Balkan endemic vascular plants of Montenegro — critical inventory with
chorological and life-form analyses
Authors: Snežana Vuksanović, Gordana Tomović, Marjan Niketić, and Vladimir Stevanović
Source: Willdenowia, 46(3) : 387-397
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
URL: https://doi.org/10.3372/wi.46.46307
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Willdenowia
Annals of the Botanic Garden and Botanical Museum Berlin-Dahlem
SNEŽANA VUKSANOVIĆ1*
, GORDANA TOMOVIĆ2
, MARJAN NIKETIĆ3
& VLADIMIR STEVANOVIĆ2
Balkan endemic vascular plants of Montenegro – critical inventory with chorological
and life-form analyses
Version of record first published online on 15 November 2016 ahead of inclusion in December 2016 issue.
Abstract: Balkan endemic taxa are those whose distribution is restricted exclusively to the Balkan Peninsula. In the
vascular flora of Montenegro there are 372 Balkan endemic taxa (at the species and subspecies levels), of which 39
taxa are distributed only in Montenegro. This paper presents an overview of the Balkan endemic taxa in Montenegro
with their families, chorological groups, life-form categories and distribution by geographical subregion. The richest
families in terms of Balkan endemic taxa are Asteraceae and Caryophyllaceae, while the richest genus is Hieracium
(Asteraceae). The chorological structure of the flora shows that the largest numbers of Balkan endemic taxa are from
the S European mountain and C European mountain chorological groups, while in the life-form spectrum of the
flora hemicryptophytes and chamaephytes are the most numerous. The presence of Balkan endemic taxa in the six
geographical subregions of Montenegro is also analysed. Floristic similarities among the geographical subregions
are shown with a cluster analysis.
Key words: Balkan Peninsula, Montenegro, endemism, vascular plants, checklist, systematics, chorology, life form
Article history: Received 6 May 2016; peer-review completed 29 June 2016; received in revised form 12 July 2016;
accepted for publication 15 July 2016.
Citation: Vuksanović S., Tomović G., Niketić M. & Stevanović V. 2016: Balkan endemic vascular plants of Montenegro
– critical inventory with chorological and life-form analyses. – Willdenowia 46: 387 – 397. doi: http://
dx.doi.org/10.3372/wi.46.46307
Introduction
The term “endemic” has many definitions and it is very
differently understood due to a lack of uniform criteria
for a more precise definition (Tomović 2007). One of
these defines an endemic as “that which occurs nowhere
except in the place in question” (Darlington 1957). According
to Dihoru & Pârvu (1987), endemics are taxonomic
units of different ranks, limited in their distribution
to a determinate territory (province, region, district, etc.).
The phenomenon of endemism characterizes those taxa
whose distribution is restricted to a certain area and is
more reduced than that of other taxa of the same rank
(Vischi & al. 2004).
Balkan endemic taxa are those that are distributed
only in the Balkan Peninsula (Turrill 1929; Strid & Tan
1997; Tomović & al. 2014). The Balkan Peninsula (or
“the Balkans”) is defined as the territory limited by the
Danube and Sava rivers in the north, the Soča (or ­Isonzo)
river in the northwest, the Black Sea in the east, the
Aegean and Ionian seas in the south, and the Adriatic Sea
1	 Natural History Museum of Montenegro, Trg Vojvode Bećir-bega Osmanagića 16, 81110 Podgorica, Montenegro; *e-mail:
vukss@t-com.me (author for correspondence).
2	 Institute of Botany and Botanical Gardens “Jevremovac”, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade,
Serbia.
3	 Natural History Museum, Njegoševa 51, 11000 Belgrade, Serbia.
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388 Vuksanović & al.: Balkan endemic vascular plants of Montenegro
in the west. Although an exhaustive list of the flora of
all the Balkan countries has not yet been completed, it
is estimated that the vascular flora of the Balkans today
includes about 8000 species and subspecies (Stevanović
2005), while the number of endemic species and subspecies
is estimated at between 2600 and 2700 (Stevanović
& al. 2007).
