. & Castroviejo, S. 2013. Taxonomic revision of
Koeleria (Poaceae) in the western Mediterranean Basin and
Macaronesia. Systematic Botany 38: 1029–1061.
Redžić, S. 1999. The syntaxonomic differentiation of the Festuco-Brometea
Br.-Bl. & R. Tx. 1943 ex Klika & Hadač1944
in the Balkans. Annali di Botanica (Rome) 57: 167–180.
Redžić, S., Lakušić, R., Muratspahić, D., Bjelčić, Z. & Omerović,
S. 1984. Struktura I dinamika fitocenoza u ekosistemima
Cincara i Vitoroga [Structure and dynamics of phytocoenoses
in the ecosystems of the mountains Cincar and Vitorog].
Godišnjak Biološkog Instituta Univerziteta u Sarajevu
37: 123–177.
Redžić, S., Trakić, S. & Barudanović, S. 2013. Patterns of vegetation
diversity of grasslands and pastures – Crvanj Mt.
(Herzegovina, Western Balkan). Scientific Research and Essays
8: 1944–1965.
Ritter-Studnička, H. 1972. Neue Pflanzengesellschaften aus den
Karstfeldern Bosniens und der Hercegovina. Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie
92: 108–154.
Royer, J.M. 1991. Synthèse eurosibérienne, phytosociologique
et phytogéographique de la classe des Festuco-Brometea.
Dissertationes Botanicae 178: 1–296.
Sanda, V., Öllerer, K. & Burescu, P. 2008. Fitocenozele din
România [Phytocoenoses of Romania]. Ars Docendi, Bucharest,
RO.
Sburlino, G., Buffa, G., Filesi, L. & Gamper, U. 2008. Phytocoenotic
originality of the N-Adriatic coastal sand dunes
(Northern Italy) in the European context: The Stipa veneta
rich communities. Plant Biosystems 142: 533–539.
Terzi, M. 2011. Nomenclatural revision for the order Scorzonero-Chrysopogonetalia.
Folia Geobotanica 46: 411–444.
Terzi, M. & Di Pietro, R. 2013. Floristic characterization of the
dry grasslands of the class Festuco-Brometea Br.-Bl. et Tx. ex
Klika et Hadac 1944 in the Adriatic Region. In: Calabrese,
G., Pacucci, C., Occhialini, W. & Russo, G. (eds) IX National
Biodiversity Conference, Vol. 3, pp. 104–111. ChieamIamb,
Bari, IT.
Terzi, M., Di Pietro, R. & D’Amico, F.S. 2010. Application of
Indicator Species Analysis to communities with Stipa austroitalica
Martinovsky and relevant syntaxonomic problems.
Fitosociologia 47: 3–28.
Tichý, L. & Chytrý, M. 2006. Statistical determination of diagnostic
species for site groups of unequal size. Journal of Vegetation
Science 17: 809–818.
Tichý, L., Chytrý, M., Hájek, M., Talbot, S.S. & Botta-Dukát,
Z. 2010. OptimClass: using species-to-cluster fidelity to determine
the optimal partition in classification of ecological
communities. Journal of Vegetation Science 21: 287–299.
Tomažic, G. 1941. Senozeti in pasniki na plitvih in suhih tleh
Slovenije [Pastures and grasslands on shallow and dry soils
in Slovenia]. Zbornik Prirodoslovnega Društva 2: 76–82.
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 29
Tomić-Stanković, K. 1970. Vegetacija Lovčena u Crnoj Gori
[Vegetation of Mt. Lovčen in Montenegro]. Zajednica Naucnih
Ustanova Kosova Studije Knjiga 17: 7–93.
Trinajstić, I. 1977. Osnovne značajke biljnog pokrova otaka
Hvara I njegov fitogeogravski položaj okviru Evropskog dijela
Sreddozemlja [Overview of the vegetation of the Island
of Hvar and its phytogeographical position in the European
part of the Mediterranean Basin]. Poljoprivreda i Šumarstvo
23: 1–36.
Trinajstić, I. 1987. Sintaksonomski pregled biljnih zajednica
planine Biokovo [Syntaxonomic revision of plant communities
in the mountains of Biokovo]. Acta Biokovica 4: 143–
174.
