HACQUETIA 5/1 • 2006, 5–23 THE WEsTErn-PonTIC sTEPPE VEgETATIon In BUlgArIA Rossen TZonEV*, Veska RoUSSAkoVA, Marius DIMITRoV** Abstract The paper presents results of a syntaxonomic analysis of the herbaceous phytocoenoses on the steep stony slopes and the plainy watersheds along the northern Black Sea Coast and in the internal part of the South Dobrudzha (north-Eastern Bulgaria). The specific patterns of the vegetation in this part of the Balkan-Moesian (Lower-Danubian) forest-steppe province of the Eurasian steppe area is discussed. The comparatively preserved relict steppe vegetation includes relatively high numbers of endemic, rare and threatened species, which is the reason for its importance for conservation of biodiversity and the specific habitats. Two new associations are described: 1) Alysso caliacrae – Atremisietum lerchianae with two subassociations – typicum and camphoresmetosum monspeliacae, and 2) Paeonio tenuifoliae-Koelerietum brevis with three variants. The associations belong to the alliance Pimpinello-Thymion zigoidi – an endemic syntaxon for Bulgarian and Romanian Black Sea coasts. Key words: syntaxonomy, Festuco-Brometea, Festucetalia valesiacae, nATURA 2000 Izvleček V članku je predstavljena sintaksonomska analiza zeliščnih fitocenoz na strmih kamnitih pobočjih in ravninskih predelih ob severnih obalah Črnega morja in v notranjosti Južne Dobrudže (severovzhodna Bolgarija). obravnava poseben vzorec vegetacije v tem delu Balkansko-Mezijske (Spodnje donavske) gozdno-stepske province Evroazijskega stepskega območja. Primerljivo ohranjen relikt stepske vegetacije vsebuje relativno veliko število endemnih, redkih in ogroženih vrst, zato je pomemben za ohranjanje biodiverzitete in posebnih habitatov. opisani sta dve novi asociaciji: 1) Alysso caliacrae-Atremisietum lerchianae z dvema subasociacijama – typicum in camphoresmetosum monspeliacae, in 2) Paeonio tenuifoliae-Koelerietum brevis s tremi variantami. Asociaciji uvrščamo v zvezo Pimpinello-Thymion zigoidi – endemni sintakson na obalah Črnega morja v Bolgariji in Romuniji. Ključne besede: sintaksonomija, Festuco-Brometea, Festucetalia valesiacae, nATURA 2000 1. InTRoDUCTIon tic) province (Stefanov 194 , gribova et al. 1980, .zenda 1994, Bozhilova 1982, 1985, Bozhilova & The object of study is the region of the Bulgarian Filipova 1986). In Bulgaria two subtypes represent Black Sea coast of Southern Dobrudzha. According typical steppes: meadow steppes (more mesophilto the flora and vegetation of this area, phyto-geo-ous and rich in Dicotyledons) and mixtoherbose – graphically it belongs to the western border of the Stipa-Festuca steppes, which include the vegetation Pontic-kazakhstanian sub-area of the vast Eurasian studied. In these steppes the species of genus Stipa steppe and forest-steppe area. It is in fact transi-are accompanied by Festuca valesiaca and many tional between the southern part of the Balkan-other cereals (Koeleria, Cleistogenes, Agropyron). The Moesian (Lower-Danubian) forest-steppe province communities have a very characteristic, rich, flow(which is more clearly presented within the inte-ering spring spectrum composed by species of the rior part of the country), and the Black Sea (Pon-genera Tulipa, Gagea, Potentilla, Astragalus, Paeonia, * Department of Ecology, Faculty of Biology, Sofia University, 8 Dragan Tzankov Blvd., Sofia 1421, Bulgaria ** Department of Dendrology, Faculty of Forestry, University of Forestry, 10 kliment ohridsky Blvd., Sofia 1756, Bulgaria 5 Hacquetia 5/1 • 2006, 5–23 Adonis, Iris etc. The subtype of Artemisia – Festuca – Stipa steppes also occurs, even though scarcely, as an edaphic phenomenon on the steep slopes. It is different than the semi-desert steppes along the northern Black Sea regions, distributed on flat and saline surfaces (gribova et al. 1980). The flora on the territory of Southern Dobrudzha – between the towns of Balchik and Shabla – was partly investigated by kozhuharov et al. (2001, 2002.). More general information about the flora and vegetation of the region could be found in the works of Davidov (1914), yordanov (19 6), Stoyanov (1940, 1941a, 1941b, 1950), Stefanov (194 ) etc. Phytocoenotic analysis of the vegetation be means of the dominant method was published by Velchev (2002). The steppes and forest-steppes along the Black Sea parts of Russia and Ukraine share similar characteristics with the studied vegetation (gribova et al. 1980), but the most similar is the vegetation in northern Dobrudzha (Romania). However, the petrophyllous steppes in Babadag (Dihoru & Donita 1970, Ivan et al. 199 ) have been formed on green shists, while those along the Bulgarian Black Sea coast are on marls and limestone. These and some other ecological conditions affected the development of different groups of phytocoenoses in northern and Southern Dobrudzha. 2. MATERIAL AnD METHoDS The region of study includes the territory starting from the northern part of Balchik and reaches northward to the village of kamen bryag and the “yaylata” locality (the whole territory of “kaliakra” reserve falls into it). To the west, in goes to the inner part of Southern Dobrudzha and reaches the surroundings of the villages Vidno and Irechek (“gyorensko dere” locality). The relief includes slopes with rocky (saltificated to a different extent Sarmatian marls as well as chalks) revelations near the Black Sea coast. They are mainly landslides (Balchik-kavarna) and flat calcareous watersheds. The latter ones are specific forms along the line over the Black Sea cliff coast (kaliakra – kamen bryag) and in the interior of the studied territory – the slopes of the dry valleys named “kayryatsi”(gyorensko dere), in Dobrudzha plateau. The climate along the Bulgarian Black Sea coast north from Balchik is transitional – Continental- Mediterranean. Temperature amplitudes are lower than inside the country and air humidity is higher, because of the sea breeze. The strong northern and eastern winter winds are typical for the region. The mean annual rainfalls are lower than in other regions of the country, which is the reason for the xerothermic peculiarities of the