bs_bs_banner Botanical Journal of the Linnean Society, 2014, 176, 173–202. With 11 figures Balkan endemic plants in Central Serbia and Kosovo regions: distribution patterns, ecological characteristics, and centres of diversity 1 Institute of Botany and Botanical Garden ‘Jevremovac’, Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia 2 Natural History Museum, Njegoševa 51, 11000 Belgrade, Serbia 3 Department of Biology and Ecology, Faculty of Natural Sciences, University of Niš, Višegradska 33, 18000 Niš, Serbia Received 26 August 2013; revised 8 May 2014; accepted for publication 23 July 2014 The aim of the present study is to determine the exact number of the Balkan endemic taxa at specific and subspecific rank in the Central Serbia and Kosovo regions, as well as their distribution, and to perform a chorological and ecological analysis of this flora. Detailed and long-term field studies confirmed the presence of 492 Balkan endemic species and subspecies in the investigated area. The most important contributors at family rank in the Balkan endemic flora are Asteraceae, Caryophyllaceae, Fabaceae, Brassicaceae, and Lamiaceae, with Hieracium, Dianthus, Centaurea, and Silene being the genera with the highest number of endemic taxa. In the chorological and life form spectra of the Balkan endemics, taxa from South European and Central European mountainous chorological groups and hemicryptophytes and chamaephytes are the most abundant. The presence of endemic plants in 11 geographical provinces in Central Serbia and Kosovo regions was also analyzed and cluster analysis was used to classify the geographical provinces according to their floristic similarities. The final analyses of distribution and magnitude of endemism include summary mapping of all the endemic taxa at 10 × 10-km Universal Transverse Mercator maps. High-mountainous areas of Mts Prokletije and Mts Šar-Planina are distinctive centres of richness and diversity of the Balkan endemic flora in Central Serbia and Kosovo regions. Most endemics were recorded on limestone and at 1500–2000 m a.s.l. Vegetation classes Asplenietea trichomanes, Festuco-Brometea, Elyno-Seslerietea, and Erico-Pinetea are the richest in endemic taxa. A catalogue of the Balkan endemic taxa in Central Serbia and Kosovo regions, with their threatened and legally protected status, is also presented. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202. ADDITIONAL KEYWORDS: ecology – endemism – flora – Peninsula. INTRODUCTION The term ‘endemic’ simply means occurring nowhere else (i.e. organisms can be endemic to a geographical location on a variety of spatial scales and at different taxonomic levels) (Brown & Lomolino, 1998). According to Peterson & Watson (1998), ‘endemic’ should be used to refer to restriction of taxon to a stated geographical region, based on either natural *Corresponding author. E-mail: gtomovic@bio.bg.ac.rs geographical features or human political boundaries. By following this definition, Balkan endemic plants can be considered such taxa that are restricted only to the territory of the Balkan Peninsula (Turrill, 1929; Strid & Tan, 1997). Kruckeberg & Rabinowitz (1985) were the first to introduce the terms narrow (local), regional and continental endemism, based on different ranges of species distribution. According to Stevanović, Tan & Iatrou (2003), some patterns of endemism for the Balkan obligate serpentine endemics can also be © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 173 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 GORDANA TOMOVIĆ1*, MARJAN NIKETIĆ2, DMITAR LAKUŠIĆ1, VLADIMIR RAND̄ELOVIĆ3 and VLADIMIR STEVANOVIĆ1 174 G. TOMOVIĆ ET AL. MATERIAL AND METHODS STUDY AREA The territory of the Central Serbia and Kosovo regions lies in the north-central part of the Balkan Peninsula and covers 88 361 km2. Extending from the rivers Sava and Danube southwards it ends with a long, almost meridionally extending massif of Mts Šar-Planina belonging to the Scardo-Pindhian mountain system. It is bordered by the mountains of the Carpathian-Balkan and Rhodope systems in the east, and the river Drina and the eastern Dinaric Alps form its western and the high Mts Prokletije (which also belongs to the Dinaric Alps) its south-western borders. In addition to lying in the Balkans, Central Serbia and Kosovo regions also occupy the southeastern part of the Pannonian Plain. Thus, these two regions are divided into two distinct geographical and orographical entities: a low-land-montane Peripannonian and a mountain-valley part of the Balkan mainland. These basic geographical and landscape entities have determined the characteristics of the flora and vegetation of Central Serbia and Kosovo regions and differences between the plants distributed in these two entities (Stevanović et al., 1999). In Central Serbia and Kosovo regions, the climate is continental in the north and south-east with semi-arid summer and cold winter periods. In the west, it is humid temperate and in the centre and the east it is semi-arid temperate-continental or subcontinental, with transitional sub-Mediterranean parts (Stevanović & Šinžar-Sekulić, 2009). There are four basic groups of geological substrata in Central Serbia and Kosovo regions: (1) silicate rocks of acidic to neutral pH (igneous, metamorphic, sedimentary); (2) silicate rocks of basic to ultra-basic reaction (serpentinites and peridotites, ophiolitic belt); (3) carbonate rocks of neutral to basic reactions (clastic, sedimentary); and (4) loess and Pleistocene sediments and deposits (sands, alluvial fans) (Stevanović & Stevanović, 1995). The main types of vegetation formations and vegetation classes are present in Central Serbia and Kosovo regions comprise: (1) coastal mud-flats and brackish waters (Ruppietea maritimae); (2) salt marsh, sand-dune and sea-cliff vegetation (Festuco-Puccinellietea); (3) rock crevice, scree and boulder-field vegetation (Asplenietea trichomanis, Thlaspietea rotundifolii); (4) freshwater aquatic vegetation (Lemnetea, Charetea fragilis, Potametea); (5) springs, shoreline and swamp vegetation (Montio-Cardaminetea, Isoëto-Nanojuncetea, Phragmito-Magnocaricetea); (6) bogs and fens (Scheuchzerio-Caricetea fuscae, OxycoccoSphagnetea); (7) temperate grasslands, heaths and fringe vegetation (Molinio-Arrhenatheretea, © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 established according to their distributional range: (1) trans-Balkan or trans-regional Balkan endemics (taxa distributed in the greater part of the serpentine areas in the Balkans); (2) regional endemics (taxa restricted to a single floristic subregion or province); and (3) local or steno-endemics (taxa distributed in a single floristic district or narrow geographical area such as a single mountain or island). A major contribution on the Balkan Peninsula flora was accomplished by Hayek (1924–1933), although the most comprehensive study of the Balkan flora was that of Turrill (1929) who numbered 1754 Balkan endemic species, representing 26.8% of the total vascular flora of the Peninsula. After Turrill, recent studies of the Balkan endemic flora were conducted by Stevanović (2005) and Stevanović, Tan & Petrova (2007), with estimates of the total number of the Balkan endemics (species and subspecies) of 2600– 2700. In addition to the entire endemic flora, 300 taxa belonging of obligate endemic serpentinophytes (Stevanović et al., 2003) and 170 Balkan endemic geophytes from the class Monocotyledones (Tan, Stevanović & Strid, 2007) have been analyzed However, besides comprehensive floristic researches of the Balkan Peninsula, to date, there has been only one detailed analysis of the Balkan plant endemics in the territory of Central Serbia and Kosovo regions (Gajić, 1984). The study recorded 197 endemic plants in Serbia, which were divided into the categories: (1) Moesian endemics (27 taxa); (2) Illyrian endemics (69 taxa); (3) Pannonian endemics (one species); (4) Endemics that are distributed in several floristic provinces (74 taxa); and (5) Local endemics (26 taxa). According to Stevanović et al. (1995), the number of Balkan endemics in Central Serbia and Kosovo regions is much larger (287 species and subspecies), representing 8.06% of the total vascular flora of this area. The aim of the present study is to present quantitative data on the taxonomy, geographical distribution (division into chorological groups and floristic elements), and ecology (life form, elevational distribution, geological substratum preference, and vegetation characteristics) of the Balkan endemic plants in Central Serbia and Kosovo regions. We also determine the degree of floristic similarity among individual provincial groups and use the results of our analyses to determine the most important centres of floristic richness of the Balkan endemics in Central Serbia and Kosovo regions. The study of the above aspects, besides revealing general patterns of endemism, can also be used for conservation purposes. The present study comprises a part of PhD thesis of the first author (Tomović, 2007) at the Faculty of Biology of the University of Belgrade. BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS SELECTION OF SPECIES As Balkan endemics, we marked all those plant species or subspecies with natural distributions delimited by the territory of the Balkan Peninsula, as defined by Turrill (1929) and Strid & Tan (1997). A list of the Balkan endemic taxa in Central Serbia and Kosovo regions was compiled based on data from Flora of Serbia (Josifović, 1970–1977; Sarić & Diklić, 1986; Sarić, 1992; Stevanović, 2012) and recently published monographs and articles (Matović & Tatić, 2002; Stevanović et al., 2003; Niketić & Stevanović, 2007; Kučera et al., 2008; Lakušić et al., 2013). Systematics and nomenclature were correlated with the latest checklists, papers, and databases (Stevens, 2001; Greuter, Burdet & Long, 1984–1989; Jalas & Suominen, 1972–1994; Jalas, Suominen & Lampinen, 1996; Jalas et al., 1999; Kurtto, Lampinen & Junikka, 2004; Kurtto, Fröhner & Lampinen, 2007; Greuter & Raab-Straube, 2008; http://www.mobot.org/MOBOT/research/APweb; http://plantnet.rbgsyd.nsw.gov.au/iopi/iopihome.htm http://ww2.bgbm.org/EuroPlusMed/) and with some older but basic floras (Pančić, 1874, 1884; Hayek, 1924–1933). To obtain field data and to make additional chorological observations of the endemic plants, field explorations were carried out in the period 2003–2012. Identification and revision of the herbarium specimens used standard botanical literature sources and monographs (Tutin et al., 1968–1980; 1993; Josifović, 1970–1977; Sarić & Diklić, 1986; Sarić, 1992). In addition to the sexual taxa, apomictic microspecies and subspecies were also included. Wherever possible, the narrow taxonomic concept was applied to apomictic taxa, particularly for representatives of Hieracium and Alchemilla. Some groups of these genera have centres of diversity in the Balkans, unlike some other apomictic complexes that are under-represented or under-explored in this part of Europe (e.g. Rubus L., Taraxacum F.H.Wigg.). Therefore, we should expect some overestimation of these genera in the analysis results. DATA AND ANALYSIS The dataset used in the present study was derived from the database with coded data on the nomenclature, taxonomy, distribution and ecology of the Balkan endemic plants in Central Serbia and Kosovo regions. It is regularly updated and currently contains 19307 literature records based on the relevant floras and another 1281 published literature sources and information from 7735 inspected and revised herbarium sheets deposited in the Collections of the Natural History Museum (BEO) and the Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade (BEOU) (Thiers, 2013). For the purpose of obtaining additional chorological and ecological observations on the endemic plants, field explorations were carried out in the period 2003– 2013. To obtain different types of statistical analysis, grouping and generalization of all the ecological data had to be carried out. Distribution data were coded using 11 (out of 15) geographical provinces of Central Serbia and Kosovo regions (Fig. 1) defined by Marković (1970). Four geographical provinces were not included in the analysis, because in the province of Pomoravlje (Po) there are no Balkan endemics, and three geographical provinces of Vojvodina region – [Banat (B), Bačka (Bč) and Srem (Sr)] do not belong to the © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Festuco-Brometea, Calluno-Ulicetea); (8) dry grasslands and semi-deserts (Helianthemetea guttati); (9) montane tall-herb, grassland, fell-field and snow-bed vegetation (Mulgedio-Aconitetea, Salicetea herbaceae, Elyno-Seslerietea, Juncetea trifidi); (10) temperate broadleaved forests and scrub (Salicetea purpureae, Populetea albae, Franguletea, Rhamno-Prunetea, Querco-Fagetea, Quercetea pubescentis, Quercetea roboris); (11) montane heaths and coniferous forests (Loiseleurio-Vaccinietea, Erico-Pinetea, VaccinioPiceetea); and (12) weed communities (Stellarietea mediae, Polygono-Poetea annuae, Artemisietea vulgaris, Galio-Urticetea, Epilobietea angustifolii, Bidentetea tripartitii) (Zupančič, 1986; Kojić, Popović & Karadžić, 1998; Lakušić, 2005). Relying on a the ten-volume edition of Flora of SR Serbia (1970–1977), Diklić (1984) cited 3115 species from Serbia. However, according to the more recent estimation (Stevanović et al., 1995), the flora of Serbia includes 3662 taxa (i.e. 3272 species and 390 subspecies assigned to 141 families) (of which the richest are Asteraceae, Poaceae, Fabaceae, Brassicaceae, Apiaceae, Ranunculaceae, Cyperaceae, and Rosaceae) and 766 genera (of which the most species-rich are Hieracium L., Carex L., Trifolium L., Silene L., Centaurea L., Ranunculus L., Dianthus L., Veronica L., Campanula L., Euphorbia L., Allium L., Viola L., and Potentilla L.). The flora of Central Serbia and Kosovo regions is chiefly affected by Central European, Alpine, and Pontic impacts because many species and genera are widely distributed and inhabit large areas of the moderate and even boreal zone of the Holarctic. However, there are some genera and species that are Mediterranean in a wider sense. This mixture of floristic elements indicates that the flora of Central Serbia and Kosovo regions shows an intermediate character between the Mediterranean and Temperate Zones of Europe (or rather Eurasia) (Stevanović et al., 1999). 175 176 G. TOMOVIĆ ET AL. territory of the Balkan Peninsula. For identification of hierarchical floristic similarities among the 11 provinces, clustering based on Sørensen distances and unweighted pair-group average hierarchical sorting strategy (unweighted pair group method with arithmetic mean) was used. The cluster analysis was performed using PC-ORD, version 6.0 (Peck, 2010). Affiliation to appropriate floristic elements was determined for each endemic taxon on the basis of geography of its range. Nomenclature of floristic elements corresponds to the phytogeographical division of the Balkan Peninsula into phytochoria of subregion and province rank given by Horvat, Glavač & Ellenberg (1974), adapted and modified by Stevanović (1992, 1996, 2012). In addition, each endemic taxon (floristic element) was classified into a broader group of floristic elements, chorological subgroup, and RESULTS TAXONOMIC SPECTRUM The total number of species and subspecies meeting the criteria of Balkan endemics sensu Turrill (1929) and Strid & Tan (1997) is 492 (Table 2), equivalent to 13.4% of the total flora of Central Serbia and Kosovo regions. The Balkan endemics in the investigated territory belong to 48 families and 167 genera. There are no endemic families, although there are two monotypic Balkan endemic genera: Paramoltkia Greuter and Halacsya Dörfler. Ferns (Pteridophyta) do not have any endemics; in gymnosperms (Gymnospermae), there are only three © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 1. The geographical provinces of the Central Serbia and Kosovo regions (Stevanović 1992): Bč, Bačka; Bt, Banat; Sr, Srem; NW, North-western Serbia; Š, Šumadija; Po, Pomoravlje; NE, North-eastern Serbia; W, Western Serbia; C, Central Serbia; E, Eastern Serbia; SW, South-western Serbia; M, Metohija); K, Kosovo); S, Southern Serbia; SE, South-eastern Serbia. chorological group corresponding to the floristic regions of Europe of Walter & Straka (1970). The list of chorological groups, subgroups, and floristic elements is presented in Table 1. For the attribution of life forms, the criteria proposed by Raunkiaer (1934), Mueller-Dombois & Ellenberg (1974) and Stevanović (1992) were followed. Taxa were classified as phanerophytes (P), nanophanerophytes (NP), chamaephytes (Ch), hemicryptophytes (H), geophytes (G), and therophytes (T). All types of geological substrata were grouped in four classes: limestone, silicate, serpentinite, miscellaneous (sand, loess, peat etc.). Elevational data were classified into six ranges: 0–500 m, 500–1000 m, 1000–1500 m, 1500–2000 m, 2000–2500 m, and 2500– 2700 m. Data on vegetation characteristics for each individual record were classified into 27 vegetation classes the nomenclature of which was largely aligned with Rodwell et al. (2002), with minor exceptions that follow syntaxonomical concepts presented in Horvat et al. (1974) and Lakušić (2005). Records without sufficient data on vegetation formations could not be classified and were therefore not included in the vegetation analysis. Analyses related to mapping and spatial calculation were performed using applications in VISUAL BASIC, version 6.3 (Microsoft) language WINWORD 2003 (author of the application: M. Niketić). The analyses of diversity and the summary distribution of Balkan endemic species in Central Serbia and Kosovo regions were presented on the grid map with squares of 10 km × 10 km, based on Military Grid Reference System and the Universal Transverse Mercator (UTM) projection (http://www.luomus.fi/ english/botany/afe/map/utm.htm; Lampinen, 2001). The list of the Balkan endemic plants in Central Serbia and Kosovo regions (Table 2) is provided in alphabetical order by families, species, and subspecies. BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS 177 Table 1. The list of chorological groups, subgroups and floristic elements of the Balkan endemic flora in the Central Serbia and Kosovo regions Chorological subgroups Floristic elements Mediterranean-Submediterranean (MED-SUBMED) Mediterranean-Submediterranean-Subcontinental Adriatic-(Illyrian) Adriatic-(Illyrian)-Ionian Adriatic-(Illyrian)-Ionian-Aegean-Macedonic-Thracian-(Moesian) Adriatic-(Illyrian)-Aegean-Macedonic-Thracian Aegean-Macedonic-Thracian-(Moesian) Macedonic-Thracian-(Moesian) Adriatic Adriatic-Ionian Adriatic-Ionian-Aegean-Macedonic-Thracian Adriatic-Ionian-Macedonic-Thracian Adriatic-Macedonic-Thracian Aegean-Macedonic-Thracian Ionian-Macedonic-Thracian Macedonic-Thracian Adriatic-(Illyrian) Adriatic-(Illyrian)-Ionian Adriatic-(Illyrian)-Ionian-Aegean-Macedonic-Thracian Adriatic-(Illyrian)-Ionian-Macedonic-Thracian Adriatic-(Illyrian)-Ionian-Macedonic-Thracian-(Moesian) Adriatic-(Illyrian)-Macedonic-Thracian-(Moesian) Adriatic-(Illyrian)-(Moesian) Adriatic-Macedonic-Thracian-(Moesian) Macedonic-Thracian-(Moesian) Illyrian Moesian Illyrian-Scardo-Pindhian-Moesian Illyrian-Scardo-Pindhian Illyrian-Moesian Illyrian Scardo-Pindhian-Moesian Scardo-Pindhian Moesian Illyrian-Scardo-Pindhian-Moesian Illyrian-Scardo-Pindhian Illyrian-Moesian Illyrian Scardo-Pindhian-Moesian Scardo-Pindhian Moesian Dinaric-Scardo-Pindhian-Moesian mountain Dinaric-Scardo-Pindhian mountain Dinaric-Moesian mountain Dinaric mountain Scardo-Pindhian-Moesian mountain Scardo-Pindhian mountain Moesian mountain Dinaric-Scardo-Pindhian-Moesian mountain Dinaric-Scardo-Pindhian mountain Dinaric-Moesian mountain Dinaric mountain Scardo-Pindhian-Moesian mountain Scardo-Pindhian mountain Moesian mountain Submediterranean Submediterranean-Subcontinental Subcontinetal-Submediterranean Central European (CEUR) Illyrian-Balkan Illyrian Balkan Pontic (PONT) Illyrian-Balkan Illyrian Balkan Central European mountainous (CEM) Dinaric-Balkan mountainous Dinaric mountainous Balkan mountainous South European mountainous (SEM) Dinaric-Balkan mountainous Dinaric mountainous Balkan mountainous endemic species and endemic monocots (Monocotyledones) are only represented by 37 taxa (7.5%). The families richest in endemic taxa are Asteraceae, Caryophyllaceae, Fabaceae, Brassicaceae, and Lamiaceae (Fig. 2). In 13 families, there is only a single endemic taxon and, in 12 families, there are only two endemic taxa. At the generic level, the highest number of endemic taxa is found in species-rich genera (Hieracium L., Dianthus L. and Centaurea L.). In addition, Silene L., Achillea L., Stachys L., and Verbascum L. make substantial contributions (Fig. 3). CHOROLOGICAL SPECTRUM The chorological analysis of the Balkan endemics in Central Serbia and Kosovo regions (Fig. 4) shows a pronounced domination of orophytes belonging to the South European mountainous chorological group (195 taxa; 40%) and Central European mountainous © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Chorological groups Threatened and legally protected status Taxon 1 Amaryllidaceae Allium melanantherum Pančić 2 Amaryllidaceae Allium meteoricum Heldr. & Hausskn. ex Halácsy 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apocynaceae Apocynaceae Aristolochiaceae Asteraceae Asteraceae Asteraceae 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Athamanta turbith (L.) Brot. subsp. haynaldii (Borbás & Uechtr.) Tutin Bupleurum apiculatum Friv. Bupleurum flavicans Boiss. & Heldr. Bupleurum karglii Vis. Chaerophyllum coloratum L. Dichoropetalum oligophyllum (Griseb.) Pimenov & Kljuykov Eryngium palmatum Pančić & Vis. Eryngium serbicum Pančić Heracleum orphanidis Boiss. Heracleum sphondylium L. subsp. verticillatum (Pančić) Brummit Laserpitium siler L. subsp. zernyi (Hayek) Tutin Ligusticum albanicum Jáv. Pastinaca hirsuta Pančić Peucedanum serpentini Andras. & Jáv. Pimpinella serbica (Vis.) Benth. & Hook.f. ex Drude Vincetoxicum hirundinaria Medik. subsp. nivale (Boiss. & Heldr.) Markgr. Vincetoxicum huteri Vis. & Asch. Aristolochia merxmuelleri Greuter & E.Mayer Achillea abrotanoides (Vis.) Vis. Achillea ageratifolia (Sibth. & Sm.) Benth. & Hook.f. subsp. ageratifolia Achillea ageratifolia (Sibth. & Sm.) Benth. & Hook.f. subsp. serbica (Nyman) Heimerl Achillea alexandri-regis Bornm. & Rudsky Achillea baldaccii Degen Achillea chrysocoma Friv. Achillea holosericea Sm. Achillea pindicola Hausskn. subsp. corabensis (Heimerl) Greuter Achillea vandasii Velen. Amphoricarpos autariatus Blečić & E.Mayer subsp. autariatus Amphoricarpos autariatus Blečić & E.Mayer subsp. bertisceus Blečić & E.Mayer Anthemis cretica L. subsp. cinerea (Pančić) Oberpr. & Greuter Anthemis macedonica Boiss. & Orph. Carduus kerneri Simonk. subsp. scardicus (Griseb.) Kazmi Carduus ramosissimus Pančić Centaurea albertii Rexhepi Centaurea chrysolepis Vis. Centaurea derventana Vis. & Pančić Centaurea deustiformis Adamović Centaurea epapposa Velen. Centaurea finazzeri Adamović Centaurea graeca Griseb. Centaurea grisebachii (Nyman) Heldr. subsp. grisebachii Centaurea ipecensis Rech.f. Centaurea kosaninii Hayek Centaurea melanocephala Pančić Centaurea nervosa Willd. subsp. josifovicii M.R.Gajic