Richness and diversity of Pontic flora on serpentine of Serbia

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1 Cent. Eur. J. Biol. 6(2) DOI: /s 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 Received 23 June 2010; Accepted 08 November 2010 Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade, Belgrade, Serbia 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. 1. Introduction Serpentine (ophiolithic, ultramafic) rock belongs to the group of siliceous rocks characterized by calcium deficiency, high concentrations of iron, magnesium, nickel, cobalt and chromium, and reduced nutrients. In contrast to other acid siliceous rocks, the ph values of serpentine substrate vary from basic to ultrabasic (ph 5.5-8). Many interesting basiphilous-calcifuge plants flourish on serpentine owing to this factor. It is one of the most important reasons why the serpentine flora, differing from that occurring on limestone or other siliceous substrates, presents taxonomical, phytogeographical and ecological phenomenon of great interest wherever it occurs [1]. The largest serpentine areas in Europe are in the Balkan peninsula (Epirus, Thessaly and Sterea Ellas in Greece, north, north-central and south-eastern Albania, central and east Bosnia, western, south-western and central Serbia, east and central Rhodopes mountains in Bulgaria and gorge of river Pcinja and south-eastern slopes of Sara mountain in north-eastern part of Republic of Macedonia) (Figure 1A). In Serbia there are several serpentine zones located primarily in central, western and south-western Serbia and in Kosovo. Some of these areas are: Brdjanska gorge, Mt Suvobor, Mt Maljen, Mt Tara, Mt Zlatibor, Mt Mokra Gora, Mt Goč, Mt Stolovi, Mt Kopaonik, the Ibar valley, Peć and surroundings, Orahovac and Koznik, Mt Ostrovica (Figure 1B). The real steppes of southeastern Europe are present in the Pannonian and Wallachian Plain, Dobruja and Thracean Plain [2,3]. On the Balkan peninsula, particularly on the skeleton soils of hills and on lower elevation mountain slopes, there are some xerophilous and steppe-like vegetation. The steppe-like vegetation is particularly well-developed in limestone and serpentine habitats of the Moesian subregion in Serbia, Bulgaria and Macedonia [4-9]. The degradation of forests and shrub formations in the temperate-continental (central and eastern) parts of the Balkan Peninsula, has only increased the extent of the steppe biome. These xerophilous grasslands include mostly Pontic, Pontic-Submediterranean and/or Mediterranean- Submediterranean-Pontic plants, but they also contain a substantial number of endemic species. Thus, they represent important centers of xerothermic flora that 260 * ksenija_jakovljevic@yahoo.com

2 K. Jakovljević et al. A C B 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 in the last glacial and postglacial period [10,11]. The occurrence of the Pontic, Pontic- Submediterranean and Mediterranean- Submediterranean-Pontic plants on serpentine is intriguing from a biogeographic standpoint because their populations in serpentine areas of the Balkans are isolated with respect to their common distribution in the steppe regions of south-eastern Europe. The open serpentine habitats are characterized by a pronounced thermophilous character and xeric conditions. The xerothermic character of serpentine plants is also enhanced by the specific chemical composition of the serpentine substrate. Among the numerous xerophytes on the serpentine of the Balkans, the species of the Pontic, Pontic-Submediterranean and Mediterranean- Submediterranean-Pontic areal-types are most prominent. The steppe relicts with Pontic distribution (for example, Astragalus dasyanthus, Oxytropis pilosa (Fabaceae), Vinca herbacea (Apocynaceae), Echium russicum (Boraginaceae), Galatella lynosiris (), Peucedanum officinale (Apiaceae), etc.), growing on serpentinite in the central part of Peninsula, have a special phytogeographic importance. The spread of Pontic flora throughout the central and east Balkans was particularly pronounced during Ice Age (Pleistocene) [2]. Serpentine areas were suitable habitats for these plants during this period, so steppe relicts could represent current remnants of this flora. The main goal of this paper is to determine the extent to which the serpentine areas of Serbia represent an important region of south-eastern Europe for the xerothermic flora of the Pontic, Pontic-Submediterranean and Mediterranean-Submediterranean-Pontic species. In south-eastern Europe serpentine areas are quite rare and unique, especially those serpentine areas that are inhabited by xerothermic flora. Therefore, these unique areas are of the great importance in phytogeographical sense. The objectives of our research were to determine the richness and distribution of species belonging to Pontic, Pontic-Submediterranean and Mediterranean- Submediterranean-Pontic species on the serpentine of Serbia and how that varies with respect to geographic, ecological and climatic factors. The degree of floristic similarity among serpentine groups will be determined, and the most important centers of diversity of Pontic flora on serpentine soils will be broadly determined. 261