Trinajstić, I. 1999. As. Saturejo-Caricetum humilis Trinajstić
(1981, nom.sol.) 1999, ass. nov. – sintaksonomska analiza flornoga
sastava [As. Saturejo-Caricetum humilis Trinajstić
(1981, nom. sol.) 1999, ass. nov. – syntaxonomical analysis of
floristic composition]. Agronomski Glasnik 1–2: 23–34.
Trinajstić, I. 2005. Travnjaci as. Koelerio macranthae-Brachypodietum
retusi Trinajstić, ass. nov. u Hrvatskom primorju
[Grasslands of the ass. Koelerio macranthae-Brachypodietum
retusi Trinajstić, ass. nov. in Croatian littoral]. Agronomski
Glasnik 5: 347–357.
Trinajstić, I. 2008. Biljne zajednice republike Hrvatske [Plant
communities of Croatia]. Akademija Šumarskih Znanosti,
Zagreb, HR.
Trinajstić, I. & Pavletić, Z. 1990. Prilog poznavanju sastava i
građe As. Carici-Centaureetum rupestris Ht. 1931 u pretplaninskom
pojasu Biokova [On the structure of the Ass. Carici-Centaureetum
rupestris Ht. 1931 in the subalpine part of
the Biokovo mountains]. Acta Botanica Croatica 49: 75–80.
Trinajstić, I. & Šugar, I. 1972. Prilog poznavanju vegetacije
suhih travnjaka na planini Dinari u Hrvatskoj [A contribution
to the knowledge of dry grasslands of the Dinarid
mountains in Croatia]. Acta Botanica Croatica 31: 165–171.
Trinajstić, I, Pavletić, Z. & Šugar, I. 1981. Prilog poznavanju
floristickog sastava i rasprostranjenosti as. Bromo-Danthonietum
alpinae Sugar u gorskim predjelima Hrvatske [A
contribution to the knowledge of floristic composition and
distribution of the ass. Bromo-Danthonietum alpinae Sugar
in the mountainous regions of Croatia]. Acta Botanica Croatica
40: 155–163.
Trinajstić, I., Pavletić, Z. & Kamenjarin, J. 1993. Fitocenološka
istraživanja As. Carici-Centaureetum rupestris Ht. 1931 (Saturejon
subspicatae) na Kozjaku povrh Splita [Plant-sociological
investigations of the ass. Carici-Centaureetum rupestris
Ht. 1931 (Saturejon subspicatae) on the mountain of
Kozjak near Split]. Acta Botanica Croatica 52: 75–80.
Tzonev, R.T. 2009. Syntaxonomy of the natural and semi-natural
vegetation of the Middle Danube Plain in Bulgaria. Biotechnology
& Biotechnological Equipment 23: 354–359.
Ubaldi, D. 2011. Le vegetazioni erbacee e gli arbusteti italiani
[Herbaceous and shrub vegetation in Italy]. Aracne Ed.,
Rome, IT.
van der Maarel, E. 1979. Transformation of cover-abundance
values in phytosociology and its effects on community similarity.
Vegetatio 39: 97–114.
Ward, J.J. 1963. Hierarchical grouping to optimize an objective
function. Journal of the American Statistical Associationc 58:
236–244.
Weber, H.E., Moravec, J. & Theurillat, J.P. 2000. International
Code of Phytosociological Nomenclature. 3
rd
edition. Journal
of Vegetation Science 11: 739–768.
Wendelberger, G. 1965. Zur Vegetationsgliederung Südosteuropas.
Mitteilungen des Naturwissenschaftlichen Vereines
für Steiermark 95: 245–286.
Westhoff, V. & van der Maarel, E. 1978. The Braun-Blanquet
approach. In: Whittaker, R.H. (ed.) Classification of plant
communities, pp. 287–399. Junk, The Hague, NL.
Willner, W., Grabherr, G., Pallas, J. & Weber, H.E. 2011. Report
of the Committee for Nomina Conservanda, Ambigua, Inversa
and Mutata:1. Phytocoenologia 41: 59–70.
Appendix 1: List of syntaxon names quoted in the text.
Abbreviations of syntaxon names used in Figures and Tables in square brackets. Syntaxon abbreviations consist of a
number, indicating the rank (1 class, 3 order, 4 suborder, 5 alliance, 6 suballiance), followed by a capital letter; abbreviations
of associations consist of 2 lower-case letters. The abbreviation “0B” indicates the surclass Festuco-Brachypodio-Brometales
described by Barbero and Loisel (1972).