vegetation (Dimitrov & Vekilska 1966, galabov 1982, Velev 1990). on the carbonate basic rocks, the soils are rendzic leptosols with xerotermic soil processes; on the molds (chernozems and haplic chernozem) the soil processes are mezothermic. (Tanov 1968, ninov 1997). The investigation of the vegetation was performed during the year 2004 according to the methods of the sigmatic school (Braun-Blanquet 1964, Westhoff & Maarel 1978). A total of 82 phytocoenotic relevés were described, but 19 of them were rejected during the process of analysis. The expanded scale of Braun-Blanquet for abundance/ dominance (Barkman et al. 1964) was used, transformed according to Maarel (1979) during the statistical processing. The cluster analysis was perfomed by means of the software Syntax (Podani 2002). Average linkage method (UPgMA) was used and floristic similarity among relevés was evaluated according to Horn’s index (krebs 1999). The taxonomic nomenclature followed kozhuharov, ed. (1992). The new syntaxa were compared with the similar syntaxa from northern Dobrudzha (Dihoru & Donita 1970, Ivan et al. 199 , Sanda et al. 1999) and were described according the rules of the International Code of Phytosociological nomenclature (Weber et al. 2000). . RESULTS AnD DISCUSSIon The cluster and syntaxonomic analysis of the steppe vegetation in north-Eastern Bulgaria revealed that the syntaxa could be included into the next hierarchical syntaxonomic classification: Festuco-Brometea Br.-Bl. et R. Tx. in Br.-Bl. 1949 Festucetalia valesiacae Br.-Bl. et R. Tx. in Br.-Bl. 1949 Pimpinello-Thymion zygoidi Dihoru 1970 Alysso caliacrae-Atremisietum lerchianae ass. nov. typicum subass. nov. Alysso caliacrae-Atremisietum lerchianae cam phoresmetosum monspeliacae subass. nov. Paeonia tenuifoliae-Koelerietum brevis ass. nov. var. typicum Teucrium chamaedrys var. nov. Rhodax canus var. nov 6 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia A B a1 a2 b1 b2 b3 Figure 1: Classification of all relevés. Average linkage method and Horn`s index of dissimilarity. A – ass. Alysso caliacrae- Atremisietum lerchianae, a1 – subass. typicum, a2 – subass. camphoresmetosum monspeliacae; B – ass. Paeonio tenuifoliae-Koelerietum brevis, b1 – var. typicum, b2 – var. Teucrium chamaedrys, b3 – var. digressive. Slika 1: Klasifikacija vseh popisov. Kopičenje na osnovi povezovanja srednjih razdalj in Hornov indeks različnosti. A – asociacija Alysso caliacrae-Atremisietum lerchianae, a1 – subasociacija typicum, a2 – subasociacija camphoresmetosum monspeliacae; B – asociacija Paeonio tenuifoliae-Koelerietum brevis, b1 – var. typicum, b2 – var. Teucrium chamaedrys, b3 – odstopajoča varianta. Western-Pontic wormwood steppes Ass. Alysso caliacrae-Atremisietum lerchianae The phytocoenoses forming this group (cluster A – Figure 1) are distributed on the steep slopes (60–90°) with carbonate rocks (Sarmatian marls), along the Black sea coast between Balchik and kavarna. The most representative of them were in the region of Balchik Tuzla and between the village of Topola and kavarna (“Chirakman” locality). The exposition is mostly eastern and the soil cover is poorly developed or lacking. Most of the phytocoenoses occur on the acting landslides (Figure 2). This type of vegetation is of natural origin. Typical chasmophyte plants (adapted for poorly developed soils) predominate: Achillea clypeolata, Agropy- Figure 2: Most of the phytocoenoses occur on the acting landslides. Slika 2: Večina rastlinskih združb se pojavlja na aktivnih plaziščih. 7 Hacquetia 5/1 • 2006, 5–23 ron cristatum ssp. brandzae, Artemisia lerchiana, Astragalus spruneri, Cephalaria uralensis, Kochia prostrata, Satureja coerulea, Ephedra distachya etc (Table 1). one interesting fact is the participation of the Pontic-Turanian, very rare (only in this region) species Matthiola odoratissima. The range of distribution of the dominant species, the halophyte Artemisia lerchiana includes the Pontic and Aralo-Caspian regions (Figure ). Another typical steppe species with a high abundance and constancy is the Pontic geo-element Agropyron cristatum ssp. brandzae. More important accompanying species are Astragalus glaucus, A. vesicarius, Jurinea stoechadifolia, Linum austriacum, L. tauricum subsp. tauricum, Echinops ritro. Species such as Alyssum caliacrae (Figure 4), Aster oleifolius, Astragalus spruneri, Festuca valesiaca, Teucrium polium, Thymus zygoides possess comparatively high constancy, but low abundance. Most of these species are typical steppe elements. The syntaxonomic analysis of the relevés allowed us to describe a new association Alysso caliacrae-Artemisietum lerchianae. Diagnostic species are Artemisia lerchiana, Agropyron cristatum ssp. brandzae, Alyssum caliacrae, Aster oleifolius, Kochia prostrata, Astragalus spruneri and Mathiola odoratissima. Two groups of relevés (Figure 1) with a high floristic similarity (more than 40%) are distinguished. The presence of differentiating species and the ecological differences define the sub-associations typicum and camphoresmetosum monspeliacae. The phytocoenoses from the typical sub-associations (subcluster a1 – Figure 1) are distributed on the slopes north from Balchik (“Tuzlata” locality), south from kavarna (the villages of Topola and Bozhurets) and those with southern exposition in the Bolata Inlet. The basis rocks are white nonsaltificated marls and chalks. This sub-association contains more autochthonous species than the sub-association camphoresmetosum monspeliacae (Table. 1), for example, the Balkan endemic Astragalus spruneri and the Balkan-Crimean species Astragalus glaucus. The main reason for the difference between the typical sub-association and subassociation camphorosmetosum monspeliacae (subcluster a2 – Figure 1) is the character of saltificated Sarmatian marls in the “Chirakman” locality, south from kavarna. Diagnostic species of the second subass. are Astragalus vesicarius, Camphorosma monspeliaca and Brassica elongata. Many species, well-presented in the typi- Figure 3 (Slika 3): Artemisia lerchiana. Figure 4 (Slika 4): Alyssum caliacrae. 