3 Richness and diversity of Pontic flora on serpentine of Serbia 2. Experimental Procedures The mapping and analyses of the distribution of Pontic species on serpentines in Serbia was based on herbarium specimens from the Herbarium of University of Belgrade (BEOU) and the Herbarium of Natural History Museum of Belgrade (BEO), published data [9,12-39] as well as on personal field observations. The nomenclature is consistent with the latest checklists, papers, monographs and databases [40-45; rbgsyd.nsw.gov.au/iopi/iopihome.htm]. The serpentine areas in Serbia were divided into the following groups according to their orographic connections: 1. north-western Serbia (including the areas of Mt Maljen, Mt Suvobor, vicinity of Gornji Milanovac); 2. western Serbia (including the areas of Mt Tara, Mt Mokra Gora, Mt Zlatibor, Priboj, and the Uvac valley); 3. central Serbia I (including the northern part of Ibar valley, with areas of Mt Stolovi, Mt Čemerno, Mt Goč, Mt Studena Planina); 4. central Serbia II (including the central and southern part of Ibar valley, with areas of Mt Željin, Mt Kopaonik, Mt Rogozna); 5. south-western Serbia (including the areas of Sjenica and Mt Ozren); 6. Metochia (Koznik, Miruša, vicinity of Đakovica, Gubavac, Milanovac); and 7. south and central Kosovo (Mt Goleš, Mt Ostrovica, Mt Brezovica, Crni Vrh peak, Koznica) (Figure 1C). In order to define areal-types [46-48] we used the concept of floristic elements made by Meusel et al. [47,49-51], Meusel, Jäger [52,53], modified for the Serbian territory by Stevanovic [54]. The species examined were grouped into the following areal-types according to their general type of distribution: 1. Pontic areal-type (P), including the species distributed in the steppe areas of Europe, reaching eastward to the Central Asian steppes 2. Pontic-Submediterranean areal-type (P-S). In addition to the Pontic region which is the main part of their range, they are also present in smaller or larger parts of the Mediterranean region, particularly in the Submediterranean subregion 3. Mediterranean-Submediterranean-Pontic arealtype (M-S-P), where the main parts of the range are situated within the Mediterranean basin, reaching to the Pontic region of southeastern and parts of eastern Europe. For each serpentine group floristic richness (as total number of species on certain groups) was calculated based on literature, herbarium and personal field data. A regression analysis was performed in order to identify the dependency level of variation in species richness, with regard to the basic geographical, ecological and bioclimatic characteristics of defined serpentine groups. The regression analysis included only the groups northwestern Serbia, western Serbia, central Serbia I and central Serbia II, which are approximately of the same size. Each serpentine group was regressed onto the following factors: area, E-long centroid, N-lat centroid, altitude, annual mean temperature, annual precipitation, precipitation of driest month, precipitation of driest quarter, diversity of potential vegetation, average distance from the steppe regions and minimal distance from the steppe regions. The bioclimatic characteristics of the groups were extracted from the WorldClim set of global climate layers [55]. The extraction of selected bioclimatic parameters was done by using DIVA-GIS 5.2 software (Version 5.2; The diversity of the vegetation that would exist at defined serpentine groups under the current climatic and geomorphologic conditions but without any human impact (potential vegetation) was established on the basis of the Map of the Natural Vegetation of Europe [4]. The diversity of potential vegetation is presented as number of different mapping units per serpentine group. Floristic similarities between each set of analyzed groups were analyzed using the clustering method based on Jaccard distances and unweighted pair-group average hierarchical sorting strategy (UPGMA). All statistical analyses were performed with the Statistica software package (version 5.1; StatSoft, 1997). The analysis of diversity and the summary distribution of Pontic species were presented on a 10 km x 10 km grid using the Universal Transverse Mercator (UTM) projection (2001, All analyses related to mapping and spatial calculation were performed by using Manifold System software (version 5.50, Manifold System, CDA International Ltd.). 3. Results According to the results of floristic studies on the serpentine areas of Serbia, there were 161 taxa at the species and subspecies level, belonging to Pontic (P), Pontic-Submediterranean (P-S) and Mediterranean- Submediterranean-Pontic (M-S-P) areal-types. Only two Pontic taxa - Bromus pannonicus subsp. pannonicus () and Stachys recta subsp. baldaccii (Lamiaceae) - belong to the group of obligate serpentinophytes on serpentine areas in Serbia, while the other plants may also be found on other types of geological substrate (limestone, silicate, etc.). 262