Classes: Brachypodio-Brometea Barbero et Loisel 1972 nom. illeg. (Art. 29c) [1L]; Brachypodio-Chrysopogonetea
Horvatić 1963 [1B]; Calluno-Ulicetea Br.-Bl. & Tx. ex Klika & Hadač 1944 [1U]; Elyno-Seslerietea Br.-Bl. 1948 [1S];
Erico carneae-Pinetea sylvestris Horvat 1959 [1E]; Festuco-Brometea Br.-Bl. & Tx. ex Klika & Hadač 1944 [1F; 1Fx
and 1Fm indicate xerophytic and mesophytic syntaxa of the class, respectively]; Molinio-Arrhenatheretea Tx. 1937
[1M]; Thero-Brachypodietea Br.-Bl. 1947 [1T; sensu lato]; Trifolio-Geranietea sanguinei T. Müller 1962 [1G].
Orders: Artemisio albae-Brometalia erecti Ubaldi ex Dengler (‘Denggler’) & Mucina in Mucina et al. 2009; Brachypodietalia
pinnati Korneck 1974 [3C]; Brachypodio-Chrysopogonetalia (Horvatić 1958) Boșcaiu 1972 nom. inval. (Art.
3m; note 1 of Art. 27); Brometalia erecti Koch 1926 [3F]; Cymbopogono hirti-Brachypodietalia ramosi Horvatić 1963;
Festucetalia valesiacae Br.-Bl. & Tx. ex Br.-Bl. 1950 [3V]; Koelerietalia splendentis Horvatić 1973 [3K]; Scorzoneretalia
villosae Kovačević 1959 [3Z]; Scorzoneretalia villosae Horvatić 1973 nom. illeg. (synonym of Scorzoneretalia villosae
Kovačević 1959); Scorzonero villosae-Chrysopogonetalia grylli Horvatić & Horvat in Horvatić 1963 (synonym of
Scorzoneretalia villosae) [3S].
Suborders: Koelerienalia splendentis (Horvatić 1973) Terzi 2015 [4K]; Mesobromenalia erecti Royer 1991 nom. inval.
(Art. 3h) [4M]; Scorzonerenalia villosae Terzi 2015;
eschweizerbart_xxx
30 Massimo Terzi
Alliances: Bromion erecti Koch 1926 [5B]; Centaureion dichroanthae Pignatti 1952 [5D]; Chrysopogono grylliDanthonion
calycinae Kojić 1959 [5T]; Chrysopogono grylli-Koelerion splendentis Horvatić 1973 (included within
Chrysopogono grylli-Saturejion subspicatae) [5K]; Chrysopogono grylli-Saturejion subspicatae Horvat & Horvatić ex
Černjavski et al. 1949 (syn. Chrysopogono grylli-Koelerion splendentis Horvatić 1973) [5C]; Cirsio-Brachypodion pinnati
Hadač & Klika in Klika & Hadač 1944; Danthonio-Brachypodion Boșcaiu 1972; Diplachnion Br.-Bl. 1961; Festucion
illyricae Trinajstić 2000 [5F]; Hypochoeridion maculatae Horvatić ex Terzi 2011 [5H]; Mesobromion erecti Oberd.
1949 (synonym of Bromion erecti) [5M]; Saturejion montanae Horvat in Horvat et al. 1974 [5J]; Saturejion subspicatae
Tomić-Stanković 1970 [5S]; Scorzonerion villosae Horvatić ex Kovačević 1959; Xerobromion erecti Zoller 1954.
Suballiances: Centaureenion dichroanthae (Pignatti 1952) Poldini & Feoli Chiapella in Feoli Chiapella & Poldini
1993 [6D]; Festucenion illyricae Horvat in Horvat et al. 1974 corr. Trinajstić 2000 [6F]; Fumano-Scabiosenion leucophyllae
Redžić 1991; Hypochoeridenion maculatae Poldini & Feoli Chiapella ex Terzi 2011 [6H]; Saturejenion montanae
Horvat ex Wendelberger 1965 [6M]; Scorzonerenion villosae Feoli Chiapella & Poldini 1993 [6V].