8 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia cal sub-association, are not presented here or else occur at very low constancy. These are: Salvia nemorosa, Alyssum caliacrae, Astragalus glaucus, Astragalus spruneri, Dianthus pseudoarmeria, Euphorbia nicaensis, Jurinea stoechadifolia, Linum austriacum. Many ruderals or semi-ruderals as Brassica elongata, Adonis flammea, Thlaspi arvense, Valerianella pumila could also be found in this subassociation. The participation of Artemisia lerchiana, Agropyron cristatum subsp. brandzae, Thymus zygoides, Dianthus pseudoarmeria, Pimpinella tragium and Cephalaria uralensis indicates that this association should be included into the alliance Pimpinello-Thymion zygoidi Dihoru 1970, described in northern Dobrudzha. Thus the range of distribution of this alliance is extended to the Southern Dobrudzha. It is of specific transitional Mediterrano-Pontic origin and is distributed on poor soils and rocky substrates. The diagnostic species for the order Festucetalia valesiacae and the class Festuco-Brometea are: Adonis flammea, Alyssum hirsutum, Astragalus glaucus, Astragalus vesicarius, Ephedra dystachia, Festuca valesiaca, Euphorbia nicaensis, Linum austriacum, Stipa capillata, Seseli tortuosum, Teucrium poliumk, Jurinea stoechadifolia, and some others. Similar to the described association are the petrophyllous communities of the associations Agropyro-Thymetum zygoidi, Koelerio-Artemisietum lerchianae, Festycetum callierii and Artemisierum caucasicae prov., established on the territory of Romania (Babadag, northern Dobrudzha) (Dihoru & Donita 1970). The floristic composition of the phytocoenoses from northern Dobrudzha indicates that most similar to the petrophyllous steppe vegetation along Bulgarian Black Sea coast are the associations Agropyro-Thymetum zygoidi and Koelerio-Artemisietum lerchianae (Babadag). Some of the main species of the phytocoenoses from the Bulgarian Black Sea (Southern Dobrudzha) and Babadag (northern Dobrudzha) are common and are diagnostic species of the alliance Pimpinello-Thymion zygoidi: Artemisia lerchiana, Agropyron cristatum subsp. brandzae, Thymus zygoides. of the other species only 6 are common for the associations Agropyro-Thymetum zygoidi and Alysso-Artemisietum lerchianae. Also, species such as Festuca callieri, Campanula romanica, Gagea callieri, Potentilla bornmuelleri, Euphorbia glareosa, Minuartia viscosa, Artemisia ucrainica do not occur or are scarcely represented in the association along the Bulgarian Black Sea coast. The differentiating species of the association Koelerio-Artemisietum lerchianae are: Alyssum caliacrae, Astragalus glaucus, A. spruneri, Jurinea stoechadifolia etc. They do not occur in the associations from Babadag (Romania). The newly described association Alysso caliacrae- Artemisietum lerchianae is endemic for one limited region of the Bulgarian Black Sea coast, mainly on the landslide terrains between Balchik and kavarna. The main characteristics of the association and the richness in rare, Pontic and endemic species increase its nature-conservation value. Western Pontic feathergrass steppes Association Paeonio tenuifoliae-Koelerietum brevis The phytocoenoses of the association Paeonio tenuifoliae-Koelerietum brevis (cluster B – Figure 1) are distributed on the plainy, calcareous belt along the Black sea coast in the regions of kavarna and Cape kaliakra, kamen bryag, Bozhuretz, Bulgarevo villages, north from Bolata Inlet, in the “yaylata” locality; and in the inner part of Southern Dobrudzha – the villages of Bilo, Vidno (“gyorensko dere” locality) (Figure 5). Most probably it occurs also on some other places inside the calcareous dry valleys of the Dobrudzha plateau, called “kayryatsi”. Figure 5: High degree of soil erosion demonstrates the clear Mediterranean physiognomy of phytocoenosis. Slika 5: Erozija nakazuje medtieransko fizionomijo te fito- cenoze. The soils are of different depth, strongly eroded and the Miocene calcareous rocky basis is disclosed on the surface in many places. other reasons for the soil erosion are the strong east and north winds, which blow along the strip near the sea, mainly in winter. Another very important factor is the continuous human impact, especially grazing pressure. The numerous typical forms of the nanorelief change in a short distance. This combination of ecological factors results in irregular, and in 9 Hacquetia 5/1 • 2006, 5–23 some cases group distribution of the plants. Therefore, Velchev (2002), using the dominant method, described 8 associations within the plant communities studied. The analysis of results of the present study provided evidence for the existence of a new association – Paeonio tenuifoliae-Koelerietum brevis (Table 1). A mixed complex of Pontic, Mediterranean and Submediterranean species, endemics, and many ruderals and semi-ruderals (some of them with a secondary distribution into the natural communities) plays an important role in the formation of this association. Typical species are: Festuca valesiaca, Koeleria brevis, Iris pumila, Thymus zygoides, Stipa lessingiana, Cerastium bulgaricum, Chamaecytisus jankae, Euphorbia nicaensis, E. myrsinites, Helianthemum salicif.lium, Erysimum diffusum, Inula oculus-christi, Paeonia tenuifolia, Potentilla bornmuelleri, Artemisia pedemontana, Tanacetum millefolium, Satureja coerulea, Scandix australis, Scutellaria orientalis subsp. pinnatifida. The participation of many therophytes is indicative for the natural climatic, soil conditions and for the influence of the pasture. Diagnostic species for the association are: Koeleria brevis (Figure 6), Cerastium bulgaricum, Paeonia tenuifolia, Chamaecytisus jankae, Helianthemum salicifolium, Potentilla bornmuelleri, Stipa lessingiana and Scandix australis. Many beautiful-flowered spring species such as Iris pumila, Adonis vernalis, Scutellaria orientalis, Paeonia tenuifolia participate within the floristic composition of this association. They form a short-term spring aspect. During the rise of the temperature at the beginning of summer this vegetation finishes its development. Many ruderals, such as Carduus acanthoides, Carthamus lanatus, Centaurea spp. and