4 K. Jakovljević et al. Family Taxon Area type NW W C I C II SW Met S&C Kos Freq Anacardiaceae Cotinus coggygria Scop. subsp. coggygria P-S Apiaceae Eryngium campestre L. P-S Apiaceae Falcaria vulgaris Bernh. P-S 1 1 Apiaceae Ferulago campestris (Besser) Grec. P-S Apiaceae Ferulago sylvatica (Besser) Reichenb. subsp. sylvatica P-S Apiaceae Orlaya grandiflora (L.) Hoffm. M-S-P Apiaceae Peucedanum officinale L. subsp. officinale M-S-P Apiaceae Seseli annuum L. subsp. annuum P 1 1 Apiaceae Trinia glauca (L.) Dumort. subsp. glauca M-S-P Apocynaceae Vinca herbacea Waldst. & Kit. P-S 1 1 Asclepiadaceae Vincetoxicum hirundinaria Medicus subsp. hirundinaria P-S Achillea aspleniifolia Vent. P 1 1 Achillea coarctata Poiret P-S 1 1 Achillea nobilis L. subsp. nobilis P-S 1 1 Carduus candicans Waldst. & Kit. subsp. candicans Carduus hamulosus Ehrh. subsp. hamulosus P-S P Carthamus lanatus L. subsp. lanatus M-S-P 1 1 Centaurea stoebe L. subsp. australis (A. Kerner) Greuter P Centaurea stoebe L. subsp. stoebe P Cota austriaca (Jacq.) Schultz-Bip. P-S 1 1 Cota tinctoria (L.) J. Gay subsp. tinctoria P-S Crupina vulgaris Cass. P-S Echinops ritro L. subsp. ruthenicus (Bieb.) Nyman Galatella linosyris (L.) Bernh. subsp. linosyris P-S 1 1 P-S Inula britannica L. subsp. britannica P-S Inula ensifolia L. P-S Inula hirta L. P Inula oculus-christi L. P-S Jurinea mollis (L.) Reichenb. subsp. mollis Lactuca viminea (L.) J. & C. Presl subsp. viminea P-S M-S-P 1 1 Leontodon crispus Vill. subsp. crispus M-S-P Picris hieracioides L. subsp. hieracioides P-S 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

5 Richness and diversity of Pontic flora on serpentine of Serbia Family Taxon Area type NW W C I C II SW Met S&C Kos Freq Pilosella bauhinii (Schultes) Arv.-Touv. subsp. bauhinii Scorzonera austriaca Willd. subsp. austriaca Scorzonera hispanica L. subsp. hispanica P-S P-S P-S Scorzonera laciniata L. subsp. laciniata P-S Tragopogon dubius Scop. P-S Xeranthemum annuum L. P Boraginaceae Anchusa officinalis L. P-S Boraginaceae Echium russicum J. F. Gmelin P Boraginaceae Echium vulgare L. subsp. vulgare P-S Boraginaceae Myosotis sparsiflora Mikan ex Pohl P 1 1 Boraginaceae Nonea erecta Bernh. subsp. erecta P 1 1 Brassicaceae Alyssum turkestanicum Regel & Schmalh. ex Regel P 1 1 Brassicaceae Berteroa incana (L.) DC. P-S 1 1 Brassicaceae Erysimum cuspidatum (Bieb.) DC. P 1 1 Brassicaceae Erysimum diffusum Ehrh. P Brassicaceae Isatis praecox Kit. ex Tratt. P Campanulaceae Asyneuma canescens (Waldst. & Kit.) Griseb. & Schenk subsp. canescens P-S 1 1 Campanulaceae Campanula bononiensis L. P 1 1 Caryophyllaceae Arenaria leptoclados (Reichenb.) Guss. P-S Caryophyllaceae Dianthus capitatus Balbis ex DC. subsp. andrzejowskianus Balbis ex DC. P Caryophyllaceae Herniaria incana Lam. subsp. incana M-S-P Caryophyllaceae Minuartia glomerata (Bieb.) Degen subsp. glomerata P Caryophyllaceae Silene otites (L.) Wibel subsp. otites P-S Cistaceae Fumana procumbens (Dunal) Gren. & Godron subsp. procumbens P-S Cyperaceae Carex caryophyllea Latourr. P-S Cyperaceae Carex humilis Leysser P-S Dipsacaceae Dipsacaceae Knautia arvensis (L.) Coulter subsp. arvensis Scabiosa ochroleuca L. subsp. ochroleuca P-S P-S Euphorbiaceae Euphorbia salicifolia Host P Euphorbiaceae Euphorbia waldsteinii (Soják) A. R. Sm. P Fabaceae Astragalus cicer L. P-S 1 1 Fabaceae Astragalus dasyanthus Palas P Table 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Continued 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.

