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15 November 2016 Balkan endemic vascular plants of Montenegro — critical inventory with chorological and life-form analyses
Snežana Vuksanović, Gordana Tomović, Marjan Niketić, Vladimir Stevanović
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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.

Version of record first published online on 15 November 2016 ahead of inclusion in December 2016 issue.

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 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), Mered'a & 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, Phragmito-Magnocaricetea, Scheuchzerio-Caricetea fuscae); (3) coastal vegetation (Cakiletea maritimae, Crithmo-Staticetea, 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 rupestris-Kobresietea 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, Molinio-Arrhenatheretea, Trifolio-Geranietea sanguinei, Festuco-Brometea); (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).

Fig. 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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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 (MED-SUBMED); 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  Supplementary Material online (wi.46.46307_Vuksanovic+al_Table1.pdf)), 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 separately as a result of its poor Balkan endemic flora compared to that of the mountain subregions.

Fig. 2.

Taxon richness at the family level in the Balkan endemic vascular flora of Montenegro.

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Fig. 3.

Taxon richness at the genus level in the Balkan endemic vascular flora of Montenegro

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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.

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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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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 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).

Fig. 6.

Distribution of Balkan endemic vascular plant taxa in the two geographical regions and six geographical subregions of Montenegro.

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Fig. 7.

Cluster analysis of floristic similarities among the six geographical sub-regions of Montenegro based on the occurrence of Balkan endemic vascular plant taxa. Using the Sørensen similarity coefficient.

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The therophytes in the life-form spectrum predominantly inhabit the Mediterranean and submediterranean areas of Montenegro. Annual plants mainly inhabit low-productivity 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 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 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.

References

1.

Blečić V. 1958: O nekim karakteristikama flore i vegetacije Crne Gore. — Zašt. Prir. 13: 1–6. Google Scholar

2.

Bräuchler C. & Cikovac P. 2007: Iris orjenii (Iridaceae) — a new species from the littoral Dinaric Alps. —  Willdenowia 37: 221–228. Google Scholar

3.

Bulić Z., Stevanović V., Lakušić D. & Hadžiablahović S. 2010: Biljni svijet NP Prokletije. [Flora of National Park Prokletije]. — Časopis Nacionalnih parkova Crne Gore 6: 18–23. Google Scholar

4.

Caković D., Stešević D., Vuksanović S. & Tan K. 2014: Colchicum cupanii Guss. subsp. glossophyllum (Heldr.) Rony, Datura innoxia Mill. and Eclipta prostrata (L.) L., new floristic records in Montenegro and western Balkan. —  Acta Bot. Croat. 73: 255–265. Google Scholar

5.

Darlington P.J. Jr 1957: Zoogeography: the geographical distribution of animals. — New York: John Wiley an Sons. Inc.; London: Chapman and Hall. Ltd. Google Scholar

6.

Dihoru G. & Pârvu C. 1987: Plante endemice în flora României. [Endemic plants in the flora of Romania]. — Bucureşti: Editura Ceres. Google Scholar

7.

Euro+Med 2006+ [continuously updated]: Euro+Med PlantBase — the information resource for Euro-Mediterranean plant diversity. — Published at  http://ww2.bgbm.org/EuroPlusMed/query.asp [accessed Apr 2015]. Google Scholar

8.

Fuštić B. & Đuretić G. 2000: Zemljišta Crne Gore. [Soils of Montenegro]. — Podgorica: Univerzitet Crne Gore, Biotehnički Institut. Google Scholar

9.

Georghiou K. & Delipetrou P. 2010: Patterns and traits of the endemic plants of Greece. —  Bot. J. Linn. Soc. 162: 130–422. Google Scholar

10.

Greuter W., Burdet H. M. & Long G. (ed.) 1984–1989: Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries 1 (1984), 3 (1986), 4 (1989). — Genève: Conservatoire et Jardin botaniques de la Ville de Genève; Berlin: Secrétariat Med-Checklist, Botanischer Garten und Botanisches Museum Berlin-Dahlem. Google Scholar

11.

Greuter W. & Raab-Straube E. von (ed.) 2008: MedChecklist. A critical inventory of vascular plants of the circum-mediterranean countries 2. — Palermo, Genève & Berlin: OPTIMA. Google Scholar

12.

Heywood V. H., Brummitt R. K., Culham A. & Seberg O. 2007: Flowering plant families of the world. — Kew: Kew Publishing. Google Scholar

13.

Horvat I., Glavač V. & Ellenberg H. 1974: Vegetation Südosteuropas. — Stuttgart: Gustav Fischer. Google Scholar

14.