Centaurea ognjanoffii Urum. Centaurea phrygia L. subsp. bosniaca (Murb.) Hayek Centaurea phrygia L. subsp. moesiaca (Urum. & J.Wagner) Hayek Centaurea stoebe L. subsp. serbica (Prodan) Ochsmann Cirsium appendiculatum Griseb. Cirsium ligulare Boiss. subsp. albanum Wettst. Cirsium ligulare Boiss. subsp. armatum (Velen.) Petr. Cirsium tymphaeum Hausskn. Crepis baldaccii Halácsy subsp. albanica Jáv. LC (IUCN) LC (ERLVP)* DD (IUCN) DD (ERLVP) ** * ** ** * ** * CR** * CR** ** * ** ** ** ** * ** ** ** * * ** ** * ** ** Life form Chorological group Distribution in geographical provinces G SEM E, SE, M G SEM M Ch T T T T H H H H H H H H H H H H G Ch Ch Ch SEM MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED SEM MED-SUBMED MED-SUBMED CEM CEM SEM CEM CEM PONT SEM SEM SEM MED-SUBMED SEM SEM SEM NW, W, SW, C, K, M E, SE, S, C, K, M E NW, W, K, M M SE, M Š, NE, E, SE, S, W, SW, C, K, M Š, E, S, C, K K E K, M M NE, E, SE, S, C M W, SW, C, K, M SE, K, M C, K, M M SW, M K, M NE, E, SE, S, NW, W, SW, K, M H H H H Ch Ch H H Ch Ch H H H H H H H H H H H H H H H H H H H H H H H SEM SEM SEM SEM SEM SEM SEM SEM SEM SEM CEM CEM MED-SUBMED SEM SEM SEM MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED PONT CEM MED-SUBMED CEUR MED-SUBMED PONT CEM SEM SEM SEM SEM K M SE, K, M K, M K, M E, C SW M SW, K, M SE, S E, SE, C, K, M M K, M NE, E, SE NW, W M E, W SE, K M E, K, M M K, M C, M K, M SE SW E, SE E, SE, S, C, K E, SE, S, K, M E, SE, S, C, K, M E M M Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Family G. TOMOVIĆ ET AL. No 178 Table 2. List of the Balkan endemic plants in the Central Serbia and Kosovo regions arranged in alphabetical order by families and taxa Asteraceae Asteraceae Asteraceae 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae CR(Srb)** CR(Srb)** ** H H H H H H Ch H H H H H H H H H H H H H H H H H H H H SEM SEM SEM SEM SEM SEM CEM PONT SEM SEM SEM SEM SEM SEM SEM CEM SEM CEUR CEUR SEM SEM SEM SEM SEM CEUR SEM SEM M M NE, E K, M SE, S E, SE, S, C, K E, M K, M M M W, SW M M M W E M M NE M W W, SW, K, M W SE W, M K, M K ** H H H SEM SEM CEM K E K, M H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H CEM SEM SEM CEM CEM SEM SEM SEM SEM SEM SEM SEM SEM CEUR CEM CEUR CEUR SEM SEM SEM CEUR CEUR CEUR CEM CEM CEM CEUR SEM SEM SEM SEM SEM CEUR E M SE, S E M M M M C, M M K, M E W SW E, SE, M M E, S NE, E, SE C, M E, NW, W, SW, C, K, M M E, S, C, K M K K K, M S M K NE, E M M M * * * * ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 84 85 86 Crepis bertiscea Jáv. Crepis macedonica Kitan. Cyanus nyssanus (Petrović) Soják Cyanus orbelicus (Velen.) Soják Cyanus tuberosus (Vis.) Soják Cyanus velenovskyi (Adamović) Wagenitz & Greuter Erigeron alpinus L. subsp. rhodopaeus (Vierh.) Kožuharov & N.Andreev Galatella albanica Degen Gnaphalium pichleri Murb. Hieracium albopellitum (Zahn) Niketić Hieracium amphithales K.Malý & Zahn Hieracium andrasovszkyi Zahn subsp. cremnophilum O.Behr, E.Behr & Zahn Hieracium andrasovszkyi Zahn subsp. doerfleri Hayek & Zahn Hieracium andrasovszkyi Zahn subsp. kobilicanum O.Behr, E.Behr & Zahn Hieracium baldaccianum Freyn Hieracium balkanum Uechtr. ex Pančić Hieracium bertisceum Niketić Hieracium bifidum Kit. ex Hornem. subsp. pallescentisimile O.Behr, E.Behr & Zahn Hieracium bifidum Kit. ex Hornem. subsp. stolanum Zahn Hieracium bjeluschae K.Malý & Zahn subsp. tommasiniiforme Schaeffer & Zahn Hieracium bosniacum Freyn subsp. bosniacum Hieracium brandisii Freyn Hieracium brevilanosum Degen & Zahn Hieracium bulgaricum Freyn Hieracium bupleuroides C.C.Gmel. subsp. malacosericeum Rech.f. & Zahn Hieracium coloriscapum Rohlena & Zahn subsp. coloriscapum Hieracium coloriscapum Rohlena & Zahn subsp. stenopyllophorum O.Behr, E.Behr & Zahn Hieracium dimoniei Zahn Hieracium divaricatum Fr. Hieracium djimilense Boiss. & Balansa subsp. brachytrichoiphyes O.Behr, E.Behr & Zahn Hieracium djimilense Boiss. & Balansa subsp. cordatifrons Zahn Hieracium durmitoricum (Rohlena & Zahn) Niketić Hieracium eriopus Boiss. & Heldr. Hieracium erythrocarpum Peter subsp. aculeatissimum Zahn Hieracium erythrocarpum Peter subsp. kurvalae Rech.f. & Zahn Hieracium gaudryi Boiss. & Orph. subsp. cernyanum Hayek & Zahn Hieracium gaudryi Boiss. & Orph. subsp. hayekianum Dörfl. & Zahn Hieracium grossianum Zahn subsp. schefferianum Zahn Hieracium guentheri-beckii Zahn subsp. guentheri-beckii Hieracium guglerianum Zahn subsp. telekianum Kümmerle & Zahn Hieracium gymnocephalum Griseb. ex Pant. subsp. gymnocephalum Hieracium heldreichii Boiss. subsp. pseudopilosissimum Zahn Hieracium heterogynum (Froel.) Gutermann subsp. heterogynum Hieracium jurassicum Griseb. subsp. papyraceum (Zahn) Greuter Hieracium macedonicum Boiss. & Orph. ex Boiss. Hieracium macrodontoides (Zahn) Zahn subsp. gigantophyllum Zahn Hieracium markovanum Arv.-Touv. Hieracium marmoreum Pančić & Vis. Hieracium mirificissimum Rohlena & Zahn Hieracium mokragorae Nägeli & Peter Hieracium murorum L. subsp. bistricense Zahn Hieracium murorum L. subsp. psilodontum Zahn Hieracium murorum L. subsp. valdecordatum Zahn Hieracium naegelianiforme (O.Behr, E.Behr & Zahn) Buttler Hieracium naegelianum Pančić subsp. ljubotenicum O.Behr, E.Behr & Zahn Hieracium naegelianum Pančić subsp. naegelianum Hieracium pallidum Biv. subsp. vranjanum Zahn Hieracium pannosum Boiss. subsp. doerflerianum Hayek & Zahn Hieracium pannosum Boiss. subsp. eumecobracchion Hayek Hieracium pannosum Boiss. subsp. trojanum Zahn Hieracium pichleri Kern. subsp. adamovicii Sagorski & Zahn Hieracium pichleri Kern. subsp. chromoneurum Zahn Hieracium praecox Sch.Bip. subsp. basilacistum Rech.f. & Zahn 179 Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 180 Table 2. Continued Taxon 120 121 122 123 124 125. 126 127. 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae 151 Asteraceae Hieracium praecurrens Vuk. subsp. megaladenophyes K.Malý & Zahn Hieracium pseuderiopus Zahn subsp. nikolae Zahn Hieracium pseuderiopus Zahn subsp. pseuderiopus Hieracium pseudobifidum Schur subsp. caesiopictum (Zahn) Zahn Hieracium pseudobifidum Schur subsp. stenolepioides (Zahn) Zahn Hieracium pseudobifidum Schur subsp. zljebense (Zahn) Zahn Hieracium pseudosparsum Uchtr. ex Zahn Hieracium racemosum Waldst. & Kit. ex Willd. subsp. chaetotrichum Zahn Hieracium racemosum Waldst. & Kit. ex Willd. subsp. semigrisescens Zahn Hieracium scardicum Bornm. & Zahn subsp. scardicum Hieracium schefferi Rech.f. & Zahn Hieracium scheppigianum Freyn subsp. scheppigianum Hieracium schultzianum Pančić & Vis. Hieracium sericophyllum Nejceff & Zahn subsp. acropolioscapum Zahn Hieracium sparsum Friv. subsp. ipekanum Rech.f. & Zahn Hieracium sparsum Friv. subsp. livadicanum O.Behr, E.Behr & Zahn Hieracium sparsum Friv. subsp. peninsulare (Zahn) Zahn Hieracium sparsum Friv. subsp. pilosifrons Zahn Hieracium sparsum Friv. subsp. staraeplaninae Zahn Hieracium sparsum Friv. subsp. subsparsiflorum (Degen & Zahn) Zahn Hieracium suborieni (Zahn) P.D.Sell & C.West Hieracium thapsiforme Uechtr. ex Asch. & Kan. Hieracium tommasinianum K.Malý subsp. setosissimum (Nägeli & Peter) Gottschl. Hieracium tommasinianum K.Malý subsp. tommasinianum Hieracium transiens (Freyn) Freyn Hieracium velenovskyi Freyn Hieracium waldsteinii Tausch subsp. nipholeucum Zahn Hieracium wiesbaurianum R.Uechtr. subsp. livadicae O.Behr, E.Behr & Zahn Hypochaeris maculata L. subsp. pelivanovicii (Velen.) Hayek Jacobaea pancicii (Degen) Vladimir. & Raab-Straube Klasea radiata (Waldst. & Kit.) Á.Löve & D.Löve subsp. cetinjensis (Rohlena) Greuter & Wagenitz Lactuca pancicii (Vis.) N.Kilian & Greuter 152 153 154 155 156 157 158 159 160 Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Pilosella serbica (F.W.Schultz & Sch.Bip.) Szela˛g Reichardia macrophylla Vis. & Pančić Scorzonera doriae Degen & Bald. Senecio hercynicus Herborg subsp. dalmaticus (Griseb.) Greuter Tanacetum larvatum (Pant.) Hayek Taraxacum pindicola (Bald.) Hand.-Mazz. Tephroseris crassifolia (Schult.) Griseb. & Schenk Tephroseris papposa (Rchb.) Schur subsp. wagneri (Degen) B.Nord. Willemetia stipitata (Jacq.) Dalla Torre subsp. albanica (Kümmerle & Jáv.) Kirschnerová Alkanna noneiformis Griseb. Alkanna pulmonaria Griseb. Alkanna scardica Griseb. Alkanna stribrnyi Velen. Halacsya sendtneri (Boiss.) Dörfl. Myosotis alpestris F.W.Schmidt subsp. suaveolens (Waldst. & Kit. ex Willd.) Strid Onosma stellulata Waldst. & Kit. Paramoltkia doerfleri (Wettst.) Greuter & Burdet Solenanthus krasniqii (Wraber) Niketić Solenanthus scardicus Bornm. Alyssum corymbosoides Formánek Alyssum montanum L. subsp. serbicum Novák Alyssum scardicum Wettst. Arabis bryoides Boiss. Aubrieta gracilis Spruner ex Boiss. subsp. scardica (Wettst.) Phitos ** ** ** ** ** ** ** CR(Srb)** ** ** ** ** ** ** ** ** CR(Srb)** LC (IUCN) LC (ERLVP)* * * ** * ** CR(Srb)** ** * ** CR** ** ** ** CR(Srb)** ** Life form Chorological group Distribution in geographical provinces H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H CEUR CEM CEM CEUR CEUR CEUR CEM CEUR CEUR SEM CEUR SEM SEM SEM CEM CEM CEM CEM CEM CEM SEM SEM CEUR CEUR CEM CEM SEM CEUR CEM CEM MED-SUBMED M E S M S, K, M M E C E K, M M M E, C, K M M K E E E C M NW, W, C, M W W, M E E, SE M K NE, E, SE, NW, W, SW, C, K, M E M H CEM E, W, SW, C, K, M H H H H Ch H H H H CEM SEM SEM CEM CEM CEM CEM CEM CEM C W, K, M M M M K, K, H H H H Ch H Ch Ch H H H Ch Ch Ch Ch MED-SUBMED SEM SEM MED-SUBMED MED-SUBMED CEM SEM MED-SUBMED SEM SEM MED-SUBMED MED-SUBMED CEM SEM SEM K S M E Š, W, SW, C, K, M NE, E, SE, S, NW, W, SW, C, K, M NW, W, SW, K, M K, M M M SE Š, NW, SW, C, K K, M K, M K, M Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Family M M M M G. TOMOVIĆ ET AL. Threatened and legally protected status No Aurinia corymbosa Griseb. Barbarea balcana Pančić 178 Brassicaceae Barbarea longirostris Velen. 179 180 Brassicaceae Brassicaceae Bornmuellera dieckii Degen Cardamine amara L. subsp. balcanica Marhold, Ančev & Kit Tan 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Campanulaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caprifoliaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Cardamine carnosa Waldst. & Kit. Cardamine pancicii Hayek Cardamine serbica Pančić Draba korabensis Kümmerle & Degen ex Jáv. Draba kuemmerlei Stevan. & D.Lakušić Erysimum korabense Kümmerle & Jáv. Erysimum kuemmerlei Jáv. Erysimum linariifolium Tausch Erysimum pusillum Bory & Chaub. subsp. microstylum (Hausskn.) Hayek Lunaria telekiana Jáv. Malcolmia orsiniana (Ten.) Ten. subsp. serbica (Pančić) Greuter & Burdet Thlaspi bellidifolium Griseb. Thlaspi dacicum Heuff. subsp. montenegrinum (F. K. Meyer) Greuter & Burdet Thlaspi microphyllum Boiss. & Orph. Asyneuma pichleri (Vis.) D.Lakušić & F.Conti Campanula hercegovina Degen & Fiala Campanula moesiaca Velen. Campanula orbelica Pančić Campanula patula L. subsp. epigaea (Janka ex Degen) Hayek Campanula persicifolia L. subsp. sessiliflora (Velen.) Fed. ex Greuter & Burdet Campanula scutellata Griseb. Campanula secundiflora Vis. & Pančić Campanula spatulata Sm. subsp. spatulata Edraianthus canescens D.Lakušić, Niketić & Stevan. Edraianthus montenegrinus Horák Edraianthus serbicus (A.Kern.) Petrović Phyteuma pseudorbiculare Pant. Centranthus longiflorus Steven subsp. junceus (Boiss. & Heldr.) I.Richardson Cephalaria flava (Sm.) Szabó subsp. flava Cephalaria pastricensis Dörfl. & Hayek Knautia ambigua Boiss. & Orph. Knautia dinarica (Murb.) Borbás subsp. dinarica Knautia midzorensis Formánek Knautia pancicii Szabó Knautia sarajevensis (Beck) Szabó Lonicera formanekiana Halácsy subsp. formanekiana Scabiosa achaeta Vis. & Pančić Scabiosa fumarioides Vis. & Pančić Scabiosa ochroleuca L. subsp. balcanica (Velen.) Stoj. & Stef. Valeriana bertiscea Pančić Valeriana pancicii Halácsy & Bald. Cerastium banaticum (Rochel) Heuff. subsp. kosaninii Georgiev Cerastium decalvans Schloss. & Vuk. subsp. decalvans Cerastium decalvans Schloss. & Vuk. subsp. leontopodium (Stoj. & Stef.) Niketić Cerastium malyi (T.Georgiev) Niketić subsp. malyi Cerastium malyi (T.Georgiev) Niketić subsp. serpentini (Novák) Niketić Cerastium neoscardicum Niketić Cerastium rectum Friv. subsp. rectum Dianthus behriorum Bornm. Dianthus cruentus Griseb. subsp. cruentus Dianthus gracilis Sm. subsp. armerioides (Griseb.) Tutin Dianthus gracilis Sm. subsp. gracilis Dianthus integer Vis. subsp. integer Dianthus integer Vis. subsp. minutiflorus (Halácsy) Bornm. ex Strid Dianthus leucophoeniceus Dörfl. & Hayek Dianthus microlepis Boiss. subsp. microlepis Dianthus moesiacus Vis. & Pančić LC (IUCN) LC (ERLVP) DD (IUCN) DD (ERLVP) ** LC (ERLVP) ** ** * * ** ** * * * * ** ** ** * CR(Srb)** ** ** * * ** ** EX** * * * * CR** CR** * * ** Ch H SEM CEM NW, W, SW, C, K, M E, K, M H SEM K, M Ch H SEM CEM K E, SE, C, K H H H H H H H Ch Ch H T Ch H Ch H Ch H H H H T Ch H Ch Ch Ch H Ch Ch H H H H H H P H H H H H Ch Ch Ch Ch Ch Ch T H H Ch Ch Ch Ch H Ch H SEM SEM SEM CEM CEM CEM SEM SEM SEM CEUR SEM CEM CEM SEM CEUR SEM CEM CEM CEUR CEUR MED-SUBMED MED-SUBMED SEM SEM SEM SEM CEM SEM MED-SUBMED CEM SEM CEM CEM CEM CEM CEM MED-SUBMED MED-SUBMED PONT CEM CEM SEM SEM SEM SEM SEM CEM SEM SEM SEM MED-SUBMED MED-SUBMED SEM SEM MED-SUBMED CEM PONT K, M C W K, M K, M M S, SW, C, K, M Š, NW, W, SW, C, K, M K, M M NE, E, K, M K, M M M W, SW, C, K, M M E, SE, W, K E, K, M E, SE, K, M E, M SE, S W, SW K, M W SW, M NE, E, SE K, M K SE, S NE, E, W, M E NW, W, SW, C, K, M E, SE W W, SW K, M C Š, E, SE, S, W, SW, C, K, M E, K M M M NE, NW, W, SW, K, M S, NW, W, SW, C, K, M NE, E, NW, W, SW, C, K, M W, SW, C, K K, M NE, E, SE, S, NW, W, SW, C, K, M M Š, E, SE, S, W, SW, C, K, M SE, S, K, M K, M M K, M M K, M NE, E, SE 181 Brassicaceae Brassicaceae BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 176 177 182 Table 2. Continued Taxon 238 Caryophyllaceae Dianthus nitidus Waldst. & Kit. subsp. lakusicii Wraber 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae 254 Caryophyllaceae 255 256 257 258 259 260 261 262 263 264 Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Cistaceae Colchicaceae Convolvulaceae Crassulaceae Crassulaceae Crassulaceae Crassulaceae Crassulaceae Crassulaceae Dioscoreaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Fabaceae Fabaceae Fabaceae Dianthus noëanus Boiss. Dianthus pelviformis Heuff. Dianthus pinifolius Sm. subsp. pinifolius Dianthus scardicus Wettst. Dianthus stenopetalus Griseb. Dianthus sylvestris Wulfen subsp. bertisceus Rech.f. Dianthus sylvestris Wulfen subsp. nodosus (Tausch) Hayek Dianthus tenuiflorus Griseb. Dianthus tristis Velen. Dianthus viridescens Clem. Gypsophila spergulifolia Griseb. Heliosperma macranthum Pančić Heliosperma nikolicii (A.Seliger & Wraber) Niketić & Stevan. Heliosperma oliverae Niketić & Stevan. Heliosperma pusillum (Waldst. & Kit.) Hoffmanns. subsp. moehringiifolium (Uechtr. ex Pančić) Niketić & Stevan. Heliosperma pusillum (Waldst. & Kit.) Hoffmanns. subsp. monachorum (Vis. & Pančić) Niketić & Stevan. Minuartia baldaccii (Halácsy) Mattf. subsp. baldaccii Minuartia bosniaca (Beck) K.Malý Minuartia bulgarica (Velen.) Graebn. Minuartia doerfleri Hayek Minuartia graminifolia (Ard.) Jáv. subsp. clandestina (Port.) Mattf. Paronychia macedonica Chaudhri subsp. macedonica Petrorhagia obcordata (Margot & Reut.) Greuter & Burdet Saponaria intermedia Simmler Silene fabarioides Hausskn. Silene flos-cuculi (L.) Greuter & Burdet subsp. subintegra (Hayek) Greuter & Burdet Silene frivaldszkyana Hampe Silene parnassica Boiss. & Spruner subsp. parnassica Silene parnassica Boiss. & Spruner subsp. serbica (Vierh. & Adamović) Greuter Silene roemeri Friv. subsp. roemeri Silene schmuckeri Wettst. Silene sendtneri Boiss. subsp. sendtneri Silene skorpilii Velen. Silene waldsteinii Griseb. Viscaria asterias (Griseb.) Frajman Fumana bonapartei Maire & Petitm. Colchicum macedonicum Košanin Convolvulus boissieri Steud. subsp. parnassicus (Boiss. & Orph.) Kuzmanov Sedum alpestre Vill. subsp. erythraeum (Griseb.) ‘t Hart Sedum stefco Stef. Sedum tuberiferum Stoj. & Stef. Sempervivum kindingeri Adamović Sempervivum kosaninii Praeger Sempervivum macedonicum Praeger Dioscorea balcanica Košanin Euphorbia glabriflora Vis. Euphorbia montenegrina (Bald.) K.Malý Euphorbia pancicii Beck Euphorbia serpentini Novák Euphorbia subhastata Vis. & Pančić Anthyllis aurea Welden Astragalus fialae Degen Astragalus glycyphylloides DC. subsp. serbicus (Rchb.) Vasić & Niketić NT (IUCN) NT (ERLVP) ** * * ** * ** * * CR(Srb)** ** ** ** ** ** ** * ** EX(Srb)** ** * CR(Srb)** ** ** ** ** * ** ** ** * * * ** ** ** Life form Chorological group Distribution in geographical provinces Ch CEM M Ch H Ch Ch H Ch Ch H H T Ch Ch Ch Ch Ch SEM PONT MED-SUBMED CEM SEM CEM CEM MED-SUBMED SEM MED-SUBMED MED-SUBMED SEM MED-SUBMED SEM SEM NE, E NE, E, SE, S, M Š, E, SE, S, NW, C, K, M K, M S, M K, M M SE, S, C, K E, K, M M NW, W, SW, M M K, M M NE, E Ch SEM NW, W, SW, C, K, M Ch H Ch Ch Ch Ch T Ch T H MED-SUBMED SEM CEM SEM SEM SEM MED-SUBMED MED-SUBMED SEM CEUR M Š, NE, E, W, SW, K, M E, SE, C M K, M K, M M K, M M SE H Ch Ch H Ch H Ch Ch H Ch G Ch Ch Ch Ch Ch Ch Ch G Ch H H H H Ch H H PONT SEM SEM SEM SEM SEM PONT SEM SEM MED-SUBMED SEM SEM CEM MED-SUBMED MED-SUBMED SEM SEM SEM MED-SUBMED MED-SUBMED SEM CEUR PONT CEUR SEM CEM CEUR SE, S, K K, M W, SW, C, K, M E, SE, S M NE, E, SE, S, NW, W, SW, C, K, M E E, SE, K, M E, SE, C, K, M W, SW, C, K, M M K K, M E, SE, S, C SE K, M K, M K, M M S, NW, W, SW, C, K, M K, M W, SW W W, SW, M SE, K, M M Š, NE, E, SE, K, M Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Family G. TOMOVIĆ ET AL. Threatened and legally protected status No Astragalus wilmottianus Stoj. Cytisus jankae Velen. Cytisus petrovicii Adamović Cytisus tommasinii Vis. Genista hassertiana (Bald.) Buchegger subsp. hassertiana Genista nissana Petrović Genista subcapitata Pančić Genista sylvestris Scop. subsp. dalmatica (Bartl.) H.Lindb. Lathyrus binatus Pančić Lathyrus pancicii (Jurišić) Adamović Lotus stenodon (Boiss. & Heldr.) Heldr. Medicago prostrata Jacq. subsp. pseudorupestris (Hayek) Micevski 304 305 306 307 308 Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae 309 310 Fabaceae Fabaceae Onobrychis montana DC. subsp. scardica (Griseb.) P.W.Ball Onobrychis pindicola Hausskn. Oxytropis dinarica (Murb.) Wettst. subsp. dinarica Oxytropis dinarica (Murb.) Wettst. subsp. weberi Chrtek & Chrtková Oxytropis halleri Bunge ex Koch subsp. korabensis (Kümmerle & Jáv.) Chrtek & Chrtková Trifolium dalmaticum Vis. Trifolium medium L. subsp. balcanicum Velen. 311 312 313 314 315 316 317 318 Fabaceae Fabaceae Fabaceae Fabaceae Gentianaceae Gentianaceae Gesneriaceae Gesneriaceae Trifolium pignantii Fauché & Chaub. Trifolium trichopterum Pančić Trifolium velenovskyi Vandas Trifolium wettsteinii Dörfl. & Hayek Gentiana pneumonanthe L. subsp. nopcsae (Jáv.) Wraber Gentianella albanica (Jáv.) J. Holub Ramonda nathaliae Pančić & Petrović Ramonda serbica Pančić 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346 347 348 349 350 Iridaceae Iridaceae Iridaceae Iridaceae Iridaceae Iridaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lentibulariaceae Liliaceae Crocus alexandri Ničić ex Velen. Crocus dalmaticus Vis. Crocus kosaninii Pulević Crocus rujanensis Rand̄elović & D.A.Hill Crocus scardicus Košanin Crocus veluchensis Herb. Ballota hispanica (L.) Benth. subsp. macedonica (Vandas) Strid & Kit Tan Micromeria albanica (K.Malý) Šilić Micromeria croatica (Pers.) Schott Micromeria longipedunculata Bräuchler Nepeta rtanjensis Diklić & Milojević Salvia officinalis L. subsp. multiflora Gajić Salvia pratensis L. subsp. pozegensis (Watzl) Diklić Sideritis scardica Griseb. Stachys anisochila Vis. & Pančić Stachys beckeana Dörfl. & Hayek Stachys milanii Petrović ex Magnier Stachys officinalis (L.) Trevisan subsp. skipetarum Jáv. Stachys plumosa Griseb. Stachys recta L. subsp. baldaccii (K.Malý) Hayek Stachys recta L. subsp. rhodopaea (Velen.) Chrtek Stachys scardica (Griseb.) Hayek Stachys serbica Pančić Teucrium arduini L. Thymus adamovicii Velen. Thymus bulgaricus (Domin & Podp.) Ronniger Thymus ciliatopubescens (Halácsy) Halácsy Thymus doerfleri Ronniger Thymus longedentatus (Degen & Urum.) Ronniger Thymus stojanovii Degen Pinguicula balcanica Casper Fritillaria gussichiae (Degen & Dörfl.) Rix 351 Liliaceae Fritillaria macedonica Bornm. ** * ** EX(Srb)** * * EX(Srb)** LC (IUCN) LC (ERLVP) ** ** * ** CR** * ** LC (IUCN) LC (ERLVP)** * ** * CR** * ** ** * CR** * CR(Srb)** * CR(Srb)** * EX(Srb)** ** * * * * DD (IUCN) DD (ERLVP) CR(Srb)** Ch Ch Ch Ch Ch Ch Ch Ch H H H Ch PONT PONT PONT SEM MED-SUBMED MED-SUBMED SEM MED-SUBMED CEUR MED-SUBMED CEM MED-SUBMED E, SE E, SW, C, K, M NE, E, S, C, K SE, S, W, SW, C, K, M M E, K NE, E, SE, S, C, K W, M NW, W, SW E SW, K, M M H Ch H H H CEM SEM CEM CEM CEM K, M M M K, M K, M T H MED-SUBMED CEUR H T H Ch H H H H MED-SUBMED SEM CEUR CEM CEUR CEM MED-SUBMED MED-SUBMED NE, E, SE, S, W, SW, C, K, M Š, NE, E, SE, S, NW, W, SW, C, K, M S, SW, C, K, M Š, E, SE, S, W, C, K, M E, SE, S, C, K, M K, M M K, M E, K NE, E, SW, K, M G G G G G G H Ch Ch Ch Ch Ch H Ch H H T H H H H H T Ch Ch Ch Ch Ch Ch Ch H G MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED SEM SEM MED-SUBMED MED-SUBMED SEM MED-SUBMED MED-SUBMED MED-SUBMED PONT SEM SEM SEM MED-SUBMED MED-SUBMED MED-SUBMED SEM PONT SEM MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED SEM MED-SUBMED SEM CEM MED-SUBMED E, SE, S, C, K SE, M S, K S K, M E, SE, S, NW, W, SW, C, K, M M K, M W, M M NE E, C, M W, K K, M NW, W W, M E, S, C M E, SE, S, K E, NW, W, SW, C, K, M SE, S NE, SE, S, W, SW, C, K, M E, M M Š, W, C NE, E K, M M NE K E, SE, K, M SE, S G SEM K, M 183 Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 292 293 294 295 296 297 298 299 300 301 302 303 184 Table 2. Continued Taxon 352 353 354 355 356 357 358 359 360 Liliaceae Liliaceae Liliaceae Liliaceae Linaceae Linaceae Linaceae Linaceae Lythraceae Fritillaria messanensis Rafin. subsp. gracilis (Ebel) Rix Lilium albanicum Griseb. Tulipa scardica Bornm. Tulipa serbica Tatić & Krivošej Linum capitatum Kit. ex Schult. subsp. capitatum Linum elegans Spruner ex Boiss. Linum hirsutum L. subsp. spathulatum (Halácsy & Bald.) Halácsy Linum tauricum Willd. subsp. serbicum (Podp.) Petrova Trapa annosa Janković 361 362 363 364 365 Malvaceae Malvaceae Nartheciaceae Oleaceae Orchidaceae Althaea kragujevacensis Pančić ex Diklić & Stevan. Althaea vranjensis Diklić & V.Nikolić Narthecium scardicum Košanin Forsythia europaea Degen & Bald. Dactylorhiza cordigera (Fr.) Soó subsp. bosniaca (Beck) Soó 366 367 368 369 370 371 372 373 374 Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Papaveraceae Papaveraceae Pinaceae Pinaceae Pinaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae 393 394 395 396 397 Plantaginaceae Plantaginaceae Plantaginaceae Plumbaginaceae Poaceae Melampyrum doerfleri Ronniger Melampyrum heracleoticum Boiss. & Orph. ex Boiss. Melampyrum hoermannianum K.Malý Melampyrum scardicum Wettst. Melampyrum trichocalycinum Vandas Orobanche esulae Pančić Pedicularis brachyodonta Schloss. & Vuk. subsp. grisebachii (Wettst.) Hayek Pedicularis brachyodonta Schloss. & Vuk. subsp. moesiaca (Stadlm.) Hayek Pedicularis brachyodonta Schloss. & Vuk. subsp. montenegrina (Janka ex Nyman) D.A.Webb Pedicularis ernesti-mayeri Stevan., Niketić & D.Lakušić Pedicularis heterodonta Pančić Pedicularis hoermanniana K.Malý Pedicularis leucodon Griseb. subsp. leucodon Pedicularis orthantha Griseb. Rhinanthus asperulus (Murb.) Soó Rhinanthus melampyroides (Borbás & Degen) Soó Corydalis solida (L.) Clairv. subsp. incisa Lidén Pseudofumaria alba (Mill.) Lidén subsp. leiosperma (Conrath) Lidén Abies borisii-regis Mattf. Picea omorika (Pančić) Purk. Pinus peuce Griseb. Digitalis viridiflora Lindl. Linaria peloponnesiaca Boiss. & Heldr. Linaria rubioides Vis. & Pančić subsp. nissana (Petrović) Niketić & Tomović Linaria rubioides Vis. & Pančić subsp. rubioides Plantago reniformis Beck Veronica barrelieri H.Schott ex Roem. & Schult. subsp. andrasovszkyi (Jáv.) M.Fisher Veronica barrelieri H.Schott ex Roem. & Schult. subsp. prodanii (Degen) M.Fisher Veronica thessalica Benth. Wulfenia blecicii Lakušić subsp. blecicii Armeria rumelica Boiss. Bromopsis moesiaca (Velen.) Holub 398 399 400 401 402 403 404 405 Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Festuca adamovicii (St.