6 K. Jakovljević et al. Family Taxon Area type NW W C I C II SW Met S&C Kos Freq Fabaceae Astragalus onobrychis L. P-S Fabaceae Chamaecytisus austriacus (L.) Link P Fabaceae Chamaecytisus heuffelii (Wierzb.) Rothm. P Fabaceae Cytisus nigricans L. subsp. nigricans P-S Fabaceae Cytisus procumbens (Waldst. & Kit. ex Willd.) Sprengel subsp. procumbens P Fabaceae Dorycnium herbaceum Vill. M-S-P Fabaceae Lathyrus nissolia L. subsp. nissolia M-S-P Fabaceae Lathyrus pallescens (Bieb.) C. Koch P-S 1 1 Fabaceae Lathyrus pannonicus (Jacq.) Garcke subsp. pannonicus P-S Fabaceae Medicago minima (L.) Bartal. P-S Fabaceae Fabaceae Medicago prostrata Jacq. subsp. prostrata Onobrychis arenaria (Kit.) DC. subsp. arenaria P-S P 1 1 Fabaceae Oxytropis pilosa (L.) DC. P 1 1 Fabaceae Trifolium alpestre L. P-S Fabaceae Trifolium montanum L. subsp. montanum P-S Fabaceae Trifolium pannonicum Jacq. subsp. pannonicum P-S Fabaceae Vicia grandiflora Scop. P-S Fabaceae Vicia sparsiflora Ten. P-S 1 1 Lamiaceae Ajuga laxmannii (L.) Bentham P-S Lamiaceae Marrubium peregrinum L. P 1 1 Lamiaceae Salvia aethiopis L. P-S 1 1 Lamiaceae Salvia pratensis L. subsp. pratensis P-S Lamiaceae Salvia sclarea L. M-S-P 1 1 Lamiaceae Salvia verticillata L. subsp. verticillata P-S Lamiaceae Sideritis montana L. subsp. montana P-S 1 1 Lamiaceae Lamiaceae Lamiaceae Stachys recta L. subsp. baldaccii (K. Malý) Hayek Stachys recta L. subsp. subcrenata (Vis.) Briq. Teucrium chamaedrys L. subsp. chamaedrys P-S P-S 1 1 P-S Lamiaceae Thymus pannonicus All. P Liliaceae Allium moschatum L. P-S Liliaceae Allium oleraceum L. subsp. oleraceum P-S 1 1 Liliaceae Allium sphaerocephalon L. subsp. sphaerocephalon P-S 1 1 Table 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Continued 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. 265

7 Richness and diversity of Pontic flora on serpentine of Serbia Family Taxon Area type NW W C I C II SW Met S&C Kos Freq Liliaceae Anthericum ramosum L. P-S Liliaceae Asparagus officinalis L. subsp. officinalis P-S 1 1 Liliaceae Gagea pusilla (F. W. Schmidt) Schultes & Schultes fil. subsp. pusilla P-S 1 1 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 Linaceae Linum hirsutum L. subsp. hirsutum P Linaceae Linum tenuifolium L. P-S Malvaceae Althaea hirsuta L. P-S 1 1 Agropyron cristatum (L.) Gaertner subsp. cristatum P Bothriochloa ischaemum (L.) Keng P-S Brachypodium pinnatum (L.) Beauv. subsp. pinnatum Bromus pannonicus Kummer & Sendtner subsp. pannonicus P-S P-S Chrysopogon gryllus (L.) Trin. P-S Elymus hispidus (Opiz) Melderis subsp. hispidus P-S Festuca rupicola Heuffel subsp. rupicola P Festuca stricta Host P 1 1 Festuca valesiaca Schleicher ex Gaudin subsp. valesiaca Koeleria macrantha (Ledeb.) Schultes subsp. macrantha Koeleria pyramidata (Lam.) Beauv. subsp. pyramidata Melica ciliata L. subsp. nebrodensis (Parl.) Husnot P-S P-S P-S M-S-P Phleum phleoides (L.) Karsten P-S Poa bulbosa L. subsp. bulbosa P-S Stipa bromoides (L.) Dörfler P-S 1 1 Stipa capillata L. P Stipa joannis Čelak. subsp. joannis P 1 1 Stipa pennata L. subsp. pennata P-S Stipa pulcherrima C. Koch P-S Stipa tirsa Steven subsp. tirsa P 1 1 Polygalaceae Polygala major Jacq. subsp. major P-S Ranunculaceae Clematis integrifolia L. P Table 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Continued 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.