other species from Asteraceae continue their development during the summer vegetation period. The seasonal dynamic resembles these petrophyllous steppes to the Mediterranean vegetation and distinguishes them from the northern Pontic steppe, which has a more continuous vegetation season and several aspects. Some specific successional processes affected the floristic composition of the dominated grass species. In the primary steppe vegetation, the role of Festuca valesiaca is more limited or similar to those of the Stipa spp. The active grazing impact on the natural vegetation and the soils resulted in Festuca valesiaca being the dominant species, and has limited the role of Stipa spp. (gribova et al. 1980). The floristic composition of the presented association and the cluster dendrogram demonstrate the subdivision into three main groups of phytocoenoses, which represent the variants of the pasture regression of the steppe vegetation. The most similar to the primary vegetation is the group of phytocoenoses (subcluster b2 – Figure 1, Table 1) described in the region between the village of Bulgarevo, Cape kaliakra and especially on the watershed north from Bolata Inlet, within the game-breeding farm “kushlata” (relevés 7–5 ) near the village of kamen bryag. Stipa lessingiana demonstrates a high constancy (IV), although the more frequently dominant species are Festuca valesiaca and Koeleria brevis. Achillea clypeolata, Scandix australis (more rare species than the ruderal Scandix pecten-veneris, which is well-distributed in the other two variants of the association), Astragalus vesicarius, Chamaecytisus jankae, Artemisia pedemontana (Figure 7) which have a high presence. We could suppose that this vegetation is most similar to the Figure 6 (Slika 6): Koeleria brevis. Figure 7 (Slika 7): Artemisia pedemontana. 10 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia primary petrophyllous steppes, distributed along the whole Black sea coast of South Dobrudzha, although it differs in the degree of anthropogenic degradation. As most representative could be accepted the phytocoenoses between Bolata Inlet and the village of kamen bryag, especially inside the enclosure region of the game-breeding farm “kushlata”, where Stipa lessingiana is dominant, but Paeonia tenuifolia, Adonis vernalis and the ruderal species are less presented. Another group of phytocoenoses are those on the “kayryak” near to the village of Bozhurets and in the interior of Dobrudzha – “gyorensko dere” near the village of Vidno (relevés 0- 6) (subcluster b1 – Figure 1, Table 1). The soils in these regions are better developed than in the other two variants, although, in spite of the strong grazing pressure and practically complete lack of Stipa lessingiana, their floristic composition is richer. Many species that are absent or have a sporadic occurrence within the other two variants, are well-presented in this one: Convolvulus cantabrica, Vinca herbacea, Artemisia austriaca, Teucrium chamaedrys, Astragalus onobrychis, Salvia argentea, Thlaspi arvense, Linum austriacum, Achillea millefolium, Bromus racemosus, Lappula barbata, Medicago minima, Poa bulbosa, Trigonella gladiata etc. Some of these species have a very limited distribution in Bulgaria, but most of them are common and demonstrate the more mesophyte character of the phytocoenoses. The reasons are different, but two are more important. They are developed on richer and deep soils, especially in the interior of Dobrudzha, where the influence of the strong winter winds is less expressed. Also, they are more similar to the communities classified as meadow steppes from the Balkan-Moesian province by gribova et al. (1980). Their distribution was expanded to the places of the former forests because they are developed on richer and more humid soils. The phytocoenoses of the third group (mainly from the intensively used pasture near the village of Bulgarevo– relevés 54–6 ) have the main characteristics of the associations. Many autochthonous species have declined from the floristic composition of these phytocoenoses and they have poorer species structures than the other two groups of communities. The high ratio of such species as Rhodax canus, Artemisia pedemontana (Figure 5) and Scutellaria orientalis ssp. pinnatifida underlines the highest degree of soil erosion, and demonstrates the clear Mediterranean physiognomy of this variant. Many diagnostic species to the alliance Festucion valesiacae occur in the floristic composition of the association Paeonio tenuifoliae-Koelerietum brevis. These are: Festuca valesiaca, Astragalus vesicarius, Inula oculus-christi, Stipa capillata. The association is composed by diagnostic species to Festucetalia valesiacae and class Festuco-Brometea. Also, some diagnostic species of the alliance Pimpinello-Thymion zigoidi are quite well represented. These are species such as Thymus zygoides, Scutellaria orientalis, Agropyrum brandzae, Potentilla bornmuelleri and some others. This fact approximates the communities to the alliance Festucion valesiacae and syntaxonomically they could be classified as belonging to it, despite their similarity to Festucion valesiacae (Table 1). Some of the above mentioned species do occur, although scarcely, in the associations established in northern Dobrudzha (Babadag) Stipo ucrainicae-Festucetum valesiacae Dihoru 1970 and Medicagini-Festucetum valesiacae Wagner 1941 (Dihoru & Donita 1970), which are most similar from the floristic and territorial point of view to the Bulgarian association. The whole composition and structure of the Babadag’s associations are different than the Bulgarian one and only a small number of the species are common for both regions of Dobrudzha. For example – Cerastium bulgaricum and the Balkan endemic Chamaecytisus janka. do not occur in the communities in northern Dobrudzha. The southern border of the distribution of Stipa ucrainica is out of the Bulgarian territory. The anthropogenic influence limited the distribution and abundance of some steppe species including the genus Stipa in Bulgaria. It allowed the penetration of the ruderals (Scandix