-Yves) Markgr.-Dann. subsp. adamovicii Festuca graeca (Hack.) Markgr.-Dann. subsp. graeca Festuca halleri All. subsp. scardica (Griseb.) Markgr.-Dann. Festuca koritnicensis Hayek & Vetter Festuca stojanovii (Acht.) Kožuharov ex Foggi & Petrova Festuca valida (R.Uechtr.) Pénzes subsp. valida Helictotrichon blavii (Asch. & Janka) C.E.Hubb. Sesleria albicans Kit. subsp. angustifolia (Hack. & Beck) Deyl * CR(Srb)** CR** CR(Srb)** * EX (ERLVP) EX** EX** EX** * ** LC (IUCN) LC (ERLVP)** * * * * ** ** ** ** * * * ** LC (IUCN)** EN (IUCN)** NT (IUCN)** ** * * ** ** ** ** DD (IUCN) DD (ERLVP) Life form Chorological group Distribution in geographical provinces G G G G H Ch H H T SEM CEM MED-SUBMED MED-SUBMED SEM SEM PONT PONT CEUR M W, SW, K, M K, M K NE, E, SE, W, SW, C, K, M K, M K, M Š, SW, C, K, M Š H H G P G CEUR MED-SUBMED SEM MED-SUBMED CEM Š SE, S K, M K, M E, SE, S, SW, C, K, M T T T T T G H H H MED-SUBMED MED-SUBMED CEUR CEM CEUR PONT CEM CEM CEM K, M SW, K, M NW, W, SW, C, M E, SE, K, M K, M E, SE, S K, M E, SE, S M H H H H H T T G G P P P H H H H H H CEM CEM CEM CEM CEM CEUR CEUR CEUR SEM SEM CEM CEM CEUR SEM MED-SUBMED MED-SUBMED CEM PONT M E, SE, S, NW, W, SW, C, K, M E, SE, C, K, M K, M E M K, M M NW, W, SW, M M W, SW K, M E, SE K, M E Š, NW, W, SW, C W, M M H Ch H H H PONT SEM CEM SEM PONT NE, E K, M M SE, S, C, K, M SE, C H H H H H H H H CEM SEM CEM CEM SEM SEM CEM SEM C, K, M M C, K, M K, M SE E, SE NW, W NW, W, SW, M Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Family G. TOMOVIĆ ET AL. Threatened and legally protected status No Sesleria comosa Velen. Sesleria korabensis (Kümmerle & Jáv.) Deyl Sesleria latifolia (Adamović) Degen Sesleria serbica (Adamović) Ujhelyi Sesleria tenerrima (Fritsch) Hayek Sesleria ujhelyii Strgar Sesleria wettsteinii Döerfl. & Hayek Stipa mayeri Martinovský Stipa novakii Martinovský Trisetum flavescens (L.) P.Beauv. subsp. serbicum (Velen.) Hayek Polygala croatica Chodat Polygala doerfleri Hayek Polygonum albanicum Jáv. Rumex balcanicus Rech.f. 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440 441 442 443 444 445 446 447 448 Primulaceae Primulaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Rhamnaceae Rhamnaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Androsace hedraeantha Griseb. Soldanella pindicola Hausskn. Aconitum burnatii Gáyer subsp. pentheri (Hayek) Jalas Aconitum toxicum Rchb. subsp. bosniacum (Beck) Niketić Aquilegia blecicii A.Podob. Aquilegia nikolicii (Niketić) Niketić & Cikovac Aquilegia pancicii Degen Consolida uechtritziana (Pančić ex Huth) Soó Delphinium balcanicum Pawł. Helleborus multifidus Vis. subsp. multifidus Helleborus multifidus Vis. subsp. serbicus (Adamović) Merxm. & Podlech Pulsatilla halleri (All.) Willd. subsp. rhodopaea (Stoj. & Stef.) K.Krause Pulsatilla montana (Hoppe) Rchb. subsp. bulgarica Rummelsp. Ranunculus concinnatus Schott Ranunculus incomparabilis Janka Ranunculus seguieri Vill. subsp. montenegrinus (Halácsy) Tutin Rhamnus orbiculata Bornm. Rhamnus rhodopea Velen. subsp. rhodopaea Alchemilla amphiargyrea Buser Alchemilla bulgarica Rothm. Alchemilla catachnoa Rothm. Alchemilla heterotricha Rothm. Alchemilla indivisa (Formánek ex Buser) Rothm. Alchemilla lanuginosa Rothm. Alchemilla pirinica Pawł. Alchemilla viridiflora Rothm. Crataegus heldreichii Boiss. Drymocallis malacophylla (Borbás) Kurtto Geum bulgaricum Pančić 449 450 451 452 453 454 455 456 457 458 459 460 461 462 463 464 465 466 467 468 Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rutaceae Rutaceae Santalaceae Sapindaceae Sapindaceae Saxifragaceae Geum rhodopeum Stoj. & Stef. Potentilla doerfleri Wettst. Potentilla montenegrina Pant. Potentilla nicicii Adamović Potentilla speciosa Willd. subsp. illyrica Soják Potentilla visianii Pančić Sanguisorba albanica András. & Jáv. Asperula aristata L.f. subsp. condensata (Heldr. ex Boiss.) Ehrend. & Krendl Asperula doerfleri Wettst. Asperula purpurea (L.) Ehrend. subsp. apiculata (Sibth. & Sm.) Ehrend. Asperula scutellaris Vis. Crucianella graeca Boiss. Galium breviramosum F. Krendl Galium firmum Tausch Haplophyllum boissieranum Vis. & Pančić Haplophyllum coronatum Griseb. Thesium auriculatum Vandas Acer heldreichii Orph. ex Boiss. subsp. visianii K.Malý Acer hyrcanum Fisch. & C.A.Mey. subsp. intermedium (Pančić) Palam. Saxifraga blavii (Engl.) Beck * * ** * ** ** DD (IUCN) DD (ERLVP)* ** * * * ** ** EX(Srb)** * ** * CR(Srb)** ** * * * * * * * * * ** ** LC (IUCN) LC (ERLVP)** ** ** * ** * * ** ** * * * * * ** ** H H H H H H H H H H Ch H T H SEM SEM SEM SEM SEM SEM SEM PONT PONT CEM CEM MED-SUBMED MED-SUBMED CEM SE, K, M K, M NE, E, SE, C, K, M Š, NW, W, SW, C, K K NW, W K, M M Š, NW, W, SW, C, K E, SE, S E, C, K, M K, M M E, SE, C, K, M Ch H H H H H H T H G G G G H H H P P H H H H H H H H P Ch H CEM CEM CEM CEM CEM CEM CEM MED-SUBMED MED-SUBMED MED-SUBMED MED-SUBMED CEM PONT CEM SEM CEM MED-SUBMED MED-SUBMED CEM CEM CEM CEM CEM CEM CEM CEM MED-SUBMED MED-SUBMED CEM E, K, M K, M E, SE, NW, W, SW, C, K, M NW, W, SW, C, K, M C, M W, SW E SE, S SE, S, M K, M Š, W, SW, C, K M NE, E, SE, S, C K, M K, M M M S M E, SE, W, SW, C, K, M M K W, C SE, K, M K, M SE, SW, C M W, SW M H Ch H H Ch H H Ch Ch Ch Ch T Ch Ch Ch Ch H P P H CEM CEM CEM PONT SEM MED-SUBMED MED-SUBMED SEM SEM SEM MED-SUBMED MED-SUBMED SEM MED-SUBMED MED-SUBMED MED-SUBMED SEM CEUR CEUR CEM SE K, M W, K, M NE K, M Š, W, SW, C, K, M M C, K, M K, M E, SE, S, K NW, W, SW, M E, SE, S M M W, C, K, M M NW, W Š, E, W, SW, C, K, M Š, NE, E, SE, S, SW, C, K, M W, SW, C, K, M 185 Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Polygalaceae Polygalaceae Polygonaceae Polygonaceae BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 406 407 408 409 410 411 412 413 414 415 416 417 418 419 186 G. TOMOVIĆ ET AL. Threatened and legally protected status No Family Taxon 469 470 471 472 473 474 475 476 477 478 479 480 481 Saxifragaceae Saxifragaceae Saxifragaceae Saxifragaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae 482 483 484 485 486 487 488 489 490 491 492 Scrophulariaceae Scrophulariaceae Scrophulariaceae Thymeleaceae Violaceae Violaceae Violaceae Violaceae Violaceae Violaceae Violaceae Saxifraga federici-augusti Biasol. subsp. federici-augusti Saxifraga federici-augusti Biasol. subsp. grisebachii (Degen & Dörfl.) D.A.Webb Saxifraga scardica Griseb. Saxifraga taygetea Boiss. & Heldr. Scrophularia aestivalis Griseb. Scrophularia bosniaca Beck Scrophularia tristis (K.Malý) Šilić Verbascum adamovicii Velen. Verbascum anisophyllum Murb. Verbascum baldaccii Degen Verbascum dieckianum Borbás & Degen Verbascum eriophorum Godr. Verbascum glabratum Friv. subsp. bosnense (K.Malý) Murb. Verbascum nicolai Rohlena Verbascum scardicola Bornm. Verbascum viridissimum Stoj. & Stef. Daphne malyana Blečić Viola aetolica Boiss. & Heldr. Viola chelmea Boiss. & Heldr. subsp. chelmea Viola dukadjinica W.Becker & Košanin Viola elegantula Schott Viola grisebachiana Vis. Viola macedonica Boiss. & Heldr. subsp. macedonica Viola orphanidis Boiss. subsp. orphanidis * * ** * * * * * * * ** ** ** * * * * Life form Chorological group Distribution in geographical provinces Ch Ch Ch H H H T-H H H H H H H SEM SEM SEM SEM CEM CEUR MED-SUBMED SEM MED-SUBMED SEM PONT MED-SUBMED MED-SUBMED C, K, M S, M K, M K, M SE, S, K, M K, M Š, W, SW, C, K, M E, SE E, SE M SE, S NE, C W, SW, M H H H Ch H H H H H H H SEM SEM MED-SUBMED SEM SEM SEM SEM SEM SEM MED-SUBMED SEM SW, K, M K, M SE W, SW M K K SW, C, K, M NE, E, K, M Š, NE, E, SE, S, NW, W, SW, C, K, M K, M IUCN (2012) threatened status and European Red List of Vascular Plants (2011) (IUCN; ERLVP – bold letters; EX – Extinct, EN – Endangered, NT – Near Threatened, LC – Least Concern, DD – Data Deficient); threatened status in Serbia (Stevanović, 1999 – regular letters; EX – Extinct, EX(Srb) – Extinct in Serbia; CR – Critically Endangered, CR(Srb) – Critically Endangered in Serbia). **Species strictly protected by the national legislation. *Species protected by the national legislation. For the explanations of life forms, chorological groups and geographical provinces of the Central Serbia and Kosovo regions, see Material and methods. Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Table 2. Continued BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS 187 Figure 3. Endemic species and subspecies richness at generic level. chorological group (122 taxa; 25%), indicating the high association level of endemic taxa to mountainous and high-mountainous areas. Also, a relatively high number of endemics belonging to the Mediterranean-Submediterranean chorological group (103 taxa; 21%) is caused by a significant influence of (sub)Mediterranean flora in investigated territory. The Central European chorological group is represented by 45 taxa (9%) and the Pontic chorological group by only 26 endemic taxa (5%) of the total © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 2. Endemic species and subspecies richness at family level. 188 G. TOMOVIĆ ET AL. Balkan endemic flora in Central Serbia and Kosovo regions. 1 The South European mountainous chorological group (SEM) is represented by 196 taxa or 40% of the total Balkan endemic flora. This chorological group is floristically richest in the total chorological spectrum of the Balkan endemic flora in Central Serbia and Kosovo regions. Endemic orophytes from this chorological group inhabit limestone rocks, screes, and rocky grounds of subalpine and alpine belts of mountains of the western and central part of the Balkan Peninsula, especially mountains that are under Mediterranean climatic influence. The relatives of such species are distributed in the mountains of southern Europe (Greece, the Apennines, Iberian Peninsula) and southern Asia Minor. Endemic orophytes of the South European mountainous group are divided into the following chorological subgroups: (1.1) DinaricBalkan mountainous (75 taxa; 38%); (1.2) Dinaric mountainous (39 taxa; 19%); and (1.3) Balkan mountainous (82 taxa; 43%). 2 The Central European mountainous chorological group (CEM) is represented by 122 taxa (25%). The endemics of this chorological group are closely related or are vicarious to the species distributed in the mountains of the Central Europe (the Alps and the Carpathians). Endemic taxa of this group are orophytes inhabiting rocks and rocky slopes of subalpine and alpine belts of both calcareous and siliceous of inland Dinaric Alps, Balkan, and ScardoPindhian mountains. Endemic orophytes of the Central European mountainous group are divided into the following chorological subgroups: (2.1) Dinaric-Balkan mountainous (53 taxa; 44%); (2.2) Dinaric mountainous (23 taxa; 18%); and (2.3) Balkan mountainous (46 taxa; 38%). 