The flora of Montenegro has been the subject of study
for nearly 200 years, during which more than 3000 papers
have been published by over 1000 authors (Pulević 2006).
However, much credit for the exploration of the flora of
Montenegro goes to the Czech botanist, Josef Rohlena,
whose Conspectus florae montenegrinae (Rohlena 1942)
remains to this day the most complete work on the flora
of Montenegro. The Conspectus includes 2623 species
and 194 subspecies of vascular plants, which makes a total
of 2817 taxa. According to Stevanović & al. (1995),
the vascular flora of Montenegro consists of 3136 species
and subspecies and, when disputed taxa are also included,
that number reaches 3336. Data that were not included in
Rohlena’s Conspectus for various reasons, as well as the
findings of botanists after World War II, were integrated in
the supplement to the Conspectus (Pulević 2005). Based
on the data collected so far, it is estimated that the flora of
Montenegro includes more than 3600 species and subspecies
(Stešević & Caković 2013). The richness and diversity
of this flora in a relatively small area of 13 812 km2
can
be explained by the position of Montenegro in the Balkans,
the complex orography, diverse climatic elements, a
heterogeneous land cover and the turbulent geological and
biological history that the flora of the Balkan Peninsula has
undergone (Stevanović & al. 1995).
Despite numerous floristic and taxonomic papers
having been published, no detailed study of the Balkan
endemic flora in Montenegro has been conducted so far.
According to Blečić (1958), who was referring to Turrill
(1929) and who also included later data, the flora of
Montenegro includes 220 taxa (species and subspecies)
endemic to he Balkan Peninsula and 20 taxa endemic to
Montenegro. According to Stevanović & al. (1995), the
flora of Montenegro includes 223 Balkan endemic taxa
accounting for 11.7 % of the total flora of the country. In
addition, papers on Balkan endemic taxa found in Montenegro
for the first time have been published recently
by the following authors: Stevanović & Lakušić (2006),
Stešević & al. (2007), Stešević & Drescher (2010) and
Caković & al. (2014); and papers on newly described
endemic taxa, which have mostly resulted from the revision
of certain genera, have been published by the following
authors: Bräuchler & Cikovac (2007), Niketić &
Stevanović (2007), Schönswetter & Schneeweiss (2009),
Surina & al. (2009), Kučera & al. (2010), Mereďa & al.
(2011), Lakušić & al. (2013), Šingliarová & al. (2013),
Janković & al. (2016) and Tomović & al. (2016).
The aim of this study is to present quantitative data on
the floristics (a critical inventory), chorology (chorological
groups) and life forms of the Balkan endemic vascular
flora of Montenegro. We also aim to determine the degree
of floristic similarity between geographical subregions of
the country.
Material and methods
Study area
Montenegro is located in the SW part of the Balkan
Peninsula at the crossroads of two major geographical
units: the Dinaric Alps and the Mediterranean Sea. The
coastal belt of Montenegro belongs to the Mediterranean,
with the Adriatic coastline extending for 316 km (Fuštić
& Đuretić 2000). Directly above the sea a range of the
coastal Dinaric Alps rises and stretches farther back as
a deep karst plateau. This area of high mountains intersected
by river canyons and gorges stretches over an
area to the north and east that belongs to the Adriatic and
Black Sea (Danube) river system. The mountainous part
of Montenegro covers the majority of the country with
an elevational range from 700 m to high-mountain peaks
above 2500 m. The continental part of the country lies
in the northeast. It is a hilly and mountainous area intersected
by the valleys of numerous large and small rivers
(Fuštić & Đuretić 2000).
