Richness and Diversity of Pontic Flora on Serpentine of Serbia

Cent. Eur. J. Biol. • 6(2) • 2011 • 260-274 DOI: 10.2478/s11535-010-0110-5

Central European Journal of Biology

Richness and diversity of Pontic flora on serpentine of Serbia

Research Article

Ksenija Jakovljević*, Dmitar Lakušić, Snežana Vukojičić, Gordana Tomović, Jasmina Šinžar-Sekulić, Vladimir Stevanović

Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade, 11000 Belgrade, Serbia

Received 23 June 2010; Accepted 08 November 2010

Abstract: Serpentine substrate in Serbia covers wide areas mainly in Kosovo and in central, western and south-western part of the country. On the serpentines of Serbia, particularly on the skeleton soils of hills and on lower elevation mountain slopes, there are some xerophilous and steppe-like vegetations. These xerophilous grasslands include mostly Pontic, Pontic- Submediterranean and/or Mediterranean- Submediterranean-Pontic plants. The distribution of 161 taxa of Pontic, Pontic- Submediterranean and Mediterranean-Submediterranean-Pontic areal - types on serpentinites of Serbia was analyzed in order to distinguish centers of richness and diversity. The distribution of taxa was analyzed with respect to geographic, ecological and climatic factors. The qualitative composition of Pontic flora on the serpentine areas was also compared. The greatest floristic richness and the center of diversity of Pontic flora on serpentine of Serbia was recorded in central part of the area investigated.

Keywords: Phytogeography • Pontic • Richness • Diversity • Flora • Serpentine • Serbia © Versita Sp. z o.o.

serpentine zones located primarily in central, western 1. Introduction and south-western Serbia and in Kosovo. Some of these Serpentine (ophiolithic, ultramafic) rock belongs to areas are: Brdjanska gorge, Mt Suvobor, Mt Maljen, Mt the group of siliceous rocks characterized by calcium Tara, Mt Zlatibor, Mt Mokra Gora, Mt Goč, Mt Stolovi, deficiency, high concentrations of iron, magnesium, Mt Kopaonik, the Ibar valley, Peć and surroundings, nickel, cobalt and chromium, and reduced nutrients. Orahovac and Koznik, Mt Ostrovica (Figure 1B). In contrast to other acid siliceous rocks, the pH values The real steppes of southeastern Europe are of serpentine substrate vary from basic to ultrabasic present in the Pannonian and Wallachian Plain, Dobruja (pH 5.5-8). Many interesting basiphilous-calcifuge and Thracean Plain [2,3]. On the Balkan peninsula, plants flourish on serpentine owing to this factor. Itis particularly on the skeleton soils of hills and on lower one of the most important reasons why the serpentine elevation mountain slopes, there are some xerophilous flora, differing from that occurring on limestone or and steppe-like vegetation. The steppe-like vegetation is other siliceous substrates, presents taxonomical, particularly well-developed in limestone and serpentine phytogeographical and ecological phenomenon of great habitats of the Moesian subregion in Serbia, Bulgaria interest wherever it occurs [1]. and Macedonia [4-9]. The degradation of forests The largest serpentine areas in Europe are in the and shrub formations in the temperate-continental Balkan peninsula (Epirus, Thessaly and Sterea Ellas in (central and eastern) parts of the Balkan Peninsula, Greece, north, north-central and south-eastern Albania, has only increased the extent of the steppe biome. central and east Bosnia, western, south-western and These xerophilous grasslands include mostly Pontic, central Serbia, east and central Rhodopes mountains Pontic-Submediterranean and/or Mediterranean- in Bulgaria and gorge of river Pcinja and south-eastern Submediterranean-Pontic plants, but they also contain slopes of Sara mountain in north-eastern part of Republic a substantial number of endemic species. Thus, they of Macedonia) (Figure 1A). In Serbia there are several represent important centers of xerothermic flora that

* E-mail: [email protected] 260 K. Jakovljević et al.