3 The Mediterranean-Submediterranean chorological group (MED-SMED) is represented by 103 taxa (21%). These endemics are distributed in Central Serbia and Kosovo regions on small surfaces or exclaves connected to the Eastern Submediterranean subregion, which includes parts of Macedonian-Thracian province in southern and south-eastern Serbia and the Adriatic province in south-western Serbia. In addition, these types of endemics are also distributed in the other parts of Central Serbia and Kosovo regions, especially inhabiting open limestone and serpentine sites. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 4. Spectrum of the basic chorological groups (corresponding to the floristic provinces) of the Balkan endemic flora in the Central Serbia and Kosovo regions: CEM, Central European mountainous; SEM, South European mountainous; PONT, Pontic; MED-SUBMED, Mediterranean-Submediterranean; CEUR, Central European. BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS LIFE FORM SPECTRUM The life form analysis of the Balkan endemics in Central Serbia and Kosovo regions is presented in Figure 5. The largest group of the Balkan endemics are hemicryptophytes (313 taxa; 64%) and chamaephytes (117 plants; 24%). Significantly less numerous are geophytes and therophytes (26 each; 5%), whereas the lowest number, only ten endemic taxa (2%) belong to phanerophytes and nanophanerophytes (trees and shrubs), which are represented by five taxa each. Of the five endemic tree species, two taxa belong to the CEUR chorological group, two species to the CEM group, and one species to the SEM group. Balkan endemic shrubs mainly belong to the MED-SUBMED group (four taxa), and one species belongs to the CEM chorological group. Hemicryptophytes are the most numerous in the SEM and CEM chorological groups (119 and 98 taxa, respectively) and less numerous in the PONT group (20 taxa). Most endemic chamaephytes (62 taxa) are attributed to the SEM chorological group and, to a lesser extent, to the MEDSUBMED group (34 taxa); in the CEUR group, there are no endemic chamaephytes. Geophytes are most numerous in the MED-SUBMED and SEM groups, with 11 and nine taxa, respectively; in other chorological groups, only one to three geophytes are recorded. Therophytes are most abundant in the MED-SUBMED chorological group (16 taxa), whereas, in the CEUR and SEM groups, they are represented by five and four species, respectively. In the CEM chorological group, there is only one endemic therophyte species and, in PONT, there are no endemic therophyte taxa recorded. ENDEMISM BY GEOGRAPHICAL PROVINCES The uneven distribution of the Balkan endemics in the 11 (out of 15) geographical provinces in Central Serbia and Kosovo regions is shown in Figure 6. In the Peripannonian part (Pomoravlje province; Po), there are no Balkan endemics, and their number increases regularly in a north–south direction (i.e. from plain to mountainous and high-mountainous areas). By far the largest numbers of endemic taxa are present in the geographical provinces of Metohija (M) (328) and in Kosovo (K) (227). The third highest number is in the province of Eastern Serbia (E) (127 taxa). Provinces of Central (C), South-eastern (SE), and Western (W) Serbia, with 102, 102, and 97 taxa, respectively, are moderately rich in endemic Balkan flora. The next group consists of provinces of Southwestern (SW) with 85 and Southern Serbia (S) with 71 taxa, characterized by a relatively small number of the endemic plants. The last groups with the lowest number of Balkan endemics are provinces of Northeastern (NE) and North-western Serbia (NW), with 41 and Šumadija (Š) with only 25 endemics. The cluster analysis of floristic similarities among the 11 geographical provinces of Central Serbia and Kosovo regions based on the Balkan endemic taxa © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 The chorological spectrum of this group consists of the following subgroups. (3.1) MediterraneanSubmediterranean-Subcontinental subgroup (10 taxa; 10%): this subgroup includes endemics with ranges that, besides the Mediterranean and Submediterranean region, spread further into the Balkan Peninsula. These species inhabit thermophilous rocky habitats of the continental parts of western and central Balkans. (3.2) Submediterranean subgroup (37 taxa; 36%): this subgroup has a largest number of the Balkan endemic taxa. Distribution of the typical Submediterranean species encompasses larger parts of the Adriatic, Ionian, Aegean and Macedonian-Thracian provinces of the Eastern Submediterranean subregion. These endemics can be found in southern, southwestern and south-eastern parts of Central Serbia and Kosovo regions; (3.3) SubmediterraneanSubcontinental subgroup (34 taxa; 33%): ranges of the endemics that belong to this subgroup, besides Submediterranean region, include the inland parts of the Balkan Peninsula. (3.4) SubcontinetalSubmediterranean subgroup (22 taxa; 21%): this subgroup mainly comprises the endemic species that are predominantly or exclusively distributed in the continental parts of the Balkans. However, its florogenetical and ecological features correspond to the Submediterranean subgroup. 4 The Central European chorological group (CEUR) is represented by 45 taxa (9%). It includes Balkan endemic taxa distributed in broad-leaved and mixed forests and mesophilous meadows of hilly and montane zones of the western and central parts of the Balkan mountains. In terms of geography and size of ranges, the endemics that belong to this chorological group are divided into the following floristic subgroups and elements: (4.1) Illyrian-Balkan (14 taxa; 31%); (4.2) Illyrian (20; 45%) and (4.3) Balkan (11 taxa; 24%). 5 The Pontic chorological group (PONT) is represented by 26 taxa (5%). These endemics are vicarious to the wide distributed Pontic relatives and they mostly inhabit serpentine and/or limestone rocky slopes of hilly and montane zones in the central, and to a lesser extent, the western part of the Balkan Peninsula. Balkan endemics of Pontic origin are divided into the following floristic subgroups and elements: (5.1) Illyrian-Balkan (four taxa; 15%); (5.2) Illyrian (six taxa; 23%) and (5.3) Balkan (16 taxa; 62%). 189 190 G. TOMOVIĆ ET AL. occurrence (Fig. 7), allows the identification of two sets of geographical provinces. The first set includes Kosovo (K) and Metohija (M) and it is well differentiated with respect to the distribution of many Balkan endemics in these two provinces. The second set includes the eastern subset (NE, E, SE, and S Serbia), reflecting the geological history and close climate characteristics of these provinces. The western subset includes the remaining provinces; Šumadija province (Š) stands separately as a result of its poor endemic flora. The close phytogeographical connections of NW, W, and SW Serbia are obvious because all three provinces are geographically close to each other and they share similar ecological and floristic features. Although the province of Central Serbia (C) belongs to this subset, it appears to be well differentiated with respect to sharing of some Balkan endemic plants with the provinces of the eastern set. CENTRES OF ENDEMISM To define centres of endemism in Central Serbia and Kosovo regions, an integrated summary distribution map for all 492 Balkan endemics was constructed on a 10 km × 10 km grid (Fig. 8). The highest number of endemic taxa is recorded in Mts Prokletije (UTM square DN32, 151 taxa), which makes this massif the most important centre of endemism (e.g. centre of the highest floristic richness of the Balkan endemic plants). Squares DN23 and DN31, also in Mts Prokletije (with 122 and 142 taxa), and DM87 and DM97, in Mts Šar-Planina (with 115 and 128 plants), belong to the second class with a large number of endemic species (115–142). Mt Paštrik (96 taxa in DM67) and parts of Mts Prokletije (107 taxa in DN33) and Mts Šar-Planina (113 taxa in DM96 and 114 taxa in EM07) belong to © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 5. Life form spectrum of the total Balkan endemic flora and within five basic chorological groups. P scap, phanerophytes scapose; NP caesp, nanophanerophytes caespitose; Ch, chamaephytes; H, hemicryptophytes; G, geophytes; T, therophytes. BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS 191 Figure 7. Cluster analysis of floristic similarities among the 11 phytogeographical provinces of the Central Serbia and Kosovo regions based on the Balkan endemic taxa occurrence. Similarity coefficient used: Sørensen. the third class by the number of endemic taxa (96–114). Squares DN21 (87) in Mts Prokletije and DM74 (94), EM06 (85), and EM17 (87) in Mts ŠarPlanina belong to the fourth class with 85–94 taxa, and DM86 (83) in Mts Šar-Planina, DM65 (79) and DM66 (82) in Mt Koritnik, and square DN89 (70) in Mt Kopaonik belong to the fifth class of 70–83 endemic representatives. These three classes are also characterized by high endemism. Medium-high levels of endemism are distinguished by the sixth, seventh, and eight classes. In the sixth class (52–69 taxa) are Mt Stara Planina with 62 endemics in square FP30 and Mt Tara and Mt Mokra Gora with 53 taxa in square CP75. Mt Suva Planina with 50 taxa in square EN98, Vlasina plateau with the surrounding mountains with 42 endemic taxa (square FN01), parts of Mt Kopaonik (square DN88), Mts Prokletije (square DN43), and Mts Šar-Planina (square DM73, DM75) belong to a seventh class with 41–50 Balkan endemics. Some parts of Mts Prokletije (squares DN20, DN23, DN43) and Mts Šar-Planina (squares DM73, DM75) also belong to these three classes. Medium levels of endemism are characteristic for classes nine (31–35 taxa), ten (25–30 taxa), and 11 (21–24 plants), which encompass 53 UTM squares. These squares are more or less related to the hilly and mountainous areas or small gorges and canyons in Central Serbia and Kosovo regions. The next three classes are characterized by a relatively small number of Balkan endemics. Namely, 17–20 taxa were recorded in 23 squares, 13–16 taxa in 32 squares, and eight to 12 taxa are present in 48 squares that are scattered throughout the Central Serbia and Kosovo regions. The lowest numbers of endemic taxa (up to seven) characterize the last three classes with 270 UTM squares. ELEVATION, GEOLOGICAL SUBSTRATUM AND VEGETATION DIFFERENTIATION Based on the distribution of endemic taxa by elevational range (Fig. 9), it is obvious that most of the endemics (287) occur in the 1500–2000 m range, with © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 6. The distribution of the Balkan endemic plants in geographical provinces of the Central Serbia and Kosovo regions. 192 G. TOMOVIĆ ET AL. Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 8. Centres of endemism of the vascular flora in the Central Serbia and Kosovo regions. Number of species per square 10 × 10 km. UTM projection map grid zone 34T. Mountains > 1500 m high are contoured. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS 193 Figure 10. Distribution of the Balkan endemic plants on different geological substrates. slightly less (249) in the 1000–1500 m range. A significant number of endemic species occurs in the 500–1000 m range (232), whereas the 2000–2500 and 0–500 m ranges, with 170 or 143 endemic plants, respectively, are much poorer. The number of endemic taxa is dramatically lower (49) above 2500 m a.s.l. (i.e. the belt encompassing only the highest mountain peaks in Central Serbia and Kosovo regions). Analysis of the distribution of the number of endemic taxa by type of geological substrate (Fig. 10) showed that most of the plants occur on limestone (376); on silicate, there are almost half of the taxa © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 9. Altitudinal distribution of the Balkan endemic vascular plant taxa shown as a number of endemics occurring in a 500-m elevational belt. 194 G. TOMOVIĆ ET AL. (232), and there are only 145 plants that grow on serpentinites. On miscellaneous geological substrates (e.g. loess, unbound Pleistocene sediments, sand, etc.), only 36 taxa were recorded. The distribution of the Balkan endemic plants in different vegetation formations and classes is shown in Figure 11. The formation of rock crevices (class Asplenietea trichomanis) is the richest in endemic plants (231), whereas in screes vegetation (class Thlaspietea rotundifolii) only 49 endemics are recorded. The formation of temperate grasslands, heaths and fringe vegetation is represented by three vegetation classes in which endemic plants are found, and the class Festuco-Brometea is the richest in the number of endemic plants (204). Within the formation of montane, tall-herb, grassland, fell-field, and snow-bed vegetation endemic plants are present in four different vegetation classes, and the richest is class Elyno-Seslerietea with 156 endemic species and subspecies. Within the formation of temperate broadleaved forests and scrub, endemic plants are recorded in seven different vegetation classes, and the richest is class Quercetea pubescentis (104 taxa). The formation of montane heaths and coniferous forests are represented with four vegetation classes, of which class Erico-Pinetea is by far the richest in © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 Figure 11. Distribution of the Balkan endemic plants in different types of vegetational formations and classes. Classes with no endemic taxa are not shown. BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS TAXONOMIC DISCUSSION GROUPS, CHOROLOGICAL GROUPS AND LIFE FORMS The floristic analysis of the endemic taxa highlighted certain families that are important contributors to the flora of the temperate regions in the Holarctic. Asteraceae have an excess of endemic genera in East Asia and North America (Qian, 2001). Caryophyllaceae show active endemism in Serbia (Niketić, Stevanović & Tomović, 2007) and, with Brassicaceae, have large concentrations of genera in the Mediterranean basin (Heywood et al., 2007). Centres of origin and diversification of some genera (e.g. Verbascum; Scrophulariaceae) is Asia Minor, although the Balkan Peninsula also represents a secondary centre of diversity for this genus (Turrill, 1929). The Mediterranean basin is the evolutionary centre of whole campanuloid lineage of Campanulaceae (Kovačić, 2004) and a centre of diversification for Lamiaceae (Heywood et al., 2007). The ten to 15 families with the highest numbers of endemic taxa fall more or less within the richest families of the total Serbian flora (Stevanović et al., 1995). A conspicuous difference between the total and Balkan endemic flora of Serbia is seen in Poaceae, which, with Asteraceae and Fabaceae, consistently figure among the three richest and most diverse families, although they rank lower in absolute number of endemics. Also, the ten to 15 families richest in endemics in Central Serbia and Kosovo regions more or less coincide with those in neighbouring Bulgaria (Petrova & Vladimirov, 2010) and Greece (Georghiou & Delipetrou, 2010). The main differences between these three countries are noted in Rubiaceae (higher rank in Greece), Lamiaceae (lower rank in Bulgaria), and Poaceae (higher rank in Bulgaria). In the chorological spectrum of the Balkan endemic flora in Central Serbia and Kosovo regions, the predominant classes are South European mountainous and Central European mountainous taxa, which together account for more than two-thirds of the endemic flora. The explanation for such a marked dominance of mountainous chorological groups and orophytic elements lies in the fact that the mountainous and high mountainous areas of Central Serbia and Kosovo regions are the centres of florogenesis and refugees of the orophytic flora of the central and southern Europe (Stevanović, 1996). The highly heterogeneous topography of the mountains favoured the emergence of new lineages, which are generally conisdered to have evolved as a result of microallopatric speciation (Hewitt, 2000; Tzedakis et al., 2002). The Ice Age was a particularly significant period for isolation and process of neospeciation, during which new infraspecific taxa were formed (mostly at subspecies or variety level), and which are related to high-mountain massifs in the investigated territory. A significant percentage of the endemics belong to MediterraneanSubmediterranean chorological group, which is a result of the presence of a large number of xerophilous and thermophilous habitats, in the southern parts of Serbia and the lower parts of the Kosovo and Metohija plains. In addition, the more intense effect of anthropogenic factors caused the spread of secondary vegetation types (grass xerophilous communities and shrub communities) in the place of the primary forest and shrub vegetation. This has led to an increase in the number of endemic taxa belonging to Mediterranean-Submediterranean chorological group and their expansion and deeper north penetration in the continental part of the territory of Central Serbia and Kosovo regions, as identified by Turrill (1929), and recent studies show that the thermophilous-xerophilous representatives are spreading through the gorges and canyons of southern Serbia (Rand̄elović et al., 2008; Zlatković et al., 2011). Endemic taxa that belong to Central European chorological group are represented by endemics distributed in broadleaved forests and mesophilous meadows mainly in the western and central parts of the Balkans. They are closely related to widely distributed Central European species and are not significant contributors in the total chorological spectrum of the Balkan endemic flora in Central Serbia and Kosovo regions. Generally, Pontic species have a significantly wider distribution outside the Balkan Peninsula, spreading westward to the peninsula along Dobruja and Wallachia to the western parts of Bulgaria and the eastern provinces of Central Serbia. The small number of their Balkan endemic vicarious relatives indicates the differentiation and speciation of populations of widely distributed Pontic species in the Balkans during the period of Pleistocene when the steppe flora was spreading along the peninsula. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 endemic plants (125). Within the formation of weed communities, the presence of endemic taxa was recorded in four vegetation classes, mostly in class Stellarietea mediae (28). All the other vegetation formations, in which endemic plants are found, are represented by a single vegetation class and < 34 endemic plants. In two vegetation formations [coastal mud-flats and brackish waters (class Ruppietea maritimae) and salt marsh, sand-dune and sea-cliff vegetation (class FestucoPuccinellietea)], Balkan endemic plants were not recorded. 195 196 G. TOMOVIĆ ET AL. FLORISTIC RICHNESS AND CENTRES OF ENDEMISM In the flora of Serbia Gajić (1984) and Diklić (1987) recorded the presence of 197 endemic species and subspecies, whereas Stevanović et al. (1995) concluded that the number of the Balkan endemic in Serbia is higher (287 taxa). According to the data, there are 492 Balkan endemic species and subspecies in Central Serbia and Kosovo regions, representing approximately 18% of the total number of 2660–2700 endemic taxa in the Balkan Peninsula (Stevanović, 2005; Stevanović et al., 2007). The highest level of endemic flora richness in the Balkans is in Greece, with 1226 single-country endemics being present (Georghiou & Delipetrou, 2010). In the flora of Bulgaria, 172 species are single-country endemics (Velchev & Kozhuharov, 2006), and 270 are Balkan endemics (Petrova & Vladimirov, 2010). Matevski et al. (2003) recorded the presence of 115 local endemic species in the Republic of Macedonia; in Croatia, there are 376 Balkan endemics (Nikolić, 2012); in Bosnia and Herzegovina, there are 298 (Lubarda et al., 2014); and, in Montenegro, Stevanović et al. (1995) found 223 Balkan endemic plants. Ruci (1997) recorded 26 ‘national’ and approximately 150 Balkan endemic taxa in Albania but, according to Turrill (1958), these numbers are much higher (50 single-country and 320 Balkan endemics). In general, areas of high species richness in Serbia coincide with those areas of endemism and the present study is in accordance with former preliminary surveys (Stevanović et al., 1995, 2002). Centres of plant endemism in Central Serbia and Kosovo regions show a marked mountainous character, falling almost entirely in the main mountain ranges and highlands of the southern and central provinces of the investigated area. Exceptional endemic flora richness of the Mts Prokletije and Mts Šar-Planina (and to a lesser extent of Mt Kopaonik) is caused by confrontation of three types of geological substrates (limestone, silicate, and serpentinite), a wide elevational range and the diverse vegetation types. Such diverse edaphic, climatic (presence of mountainous, temperatecontinental, and transitional Submediterranean climate at lower elevations and in river gorges), and habitat conditions enabled the survival and development of many high-mountain plant taxa that were differentiated during the long process of speciation (Stevanović et al., 1995). According to Stebbins & Major (1965), the richness and diversity of a flora, particularly in respect to endemic species, may be a result of either the presence of a large number of ancient, relict species, or active speciation in relatively recent times, or both factors. In regions on the borderline between zones of adequate and deficient mois- © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 The life form spectrum of floras of certain areas is an indicator of a complex interdependence of plant life forms and the environmental (especially climatic and edaphic) features of such areas. It still contains a historical dimension and reflects adaptation of plants through time and the current conditions, in agreement with the environmental conditions in which these plants survive (Diklić, 1984). The life form spectrum of the endemics in Central Serbia and Kosovo regions, with a predominance of perennial hemicryptophytes and chamaephytes and low proportions of annuals and geophytes, is closer to the high-mountain spectra and compatible with the results of biological analysis of Greek endemic plants (Georghiou & Delipetrou, 2010) and Balkan endemics in Bulgaria (Petrova & Vladimirov, 2010). In the comparative life form spectra, the endemic flora follows trends for the total flora of Serbia (Diklić, 1984) and the Balkan flora (Turrill, 1929) (although the highest percentage of hemicryptophytes was recorded in the endemic flora of Central Serbia and Kosovo regions). All three floras are of hemicryptophytic character, which is generally characteristic of the floras of most regions of temperate zone. Chamaephytes account for approximately 27% of the endemic flora, which is almost three times more than the total flora of Central Serbia and Kosovo regions and almost twice the amount in the Balkan total flora. An increased proportion of chamaephytic members compared with two other floras indicates that the majority of the endemics are inhabitants of mountain and high-mountain rocks, cliffs and screes. Floras that develop on these habitat types are characterized by slow-growing, desiccation tolerant or avoiding, perennial plants (Larson, Matthes & Kelly, 2000). The greatest discrepancy among the life form spectra of these three floras was noticed in the representation of annual plants (therophytes). Therophytes in the Balkan Peninsula and total Serbian flora represent approximately one-fifth of the whole floras, whereas, in