In Montenegro three main climatic zones can be distinguished:
(1) Mediterranean; (2) temperate continental;
and (3) mountain (Horvat & al. 1974). The Mediterranean
climate in the narrow sense is restricted to the
coastal region of the Adriatic Sea, and along the slopes
of the Dinaric Alps up to an elevation of 300 – 400 m.
The modified Mediterranean climate or the submediterranean
perhumid variant is present at elevations above
400 m and deeper inland in the gorges and canyons as
well as in some valleys exposed to Mediterranean influences.
The modified Mediterranean climate in Montenegro
belongs to the perhumid variant due to high annual
precipitation of between 1500 and 2200 mm. The temperate
continental climate is characteristic in the NW,
N and E parts of Montenegro. In the mountains in the
area of temperate continental climate, two subtypes of
mountain climate can be distinguished: (3a) the perhumid
Mediterranean-submediterranean mountain climate
with a precipitation of over 4500  mm, pronounced in
the coastal Dinaric Alps of Montenegro; and (3b) a humid
mountain climate of the alpine type, characteristic
of the mountains of the NW, N and E of Montenegro
(Stevanović & Stevanović 1995).
According to the Conspectus of classes of European
vegetation (Mucina 1997), the following vegetation
types and classes are present in Montenegro: (1)
aquatic vegetation (Lemnetea, Charetea fragilis, Potametea,
Ruppietea maritimae, Zosteretea); (2) vegetation
of freshwater marshes and fens (Isoëto-Littorelletea,
Isoëto-Nanojuncetea, Montio-Cardaminetea, PhragmitoMagno­caricetea,
Scheuchzerio-Caricetea fuscae); (3)
coastal vegetation (Cakiletea maritimae, Crithmo-StatiDownloaded
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389Willdenowia 46 – 2016
cetea, Thero-Salicornietea, Salicornietea fruticosae,
Juncetea maritimi, Ammophiletea); (4) chasmophytic
vegetation (Asplenietea trichomanis, Adianthetea capilliveneris,
Epipetrea lichenosa, Thlaspietea rotundifolii);
(5) arctic and alpine vegetation (Loiseleurio-Vaccinietea,
Salicetea herbaceae, Juncetea trifidi, Carici rupestrisKobresietea
bellardii, Elyno-Seslerietea, Mulgedio-Aconitetea);
(6) synanthropic vegetation (Bidentetea tripartiti,
Polygono arenastri-Poetea annuae, Stellarietea
mediae, Artemisietea vulgaris, Galio-Urticetea, Epilobietea
angustifolii); (7) temperate heathlands and grasslands
(Calluno-Ulicetea, Koelerio-Corynephoretea, MolinioArrhenatheretea,
Tri­folio-Geranietea sanguinei, FestucoBrometea);
(8) temperate and boreal woodlands and scrub
(Rhamno-Prunetea, Salicetea purpureae, Alnetea glutinosae,
Populetea albae, Franguletea, Querco-Fagetea,
Quercetea pubescentis, Quercetea roboris, Erico-Pinetea,
Vaccinio-Piceetea); (9) oromediterranean grasslands
and scrub (Daphno-Festucetea); and (10) Mediterranean
vegetation (Thero-Brachypodietea ramosi, Helianthemetea
guttati, Cisto cretici-Micromerietea julianae; Euphorbietea
dendroidis, Nerio-Tamaricetea, Quercetea ilicis).
Selection of taxa
“Taxa” are here counted as species and subspecies. A
species with no or one subspecies is counted as one taxon,
while a species with two subspecies is counted as two
taxa, and so on. A checklist of Balkan endemic vascular
plant taxa in Montenegro was made based on data from
studies on floristics, taxonomy, morphology, phytocoenology
and karyology. The Conspectus florae montenegrinae
(Rohlena 1942) and its supplement (Pulević 2005)
and the catalogue of the vascular flora of Montenegro
(Stešević & Caković 2013) were used as the basis for the
inventory.