A C

Figure 1. Distribution of serpentine bedrock: (A) in Balkans [1] (Cro-Croatia, B&H-Bosnia &Hercegovina, Mg-Montenegro, Sr-Serbia, Al-Albania, RM-Romania, BU-Bulgaria, GR-Greece, FYRMa-Former Yugoslav Republic of Macedonia, Tu-Turkey); (B) in Serbia; (C) serpentine groups studied in Serbia (1-north-western Serbia, 2-western Serbia, 3-central Serbia I, 4-central Serbia II, 5-south-western Serbia, 6-Metochia, 7-south & central Kosovo)). used to be widely distributed throughout the Balkan Peninsula, have a special phytogeographic importance. peninsula in the last glacial and postglacial period The spread of Pontic flora throughout the central and [10,11]. east Balkans was particularly pronounced during Ice The occurrence of the Pontic, Pontic- Age (Pleistocene) [2]. Serpentine areas were suitable Submediterranean and Mediterranean- habitats for these plants during this period, so steppe Submediterranean-Pontic plants on serpentine is relicts could represent current remnants of this flora. intriguing from a biogeographic standpoint because The main goal of this paper is to determine the their populations in serpentine areas of the Balkans extent to which the serpentine areas of Serbia represent are isolated with respect to their common distribution in an important region of south-eastern Europe for the the steppe regions of south-eastern Europe. The open xerothermic flora of the Pontic, Pontic-Submediterranean serpentine habitats are characterized by a pronounced and Mediterranean-Submediterranean-Pontic species. thermophilous character and xeric conditions. The In south-eastern Europe serpentine areas are quite rare xerothermic character of serpentine plants is also and unique, especially those serpentine areas that are enhanced by the specific chemical composition of the inhabited by xerothermic flora. Therefore, these unique serpentine substrate. Among the numerous xerophytes areas are of the great importance in phytogeographical on the serpentine of the Balkans, the species of the sense. The objectives of our research were to determine Pontic, Pontic-Submediterranean and Mediterranean- the richness and distribution of species belonging to Submediterranean-Pontic areal-types are most Pontic, Pontic-Submediterranean and Mediterranean- prominent. The steppe relicts with Pontic distribution Submediterranean-Pontic species on the serpentine of (for example, Astragalus dasyanthus, Oxytropis Serbia and how that varies with respect to geographic, pilosa (Fabaceae), Vinca herbacea (Apocynaceae), ecological and climatic factors. The degree of floristic Echium russicum (Boraginaceae), Galatella lynosiris similarity among serpentine groups will be determined, (Asteraceae), Peucedanum officinale (Apiaceae), and the most important centers of diversity of Pontic etc.), growing on serpentinite in the central part of flora on serpentine soils will be broadly determined.

261 Richness and diversity of Pontic flora on serpentine of Serbia

2. Experimental Procedures A regression analysis was performed in order to identify the dependency level of variation in species richness, The mapping and analyses of the distribution of with regard to the basic geographical, ecological and Pontic species on serpentines in Serbia was based on bioclimatic characteristics of defined serpentine groups. herbarium specimens from the Herbarium of University The regression analysis included only the groups north- of Belgrade (BEOU) and the Herbarium of Natural western Serbia, western Serbia, central Serbia I and History Museum of Belgrade (BEO), published data central Serbia II, which are approximately of the same [9,12-39] as well as on personal field observations. size. Each serpentine group was regressed onto the The nomenclature is consistent with the latest check- following factors: area, E-long centroid, N-lat centroid, lists, papers, monographs and databases [40-45; altitude, annual mean temperature, annual precipitation, http://ww2.bgbm.org/EuroPlusMed/; http://plantnet. precipitation of driest month, precipitation of driest rbgsyd.nsw.gov.au/iopi/iopihome.htm]. quarter, diversity of potential vegetation, average The serpentine areas in Serbia were divided into distance from the steppe regions and minimal distance the following groups according to their orographic from the steppe regions. The bioclimatic characteristics connections: of the groups were extracted from the WorldClim set of 1. north-western Serbia (including the areas of global climate layers [55]. The extraction of selected Mt Maljen, Mt Suvobor, vicinity of Gornji Milanovac); bioclimatic parameters was done by using DIVA-GIS 2. western Serbia (including the areas of Mt Tara, 5.2 software (Version 5.2; http://www.diva-gis.org). Mt Mokra Gora, Mt Zlatibor, Priboj, and the Uvac valley); The diversity of the vegetation that would exist at 3. central Serbia I (including the northern part of Ibar defined serpentine groups under the current climatic valley, with areas of Mt Stolovi, Mt Čemerno, Mt Goč, and geomorphologic conditions but without any human Mt Studena Planina); impact (potential vegetation) was established on the 4. central Serbia II (including the central and southern basis of the Map of the Natural Vegetation of Europe part of Ibar valley, with areas of Mt Željin, Mt Kopaonik, [4]. The diversity of potential vegetation is presented as Mt Rogozna); number of different mapping units per serpentine group. 5. south-western Serbia (including the areas of Floristic similarities between each set of analyzed groups Sjenica and Mt Ozren); were analyzed using the clustering method based on 6. Metochia (Koznik, Miruša, vicinity of Đakovica, Jaccard distances and unweighted pair-group average Gubavac, Milanovac); and hierarchical sorting strategy (UPGMA). All statistical 7. south and central Kosovo (Mt Goleš, Mt Ostrovica, analyses were performed with the Statistica software Mt Brezovica, Crni Vrh peak, Koznica) (Figure 1C). package (version 5.1; StatSoft, 1997). The analysis of In order to define areal-types [46-48] we used the diversity and the summary distribution of Pontic species concept of floristic elements made by Meusel et al. were presented on a 10 km x 10 km grid using the [47,49-51], Meusel, Jäger [52,53], modified for the Universal Transverse Mercator (UTM) projection (2001, Serbian territory by Stevanovic [54]. http://www.luomus.fi/english/botany/afe/map/utm.htm). The species examined were grouped into the All analyses related to mapping and spatial calculation following areal-types according to their general type of were performed by using Manifold System software distribution: (version 5.50, Manifold System, CDA International Ltd.). 1. Pontic areal-type (P), including the species distributed in the steppe areas of Europe, reaching eastward to the Central Asian steppes 3. Results 2. Pontic-Submediterranean areal-type (P-S). In addition to the Pontic region which is the main part of According to the results of floristic studies on the their range, they are also present in smaller or larger serpentine areas of Serbia, there were 161 taxa at the parts of the Mediterranean region, particularly in the species and subspecies level, belonging to Pontic (P), Submediterranean subregion Pontic-Submediterranean (P-S) and Mediterranean- 3. Mediterranean-Submediterranean-Pontic areal- Submediterranean-Pontic (M-S-P) areal-types. type (M-S-P), where the main parts of the range are Only two Pontic taxa - Bromus pannonicus subsp. situated within the Mediterranean basin, reaching to the pannonicus (Poaceae) and Stachys recta subsp. Pontic region of southeastern and parts of eastern Europe. baldaccii (Lamiaceae) - belong to the group of obligate For each serpentine group floristic richness (as total serpentinophytes on serpentine areas in Serbia, while number of species on certain groups) was calculated the other plants may also be found on other types of based on literature, herbarium and personal field data. geological substrate (limestone, silicate, etc.).