Associations: Achilleo nobilis-Dorycnietum herbacei Redžić & Lakušić in Redžić 1999; Artemisio albae-Rutetum
graveolentis Redžić & Lakušić in Lakušić & Redžić 1991 [ar]; Artemisio albae-Salvietum officinalis Grebenščikov
1950 [ao]; Asphodelo microcarpi-Chrysopogonetum grylli Horvatić 1963 (cf. Bromo erecti-Chrysopogonetum grylli
Horvatić 1934) [ac]; Astragalo croatici-Seslerietum robustae Trinajstić ex Terzi 2011 [as]; Avenulo praeustae-Brometum
erecti Poldini & Feoli Chiapella ex Terzi 2011 [ab]; Avenulo-Brometum erecti sensu Lasen 1995 non Feoli Chiapella
& Poldini 1993 (= Thlaspidi-Trifolietum pratensis Zanotto 1960 according to Pignatti & Pignatti 2014) [ts];
Bromo condensati-Stipetum eriocaulis Lasen ex Terzi 2015 [si]; Bromo erecti-Chrysopogonetum grylli Horvatić 1934
(incl. Asphodelo microcarpi-Chrysopogonetum grylli Horvatić 1963) [bc]; Bromo erecti-Festucetum lapidosae Trinajstić
1992 [bf]; Bromo erecti-Plantaginetum mediae Horvat 1949; Bromo erecti-Seslerietum interruptae Trinajstić ex Terzi
2011 [bs]; Bupleuro ranunculoidis-Brometum condensati Terzi 2015 (= Bupleuro-Brometum condensati Poldini &
Feoli Chiapella in Feoli Chiapella & Poldini 1993 nom. inval.) [bb]; Carici vernae-Scabiosetum leucophyllae Redžić et
al. ex Terzi 2012 [cs]; Centaureo cristatae-Chrysopogonetum grylli Ferlan & Giacomini 1957 [cc]; Centaureo dichroanthae-Globularietum
cordifoliae Pignatti 1952 [cg]; Centaureo rupestris-Caricetum humilis Horvat 1931 [cr]; Chamaecytiso
hirsuti-Chrysopogonetum grylli Pignatti ex Feoli Chiapella & Poldini 1993 [ch]; Danthonio-Chrysopogonetum
grylli Boșcaiu 1972 [zd]; Danthonio-Scorzoneretum villosae Horvat & Horvatić in Horvatić 1963 (synonym of Scorzonero
villosae-Danthonietum calycinae Kovačević 1959); Dichanthio ischaemum-Cleistogenetum serotinae Horvatić
ex Trinajstić in Poldini 1975 nom. mut. propos. [dc]; Euphorbio nicaeensis-Chrysopogonetum grylli Horvatić 1963
[ec]; Festucetum illyrico-valesiacae Horvat in Horvat et al. 1974 corr. Trinajstić 2000 [fi]; Festuco illyricae-Poetum
bulbosae Horvat in Horvat et al. 1974 corr. Terzi 2011 [fp]; Festuco rubrae-Danthonietum calycinae Csűrös et al. 1968
[zr]; Festuco valesiacae-Danthonietum calycinae Boșcaiu 1972 [zf]; Festuco-Armerietum canescentis Trinajstić & Sugar
1972 [fa]; Festuco-Linetum flavi Ritter-Studnička 1972 [fl]; Genisto sericeae-Seslerietum kalnikensis Dakskobler &
Peljhan ex Terzi 2015 [gk]; Genisto sericeae-Seslerietum tenuifoliae Poldini 1980 [gs]; Genisto-Caricetum mucronatae
Horvat in Horvat et al. 1974 [gc]; Genisto-Globularietum bellidifoliae Tomić-Stanković 1970 [gg]; Gentianello pilosae-Brometum
erecti Dakskobler & Završnik 2009 [gb]; Gladiolo palustris-Molinietum arundinaceae Poldini & Feoli
Chiapella in Feoli Chiapella & Poldini 1993 [gm]; Globulario cordifoliae-Scabiosetum leucophyllae Redžić et al. ex
Terzi 2011 [gl]; Globulario elongatae-Chrysopogonetum grylli Ilijanić et al. ex Terzi 2011 [ge]; Helichryso italiciArmerietum
dalmaticae Horvatić 1963 [ha]; Koelerio macranthae-Brachypodietum retusi Terzi 2015 (= Koelerio macranthae-Brachypodietum
retusi Trinajstić 2005 nom. inval., Art. 10b) [kb]; Koelerio splendentis-Festucetum illyricae
Horvatić 1963 corr. Trinajstić 1992 [kf]; Lactuco vimineae-Bothriochloetum ischaemum Poldini 1975 corr. Terzi 2011
[lb]; Laserpitido-Festucetum alpestris Pedrotti 1970; Leontodonto brumatii-Seslerietum calcariae Dakskobler et al.