8 K. Jakovljević et al. Family Taxon Area type NW W C I C II SW Met S&C Kos Freq Ranunculaceae Nigella arvensis L. subsp. arvensis M-S-P Ranunculaceae Ranunculus polyanthemos L. P-S Rosaceae Agrimonia eupatoria Ledeb. subsp. eupatoria P-S Rosaceae Filipendula vulgaris Moench P-S Rosaceae Potentilla alba L. P Rosaceae Potentilla argentea L. P-S Rosaceae Rosaceae Potentilla chrysantha Trev. subsp. amphibola (Schur) Soják Potentilla incana P. Gaertner, B. Meyer & Scherb. P P Rosaceae Potentilla patula Waldst. & Kit. P 1 1 Rosaceae Prunus mahaleb L. P-S Rosaceae Rosa spinosissima L. P-S Sanguisorba minor Scop. subsp. Rosaceae balearica (Bourgeau ex Nyman) Munoz P-S Garmendia & C. Navarro Rosaceae Sanguisorba minor Scop. subsp. minor M-S-P Rubiaceae Asperula cynanchica L. P-S Rubiaceae Cruciata pedemontana (Bellardii) Ehrend. P-S Rubiaceae Galium glaucum L. subsp. glaucum P-S Rubiaceae Galium verum L. subsp. verum P-S Rutaceae Dictamnus albus L. P-S Santalaceae Comandra elegans (Rochel ex Reichenb.) Reichenb. fil. P 1 1 Santalaceae Thesium arvense Horvátovszky P-S Scrophulariaceae Euphrasia tatarica Fischer ex Sprengel P Scrophulariaceae Melampyrum arvense L. P-S Scrophulariaceae Melampyrum cristatum L. P Scrophulariaceae Rhinanthus rumelicus Velen. P Scrophulariaceae Verbascum banaticum Schrader P-S Scrophulariaceae Verbascum lychnitis L. subsp. lychnitis P-S Scrophulariaceae Verbascum nigrum L. P Scrophulariaceae Verbascum phoeniceum L. P Scrophulariaceae Veronica austriaca L. P Scrophulariaceae Veronica chamaedrys L. subsp. chamaedrys P-S Scrophulariaceae Veronica jacquinii Baumg. P Scrophulariaceae Veronica praecox All. subsp. praecox P-S Scrophulariaceae Veronica spicata L. subsp. spicata P Table 1. Distribution of Pontic taxa sensu lato on serpentine groups of Serbia (NW North-western Serbia (Mt Maljen, Mt Suvobor, the vicinity of Continued 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. 267