262 K. Jakovljević et al.

Area S&C Family Taxon NW W C I C II SW Met Freq type Kos

Cotinus coggygria Scop. subsp. Anacardiaceae P-S 1 1 1 1 4 coggygria Apiaceae Eryngium campestre L. P-S 1 1 1 3

Apiaceae Falcaria vulgaris Bernh. P-S 1 1

Apiaceae Ferulago campestris (Besser) Grec. P-S 1 1 2 Ferulago sylvatica (Besser) Reichenb. Apiaceae P-S 1 1 2 subsp. sylvatica Apiaceae Orlaya grandiflora (L.) Hoffm. M-S-P 1 1 2 Peucedanum officinale L. subsp. Apiaceae M-S-P 1 1 1 3 officinale Apiaceae Seseli annuum L. subsp. annuum P 1 1

Apiaceae Trinia glauca (L.) Dumort. subsp. glauca M-S-P 1 1 1 1 1 1 1 7

Apocynaceae Vinca herbacea Waldst. & Kit. P-S 1 1 Vincetoxicum hirundinaria Medicus Asclepiadaceae P-S 1 1 1 1 1 5 subsp. hirundinaria Asteraceae Achillea aspleniifolia Vent. P 1 1

Asteraceae Achillea coarctata Poiret P-S 1 1

Asteraceae Achillea nobilis L. subsp. nobilis P-S 1 1 Carduus candicans Waldst. & Kit. Asteraceae P-S 1 1 1 3 subsp. candicans Carduus hamulosus Ehrh. subsp. Asteraceae P 1 1 2 hamulosus Asteraceae Carthamus lanatus L. subsp. lanatus M-S-P 1 1 Centaurea stoebe L. subsp. australis (A. Asteraceae P 1 1 1 3 Kerner) Greuter Asteraceae Centaurea stoebe L. subsp. stoebe P 1 1 1 1 4

Asteraceae Cota austriaca (Jacq.) Schultz-Bip. P-S 1 1

Asteraceae Cota tinctoria (L.) J. Gay subsp. tinctoria P-S 1 1 1 3

Asteraceae Crupina vulgaris Cass. P-S 1 1 2 Echinops ritro L. subsp. ruthenicus Asteraceae P-S 1 1 (Bieb.) Nyman Galatella linosyris (L.) Bernh. subsp. Asteraceae P-S 1 1 1 1 4 linosyris Asteraceae Inula britannica L. subsp. britannica P-S 1 1 2

Asteraceae Inula ensifolia L. P-S 1 1 1 1 1 5

Asteraceae Inula hirta L. P 1 1 1 3

Asteraceae Inula oculus-christi L. P-S 1 1 2 Jurinea mollis (L.) Reichenb. subsp. Asteraceae P-S 1 1 1 3 mollis Lactuca viminea (L.) J. & C. Presl subsp. Asteraceae M-S-P 1 1 viminea Asteraceae Leontodon crispus Vill. subsp. crispus M-S-P 1 1 1 1 1 5

Asteraceae Picris hieracioides L. subsp. hieracioides P-S 1 1 1 3

Table 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW – North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Gornji Milanovac), W − Western Serbia (Mt Tara, Mt Mokra Gora, Mt Zlatibor, Priboj, Uvac valley), C I − Central Serbia I (northern part of Ibar valley, Mt Stolovi, Mt Čemerno, Mt Goč, Mt Studena Planina), C II − Central Serbia II (central and southern part of Ibar valley, Mt Željin, Mt Kopaonik, Mt Rogozna), SW – South-western Serbia (Sjenica, Mt Ozren), Met − Metochia (Mt Koznik, Miruša, vicinity of Đakovica, Gubavac, Milanovac), S&C Kos – South & Central Kosovo (Mt Goleš, Mt Ostrovica, Mt Brezovica, Crni Vrh peak, Koznica), Freq − frequency of occurrence in defined serpentines groups, P-S- Pontic-Submediterranean, P-Pontic, M-S-P- Mediterranean-Submediterranean-Pontic.