pecten-veneris, Marrubium peregrinum, Valerianella pumila, Carduus acanthoides, Erodium cicutarium, Bromus racemosus, Thlaspi arvense, Adonis flammea, Achillea millefolium etc.) in the plant communities. The increased participation of some poisonous species, such as Paeonia tenuifolia, Adonis spp. etc. is probably also a result of the pasture regression of these communities. The evidence of such development is the more restricted participation of these species especially in the region north of Bolata Inlet, which is the most similar to the primary vegetation in the region and there is a clearer dominance of Stipa lessingiana. The association Paeonio tenuifoliae-Koelerietum brevis is a successional stage of the pasture regression of the natural steppe vegetation. Such are the associations in northern Dobrudzha. Many preserved fragments of the forest vegetation, having wilder distribution in the past, are presented in the region of study. They are not only on the northern slopes, but also on the watersheds. Single individu 11 Hacquetia 5/1 • 2006, 5–23 als or groups of trees (Quercus pubescens, Fraxinus ornus etc.) and shrubs (Crataegus monogyna, Cornus mas, Carpinus orientalis, Paliurus spina-christis, Cotinus coggygria etc.) represent refugees of the more mesophylous species as Poa angustifolia, Phleum phleoides, Fragaria viridis, Oryganum vulgare etc. This fact could be better established especially in the variant with Teucrium chamaedrys, which is the most similar to the more mesophylous meadow steppes. A secondary phenomenon is the distribution of some mono-dominant phytocoenoses of Asphodeline lutea in many places in the region of study. An interesting fact is that they are common near the ancient settlements – cape kaliakra and the region of “yaylata”. The anthropogenic degradation very probably is connected to the mesophytisation and nitration of the terrain near the former settlements, which was favorable for the development of the phytocoenoses of these species. These communities occur naturally within the open bushes. Similar processes are the reasons for the good population of Paeonia peregrina around the Byzantine castle on “yaylata”. This species is typical of the open oak woods, widely available on the seaward facing terraces of “Taukliman” (“Rusalka” Resort). Such forests probably have occurred on “yaylata” too, which is a big terrace over the sea. After deforestation Paeonia peregrina and Asphodeline lutea survived around the ruins as semi-ruderals. They are forming now mixed communities with some mesoxerophyllous ruderals, which inhabit within the open forests and some of them are nitrophyllous. Such are Dactylis glomerata, Galium aparine, Hordeum bulbosum, Tordylium maximum, Ornithogalum narbonense, Vicia grandiflora, Geranium rotundifloium etc. The positive influence of the nitration of the soils, especially for Paeonia peregrina has been established experimentally in Bulgaria (Radulovski 2004). Phyto-geographic character and history of the Western Pontic steppe vegetation. The results of the present work confirm the hypothesis of the close connection between the steppes in Southern Dobrudzha and the Pontic steppes, as well as the dominant role of some Submediterranean geo-elements for the floristic composition of the steppe vegetation in the investigated region (Stoyanov 1940, 1941., 1941b, kozhuharov et al. 2001, 2002). The species, participating in the discussed phytocoenoses, are of southern origin. Their ranges of distribution are in Southern Europe, the Mediterranean region, Cau casus, Southwestern Asia, but many of them have a clear Pontic origin. They practically belong to the flora of the so named “Ancient Mediterranean region” according to the concept of Popov (196 ). This idea strongly corresponds to the earlier work of Stoyanov (1925), where he discuses the origin of many so called “steppe species” with a connection with the refugees of Mediterranean flora during the Pleistocene and their invasion during the Holocene. Some of the species reported by the authors do not occur in the relevés studied in this work, and conversely, some species we found are absent in these works. This fact does not change the common composition of the geoelements within the new associations, which confirms the facts established by the cited authors. According to kozhuharov et al. (2002) inside the biggest group of Sub-Mediterranean geo-elements are the following species: Aegilops geniculata, Crepis sancta, Nonea pulla, Sideritis montana, Valerianella pumila, Xeranthemum annuum, Ligustrum vulgare, Cornus mas, Carpinus orientalis, Acinos suaveolens, Campanula sibirica, Euphorbia nicaensis, Festuca valesiaca, Linum austriacum, Salvia nemorosa, Scutellaria orientalis, Teucrium chamaedrys, Digitalis lanata, Vicia grandiflora etc. The second position keep the Eurasian (s. lat.) species: Adonis wolgensis, Daucus carota, Dactylis glomerata, Hyacinhtella leucophaea, Matthiola odoratissima (rather Pontic-Turanian species), Medicago falcata, M. minima, Poa bulbosa, Scandix pecten-veneris etc. Euro-Mediterranean geoelement are: Asperula cynanchica, Astragalus vesicarius, Carduus acanthoides, Filipendula vulgaris, Stachys recta etc. Pontic-Mediterranean and Mediterranean species are: Carduus pycnocephalus, Crupina vulgaris, Iris pumila, Medicago disciformis, Satureja coerulea, Ziziphora capitata, Ajuga chamaepytis, Asphodeline lutea, Centaurea napulifera subsp. thirkei, Cephalaria uralensis, Eryngium campestre, Euphorbia myrsinites, Fraxinus ornus, Haplophyllum suaveolens, Jasminum fruticans, Potentilla bornmuelleri, Salvia argentea, Stipa capillata, Tanacetum milleflium, Trigonella gladiata ect. The species belonging to the Euro-Siberian group and other geo-elements are less represented. The Sub-Mediterranean aspect of the investigated vegetation is the reason that Rodwell et al. (2002) defined the alliance Pimpinello-Thymion zigoidi as “semi-evergreen ‘phrygana’-like vegetation of the east Balkan steppes”. Many rare and endemic species to Bulgaria or the Balkan Peninsula participate in the floristic composition of the investigated phytocoenoses. This fact