Taxonomy and nomenclature follow contemporary
checklists and Floras (Tutin & al. 1964 – 1980, 1993; Jalas
& Suominen 1972 – 1994; Greuter & al. 1984 – 1989; Jalas
& al. 1996, 1999; Strid & Tan 1997; Kurtto & al. 2004,
2007; Greuter & Raab-Straube 2008), online databases
(IOPI 1996 – 2007; Euro+Med 2006+; The Plant List
2013; IPNI 2013+) and in some cases older basic floras
(Pančić 1875; Rohlena 1942).
Data analysis
To define the chorological groups, the methods of Meusel
& al. (1965, 1978), Meusel & Jäger (1992) and Stevanović
(1992) were used, modified and adapted to Montenegro
as proposed by Tomović & al. (2014). The taxa were
classified into the following chorological groups: central
European mountain (CEM); southern European mountain
(SEM); Mediterranean-submediterranean (MEDSUBMED);
central European (CEUR); and Pontic
(PONT).
The classification of taxa as particular life forms is
based on the Raunkiaer (1934) system, supplemented by
Mueller-Dombois & Ellenberg (1974) and modified by
Stevanović (1992). The taxa were classified as phanerophytes
(P), chamaephytes (Ch), hemicryptophytes (H),
geophytes (G), and therophytes (T).
The geographical regionalization was conducted according
to Marković (1970) and Stešević & Caković
(2013) with some minor modifications (Fig. 1). Consequently,
Montenegro is divided into two regions – maritime
and mountain – and six subregions. Within the maritime
region there are two subregions: Mediterranean (M)
and submediterranean (SM); within the mountain region
there are four subregions: central (C); eastern (E); northern
(N); and western (W). The distribution of Balkan endemic
taxa within each of these geographical subregions
was established.
The clustering method based on Sørensen distances
and the unweighted pair-group average hierarchical sorting
strategy (unweighted pair-group method with the arithmetic
mean) was used for identification of the hierarchical
floristic similarities among the six geographical subregions.
The cluster analysis was performed using the package
Statistica 5.1 by StatSoft (http://www.statsoft.com/).
Results
The total number of vascular plant taxa that meet the
criteria of Balkan endemics according to Turrill (1929)
and Strid & Tan (1997) is 372 (Table 1, see SupplemenFig.
1. Geographical subregions of Montenegro (modified after
Stešević & Caković 2013). – C: central; E: eastern; M: Mediterranean;
N: northern; SM: submediterranean; W: western.
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390 Vuksanović & al.: Balkan endemic vascular plants of Montenegro
tary Material online), which is equivalent to
10.4 % of the total vascular flora of Montenegro.
The Balkan endemic taxa in the
study area have been classified into 46 families
and 147 genera. Pteridophytes have
no endemic representatives, while gymnosperms
are represented by only one taxon
(Pinus peuce Griseb.). Angiosperms make
up 99.7 % of the Balkan endemic taxa. The
Balkan endemic flora of Montenegro has
no endemic families, but there is one unispecific
Balkan endemic genus: Petteria C.
Presl (Fabaceae). The families richest in
Balkan endemic taxa (Fig. 2) are Asteraceae
with 108 taxa, followed by Caryophyllaceae
with 34 taxa, while the richest
genus (Fig. 3) is Hieracium (Asteraceae)
with 71 taxa.
The chorological spectrum (Fig. 4)
shows that almost two-thirds of the Balkan
endemic taxa are made up of orophytes,
which belong to two chorological groups:
the S European mountain group (145 taxa,
39.0 %) and the C European mountain group
(100 taxa, 26.9 %). Thus, we can state that
mountain and alpine endemism dominates
the flora of Montenegro. A proportionally
large number of taxa (81, 21.8 %) is represented
by the Mediterranean-submediterranean
chorological group, which is an
indicator of the strong Mediterranean and
submediterranean influences deep into the
interior of the country. The C European
chorological group is represented by 45
taxa (12.1 %), mostly represented by elements
of C European deciduous forests.