the endemic flora of Central Serbia and Kosovo regions, it is only 5%. This could be explained by the fact that annual plants mostly inhabit dry grasslands and rocky grounds that are often characterized by low productivity and intense drought stress combined with low- to high-intensity, unpredictable yet recurrent or even continuous, anthropogenic or natural disturbance (Noble & Slatyer, 1980). The life form spectrum of the endemics cannot be explained only by elevational or geological substratum distribution but appears to be more indicative of their habitat and adaptive strategy and speciation processes (Georghiou & Delipetrou, 2010). BALKAN ENDEMIC PLANTS IN CENTRAL SERBIA AND KOSOVO REGIONS ELEVATION, GEOLOGICAL SUBSTRATE, AND VEGETATION DIFFERENTIATION The distribution curve of endemic taxa in relation to elevational range has an almost normal distribution, which means that, going to the lowest and highest elevations, the number of endemics is reduced, with the largest number of endemic taxa noted in the zone just above the upper forest border (1500–2000 m a.s.l). In the mountains, an increase in percent of endemism is reported to increase with elevation up to the subalpine zone (Major, 1988), and a high level of endemism at high elevations (above the tree line) compared to the lowlands is a general phenomenon in Eurasian mountain systems (Dhar, 2002). A number of surveys have shown that species richness, either of particular taxonomic groups (Hemp, 2002; Tribsch & Schönswetter, 2003; Essl et al., 2009) or of the whole flora (Rahbek, 1995), reaches its highest values at medium elevations. This can be explained by the fact that many species can adapt more easily to moderate environments but not to extremely adverse ones (Grace, 1999; Van der Meulen, Hudson & Scheiner, 2001). In some other mountainous regions of the world, most endemics are found in low elevation mountainous areas and medium elevations of the subalpine zone, and less in the alpine zone (Borchsenius, 1997; Médail & Verlaque, 1997; Giménez et al., 2004; Van der Werff & Consiglio, 2004; Nowak, Nowak & Nobis, 2011). Analyses of the distribution of the number of endemic taxa in relation to the types of geological substrates showed that approximately 75% Balkan endemics in Central Serbia and Kosovo regions grow on limestone. On serpentinites, 145 endemic plants were recorded, which is almost half of the endemic serpentinophytes of the entire Balkans (Stevanović et al., 2003). According to Stebbins (1980), generally neutral and basic geological substrates are ideal for the survival and development of endemic taxa, which was confirmed at the level of the endemic flora of the Caucasus (Kikvidze & Oshawa, 2001), the eastern Alps (Tribsch & Schönswetter, 2003) and the flora of Bulgaria (Velchev, 1998). According to Tribsch & Schönswetter (2003), it is impossible to exclude that higher habitat diversity may also cause higher endemism in calcareous regions. Silicate areas of Central Serbia and Kosovo regions are characterized by a far lower number of endemics (N = 232) than limestone areas. Although this type of substrate is widely represented in hills and mountains of South-eastern Serbia, and in the high-mountain areas of Central Serbia and Kosovo, silicate habitats of the Balkan Peninsula are much poorer in endemic and nonendemic plants than limestone (Velchev, 1998). Similar results regarding the less presence of © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 ture, even small climatic shifts will change local conditions beyond the limits of tolerance of the resident species, so that these must either migrate or evolve new ranges of tolerance. In this way, climatic and edaphic diversity, occurring on ecotones or border regions between different biotic provinces, represent factors that most actively promote the evolution and differentiation of higher plant species. In addition to high-mountain endemism, edaphic endemism is also expressed in Central Serbia and Kosovo regions. Mutually conditional, highmountainous and edaphic endemism often occur in combination. Limestone mountains of Central Serbia and Kosovo regions such as Mt Paštrik, Mt Koritnik, and Mt Suva Planina represent centres of richness of the Balkan endemics. In the serpentinite habitats in the Western and Central Serbia, Kosovo and Metohija provinces, a specific ophiolitic endemic flora develops. Parts of Mt Tara, Mt Zlatibor, Mt Stolovi, Mt Studena, the southern slopes of Mt Kopaonik, the valley of the river Ibar, and the Metohija plain (surroundings of the cities Ðakovica and Klina) are important centres of serpentinite endemic flora richness. Vlasina plateau with relatively high surrounding mountains (up to 1900 m a.s.l.) and Mt Stara Planina in the Eastern Serbia represent important centres of siliceous endemic flora. Areas of endemism located outside the highmountainous ranges include river valleys and gorges of the eastern (Sićevačka gorge) and south-eastern Serbia (gorge of the river Pčinja) and some hilly and lowland parts of southern Serbia (surrounding of Vranje city and Mt Rujan). By contrast to mountains, lowland habitats rarely form suitable refuges (at least in temperate latitudes) because of their much harsher and less stable eco-climatic conditions (Tribsch & Schönswetter, 2003; Tribsch, 2004). In addition, it should be noted that lowland areas in Central Serbia and Kosovo regions have been severely transformed by human activity (cultivation, human settlements, infrastructure), especially those areas in the large river valleys (Danube, Sava, Morava, etc.). However, it is already known that areas of high endemism do not always correspond with those of high species richness (Prendergast et al., 1993; Trigas, Iatrou & Karetsos, 2007). This ‘rule’ is confirmed for Mt Suva Planina in the Eastern Serbia, which ranks second by its diversity and richness of the total flora (1250–1500 plants at UTM 20 squares), at the same time as having moderate level of endemism. Also, Ðerdap gorge in north-eastern Serbia was not found to be a centre of Balkan flora endemism, although it has been confirmed as a significant centre of total species richness in the country (Stevanović et al., 2002). 197 198 G. TOMOVIĆ ET AL. habitats tend to have much lower levels of endemism than forest, scrub, and alpine habitats. Weed communities are also characterized by low number of endemics, which is a consequence of the intensive human activity and lack of adequate habitats for the development of such plants (Stešević, Jovanović & Šćepanović, 2009). CONSERVATION STATUS The lack of more recent data is one of the reasons that Balkan endemics in Central Serbia and Kosovo regions rank so low in the world Red Data Book of Plants (IUCN, 2012), which contains only 16 taxa from the country, and in the European Red List of Vascular Plants (Bilz et al., 2011), which contains only 17 plants (Table 2). Until now, only 33 endemics had a nationally threatened status; five plants were extinct in Serbia, EX(Srb), four of which were globally extinct, EX, because they were local endemic taxa. Of the 24 critically endangered taxa, 15 plants are critically endangered in Serbia, CR(Srb), of which nine are globally critically endangered, CR, given that these plants are distributed only in the Central Serbia and Kosovo regions (Stevanović, 1999). Three hundred and four endemic species and subspecies are protected by national legislation (Anonymous, 2010); 109 are strictly protected and 113 protected (Table 2), representing 62% of all the Balkan endemic vascular plants for the country. CONCLUSIONS: IMPLICATION FOR CONSERVATION Potentially, some of our results would be of major interest from a conservation point of view. The area of nature conservation in Central Serbia and Kosovo regions is regulated by the Law on Nature Conservation (‘Official Gazette RS’ no. 36/2009 and 88/2010) and other legal acts and bylaws that directly or indirectly relate to nature and natural resources. So far 464 natural assets have been legally protected: five national parks, 16 nature parks, 17 landscapes of outstanding features, 72 nature reserves, 313 natural monuments, and 43 surroundings of immobile cultural assets and significant places (Amidžić, Krasulja & Belij, 2007). In addition, there are 62 areas that are proposed for proclamation as ‘Important Plant Areas’ (IPAs). Almost 56% of Serbian IPAs are nationally protected in full or in part, and almost half are at a higher level of protection (Stevanović & Šinžar-Sekulić, 2009). An important proportion of the legally protected natural assets and of proposed IPAs are currently located in the main mountainous ranges. As stated above, there is a general coincidence between the areas of high level of endemicity and mountainous ranges. These areas usually include the © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 173–202 Downloaded from https://academic.oup.com/botlinnean/article/176/2/173/2416323 by guest on 08 November 2023 endemics in acidophilous areas of Austria and in Iberia and the Balearic islands were reported by Essl et al. (2009) and Domínguez Lozano et al. (2000), respectively. Endemics are known to occupy different habitat ranges. Their occurrence is associated with rocky cliffs and rapidly eroding sites in most of the mountain systems (Major, 1988). In the present study, vegetation of rock crevices (class Asplenietea trichomanis) is the richest in number of endemic plants as a result of its wide elevation range and the variety of geological substrates in which this type of vegetation can be found. Also, this class encompasses many vegetation orders, alliances, and plant communities (including many endemic associations described from Central Serbia and Kosovo regions) that are especially rich in endemic plants. On the basis of the analysis of high-elevation Himalayan endemic plants (Dhar, 2002), it was concluded that most of the narrow endemics occupy the drier habitats and this relationship is more prominent among alpine endemics. Plant communities included in the vegetation class Festuco-Brometea predominantly occupy xerophilous grasslands in which many of endemics can also be found. Similar trends of distribution are noticed in south-eastern France (Médail & Verlaque, 1997), with the majority of endemic species occupying communities with high stress levels, such as rocky xerophilous grasslands, screes, cliffs, and rock habitat at mid- or high elevations. Nowak et al. (2011) stated that habitats with wide elevational ranges (e.g. screes) are also characterized by rich endemic flora, although the study was not specific in terms of elevation. By contrast, in the present study, screes vegetation of the class Thlaspietea rotundifolii are quite poor in number of endemics, which is probably a result of the limited and scattered distribution of this vegetation type in Central Serbia and Kosovo regions. Mountainous grasslands (class Elyno-Seslerietea), which represent the climax and zonal vegetation types, are also inhabited by a large number of endemics. It has been argued (Essl et al., 2009) that patches of these rare habitats offer favourable conditions for speciation and long-term survival; extreme abiotic conditions give rise to high selection pressure and populations are effectively separated from each other. Mountainous coniferous forests (class Erico-Pinetea) and temperate broadleaved forests (class Quercetea pubescentis), which have wide altitudinal distribution in Central Serbia and Kosovo regions, rank fourth and fifth by the number of endemics, respectively. Generally, in fresh-water aquatic vegetation, no endemic plants were found. 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