263 Richness and diversity of Pontic flora on serpentine of Serbia

Area S&C Family Taxon NW W C I C II SW Met Freq type Kos

Pilosella bauhinii (Schultes) Arv.-Touv. Asteraceae P-S 1 1 1 1 1 5 subsp. bauhinii Scorzonera austriaca Willd. subsp. Asteraceae P-S 1 1 1 1 4 austriaca Scorzonera hispanica L. subsp. Asteraceae P-S 1 1 2 hispanica Asteraceae Scorzonera laciniata L. subsp. laciniata P-S 1 1 1 1 4

Asteraceae Tragopogon dubius Scop. P-S 1 1 2

Asteraceae Xeranthemum annuum L. P 1 1 2

Boraginaceae Anchusa officinalis L. P-S 1 1 2

Boraginaceae Echium russicum J. F. Gmelin P 1 1 1 1 1 1 6

Boraginaceae Echium vulgare L. subsp. vulgare P-S 1 1 1 1 4

Boraginaceae Myosotis sparsiflora Mikan ex Pohl P 1 1

Boraginaceae Nonea erecta Bernh. subsp. erecta P 1 1 Alyssum turkestanicum Regel & Brassicaceae P 1 1 Schmalh. ex Regel Brassicaceae Berteroa incana (L.) DC. P-S 1 1

Brassicaceae Erysimum cuspidatum (Bieb.) DC. P 1 1

Brassicaceae Erysimum diffusum Ehrh. P 1 1 1 1 1 5

Brassicaceae Isatis praecox Kit. ex Tratt. P 1 1 1 3 Asyneuma canescens (Waldst. & Kit.) Campanulaceae P-S 1 1 Griseb. & Schenk subsp. canescens Campanulaceae Campanula bononiensis L. P 1 1

Caryophyllaceae Arenaria leptoclados (Reichenb.) Guss. P-S 1 1 2 Dianthus capitatus Balbis ex DC. subsp. Caryophyllaceae P 1 1 2 andrzejowskianus Balbis ex DC. Caryophyllaceae Herniaria incana Lam. subsp. incana M-S-P 1 1 1 3 Minuartia glomerata (Bieb.) Degen Caryophyllaceae P 1 1 2 subsp. glomerata Caryophyllaceae Silene otites (L.) Wibel subsp. otites P-S 1 1 1 3 Fumana procumbens (Dunal) Gren. & Cistaceae P-S 1 1 2 Godron subsp. procumbens Cyperaceae Carex caryophyllea Latourr. P-S 1 1 1 1 1 5

Cyperaceae Carex humilis Leysser P-S 1 1 1 1 1 5 Knautia arvensis (L.) Coulter subsp. Dipsacaceae P-S 1 1 1 3 arvensis Scabiosa ochroleuca L. subsp. Dipsacaceae P-S 1 1 1 1 1 5 ochroleuca Euphorbiaceae Euphorbia salicifolia Host P 1 1 2

Euphorbiaceae Euphorbia waldsteinii (Soják) A. R. Sm. P 1 1 2

Fabaceae Astragalus cicer L. P-S 1 1

Fabaceae Astragalus dasyanthus Palas P 1 1

ContinuedTable 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW – North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Gornji Milanovac), W − Western Serbia (Mt Tara, Mt Mokra Gora, Mt Zlatibor, Priboj, Uvac valley), C I − Central Serbia I (northern part of Ibar valley, Mt Stolovi, Mt Čemerno, Mt Goč, Mt Studena Planina), C II − Central Serbia II (central and southern part of Ibar valley, Mt Željin, Mt Kopaonik, Mt Rogozna), SW – South-western Serbia (Sjenica, Mt Ozren), Met − Metochia (Mt Koznik, Miruša, vicinity of Đakovica, Gubavac, Milanovac), S&C Kos – South & Central Kosovo (Mt Goleš, Mt Ostrovica, Mt Brezovica, Crni Vrh peak, Koznica), Freq − frequency of occurrence in defined serpentines groups, P-S- Pontic-Submediterranean, P-Pontic, M-S-P- Mediterranean- Submediterranean-Pontic. 264 K. Jakovljević et al.