increases their value for the conservation 12 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia of biodiversity on national and international level. The group of the rare species includes: Adonis wolgensis, Matthiola odoratissima, Avena eriantha and Bellevalia ciliata (they both have only one locality in the country in “kaliakra” Reserve), Scandix australis, Artemisia pedemontana, A. lerchiana, Ruta graveolens, Convolvulus lineatus, Anchusa stylosa, Nepeta parviflora, Scabiosa atropurpurea, Koeleria brevis, Paeonia tenuifolia, Stipa lessingiana, Goniolimon besseranum. Some of these species, not occurring or being very rare in other places in Bulgaria, are dominants in the part of the phytocoenoses in Dobrudzha. The percentage of the endemics and sub-endemics is high, too: Alyssum caliacrae, Astragalus spruneri, Achillea clypeolata, Chamaecytisus jankae, Satureja coerulea, Koeleria davidovii, Jurinea stoechadifolia. Another interesting question is that of the origin and the phyto-geographycal relations of this vegetation with the steppes north from it. These steppes are a continuation of the cold, often saltificated steppes distributed during the Ice Age, which were dominated by Artemisia, Chenopodiaceae. Contrary to other places in Bulgaria, the forest invasion during the Holocene with the improvement of the climatic conditions was not so active, and the dominated landscape was the forest-steppes. (see Bozhilova 1982, 1985, Bozhilova & Filipova 1986). Inside the interior of Dobrudzha, probably during the improvement of the climatic and soil conditions, the forest vegetation increased. Therefore, Davidov (1914) considered wrongly that the whole Dobrudzha was covered by forests in the past, which were destroyed by the humans. The most deforested one is the strip near the sea, where the forest vegetation has survived only on some wet slopes and valleys (“Taukliman”, “kushlata”). only strongly xerophytic species participate in their floristic composition. The main reason for this condition is the poor soil cover, which was caused by the strong winds and the human influence over the vegetation. As was already mentioned, according to gribova et al. (1980) the Bulgarian steppes (and forest-steppes) belong to Balkan-Moesian (Lower-Danubian) meadow and various-grass – Festuca - Stipa steppes. However, we should note also their high similarity to the typical Black sea steppes (Pontic province) and especially to their edaphic (petrophyllous) subtype. The main difference is the strong Mediterranean influence in Bulgaria. We could presume that the invasion of Mediterranean species (mainly aromatic semi-shrubs, terophytes and ruderals) happened during the improvement of the climatic conditions in Holocene under the increase of the anthropogenic degradation of the vegetation. The successional changes of this vegetation were connected with the exchange of the steppes dominated by Stipa spp. with these rich in Festuca spp., as well as ruderal, thorny and poisonous species, mainly as a result of the grazing of domestic animals. Similar processes have been established not only in Russia (keller 192 , gribova et al. 1980), but in Bulgaria too. yordanov (19 6) notes that the exchange of the communities of Stipa spp. has been realized by Sub-Mediterranean “andropogonide” grasses – Chrysopogon gryllus, Dichanthium ischaemum, and the reasons are mainly climatic. This process is more typical for the forest-steppe vegetation in the interior of Bulgaria. The disappearance or limitation of the distribution of some species from genus Stipa in Southern Dobrudzha and their replacement by the more flexible Festuca spp. was connected also to the human influence as well as to the climatic changes during the Holocene. 4. ConCLUSIonS The steppe vegetation had a wider distribution on the territory Bulgaria in the past. only small fragments have survived today. Many typical steppe species, like those of the genus Stipa, have vanished from the studied territory as a result of the pasture pressure. The recent steppe vegetation is presented by different successional stages of development. However, as a whole, the steppe character of the grass vegetation in the investigated regions of Southern Dobrudzha remained. Despite the human degradation, the survived fragments of the steppe vegetation not only demonstrate the evolution of steppe vegetation in the region, but they have special importance because of the participation of many rare, threatened species, Bulgarian and Balkan endemics. This fact increases their value for the conservation of biodiversity in Bulgaria and Europe. The steppe syntaxa in Bulgarian Dobrudzha are represented into the Directive 92/4 / EEC by the unique code for the Pontic bio-geographical region – 6290 Western Pontic Paeonia steppes. The results of this study provide the reasons for the following conclusions: The steppe phytocoenoses on the steep slopes and the plane watersheds along the Bulgarian Black-sea coast as well as in the interior of Southern Dobrudzha have special peculiarities. They differ 1 Hacquetia 5/1 • 2006, 5–23 to the mixed Pontic-Submediterranean origin, but they have a strong connection with the steppes of the Pontic province. Two associations were distinguished from a syntaxonomical point of view. The syntaxa from the higher ranks are common with the steppe vegetation in northern and Southern Dobrudzha. The results from this work could be used in the classification of the natural habitats and for the purposes of the European ecological network nATURA 2000 in Bulgaria. 5. ACknoWLEDgEMEnTS. The authors would like to express their thanks to WWF – Bulgaria (Danube-Carpathian Programme) for the financial and logistic support of the field investigations. We would also like to thank “Sylvica” Foundation (BSFP) for giving permission to use the programme Syntax, and Dr. Rumen Beleliev for the technical support. 