9 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 part of Ibar valley (DN78, DN85). Analysis of diversity of species belonging to P areal-type has, in addition to Mt Kopaonik (DN79), another center of diversity at Mt Stolovi (DP72), although the region of Ibar valley (DN69, DN86) is also characterized by a high number of species. The summary of the distributions of P, P-S and M-S-P flora on serpentine in Serbia (Figure 2D) shows that the centers of diversity are situated in the S&C part of Ibar valley (UTM squares DN85, DN86, DN78, DN87 and on Mt Kopaonik - DN88, DN79, DN97, DN89, DN98). Apart from these two broad areas, additional centers of diversity were found in parts of Mt Goč (DP82), Mt Stolovi (DP62 and northern part of DP72), Mt Studena Planina - southern part of DP72 and Mt Zlatibor - CP93, CP94 and CP82. Most taxa were recorded in only one (43 taxa or 26.7%) or two serpentine groups (33 taxa or 20.5%). Therefore almost 50% of investigated plants occur infrequently on serpentine substrate in Serbia. Among the rarest Pontic plants in the area studied were Figure 3. The linear regression for dependency of variation of number of species with regard to the frequency of occurrence in all defined serpentine groups. Alyssum turkestanicum (Brassicaceae), Astragalus dasyanthus, Onobrychis arenaria, Oxytropis pilosa, (Fabaceae), Nonea erecta (Boraginaceae), Seseli annum (Apiaceae), and Stipa joanis aggr. () (Table 1). Only 7 taxa (4.3%) were represented in all serpentine groups: Cytisus nigricans subsp. nigricans, Trifolium alpestre (Fabaceae), Filipendula vulgaris, Sanguisorba minor subsp. minor (Rosaceae), Melica ciliata subsp. nebrodensis (), Trinia glauca subsp. glauca (Apiaceae) and Veronica jacquinii (Scrophulariaceae), while eight taxa (5%) were recorded within six geographic groups. This shows that less than 10% of the total analyzed flora consists of the highly frequent taxa. Some of the other common species in the area studied were Astragalus onobrychis (Fabaceae), Echium russicum (Boraginaceae), Koeleria macrantha subsp. macrantha () and Veronica chamaedrys (Scrophulariaceae). 268

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

11 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). Only three unique taxa (Achillea nobilis subsp. nobilis (), Clematis integrifolia (Ranunculaceae), Stipa tirsa subsp. tirsa ()) were found in Metochia that did not occur within the first group. 4. Discussion The floristic studies on the serpentine areas of Serbia have shown the presence of 161 P, P-S and M-S-P taxa at species and subspecies level (Table 1), representing almost 40% of the total number of recorded Pontic plants in Serbia [56]. As the area studied is situated in the western, southwestern and central, mostly mountainous areas of Serbia, it is relatively distant from the real steppe areas of Pannonian plain, Danube area and Thrace. Such a high percentage of the analyzed flora, particularly typical steppe relicts such as Astragalus dasyanthus, Chamaecytisus heuffelii, Oxythropis pilosa (Fabaceae), Peucedanum officinale (Apiaceae), Vinca herbacea (Apocynaceae), Dictamnus albus (Rutaceae), Comandra elegans (Santalaceae), Linum hirsutum (Linaceae) and Echium russicum (Boraginaceae) could indicate the importance of serpentine areas in Serbia as refugia for xerothermic flora in the Balkan peninsula. In addition the number of paleoendemics belonging to the group of obligate serpentinophytes (e.g. Halacsya sendtneri, Paramoltkia doerfleri (Boraginaceae), Gypsophila spergulifolia, 270