Area S&C Family Taxon NW W C I C II SW Met Freq type Kos

Fabaceae Astragalus onobrychis L. P-S 1 1 1 1 1 1 6

Fabaceae Chamaecytisus austriacus (L.) Link P 1 1 1 3 Chamaecytisus heuffelii (Wierzb.) Fabaceae P 1 1 2 Rothm. Fabaceae Cytisus nigricans L. subsp. nigricans P-S 1 1 1 1 1 1 1 7 Cytisus procumbens (Waldst. & Kit. ex Fabaceae P 1 1 1 1 1 5 Willd.) Sprengel subsp. procumbens Fabaceae Dorycnium herbaceum Vill. M-S-P 1 1 1 1 1 1 6

Fabaceae Lathyrus nissolia L. subsp. nissolia M-S-P 1 1 2

Fabaceae Lathyrus pallescens (Bieb.) C. Koch P-S 1 1 Lathyrus pannonicus (Jacq.) Garcke Fabaceae P-S 1 1 1 1 4 subsp. pannonicus Fabaceae Medicago minima (L.) Bartal. P-S 1 1 2 Medicago prostrata Jacq. subsp. Fabaceae P-S 1 1 1 1 1 5 prostrata Onobrychis arenaria (Kit.) DC. subsp. Fabaceae P 1 1 arenaria Fabaceae Oxytropis pilosa (L.) DC. P 1 1

Fabaceae Trifolium alpestre L. P-S 1 1 1 1 1 1 1 7

Fabaceae Trifolium montanum L. subsp. montanum P-S 1 1 1 1 1 5 Trifolium pannonicum Jacq. subsp. Fabaceae P-S 1 1 1 1 1 5 pannonicum Fabaceae Vicia grandiflora Scop. P-S 1 1 2

Fabaceae Vicia sparsiflora Ten. P-S 1 1

Lamiaceae Ajuga laxmannii (L.) Bentham P-S 1 1 2

Lamiaceae Marrubium peregrinum L. P 1 1

Lamiaceae Salvia aethiopis L. P-S 1 1

Lamiaceae Salvia pratensis L. subsp. pratensis P-S 1 1 2

Lamiaceae Salvia sclarea L. M-S-P 1 1

Lamiaceae Salvia verticillata L. subsp. verticillata P-S 1 1 1 1 4

Lamiaceae Sideritis montana L. subsp. montana P-S 1 1 Stachys recta L. subsp. baldaccii (K. Lamiaceae P-S 1 1 1 1 1 1 6 Malý) Hayek Stachys recta L. subsp. subcrenata Lamiaceae P-S 1 1 (Vis.) Briq. Teucrium chamaedrys L. subsp. Lamiaceae P-S 1 1 1 3 chamaedrys Lamiaceae Thymus pannonicus All. P 1 1 2

Liliaceae Allium moschatum L. P-S 1 1 2

Liliaceae Allium oleraceum L. subsp. oleraceum P-S 1 1 Allium sphaerocephalon L. subsp. Liliaceae P-S 1 1 sphaerocephalon

Area S&C Family Taxon NW W C I C II SW Met Freq type Kos

Liliaceae Anthericum ramosum L. P-S 1 1 1 3

Liliaceae Asparagus officinalis L. subsp. officinalis P-S 1 1 Gagea pusilla (F. W. Schmidt) Schultes & Liliaceae P-S 1 1 Schultes fil. subsp. pusilla Liliaceae Ornithogalum kochii Parl. M-S-P 1 1

Liliaceae Ornithogalum refractum Kit. ex Schlecht. P-S 1 1

Linaceae Linum austriacum L. subsp. austriacum P-S 1 1

Linaceae Linum flavum L. subsp. flavum P-S 1 1 1 1 4

Linaceae Linum hirsutum L. subsp. hirsutum P 1 1 2

Linaceae Linum tenuifolium L. P-S 1 1 1 1 4

Malvaceae Althaea hirsuta L. P-S 1 1 Agropyron cristatum (L.) Gaertner Poaceae P 1 1 1 3 subsp. cristatum Poaceae Bothriochloa ischaemum (L.) Keng P-S 1 1 1 3 Brachypodium pinnatum (L.) Beauv. Poaceae P-S 1 1 1 1 4 subsp. pinnatum Bromus pannonicus Kummer & Poaceae P-S 1 1 1 1 4 Sendtner subsp. pannonicus Poaceae Chrysopogon gryllus (L.) Trin. P-S 1 1 1 1 4 Elymus hispidus (Opiz) Melderis subsp. Poaceae P-S 1 1 2 hispidus Poaceae Festuca rupicola Heuffel subsp. rupicola P 1 1 1 1 1 5

Poaceae Festuca stricta Host P 1 1 Festuca valesiaca Schleicher ex Gaudin Poaceae P-S 1 1 1 1 1 5 subsp. valesiaca Koeleria macrantha (Ledeb.) Schultes Poaceae P-S 1 1 1 1 1 1 6 subsp. macrantha Koeleria pyramidata (Lam.) Beauv. Poaceae P-S 1 1 1 1 1 5 subsp. pyramidata Melica ciliata L. subsp. nebrodensis Poaceae M-S-P 1 1 1 1 1 1 1 7 (Parl.) Husnot Poaceae Phleum phleoides (L.) Karsten P-S 1 1 1 1 4