2012 [ls]; Lino linearifolii-Gypsophiletum glomerati Tzonev 2009 nom. inval. (Art. 1); Minuartio capillaceae-Genistetum
pulchellae Šegulja & Bedalov ex Terzi 2011 [mg]; Narcisso tazettae-Asphodeletum microcarpi Šegulja 1969 [na];
Onobrychido arenariae-Brometum erecti Poldini & Feoli Chiapella in Feoli Chiapella & Poldini 1993 [ob]; Onobrychido-Brometum
erecti T. Müller 1966; Ononido antiquorum-Brometum condensati (Horvatić 1934) Horvatić 1963
[oa]; Pediculari acaulis-Caricetum humilis Horvat in Horvat et al. 1974 [pc]; Peucedano oreoselini-Lathyretum filiformis
Ritter-Studnička 1972 [pl]; Physospermo-Saturejuetum montanae Redžić & Lakušić in Lakušić & Redžić 1991;
Poo bulbosae-Saturejetum subspicatae Černjavski et al. 1949 [pb]; Potentillo pilosae-Achilleetum clypeolatae Tzonev
2009 nom. inval. (Art. 1); Salvio officinalis-Euphorbietum fragiferae Lausi & Poldini 1962 [se]; Salvio officinalisSeslerietum
juncifoliae Trinajstić 1977 [ss]; Saturejo montanae-Dichanthietum ischaemum Horvat in Horvat et al. 1974
corr. Terzi 2011 [sd]; Saturejo subspicatae-Caricetum humilis Trinajstić 1999 [sm]; Saturejo subspicatae-Edraianthetum
tenuifolii Horvat in Horvat et al. 1974 [su]; Saturejo subspicatae-Festucetum dalmaticae Redžić & Lakušić in
Lakušić & Redžić 1991 [sf]; Saturejo variegatae-Brometum condensati Poldini & Feoli Chiapella in Feoli Chiapella &
Poldini 1993 [sb]; Schoeno nigricantis-Chrysopogonetum grylli Pignatti ex Feoli Chiapella & Poldini 1993 [sn]; Scorzonero
villosae-Danthonietum calycinae Kovačević 1959 [ds]; Scorzonero villosae-Hypochoeridetum maculatae
eschweizerbart_xxx
Numerical analysis of the order Scorzoneretalia villosae 31
Horvatić 1949 nom. nud. [sh]; Seseli gouanii-Artemisietum albae Poldini 1980 [sa]; Seslerio robustae-Juniperetum sibiricae
Domac 1962 [sj]; Seslerio tenuifoliae-Caricetum humilis Horvat 1930 [sc]; Stipo bromoidis-Salvietum officinalis
Horvatić 1963 [so]; Stipo eriocaulis-Caricetum humilis Trinajstić ex Terzi 2011 [st]; Stipo pennatae-Genistetum dalmaticae
Redžić et al. ex Terzi 2011 [sg]; Teucrio capitati-Chrysopogonetum grylli Sburlino et al. 2008 [tc]; ThlaspioTrifolietum
pratensis Zanotto 1960; Thymo serpylli-Teucrietum chamaedryos Redžić & Lakušić in Redžić 1999.
Appendix 2: Nomenclatural notes, including descriptions of new syntaxa and typifications.
[1] The names Scorzoneretalia villosae and Scorzonero-Danthonietum calycinae were validly published for the first
time by Kovačević (1959) with five relevés from Bosanski Petrovac (Bosnia). Kovačević (1959) also validated the Scorzonerion
villosae (Horvatić 1949). Order and alliance, with only one subordinated syntaxon in their original diagnoses,
were automatically typified.
[2] The Scorzonero-Danthonietum calycinae is typified as follows: lectotypus is relevé 2 of table 7 in Kovačević (1959:
30). The Scorzonero-Danthonietum calycinae ranges from Bosnia and Herzegovina to Croatia, Slovenia and Italy.