The Pontic group is represented by only
one taxon: Stipa joannis subsp. balcanica
Martinovský (Poaceae).
Analysis of the life-form spectrum (Fig.
5) indicates hemicryptophytes are prevalent
with 234 Balkan endemic taxa (62.9 %), followed
by chamaephytes (88 taxa, 23.7 %),
geophytes (22 taxa, 5.9 %), therophytes (19
taxa, 5.1 %) and phanerophytes (nine taxa,
2.4 %).
The largest number of Balkan endemic taxa is recorded
in the eastern (224) and northern (205) geographical
subregions (Fig. 6). A large number is also present in the
submediterranean (162) and western (94) subregions.
The central and Mediterranean subregions are characterized
by a relatively small number of taxa (60 and 57,
respectively).
Cluster analysis of the floristic similarities between
the six geographical subregions of Montenegro based
on the presence of Balkan endemic taxa (Fig. 7) permits
the identification of two groups of subregions. The first
group comprises the eastern and northern subregions
and is well differentiated as a result of the largest number
of Balkan endemic taxa occurring therein. The second
group comprises the submediterranean and western subregions,
between which the close phytogeographical
connection is obvious, because both are geographically
close to each other and both share a similar climate as
well as ecological and floristic features. The central subregion
shows some floristic, climatic, geological and
ecological similarities with the four aforementioned
sub­regions, whereas the Mediterranean subregion stands
Fig. 2. Taxon richness at the family level in the Balkan endemic vascular flora
of Montenegro.
Fig. 3. Taxon richness at the genus level in the Balkan endemic vascular flora
of Montenegro.
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391Willdenowia 46 – 2016
separately as a result of its poor Balkan endemic flora
compared to that of the mountain subregions.
Discussion
The floristic analysis of Balkan endemic taxa singled out
some families that represent an important contribution to
the flora of the Mediterranean basin. Asteraceae are the
largest family in the area stretching from N Europe to the
Mediterranean, over which range the taxonomic diversity
of the family increases from north to south (Stevanović
& al. 1995). The families Caryophyllaceae and Brassicaceae
have a large number of genera in the Mediterranean
basin (Heywood & al. 2007), and the Mediterranean
is also a centre of diversity for the
families Campanulaceae (Kovačić 2004)
and Lamiaceae (Heywood & al. 2007).
Campanulaceae, with a large number of
Balkan endemic taxa in Montenegro (19),
occupies fourth place, which suggests
strong influences from the Mediterranean
climate, despite the high-mountain barriers
(Stevanović & al. 1995). Also, the 10 to
12 families richest in Balkan endemic taxa
more or less coincide with those in Greece
(Georghiou & Delipetrou 2010), Bosnia and
Herzegovina (Lubarda & al. 2014) and the
regions of C Serbia and Kosovo (Tomović
& al. 2014). The main differences between
these four regions are noted in the number
of Balkan endemic taxa that belong to, e.g.,
Rubiaceae (a higher ranking in Greece),
Poaceae (lower ranking in Montenegro),
Lamia­ceae (higher ranking in Bosnia and
Herzegovina) and Campanulaceae (lower
ranking in C Serbia and Kosovo).
In the chorological spectrum of the Balkan
endemic flora of Montenegro, S European
mountain and C European mountain
taxa are predominant, and these together
account for just about two-thirds of the
Balkan endemic flora. The explanation for
such a pronounced predominance of orophytic
elements lies in the mountain and
high-mountain areas of Montenegro being
centres of florogenesis and of refugees
from the orophytic flora of C and S Europe
(Stevanović 1996a; Tomović & al. 2014).
The Pleistocene periods of glaciation, and
especially interglaciation, enabled a more
or less unrestricted migration of species
and the exchange of indigenous elements
between the high-mountain flora of the Dinaric
Alps and the Balkan mountains, and
the Alps and the Carpathians (Kramp & al.