Huck S., Büdel B., Kadereit W. J. & Printzen C. 2009: Range-wide phylogeography of the European temperate-montane herbaceous plant Meum athamanticum Jacq.: evidence for periglacial persistence. —  J. Biogeogr. 36: 1588–1599. Google Scholar

15.

IOPI 1996–2007: The International Organization for Plant Information. Provisional global plant checklist. — Published at  http://bgbm3.bgbm.fu-berlin.de/iopi/gpc/default.asp [accessed 23 Nov 2007] Google Scholar

16.

IPNI 2013+ [continuously updated]: The International Plant Names Index. — Published at  http://www.ipni.org [accessed 15 Mar 2013]. Google Scholar

17.

Jalas J. & Suominen J. (ed.) 1972–1994: Atlas florae europaeae. Distribution of vascular plants in Europe 1 (1972), 2 (1973), 3 (1976), 4 (1979), 5 (1980), 6 (1983), 7 (1986), 8 (1989), 9 (1991), 10 (1994). — Helsinki: Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. Google Scholar

18.

Jalas J., Suominen J. & Lampinen R. (ed.) 1996: Atlas florae europaeae. Distribution of vascular plants in Europe 11. — Helsinki: Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. Google Scholar

19.

Jalas J., Suominen J., Lampinen R. & Kurtto A. (ed.) 1999: Atlas florae europaeae. Distribution of vascular plants in Europe 12. — Helsinki: Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. Google Scholar

20.

Janković I., Šatović Z., Liber Z., Kuzmanović N., Radosavljević I. & Lakušić D. 2016: Genetic diversity and morphological variability in the Balkan endemic Campanula secundiflora s.l. (Campanulaceae). —  Bot. J. Linn. Soc. 180: 64–88. Google Scholar

21.

Jeanmonod D., Naciri Y., Schlüssel A. & Gamisans J. 2015: Floristic analyses of the Corsican flora: biogeographical origin and endemism. —  Candollea 70: 21–41. Google Scholar

22.

Kovačić S. 2004: The genus Campanula L. (Campanulaceae) in Croatia, circum-Adriatic and west Balkan region. — Acta Bot. Croat. 63: 171–202. Google Scholar

23.

Kramp K., Huck S., Niketić M., Tomović G. & Schmitt T. 2008: Multiple glacial refugia and complex post-glacial range shifts of the obligatory woodland plant Polygonatum verticillatum (Convallariaceae). —  Pl. Biol. (Stuttgart) 11: 392–404. Google Scholar

24.

Kučera J., Marhold K. & Lihová J. 2010: Cardamine maritima group (Brassicaceae) in the amphi-Adriatic area: a hotspot of species diversity revealed by DNA sequences and morphological variation. — Taxon 59: 148–164. Google Scholar

25.

Kurtto A., Fröhner S. E. & Lampinen R. (ed.) 2007: Atlas florae europaeae. Distribution of vascular plants in Europe 14. — Helsinki: Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. Google Scholar

26.

Kurtto A., Lampinen R. & Junikka L. (ed.) 2004: Atlas florae europaeae. Distribution of vascular plants in Europe 13. — Helsinki: Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. Google Scholar

27.

Lakušić D., Liber Z., Nikolić T., Surina B., Kovačić S., Bogdanović S. & Stefanović S. 2013: Molecular phylogeny of the Campanula pyramidalis species complex (Campanulaceae) inferred from chloroplast and nuclear non-coding sequences and its taxonomic implications. —  Taxon 62: 505–524. Google Scholar

28.

Lakušić R. 1985: Nov rod i nove vrste za floru Crne Gore. [A new genus and new species for the flora of Montenegro]. — God. Biol. Inst. Univ. Sarajevu 38: 73–75. Google Scholar

29.

Larson D. W., Matthes U. & Kelly P. E. 2000: Cliff ecology: pattern and process in cliff ecosystems. —  Cambridge: Cambridge University Press. Google Scholar

30.

Lubarda B., Stupar V., Milanović Đ. & Stevanović V. 2014: Chorological characterization and distribution of the Balkan endemic vascular flora in Bosnia and Herzegovina. — Bot. Serbica 38: 167–184. Google Scholar

31.

Marković M. 1970: Geografske oblasti Socijalističke Federativne Republike Jugoslavije. [The geographical areas of the Socialist Federal Republic of Yugoslavia]. — Beograd: Zavod za udžbenike i nastavna sredstva Srbije. Google Scholar

32.