Poldini (1989) and Poldini & Kaligaric (1997) extensively dealt with this association in Italy and Slovenia and Horvatić
(1963) in Croatia. However, the diagnoses given by these authors are quite different from those given for Bosnia
(Kovačević 1959, Bajić 1960), lacking many important taxa, such as Ferulago campestris, Knautia illyrica, Centaurea
jacea subsp. weldeniana, and Dianthus ferrugineus subsp. liburnicus, while including others with higher frequencies,
such as Veronica austriaca subsp. jacquinii, Scabiosa ochroleuca, Phleum phleoides, Genista sagittalis (acidophilous), as
well as Rhinanthus alectorolophus, Scorzoneroides autumnalis, Cynosurus cristatus and Lotus tenuis shared with the
Molinio-Arrenatheretea. This floristic dissimilarity mirrors different ecological conditions and maybe two different
associations could be differentiated.
[3] The Saturejion subspicatae was validated by Tomić-Stanković (1970) with the valid publication of the GenistoGlobularietum
bellidifoliae, containing Satureja subspicata (see Terzi 2011).
[4] The Centaureo-Caricetum humilis (nomenclatural type of Saturejion subspicatae) was invalidly typified in Terzi
(2011) due to the erroneous reference; the lectotypus of the Centaureo-Caricetum humilis is relevé A in Horvat (1931:
81).
[5] The name of the Chrysopogono-Saturejion subspicatae was validated by Černjavski et al. (1949) with the valid publication
of the Poo-Saturejetum subspicatae, containing Satureja subspicata (the other alliance name-giving taxon,
Chrysopogon gryllus, being within the original diagnosis of Bromo-Chrysopogonetum grylli). The Chrysopogono-Saturejion
subspicatae was thus validly published before the Chrysopogono-Koelerion splendentis and it can be no longer
considered a superfluous name (see Terzi 2011). The Bromo-Chrysopogonetum grylli Horvatić 1934 (table 28 in
Horvatić 1934) is designated as the lectotypus of the Chrysopogono-Saturejion subspicatae Horvat & Horvatić ex
Černjavski et al. 1949. With this choice, I intend to mantain the distinction, largely used until now (e.g., Jovanović et
al. 1986; Trinajstić 2008; Poldini 2009), between the Saturejion subspicatae and the Chrysopogono-Saturejion subspicatae
(Recommendation 19A of ICPN).
[6] The type-relevé of the Bromo-Chrysopogonetum grylli is very similar to the one of the Asphodelo-Chrysopogonetum
grylli even though the former has a lower number of species. The two associations can be thus unified under the
earliest name.
[7] Following Poldini (1989), the Lactuco-Bothriochloetum ischaemum and Seseli-Artemisietum albae are reduced to
the rank of subassociations of the Centaureo-Chrysopogonetum grylli (subass. sedetosum sexangulari Poldini (1975)
1989 and subass. genistetosum sericeae Poldini (1980) 1989, respectively).
[8] The following associations are validated: Bromo condensati-Stipetum eriocaulis Lasen ex Terzi 2015 (invalidly described
due to Art. 3b), holotypus is rel. 7, table 1, in Lasen (1995: 193); Genisto sericeae-Seslerietum kalnikensis
Dakskobler & Peljhan ex Terzi 2015, holotypus is rel. 2, table 4, in Dakskobler & Peljhan (2007: 164).
[9] Two other invalidly described associations (Art. 3g and 10b) can not be validated and must be renamed (Art. 6).
The first is Bupleuro ranunculoidis-Brometum condensati Terzi 2015, holotypus is rel. 9 in Feoli Chiapella & Poldini
(1993: 30); according to Feoli Chiapella & Poldini (1993), differential species of the association are Calamagrostis varia
and Bupleurum ranunculoides (indicated as Bupleurum ranunculoides s.l. in Feoli Chiapella & Poldini [1993: 7 and
30]). The other is Koelerio macranthae-Brachypodietum retusi Terzi 2015 (holotypus is rel. 5, table 1, in Trinajstić
[2005: 351]), whose differential species are those indicated by Trinajstić (2005): Koeleria macrantha and Brachypodium
retusum.
eschweizerbart_xxx
32 Massimo Terzi
Electronic Supplements
Supplementary material associated with this article is embedded in the pdf of this article. The online version of Phytocoenologia
is hosted at the journal’s website www.schweizerbart.com/journals/phyto. The publisher does not bear any
liability for the lack of usability or correctness of Supplementary material.
Supplement S1: Relevés included in the data set.
Supplement S2: Indicator species and their diagnostic role.
Supplement S3: Life-form and chorological spectra.
Please download the electronic supplement and rename the file extension to .zip (For security reasons Adobe does not
allow to embed .exe, .zip, .rar etc. files).
eschweizerbart_xxx