2008; Schmitt & Haubrich 2008; Huck &
al. 2009; Stachurska-Swakon & al. 2013; Surina & al.
2014). Parallel to these processes, speciation occurred
(in the indigenous Tertiary oromediterranean and newly
coming Alpine/Carpathian floras) due to the separation
between the mountains creating genetically isolated
populations (Stevanović & Lakušić 1996; Stefanović &
al. 2008; Surina & al. 2009; Kučera & al. 2010; Mereďa
& al. 2011; Lakušić & al. 2013; Janković & al. 2016).
A high percentage of Balkan endemic taxa belong to
the Mediterranean-submediterranean chorological group
as a result of the presence of a large number of xerophytic
and thermophytic habitats in the Mediterranean, submediterranean,
central and western subregions of Montenegro.
As thermophytic ecological elements, the Balkan
endemic members of this chorological group are also
Fig. 4. Spectrum of chorological groups in the Balkan endemic vascular flora
of Montenegro. – SEM: southern European mountain; CEM: central European
mountain; MED-SUBMED: Mediterranean-submediterranean; CEUR: central
European; PONT: Pontic.
Fig. 5. Spectrum of life forms in the Balkan endemic vascular flora of Montenegro.
– H: hemicryptophytes; Ch: chamaephytes; G: geophytes; T: therophytes;
P: phanerophytes.
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392 Vuksanović & al.: Balkan endemic vascular plants of Montenegro
present in the mountain massifs of the eastern
and northern subregions of Montenegro,
usually in the lowest mountain belt on
the S slopes of mountains, and also in river
canyons under the influence of the Mediterranean
climate. Furthermore, the more
intense impact of anthropogenic factors has
caused the spread of secondary vegetation
types (xerophytic grass and shrub communities)
at the expense of the primary forest
and shrub vegetation. This has allowed the
endemic taxa belonging to the Mediterranean-submediterranean
chorological group
to penetrate deeper into the continental
part of Montenegro, as identified by Turrill
(1929), and has thus caused an increase in
the number of Balkan endemic taxa in this
area. This phenomenon was also demonstrated
by Zlatković & al. (2011) for some
Mediterranean taxa that had extended their
occurrence deep into the C part of the Balkan
Peninsula (Serbia) from the N part of
the Republic of Mace­donia.
The Balkan endemic taxa that inhabit
broad-leaved forests and mesophytic meadows
in the W and C parts of the Balkan
Peninsula belong to the C European chorological
group (Tomović & al. 2014). They
are less important contributors to the chorological
spectrum of Balkan endemic flora
in Montenegro.
The life-form spectrum of floras reflects
the basic features of habitats as well as the
continuum of changes in time and is the
result of adaptation during the evolution of
species (Stevanović & Janković 2001). Most
holarctic plants or plants of the temperate
zone, and of the Balkan Peninsula, belong
to the life-form category of hemicryptophytes.
In addition, hemicryptophytes and
chamaephytes inhabit hill and mountain areas,
and over 80 % of the territory of Montenegro
is represented by hills and mountains. The high
percentage of chamaephytes (as many as 23.4 %) inhabiting
high-mountain and cold habitats, river canyons and
gorges attests to the abundance of these habitats. Endemic
chamaephyte dwarf shrubs are also present in the submediterranean
subregion of Montenegro, where they usually
inhabit thermophytic limestone rocks and rocky ground.
Perennials that develop in such habitats are usually resistant
to drought and are characterized by slow growth (Larson
& al. 2000).
The geophytes in the life-form spectrum of the Balkan
endemic flora of Montenegro inhabit thermophytic and
xerophytic habitats (Mediterranean and temperate woodlands
and scrub, as well as rocky ground) of the submediterranean
and western subregions, but are mostly present
in the high-mountain areas of the eastern and northern subregions
(on rocky ground, screes, high-mountain grasslands
and wet meadows).