Matevski V., Petkovski S., Andonov S., Melovski L. & Krstić S. 2003: Country study for biodiversity of the Republic of Macedonia. — Skopje: MoEPP. Google Scholar

33.

Mered'a P. Jr , Hodálová I., Kučera J., Zozomová-Lihová J., Letz R. D. & Slovák M. 2011: Genetic and morphological variation in Viola suavis s.l. (Violaceae) in the western Balkan Peninsula: two endemic subspecies revealed. —  Syst. Biodivers. 9: 211–231. Google Scholar

34.

Meusel H., Jäger E. & Weinert E. 1965: Vergleichende Chorologie der zentraleuropäischen Flora 1. Karten. — Jena: Gustav Fischer. Google Scholar

35.

Meusel H., Jäger E. & Weinert E. 1978: Vergleichende Chorologie der zentraleuropäischen Flora 2. Karten. — Jena: Gustav Fischer. Google Scholar

36.

Meusel H. & Jäger E. 1992: Vergleichende Chorologie der zentraleuropäischen Flora 3. Karten, Literatur, Register. — Jena, Stuttgart, New York: Gustav Fischer. Google Scholar

37.

Mucina L. 1997: Conspectus of classes of European vegetation. —  Folia Geobot. Phytotax. 32: 117–172. Google Scholar

38.

Mueller-Dombois D. & Ellenberg H. 1974: Aims and methods of vegetation ecology. — New York: John Wiley & Sons. Google Scholar

39.

Niketić M. & Stevanović V. 2007: A new species of Heliosperma (Caryophyllaceae) from Serbia and Montenegro. —  Bot. J. Linn. Soc. 154: 55–63. Google Scholar

40.

Nikolić T. & Milašinović B. 2012: Distribution pattern, range, ecology, hotspots and conservation of Croatian endemic plants. — Pp. 35–36 in: International Symposium on “Evolution of Balkan Biodiversity”. Book of abstracts. — Zagreb: BalkBioDiv Consortium and Croatian Botanical Society. Google Scholar

41.

Noble I. R. & Slatyer R. O. 1980: The use of vital attributes to predict successional changes in plant communities subject to recurrent disturbances. —  Veg. Hist. & Archaeobot. 43: 5–21. Google Scholar

42.

Pančić J. 1875: Elenchus plantarum vascularium quas aestate a. 1873 in Crna Gora. — Belgradi: Edidit Societas Erudita Serbica. Google Scholar

43.

Petrova A. & Vladimirov V. 2010: Balkan endemics in the Bulgarian flora. — Phytol. Balcan. 16: 293–311. Google Scholar

44.

Pulević V. 2005: Građa za vaskularnu floru Crne Gore. Dopuna “Conspectus Florae Montenegrinae”. [Material for the vascular flora of Montenegro. Supplementum to “Conspectus Florae Montenegrinae”]. — Podgorica: Posebno izdanje Republičkog zavoda za zaštitu prirode Crne Gore. Google Scholar

45.

Pulević V. 2006: Botaničari i Crna Gora. [Botanists and Montenegro]. — Podgorica: Prirodnjački muzej Crne Gore, posebna izdanja, knj. 2. Google Scholar

46.

Raunkiaer C. 1934: The life forms of plants and statistical plant geography. — London: Clarendon Press. Google Scholar

47.

Rohlena J. 1942: Conspectus florae montenegrinae. — Preslia 20–21: 1–506. Google Scholar

48.

Ruci B. 1997: An overview of Albanian flora and Albanian plant endemism. First Balkan Botanical Congress. Abstracts, Thessaloniki, Greece. — Thessaloniki: Greek Ministry of Macedonia-Thrace. Google Scholar

49.

Schmitt T. & Haubrich K. 2008: The genetic structure of the mountain forest butterfly Erebia euryale unravels the late Pleistocene and postglacial history of the mountain coniferous forest biome in Europe. —  Molec. Ecol. 17: 2194–2207. Google Scholar

50.

Schönswetter P. & Schneeweiss G. M. 2009: Androsace komovensis sp. nov., a long mistaken local endemic from the southern Balkan Peninsula with biogeographic links to the eastern Alps.— Taxon 58: 544–549. Google Scholar

51.

Šingliarová B., Šuvada R. & Mráz P. 2013: Allopatric distribution, ecology and conservation status of the Pillosela alpicola group (Asteraceae). —  Nordic J. Bot. 31: 122–128. Google Scholar

52.

Stachurska-Swakon A., Cielślak E. & Ronikier M. 2013: Phylogeography of a subalpine tall-herb Ranunculus platanifolius (Ranunculaceae) reveals two main genetic lineages in the European mountains. —  Bot. J. Linn. Soc. 171: 413–428. Google Scholar

53.