8. REFEREnCES: Barkman, J., Doing, H. & Segal, S. 1964: kritische bemerkungen und vorschlage zur quantitativen vegetationanalyse. Act. Bot. neederland., 1 : 94–419. Bozhilova, E. 1982: Holocene Chronostratigraphy in Bulgaria. Stiae, 16: 88–90. Bozhilova, E. 1985: Palaeoecological Studies in Lake Durankulak. Ann. Univ. Sofia, Biol. fac., 2(75): 88–95. Bozhilova, E., & Filipova, M. 1986: Palaeoecological environment in northeast Black Sea area during neolithic, Eneolithic and Bronze periods. Studia Praehistorica, 8: 160–165. Braun–Blanquet,J.1964:Pflanzensoziologie.grundzuege der Vegetationkunde. Aufl. Spinger Verlag, Wien-new york, 865 pp. Davidov, B. 1914: Phytogeographical characters of Bulgarian Dobrudzha. Trud. Bulg. prip. izp. drvo, 6: 17–28. (in Bulgarian) Dihoru, g. 1970: Vegetatia ierbosa a padisului Babadag. In: Dihoru, g., & Donita, n.: Flora si vegetatia podisului Babadag. Edit. A.S.R., Bucuresti, pp. 16 –297 Dimitrov, D., & Vekilska, B. 1966: Po vyprosa za teritorialnoto razpredelenie na valezhite v Dunavskata ravnina. god. SU, ggF, 2(59): 47–66. (in Bulgarian) galabov, J. (ed.) 1982: geography of Bulgaria. Physical geography. BAS. Sofia, 247 pp. (in Bulgarian) gribova, S., Isachenko, T. & Lavrenko, E. (eds.) 1980: Rastitel’nost’ evropejskoj chasti SSSR. nauka, Leningrad, 429 pp. (in Russian) keller, B. 192 : Rastitel’nuj mir russkih stepej, polupustujn’ i pustin’. T. 1, gIZo, Voronezh. (in Russian) Ivan, D., Donita, n., Coldea, g., Sanda, V., Popescu, A., Chifu, T., Boscaiu, n., Mititelu, D. & Pauca- Comanescu, M. 199 : Vegetation potentielle de la Roumanie. 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Proceedings of Scientific Reports, HAI, Press, Plovdiv: 21–26. ozenda, P. 1994: Végétation du Continent Européen. Delachaux et niestlé, Lausanne, 280 pp. Podani, J. 2001: Syn-TAx 2000. Computer Program for Data Analysis in Ecology and Systematics. User’s Manual. Scientia Publ., Budapest, 5 pp. Popov, M. 196 : osnovuj florogenetiki. Izd. An na SSSR, Moskva, 1 5 pp. (in Russian) Radulovski, S. 2004: Ekologichna ocenka na populacii na cherven bozhur (Paeonia peregrina Mill.) v Plevenski i Velikoturnovski rajoni. Diploma thesis, Sofia, 46 pp. (in Bulgarian) Rodwell, J., Schamenee, J., Mucina, L., Pignatti, S., Dring, J. & Moos, D. 2002: The Diversity of European Vegetation. An overview of phytosociological Alliances and their relationships to 14 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia EUnIS habitats. Landbouw, natuurbeheer en visserij. 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Velchev, V. 2002: Phytocoenological and ecological characteristics of the communities of Koeleria brevis Stev. and Festuca pseudovina Hack ex Wiesd. in the Balchik-kavarna region of north- Eastern Bulgaria. Ann. of Sofia Univ., Fac. of Biology, 92(2): 9 29. Velev, S. 1990: klimatyt na Bylgariia. narodna pros- veta, Sofia, 179 pp. (in Bulgarian) yordanov, D. 19 6: on the distribution of the steppes vegetation in Bulgaria. Bull. BAS, 2(5): 1–105. (in Bulgarian) Weber, H. e., Moravec, J., & theurillat, J.-P. 2000: International Code of Phytosociological nomenclature. rd edition. – J. Veg. Sci., 11: 7 9– 768 Westhoff, V. & van der Maarel, E. 1978: The Braun- Blanquet approach. In: Whittaker, R.H. (ed.): Classification of plant communities. Junk, The Hague.pp. 287– 99. Recieved 16. 8. 2005 Revision recieved 9. 1. 2006 Accepted 2. 6. 2006 15 Hacquetia 5/1 • 2006, 5–23 Table 1: The syntaxa of Western - Pontic Steppes in Bulgaria: association Alysso caliacrae-Artemisietum lerchianae and association Paeonio tenuifoliae-Koelerietum brevis. Tabela 1: Sintaksoni Zahodno-Pontske stepe v Bolgariji: asociacija Alysso caliacrae-Artemisietum lerchianae in asociacija Paeonio tenuifoliae-Koelerietum brevis. Number of relevé 1 2 3 4 5 6 7 8 9 1011121314151617181920 2122232425 Date 7V 7V 7V 7V 7V 7V 7V 8V 8V 8V 8V 8V 8V 8V 7V 7V 7V 7V 8V 9V 8V 8V 8V 8V 8V Relevé area, m2 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 50 100 100 100 100 100 Exposition . . . . . . .. . . . .. .. . . . . . . .. . . . . . .. Slope, degrees 70 80 60 70 70 60 20 50 70 60 15 50 30 70 50 80 70 70 20 90 60 70 60 30 50 Total cover, % 80 60 80 80 70 60 80 40 60 80 40 80 50 60 80 85 80 70 90 10 60 70 50 40 30 Number of taxa 16 18 22 28 21 22 18 25 22 21 14 23 11 17 19 14 15 18 28 7 21 26 30 18 11 Diagnostic taxa of associations, subassociations and variants Ass. Alysso caliacrae-Artemisietum lerchianae ass. nova, type nom. rel. 10 holotype Agropyron cristatum (L.)Gaertn. ssp. brandzae P.et S. 2b 2a 3 3 3 1 2a 1 2b 2b 2a 2b 2a 3 3 2a 3 3 + + V 3 3 3 3 3 Artemisia lerchiana Weber. 3 3 2a 2a 3 2a 2a 3 1 3 2a 2a 3 2a 2a 3 2b 1 + + V 3 2b 2a 2a 2b Alyssum caliacrae Nyar. + . . 2m + 2m 2a + 2m 2m + 2a . 2m 2a 2m 2m 2m + + V . + . . . Astragalus spruneri Boiss. + 1 2a 2a 2a 1 1 2a 2a 1 . 1 . + 1 1 + 2a 1 . V . . . + . Kochia prostrata (L.) Schrader. . 3 2b 1 2a 2a . 2a . 2a 2b 3 2a 2a . . . . . + III 2a 2a 1 . . Aster oleifolius (Lam.) Wagenitz . . + . . 1 1 2a 2b 1 . 2a 2a 2a . . 1 1 2a . III . 2a 1 . 1 Mathiola odoratissima (Bieb.)R.Br. . . . . . . . . . 1 + . . . . . . . . . I . . . 2a 2a Subass. Alysso caliacrae-Artemisietum lerchianae camphorosmetosum monspeliacae subass. nova, type nom. rel. 22 holotype Astragalus vesicarius L. . . . . . . . . . . . . . . . . . . . . 2a 2b 2b 2a 2b Camphorosma monspeliaca L. . . . . . . . . . . . . . . . . . . . . 2a 2a 2a 2a . Brassica elongata Ehrh. 1 + . . . + . . 1 . . . . . . . . . 1 . II + 1 + + + Ass. Paeonio tenuifoliae-Koelerietum brevis ass. nova, type nom. rel. 39 holotype Potentilla bornmuelleri Borbas . . . . . . . . . . . . . . . . . . 1 . I . . . . . Helianthemum salicifolium (L.) Miller. . . . . . . . . . . . . . . . . . . . . . . . . . Koeleria brevis Steven. . . . . . . . . . . . . . . . . . . . . . . . . . Cerastium bulgaricum Uechtr. . . . . . . . . . . . . . . . . . . . . . . . . . Paeonia tenuifolia L. . . . . . . . . . . . . . . . . . . . . . . . . . Scandix australis L. . . . . . . . . . . . . . . . . . . . . . + . . . Stipa lessingiana Trin. et Rupr. . . . . . . . . . . . . . . . . . . . . . . . . . Chamaecytisus jankae (Velen.) Rothm. . . . . . . . . . . . . . . . . . . . . . . . . . var. Teucrium chamaedrys Teucrium chamaedrys L. . . . . . . . . . . . . . . . . . . + . I . . . . . Scandix pecten-veneris L. . . . . . . . . . . . . . . . . . . . . . . + . . Alyssum alyssoides (L.) L. . . . . . . . . . . . . . . . . . . . . . . . . . Salvia argentea L. . . . . . . . . . . . . . . . . . . . . . . . . . Lappula barabata (Bieb.) Gurke + + + . . + . . . . . + . . . . . . + . II . . + + . var. Rhodax canus Rhodax canus (L.) Fuss . . . . . . . . . . . . . . . . . . . . . . . . . Scutellaria orientalis L. subsp. pinnatifida . . . . . . . . . . . . . . . . . . . . . . . . . Artemisia pedemontana Balbis . . . . . . . . . . . . . . . . . . . . . . . . . Diagnostic taxa of higher units Al. Pimpinello-Thymion zygoidi Thymus zygoides Griseb. . 