The therophytes in the life-form spectrum predominantly
inhabit the Mediterranean and submediterranean
areas of Montenegro. Annual plants mainly inhabit lowproductivity
habitats, such as dry meadows and rocky
ground, where they are exposed to droughts of greater
or lesser intensity during the summer, as well as habitats
that are frequently exposed to anthropogenic and
natural distortions (Noble & Slatyer 1980). The number
of therophytes present in the high-mountain regions of
Montenegro is proportionally lower, while their presence
can be explained by Mediterranean influences that
penetrate deep into the interior of the country through
Fig. 6. Distribution of Balkan endemic vascular plant taxa in the two geographical
regions and six geographical subregions of Montenegro.
Fig. 7. Cluster analysis of floristic similarities among the six geographical subregions
of Montenegro based on the occurrence of Balkan endemic vascular
plant taxa. Using the Sørensen similarity coefficient.
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393Willdenowia 46 – 2016
river canyons and gorges. Phanerophytes are represented
in the Balkan endemic flora of Montenegro by only nine
taxa.
In Montenegro, Stevanović & al. (1995) recorded the
presence of 223 Balkan endemic taxa (i.e. species and
subspecies). In the Balkans, Greece is far ahead in first
place with 1226 single-country endemic taxa (Georghiou
& Delipetrou 2010). In other Balkan countries, the
number of endemic taxa, whether single-country endemic
or Balkan endemic, is much lower. In the flora of Bulgaria,
172 taxa are single-country endemics (Velchev &
Kozhuharov 2006) and 270 are Balkan endemics (Petrova
& Vladimirov 2010). Matevski & al. (2003) recorded
115 single-country endemic species (i.e. excluding subspecies)
in the Republic of Macedonia. In the regions of
C Serbia and Kosovo there are 492 Balkan endemic taxa
(Tomović & al. 2014), in Croatia there are 376 Balkan
endemic taxa (Nikolić & Milašinović 2012), and in Bosnia
and Herzegovina Lubarda & al. (2014) recorded 298
Balkan endemic taxa. Ruci (1997) recorded 26 singlecountry
endemic taxa and c. 150 Balkan endemic taxa in
Albania, but, according to Tomović & al. (2014, quoting
Turrill 1958), the actual numbers are much higher (50
and 320, respectively).
The fact that most of the territory of Montenegro consists
of hills and mountains, except for the narrow coastal
belt, which belongs to the Mediterranean, is reflected by
its Balkan endemic vascular flora. The distribution of
Balkan endemic taxa by subregions shows that the highmountain
type of endemism is dominant in the flora, and
there is also edaphic (limestone) endemism, and a combination
of high-mountain and edaphic endemism is consistent
with previous studies (Lakušić 1985; Stevanović
& al. 1995; Tribsch & Schönswetter 2003; Georghiou
& Delipetrou 2010; Tomović 2014; Jeanmonod & al.