Poaceae Poa bulbosa L. subsp. bulbosa P-S 1 1 2

Poaceae Stipa bromoides (L.) Dörfler P-S 1 1

Poaceae Stipa capillata L. P 1 1 1 3

Poaceae Stipa joannis Čelak. subsp. joannis P 1 1

Poaceae Stipa pennata L. subsp. pennata P-S 1 1 1 1 1 5

Poaceae Stipa pulcherrima C. Koch P-S 1 1 1 1 4

Poaceae Stipa tirsa Steven subsp. tirsa P 1 1

Polygalaceae Polygala major Jacq. subsp. major P-S 1 1 1 1 1 5

Ranunculaceae Clematis integrifolia L. P 1 1

Area S&C Family Taxon NW W C I C II SW Met Freq type Kos

Ranunculaceae Nigella arvensis L. subsp. arvensis M-S-P 1 1 2

Ranunculaceae Ranunculus polyanthemos L. P-S 1 1 1 1 1 5 Agrimonia eupatoria Ledeb. subsp. Rosaceae P-S 1 1 1 3 eupatoria Rosaceae Filipendula vulgaris Moench P-S 1 1 1 1 1 1 1 7

Rosaceae Potentilla alba L. P 1 1 1 1 4

Rosaceae Potentilla argentea L. P-S 1 1 1 1 1 5 Potentilla chrysantha Trev. subsp. Rosaceae P 1 1 1 3 amphibola (Schur) Soják Potentilla incana P. Gaertner, B. Meyer & Rosaceae P 1 1 1 1 1 5 Scherb. Rosaceae Potentilla patula Waldst. & Kit. P 1 1

Rosaceae Prunus mahaleb L. P-S 1 1 1 1 4

Rosaceae Rosa spinosissima L. P-S 1 1 1 1 1 5 Sanguisorba minor Scop. subsp. Rosaceae balearica (Bourgeau ex Nyman) Munoz P-S 1 1 1 1 1 1 6 Garmendia & C. Navarro Rosaceae Sanguisorba minor Scop. subsp. minor M-S-P 1 1 1 1 1 1 1 7

Rubiaceae Asperula cynanchica L. P-S 1 1 1 1 1 1 6 Cruciata pedemontana (Bellardii) Rubiaceae P-S 1 1 2 Ehrend. Rubiaceae Galium glaucum L. subsp. glaucum P-S 1 1 1 3

Rubiaceae Galium verum L. subsp. verum P-S 1 1 1 1 1 5

Rutaceae Dictamnus albus L. P-S 1 1 1 1 4 Comandra elegans (Rochel ex Santalaceae P 1 1 Reichenb.) Reichenb. fil. Santalaceae Thesium arvense Horvátovszky P-S 1 1 1 3

Scrophulariaceae Euphrasia tatarica Fischer ex Sprengel P 1 1 1 3

Scrophulariaceae Melampyrum arvense L. P-S 1 1 1 3

Scrophulariaceae Melampyrum cristatum L. P 1 1 1 1 4

Scrophulariaceae Rhinanthus rumelicus Velen. P 1 1 1 1 4

Scrophulariaceae Verbascum banaticum Schrader P-S 1 1 2

Scrophulariaceae Verbascum lychnitis L. subsp. lychnitis P-S 1 1 1 3

Scrophulariaceae Verbascum nigrum L. P 1 1 2

Scrophulariaceae Verbascum phoeniceum L. P 1 1 1 3

Scrophulariaceae Veronica austriaca L. P 1 1 1 1 1 5 Veronica chamaedrys L. subsp. Scrophulariaceae P-S 1 1 1 1 1 1 6 chamaedrys Scrophulariaceae Veronica jacquinii Baumg. P 1 1 1 1 1 1 1 7

Scrophulariaceae Veronica praecox All. subsp. praecox P-S 1 1 2

Scrophulariaceae Veronica spicata L. subsp. spicata P 1 1 1 1 4

71 76 76 148 31 38 43

267 Richness and diversity of Pontic flora on serpentine of Serbia

A B C D

Figure 2. Distribution of species on serpentine areas of Serbia shown on a UTM 10 x 10 km grid. (A) Mediterranean-Submediterranean-Pontic area type; (B) Pontic-Submediterranean area type; (C) Pontic area type; (D) Summary map of the distribution of all study species.