Stefanović S., Lakušić D., Kuzmina M., Međedović S., Tan K. & Stevanović V. 2008: Molecular phylogeny of Edraianthus (grassy bells; Campanulaceae) based on non-coding plastid DNA sequences. — Taxon 57: 452–475. Google Scholar

54.

Stešević D. & Caković D. 2013: Katalog vaskularne flore Crne Gore 1. [Catalogue of vascular flora of Montenegro 1]. — Podgorica: Crnogorska akademija nauka i umjetnosti-Odjeljenje prirodnih nauka. Knjiga 7. Google Scholar

55.

Stešević D. & Drescher A. 2010: Additions to the vascular flora of Montenegro (new taxa and new records). -Nat. Montenegr. 10: 7–16. Google Scholar

56.

Stešević D., Petrović D., Bubanja N., Vuksanović S. & Biberdžić V. 2007: Contribution for the flora of Montenegro (Supplementum to the material for vascular flora of Montenegro). — Nat. Montenegr. 7: 463–480. Google Scholar

57.

Stevanović B. & Janković M. 2001: Ekologija biljaka sa osnovama fiziološke ekologije biljaka. [Plant ecology with a basis of plant physiological ecology]. — Beograd: NNK International. Google Scholar

58.

Stevanović V. 1992: Floristička podela teritorije Srbije sa pregledom viših horiona i odgovarajućih flornih elemenata. [Floristic division of the territory of Serbia with an overview of higher chorions and appropriate floristic elements]. — Pp. 47–56 in: Sarić M. R. (ed.), Flora Srbije 1. — Beograd: Srpska akademija nauka. Google Scholar

59.

Stevanović V. 1996a: Analysis of the central European and Mediterranean orophytic element on the mountains of the west and central Balkan Peninsula, with special reference to endemics. — Bocconea 5: 77–97. Google Scholar

60.

Stevanović V. 1996b: Fitogeografska analiza flore Durmitora. [Phytogeographical analysis of the flora of Mt Durmitor]. — Pp. 185–205 in: Lješević M. (ed.), Priroda Nacionalnog parka “Durmitor”. — Beograd: Geografski fakultet, Posebna izdanja 8. Google Scholar

61.

Stevanović V. 2005: Procena biodiverziteta — od interpretacije do konzervacije. Primer endemične vaskularne flore Balkanskog poluostrva. — Pp. 53–73 in: Anđelković M. (ed), “Biodiverzitet na početku novog milenijuma”, Zbornik radova sa naučnog skupa. — Beograd: Srpska Akademija nauka i umetnosti, Naučni skupovi CXI. Odeljenje hemijskih i bioloških nauka 2. Google Scholar

62.

Stevanović V., Jovanović S., Lakušić D. & Niketić M. 1995: Diverzitet vaskularne flore Jugoslavije sa pregledom vrsta od međunarodnog značaja. [Diversity of vascular flora with an overview of internationally important species]. — Pp. 183–217 in: Stevanović V. & Vasić V. (ed.), Biodiversity of Yugoslavia with an overview of internationally important species. – Beograd: Biološki fakultet, Ecolibri. Google Scholar

63.

Stevanović V. & Lakušić D. 1996: Florističke i florogenetske karakteristike visokoplaninske endemične flore Durmitora. [Floristic and florogenetic characteristics of the high-mountain endemic flora of Mt Durmitor]. — Pp. 206–219 in: Lješević M. (ed), Priroda Nacionalnog parka “Durmitor”. — Beograd: Geografski fakultet, Posebna izdanja 8. Google Scholar

64.

Stevanović V. & Lakušić D. 2006: Reports 72. Lactuca visianii and 73. Asperula hercegovina. — Pp. 291–292 in: Vladimirov V., Dane F., Nikolić T., Stevanović V. & Tan K., New floristic records in the Balkans 2. — Phytol. Balcan. 12: 279–301. Google Scholar

65.

Stevanović V. & Stevanović B. 1995: Osnovni klimatski, geološki i pedološki činioci biodiverziteta kopnenih ekosistema Jugoslavije. [Basic climatic, geological and pedological factors of biodiversity of the terrestrial ecosystems of Yugoslavia]. — Pp. 75–95 in: Stevanović V. & Vasić V. (ed.), Biodiversity of Yugoslavia with an overview of internationally important species. — Beograd: Biološki fakultet, Ecolibri. Google Scholar

66.