2015). Almost the entire mountain part of Montenegro,
starting from the coastal Dinaric Alps, which belong to
the submediterranean and western subregions, to the
mountains of the eastern and northern subregions, including
the canyons and gorges between the massifs, can
be characterized as a unique centre of diversity of taxa
endemic both to the Balkan Peninsula and to Montenegro
(Stevanović & al. 1995). In the eastern subregion,
the Prokletije mountain range stands out on account of
its extraordinary richness in endemic taxa. This range
of mountains is located at the intersection of the Dinaric
Alps and Scardo-Pindic mountains and is characterized
by the different orientations of the mountain branches,
their massiveness, high average elevation and a complex
relief. The mosaic of geological substrata (limestone and
silicate), exposure to different climates, and diversity of
habitats in this range have enabled the survival and development
of a number of high-mountain endemic taxa that
have differentiated during a long process of speciation
(Stevanović & al. 1995). The endemic flora of the Montenegro
part of the Prokletije mountains represents a link
between the ancient orophytic flora of the Alps and the
Carpathians with the oromediterranean tertiary flora. The
current composition of the flora of the Prokletije massif
largely bears the features of these dynamic processes of
florogenesis (Bulić & al. 2010). The mountain Bjelasica
also belongs to the eastern subregion, and this silicate “island”
of limestone is also an important centre of silicate
endemic flora. The combination of high-mountain and
edaphic endemism occurs exclusively in the eastern subregion,
which is also the richest in Balkan endemic taxa
in the flora of Montenegro. In the northern subregion, the
mountain of Durmitor dominates with its rich flora and
its endemic taxa, together with the canyons of the Piva
and Tara rivers. Despite its uniform limestone geological
composition, Durmitor is one of the most important
developmental and also refuge centres of endemic alpine
flora in the whole Balkan Peninsula (Stevanović &
Lakušić 1996). Also, the canyons of the Piva and Tara
rivers represented an important refuge area of nemoral
and chasmophytic tertiary flora during the Pleistocene
glaciations in the Dinaric Alps. In the submediterranean
subregion, the centres of endemism are the limestone
coastal mountains of Lovćen and Rumija, and in the
western subregion they are the coastal mountain of Orjen
and its branches surrounded by an extreme karst area. In
the central subregion, the alpine type of endemism also
prevails, in the Morača Mountains along the upper course
of the River Morača and its tributaries.
The main generators of endemism in the high mountains
of the eastern and northern subregions and in the
coastal Dinaric Alps are a combination of edaphic, climatic
(the presence of mountain, temperate continental
and modified Mediterranean climates in the mountain
belt and canyons) and habitat conditions, which have
also been noticed in the Balkan endemic flora of C Serbia
and Kosovo (Tomović & al. 2014). The differentiation of
endemic taxa occurred mostly during the glaciation and
interglaciation of the indigenous tertiary oromediterranean
and alpine floras (Stevanović 1996b). The areas of
endemism located outside the high-mountain ranges include
limestone river canyons (the Cijevna river and the
middle course of the Morača river) and some karst areas
in the submediterranean subregion. A relatively small
number of Balkan endemic taxa in the narrow coastal
area of Montenegro (the Mediterranean subregion) can
be explained by the absence of a large number of islands
and the dominance of Adriatic taxa in lowland habitats.
Lowland habitats rarely form refuges suitable for endemics
(at least in temperate latitudes) because of their much
harsher and less stable ecoclimatic conditions, which is
not the case in the mountains (Tribsch & Schönswetter
2003; Tribsch 2004; Jeanmonod & al. 2015). Also, it
should be noted that lowland areas in the Mediterranean
subregion have been intensely transformed by human activity
(urbanization, cultivation and infrastructure).
Cluster analysis clearly demonstrates that the floristically
most similar subregions of Montenegro are (1)
the two richest in Balkan endemic taxa: the eastern and
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394 Vuksanović & al.: Balkan endemic vascular plants of Montenegro
northern subregions; and (2) the two closest geographically
and climatically: the submediterranean and western
subregions.
Acknowledgements
We would like to thank Dmitar Lakušić and Ivana
Janković (Institute of Botany and Botanical Garden, Faculty
of Biology, University of Belgrade) for helping us to
resolve the taxonomic status of species of Edraianthus
(DL) and for making databases of species of Campanula
available (IJ); our colleagues Danka Caković and Danijela
Stešević (Faculty of Sciences of the University of
Montenegro) for providing us with certain herbarium and
literature data; and the Ministry of Education, Science
and Technological Development of the Republic of Serbia
(Grant 173030), which supported this research. We
would also like to thank Boštjan Surina (Natural History
Museum Rijeka, Croatia and University of Primorska,
Slovenia) and an anonymous reviewer for their comments
on an earlier draft of this paper.
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