The greatest floristic richness was recorded in groups C Serbia II (148 taxa), W Serbia (76) and C Serbia I (76), while the lowest numbers were recorded in groups Metochia province (38) and SW Serbia (31) (Table 1). Therefore, most recorded species belong to the P-S areal-type (99 taxa), followed by P (48), while M-S-P species were the least prominent in the area studied (14). The analysis of distribution of taxa belonging to particular areal types is presented in Figure 2A-C. High numbers of M-S-P species were recorded in south and central parts of the Ibar valley (UTM squares DN78, DN69), Mt Kopaonik (DN79, DN97), Mt Stolovi (DP62) and parts of Mt Šar Planina foothill (DM97). The center

of diversity of species of P-S areal-type was in the S&C Figure 3. The linear regression for dependency of variation of part of Ibar valley (DN78, DN85). Analysis of diversity number of species with regard to the frequency of of species belonging to P areal-type has, in addition occurrence in all defined serpentine groups. to Mt Kopaonik (DN79), another center of diversity at Mt Stolovi (DP72), although the region of Ibar valley Alyssum turkestanicum (Brassicaceae), Astragalus (DN69, DN86) is also characterized by a high number dasyanthus, Onobrychis arenaria, Oxytropis pilosa, of species. (Fabaceae), Nonea erecta (Boraginaceae), Seseli The summary of the distributions of P, P-S and annum (Apiaceae), and Stipa joanis aggr. (Poaceae) M-S-P flora on serpentine in Serbia (Figure 2D) shows (Table 1). Only 7 taxa (4.3%) were represented in all that the centers of diversity are situated in the S&C serpentine groups: Cytisus nigricans subsp. nigricans, part of Ibar valley (UTM squares DN85, DN86, DN78, Trifolium alpestre (Fabaceae), Filipendula vulgaris, DN87 and on Mt Kopaonik - DN88, DN79, DN97, DN89, Sanguisorba minor subsp. minor (Rosaceae), Melica DN98). Apart from these two broad areas, additional ciliata subsp. nebrodensis (Poaceae), Trinia glauca centers of diversity were found in parts of Mt Goč subsp. glauca (Apiaceae) and Veronica jacquinii (DP82), Mt Stolovi (DP62 and northern part of DP72), (Scrophulariaceae), while eight taxa (5%) were recorded Mt Studena Planina - southern part of DP72 and Mt within six geographic groups. This shows that less than Zlatibor - CP93, CP94 and CP82. 10% of the total analyzed flora consists of the highly Most taxa were recorded in only one (43 taxa or frequent taxa. Some of the other common species in the 26.7%) or two serpentine groups (33 taxa or 20.5%). area studied were Astragalus onobrychis (Fabaceae), Therefore almost 50% of investigated plants occur Echium russicum (Boraginaceae), Koeleria macrantha infrequently on serpentine substrate in Serbia. Among subsp. macrantha (Poaceae) and Veronica chamaedrys the rarest Pontic plants in the area studied were (Scrophulariaceae).

268 K. Jakovljević et al.

Variation in number of species with diverse recording Species number varied significantly with elevation frequencies has shown a pronounced negative trend (R2=0.814, F(2,6)=13.150, P=0.006) (Figure 5). Most described through the high factor of linear regression species were recorded at elevations between 750-1000 m R2=0.904, F(1,6)=56.817, P=0.000 (Figure 3). This (101 taxa), 500-750 m (90 taxa) and 250-500 m result supports the results from the distribution analysis, (73 taxa). The smallest number of species was recorded which showed that the P, P-S and M-S-P flora of the both at the highest and lowest altitudes of the area serpentine of Serbia are composed mostly of rare plants. studied. Between 2000 and 2250 m only one taxon Floristic richness is strongly influenced by was recorded, between 1750 and 2000 m there were precipitation in the driest month (R2=0.656, 14 taxa, while at 250 m and below there were 40 taxa. F(2,7)=6.659, P=0.024), annual total precipitation Total flora on the serpentinian subregions clusters into (R2=0.646, F(2,7)=6.393, P=0.026), centroids of N three units (Figure 6). The first unit includes the groups latitude (R2=0.897, F(1,7)=60.971, P=0.000) and north-western Serbia, western Serbia, central Serbia II E longitude (R2=0.800, F(2,5)=10.009, P=0.018). and central Serbia I. The second unit includes south- Proximity to steppe regions of Pannonian plain, western Serbia and south and central Kosovo, while the Danube area and Thrace, as well as the diversity of third isolated unit is the group of Metochia. The first unit potential vegetation, were not significant predictors of (NW Serbia – W Serbia – C Serbia II – C Serbia I) is floristic richness of individual serpentine groups (not the most compact, most complex and richest in floristic shown). While the changes in number of species at the diversity. Almost all taxa of the whole area studied (98%) latitudinal gradients was described by linear types of were recorded in this unit. 56.5% (91 taxa) were recorded regression, the changes in longitudinal gradient and exclusively in this unit. The second unit (SW Serbia – humidity gradients were described by the polynomial S & C Kosovo) had 55 recorded taxa (34.16%) and is type of regression (Figure 4). therefore much poorer, while the smallest number (38)

Figure 4. The linear regression for dependency of variation of number of species in regard to different geographical and bioclimatic factors (only for the largest serpentine groups: north-western Serbia, western Serbia, central Serbia I, central Serbia II). A-dependency of N-latitude, B-dependency of E-longitude, C- dependency of precipitation of driest month, D-dependency of annual precipitation.

269 Richness and diversity of Pontic flora on serpentine of Serbia

Figure 5. Species richness at different altitudes, showing their preferences for certain altitude ranges.