Stevanović V., Tan K. & Petrova A. 2007: Mapping the endemic flora of the Balkans — a progress report. — Bocconea 21: 131–137. Google Scholar

67.

Strid A. & Tan K. (ed.) 1997: Flora hellenica 1. — Königstein: Koeltz Scientific Books. Google Scholar

68.

Surina B., Pflanzelt S., Einzmann H. J. R. & Albach D. C. 2014: Bridging the Alps and the Middle East: evolution, phylogeny and systematics of the genus Wulfenia Jacq. (Plantaginaceae). —  Taxon 63: 843–858. Google Scholar

69.

Surina B., Rakić T., Stefanović S., Stevanović V. & Lakušić D. 2009: One new species of the genus Edraianthus, and a change in taxonomic status for Edraianthus serpyllifolius f. pilosulus (Campanulaceae) from the Balkan Peninsula. —  Syst. Bot. 34: 602–608. Google Scholar

70.

The Plant List 2013: The Plant List. Version 1.1. — Kew: Royal Botanic Gardens. — Published at  http://www.theplantlist.org [accessed Sep 2013]. Google Scholar

71.

Tomović G. 2007: [Phytogeographical reference, distribution and diversity centres of the Balkan endemic flora in Serbia]. — Belgrade: Ph.D. thesis, University of Belgrade. Google Scholar

72.

Tomović G., Niketić M., Lakušić D., Ranđelović V. & Stevanović V. 2014: Balkan endemic plants in central Serbia and Kosovo regions: distribution patterns, ecological characteristics and centres of diversity. —  Bot. J. Linn. Soc. 176: 173–202. Google Scholar

73.

Tomović G., Niketić M., Lazarević M. & Melovski L. J. 2016: Taxonomic reassessment of Viola aetolica and Viola elegantula (V. sect. Melanium, Violaceae), with descriptions of two new species from the Balkan Peninsula. —  Phytotaxa 253: 237–265. Google Scholar

74.

Tribsch A. 2004: Areas of endemism of vascular plants in the eastern Alps in relation to Pleistocene glaciation. —  J. Biogeogr. 31: 747–760. Google Scholar

75.

Tribsch A. & Schönswetter P. 2003: Patterns of endemism and comparative phylogeography confirm palaeoenvironmental evidence for Pleistocene refugia in the eastern Alps. —  Taxon 52: 477–497. Google Scholar

76.

Turrill W. B. 1929: The plant life of the Balkan Peninsula. A phytogeographical study. — Oxford: Clarendon Press. Google Scholar

77.

Turrill W. B. 1958: The evolution of floras with special reference to those of the Balkan Peninsula. — J. Linn. Soc., Bot. 56: 136–152. Google Scholar

78.

Tutin T. G., Heywood V. H., Burges N. A., Moore D. M., Valentine D. H., Walters S. M. & Webb D. A. (ed.) 1964–1980: Flora europaea 1 (1964), 2 (1968), 3 (1972), 4 (1976), 5 (1980). — Cambridge: Cambridge University Press. Google Scholar

79.

Tutin T. G., Burges N. A., Chater A. O., Edmondson J. R., Heywood V. H., Moore D. M., Valentine D. H., Walters S. M. & Webb D. A. (ed.) 1993: Flora europaea 1 Psilotaceae to Platanaceae, ed. 2. — Cambridge: Cambridge University Press. Google Scholar

80.

Velchev V. & Kozhuharov S. 2006: Bulgarian endemic higher plants gene pool, origin and distribution. — Pp. 14–33 in: Petrova A. (ed.), Atlas of Bulgarian endemic plants. — Sofia: Gea-Libris Publishing House. Google Scholar

81.

Vischi N., Natale E. & Villamil C. 2004: Six endemic plants species from central Argentina: an evaluation of then conservation status. —  Biodivers. & Conservation 13: 997–1008. Google Scholar

82.

Zlatković B., Randelović V., Lakušić D. & Stevanović V. 2011: Novelties for the vascular flora of Serbia. — Bot. Serbica 35: 103–110. Google Scholar
© 2016 The Authors · This open-access article is distributed under the CC BY 4.0 licence
Snežana Vuksanović, Gordana Tomović, Marjan Niketić, and Vladimir Stevanović "Balkan endemic vascular plants of Montenegro — critical inventory with chorological and life-form analyses," Willdenowia 46(3), 387-397, (15 November 2016). https://doi.org/10.3372/wi.46.46307
Received: 6 May 2016; Accepted: 1 July 2016; Published: 15 November 2016
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KEYWORDS
Balkan Peninsula
checklist
chorology
Endemism
life form
Montenegro
systematics
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