12 K. Jakovljević et al. Saponaria intermedia (Caryophyllaceae), Drymocallis malacophila, Potentilla visianii, Sanquisorba albanica (Rosaceae), Haplophyllum boisserianum (Rutaceae), Fumana bonapartei (Cistaceae), Euphorbia glabriflora (Euphorbiaceae), Bornmuellera dieckii (Brassicaceae), Eryngium serbicum (Apiaceae)) confirm serpentine areas of the central Balkans as an important refugium and possibly a center of speciation [1]. The analysis of dependence of number of species on main geographic, ecological and bioclimatic factors has shown that the factors such as latitude and longitude, altitude, the precipitation rate in the driest month, and the annual precipitation rate, have a high impact on species richness of the study region (Figure 4, 5). In addition, the highest number of Pontic species is recorded at the driest month precipitation of the value 58 mm and at the annual precipitation of 900 mm. The factors related to distance from the steppe regions of southeastern Europe and diversity of potential vegetation did not impact floristic richness. An important result of these analyses is the fact that the changes in the number of species at both the latitudinal and the longitudinal gradient show opposite trends. Along the latitudinal gradient (temperature gradient) there was a statistically important increase in number of species. In contrast, the longitudinal gradient (gradient of humidity) shows a significant decrease in number of species until the 20 longitude and then there is a marked increase. As the closest steppe areas of Pannonian and Wallachian Plains are more toward north and east than the serpentine areas of western Serbia, it was expected that the greatest values of richness would be recorded in serpentine groups close to these areas. This result indicates that historical factors play an important role in the distribution of Pontic species in the study area. These factors may be more important than other ecological and biogeographic variables, like the deforestation that has created numerous open habitats that favour xerophyte colonization. 5. Conclusions Pontic, Pontic-Submediterranean and Mediterranean- Submediterranean-Pontic types of distribution of the serpentine areas of Serbia are represented with 161 taxa on species and subspecies level. The centers of diversity of Pontic, Pontic- Submediterranean and Mediterranean- Submediterranean-Pontic flora on serpentine of Serbia, both as a whole and of particular areal-types, are situated in Ibar valley and southern slopes of Mt Kopaonik. Latitude, longitude, altitude, the precipitation level of the driest month and the annual precipitation also show a pronounced impact on species richness. Annual mean temperature, distance from the steppe regions of southeastern Europe and the diversity of potential vegetation have shown a very low impact on floristic richness. The regression types at the latitudinal and longitudinal gradient show that the recent distribution of Pontic taxa sensu lato in the area studied was greatly influenced by historical factors. Serpentine areas in W&C Serbia, together with those in N Albania and N Macedonia represent important refugia for numerous plants of different age and chorology. Occurrence of endemic plants, particularly relict ones, strongly support this fact. In addition, certain number of widespread Pontic, Pontic-Submediterranean and Mediterranean- Submediterranean-Pontic species is distributed far from their main range of distribution in serpentine areas of Serbia and neighboring countries. Such taxa could be also considered as relicts of former steppe flora widespread in the Balkans during Pleistocene. The importance of this study lies in the fact that it summarizes, for the first time, a detailed distribution data of the Pontic flora in serpentine areas in the territory of Serbia. Acknowledgements The Ministry of Science and Technological Development of the Republic of Serbia, Grant : Diversity of flora and vegetation of the central part of Balkan peninsula - Ecology, Chorology and Conservation, supported this research. References [1] Stevanović V., Tan K., Iatrou G., Distribution of the endemic Balkan flora on serpentine I. obligate serpentine endemics, Plant Syst. Evol., 2003, 242, [2] Horvat I., Glavac V., Ellenberg H., Vegetation Südosteuropas (South Europe vegetation), In: Tüxen R., (Ed.), Geobotanica selecta, Gustav Fischer, Jena/ Stuttgart, 1974, 4, (I-XXXII), 1-768, (in German) 271

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15 Richness and diversity of Pontic flora on serpentine of Serbia [49] Meusel H., Jager E., Weinert E., Vergleichende Chorologie der zentraleuropaischen Flora (Comparative chorology the Central European flora) [1], Karten (Maps), Gustav Fischer, Jena, 1965, (in German) [50] Meusel H., Jager E., Weinert E., Vergleichende Chorologie der zentraleuropaischen Flora (Comparative chorology the Central European flora) [2], Karten (Maps), Gustav Fischer, Jena, 1978, (in German) [51] Meusel H., Jager E., Weinert E., Vergleichende Chorologie der zentraleuropaischen Flora (Comparative chorology the Central European flora) [2], Text, Gustav Fischer, Jena, 1978, (in German) [52] Meusel H., Jager E., Vergleichende Chorologie der zentraleuropaischen Flora (Comparative chorology the Central European flora) [3], Karten, Literatur, Register (Maps, Literature, Register), Gustav Fischer, Jena, Stuttgart, New York, 1992, (in German) [53] Meusel H., Jager E., Vergleichende Chorologie der zentraleuropaischen Flora (Comparative chorology the Central European flora) [3], Text, Gustav Fischer, Jena, 1992, (in German) [54] Stevanovic V., Floristicka podela teritorije Srbije sa pregledom visih horiona i odgovarajucih flornih elemenata (Floristic division of the territory of Serbia with an overview of higher chorion and appropriate floral elements), In: Saric M.R., (Ed.), Flora Srbije (Flora of Serbia) 1, Serbian Academy of Science and Art, Belgrade, 1992, 47-56, (in Serbian) [55] Hijmans R. J., Cameron S.E., Parra J. L., Jones P. G., Jarvis A., Very high resolution interpolated climate surfaces for global land areas, International Journal of Climatology, 2005, 25, [56] Gajic M., Florni elementi SR Srbije (Floristic elements in the Republic of Serbia), In: Jankovic M., Pantic N., Misic V., Diklic N., Gajic M., Vegetacija SR Srbije (Vegetation of the Republic of Serbia) 1, Serbian Academy of Science and Art, Belgrade, 1984, (in Serbian) 274