Figure 6. Unweighted pair-group average of Jaccard distances between Pontic taxa sensu lato on serpentine groups of Serbia.

of taxa was recorded at Metochia province (third unit). mountainous areas of Serbia, it is relatively distant Only three unique taxa (Achillea nobilis subsp. nobilis from the real steppe areas of Pannonian plain, Danube (Asteraceae), Clematis integrifolia (Ranunculaceae), area and Thrace. Such a high percentage of the Stipa tirsa subsp. tirsa (Poaceae)) were found in Metochia analyzed flora, particularly typical steppe relicts such that did not occur within the first group. as Astragalus dasyanthus, Chamaecytisus heuffelii, Oxythropis pilosa (Fabaceae), Peucedanum officinale (Apiaceae), Vinca herbacea (Apocynaceae), Dictamnus 4. Discussion albus (Rutaceae), Comandra elegans (Santalaceae), Linum hirsutum (Linaceae) and Echium russicum The floristic studies on the serpentine areas of Serbia (Boraginaceae) could indicate the importance of have shown the presence of 161 P, P-S and M-S-P taxa serpentine areas in Serbia as refugia for xerothermic at species and subspecies level (Table 1), representing flora in the Balkan peninsula. In addition the number almost 40% of the total number of recorded Pontic of paleoendemics belonging to the group of obligate plants in Serbia [56]. As the area studied is situated serpentinophytes (e.g. Halacsya sendtneri, Paramoltkia in the western, southwestern and central, mostly doerfleri (Boraginaceae), Gypsophila spergulifolia,

270 K. Jakovljević et al.

Saponaria intermedia (Caryophyllaceae), Drymocallis the serpentine areas of Serbia are represented with malacophila, Potentilla visianii, Sanquisorba albanica 161 taxa on species and subspecies level. (Rosaceae), Haplophyllum boisserianum (Rutaceae), The centers of diversity of Pontic, Pontic- Fumana bonapartei (Cistaceae), Euphorbia glabriflora Submediterranean and Mediterranean- (Euphorbiaceae), Bornmuellera dieckii (Brassicaceae), Submediterranean-Pontic flora on serpentine of Eryngium serbicum (Apiaceae)) confirm serpentine areas Serbia, both as a whole and of particular areal-types, of the central Balkans as an important refugium and are situated in Ibar valley and southern slopes of Mt possibly a center of speciation [1]. Kopaonik. The analysis of dependence of number of species Latitude, longitude, altitude, the precipitation level of on main geographic, ecological and bioclimatic factors the driest month and the annual precipitation also show has shown that the factors such as latitude and a pronounced impact on species richness. longitude, altitude, the precipitation rate in the driest Annual mean temperature, distance from the steppe month, and the annual precipitation rate, have a high regions of southeastern Europe and the diversity of impact on species richness of the study region (Figure potential vegetation have shown a very low impact on 4, 5). In addition, the highest number of Pontic species floristic richness. is recorded at the driest month precipitation of the value The regression types at the latitudinal and 58 mm and at the annual precipitation of 900 mm. The longitudinal gradient show that the recent distribution of factors related to distance from the steppe regions Pontic taxa sensu lato in the area studied was greatly of southeastern Europe and diversity of potential influenced by historical factors. vegetation did not impact floristic richness. An important Serpentine areas in W&C Serbia, together with result of these analyses is the fact that the changes in those in N Albania and N Macedonia represent the number of species at both the latitudinal and the important refugia for numerous plants of different longitudinal gradient show opposite trends. Along the age and chorology. Occurrence of endemic plants, latitudinal gradient (temperature gradient) there was a particularly relict ones, strongly support this fact. statistically important increase in number of species. In In addition, certain number of widespread Pontic, contrast, the longitudinal gradient (gradient of humidity) Pontic-Submediterranean and Mediterranean- shows a significant decrease in number of species Submediterranean-Pontic species is distributed far until the 20° longitude and then there is a marked from their main range of distribution in serpentine increase. As the closest steppe areas of Pannonian and areas of Serbia and neighboring countries. Such taxa Wallachian Plains are more toward north and east than could be also considered as relicts of former steppe the serpentine areas of western Serbia, it was expected flora widespread in the Balkans during Pleistocene. that the greatest values of richness would be recorded The importance of this study lies in the fact that it in serpentine groups close to these areas. This result summarizes, for the first time, a detailed distribution indicates that historical factors play an important role in data of the Pontic flora in serpentine areas in the the distribution of Pontic species in the study area. These territory of Serbia. factors may be more important than other ecological and biogeographic variables, like the deforestation that has created numerous open habitats that favour xerophyte Acknowledgements colonization. The Ministry of Science and Technological Development 5. Conclusions of the Republic of Serbia, Grant 143015: “Diversity of flora and vegetation of the central part of Balkan Pontic, Pontic-Submediterranean and Mediterranean- peninsula - Ecology, Chorology and Conservation”, Submediterranean-Pontic types of distribution of supported this research.

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