2a . . . 3 2b 2a 2m + 2a . . 2a . . 2a 2a + . III 2a 2a 2a 2b 3 Dianthus pseudoarmeria Bieb. . . + 1 2a . . 1 1 1 . 2a 1 + 1 3 2a 1 . . IV . . . . . Pimpinella tragium Vill.ssp. titanophila (Wor.)Tutin . + . . . . . 2a 1 + 2a . . . . . . . + . II . . . 1 2a Satureja coerulea Janka . . . . . . . . . . . . . . . . . . . . . . . . . Scorzonera mollis Bieb. . . . . . . . . . . . . . . . . . . . . . . . . . 16 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia 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 57 58 59 60 61 62 63 8V 8V 8V 8V 8V 8V 10V 10V 10V 10V 10V 8V 8V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 9V 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 200 100 100 100 100 100 100 100 100 100 100 100 .. . . . 0 0 .. 0 0 . . 0 0 0 0 0 0 0 0 0 0 . 0 . 0 0 0 0 0 0 . 0 0 0 . 0 0 0 50 70 80 80 0 0 5 0 0 10 5 0 0 0 0 0 0 0 0 0 0 10 0 5 0 0 0 0 0 0 10 0 0 0 5 0 0 0 50 70 60 60 90 95 95 90 90 70 85 90 90 90 80 100 90 80 95 100 80 60 90 70 70 90 95 60 90 90 70 80 90 75 70 95 60 80 11 21 11 14 43 36 52 55 50 50 45 24 22 30 32 33 36 49 35 27 27 32 36 32 37 37 34 33 37 34 33 35 31 25 28 26 23 24 2a3 3 3VV 2a2b 3 3 VV . . . .IIV . . . .IIV . 2a . . IIIIII 1 . . . IIIIII 2a. . .III 1 . 1+VII . 32b3IVII + . . .IVII .... I .... .... .... .... .+. .III .... .... .+. .II .+. .III .... .... . .+.IIII ... . ... . ... . 2a. . .IVIII .... III 2a. . .IIII .... .... . 1 .2a. . 1III. . 12a+ . 3 . . . 1 . 3 . 12a1III12a. 11 .2a. 12aIVIII ....... ................. ......... . ....... ................. ......... . ...1++.III...........+..+..I.......... I ....... ................. ......... . +.2a11..III..+...........12a.I.......... II ....... ..............1..I.......... I . . . . . . . 2a11++2a2a . .1.2a2a .+1.IV2a2a2b .2a12a1+.IVIII ....... ................. .......... ....... ................. .......... +2a12a131V2a2a2a2a +2a2a .2a2a2a2a 11112aV1112a++1. .+IVV . . +2m2a2m+ IV2a+2m2a2m2m+ 2a2m2b2m32m2b2m . + V2m. + +2a .2m+ .2mIVV . . .2a3. .II2b2a . .2m+ 2b. .32b2b2b2a2a2a .IV32b332b2a 333.VIV 2m2a2a2a2a . . IV+ 2a2m2b2a2m+2m2a2m . . 2a + . . . IV2m2m2m2a . + . 2m . . IIIIV 12a2a2a 112bV. .2b1.2b+. . . . . . . . . .II2b32a2b 332a.+.IVIII . . . . . . . . .2b2a 3+3. .2m2a +3.2m.III.++2m+ +++. .IVIII . . . . .1.I. .2b.3.11412a32a1333IV+. . . . . . . . .IIII .+. . . . .I2a2b2b 1. . . . .33312a2b 31IV. .11. .1. .1IIIII 1 1 2b 2b 2a . 2a V . . . 2a . . + 1 + . . . . . . . . II . . . 2a . . . 1 . . I II 2a + 2b 2a 1 2b 2a V . . . . . . . . . . . 3 . . . . . I . . . . . . . . . 1 I II 2m 2m 2m + + 2m . V . . . . . . . . . . . . . 2m . . . I . + . . . . . . . . I II + + + r + 1 1 V . . + . + . . . . + . . . + . . . II . . . . . . . . . . II . . + + + + + IV . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . 3 2a + 3 2b 2b 2a 2m 3 . V II . . . . . . . . . . . . . . . . . . 2a 2a . . . . I 2b 2a 3 2b 2b 2a 1 + 2a . V II . . . . . . . . . . 2a . . + . . . . . 2a . 1 1 1 II + 1 2a . 1 1 2a 1 + . IV III . . . . . . . . . . . . . . . . 3 3 3 3 2a . 3 3 III 3 3 3 3 3 2a 3 3 3 3 V III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + 1 1 . . . . 2b . III I . . . 2b 1 3 1 III . . . . . . . . . 2a 3 3 2a . 2b . . II . 2a 2a 2a 1 2a 3 . 1 2b IV III . . . . . . . . . . . . . 1 . . . 1 1 . . + . 1 II . . . . . . . . . . I 17 Hacquetia 5/1 • 2006, 5–23 Number of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Goniolimon besseranum (Schult.ex Reichenb.) Kusn. . . . . . . . . . + . + . . . . . . . . I . . . . . Al. Festucion valesiacae Festuca valesiaca Schleich..x Gaudin . . . . 2m . 2a 2m 2a 2m 2a 2a 2m 2a 2a 2a 3 3 + . IV 2b 2a 2m + . Inula oculus-christi L. . . . . . . . . . . . . . . . . . . . . . . . . . Convolvulus cantabrica L. . . . . . . . . . . . . . . . . . . . . . . . . . Salvia nutans L. . . . . . . . . . . . . . . . . . . + . I . . . . . Adonis vernalis L. . . . . . . . . . . . . . . . . . . . . . . . . . Medicago minima (L.) Bart. . . 2a . 2b . . . . . . . . . . . . . . . I . . . . . Chrysopopon gryllus (L.) Trin. . . . . . . . . . . . . . . . . . . . . . . . . . Vinca herbacea Waldst. et Kit. . . . . . . . . . . . . . . . . . . . . . . . . . Scorzonera hispanica L. . . . . . . . . . . . . . . . . . . . . . . . . . Ord. Festucetalia valesiacae Teucrium polium L. . . . 1 + . . 1 2a 2a 1 1 . + 2a 2a 2a 1 1 . IV . 2a . . . Cephalaria uralensis (Murr.) Roemer et Schultes . . . . + 2a 2a 1 1 1 2a 2a 1 + . . . 2a 1 . III 2a . . 1 2b Iris pumila L. . . . . . . . . . . . . . . . . . . + . I . . . . . Stipa capillata L. . . . . 2a . . . . . . 3 . . . . . . . . I . . . . . Poa bulbosa L. . . . . . . . . . . . + . . . . . . . . I . . . . . Jurinea mollis (L.) Reichenb. . . . . . . . . . . . . . + . . . . . . I . . . . . Tragopogon dubius Scop. . . . . . . . . . . . . . . . . . . + . I . . + . . Coronilla varia L. . . . . . . . . . . . . . . . . . . . . . . . . . Campanula sibirica L. . . . . . . . . . . . . . . . . . . . . . . . + 1 Cl. Festuco-Brometea Astragalus glaucus Bieb. + 3 3 2a 2b 2b 3 3 3 2b 3 2b 2b 2a 2b 1 1 1 + + V . . . . . Echinops ritro L. 1 + . 1 + . . . . + . + . . . . . 1 . . II 2a 2a + . 1 Jurinea stoechadifolia (Bieb.) DC. 1 2a 1 2a 1 2b 3 2b . 2a 3 3 + . 3 1 2b 2a 4 . IV . . . . . Linum austriacum L. . . 1 2a 2a + 1 2a 2a + + 1 . . 1 1 1 + + . IV . . . . . Crupina vulgaris Cass. . . 1 2a . + 1 . . . . + . . . . . + 1 . II + + + . . Alyssum hirsutum Bieb. subsp. hirsutum 2a 2a 2m 2m . . + . . . . 2m . . 2m . . . . . II . . + . . Ephedra distachya L. . . 2a 2a . . 3 . . . . . . + 1 2b + 2a . . II . . . . . Euphorbia nicaeensis All. subsp. nicaeensis . 2a . 1 + + 1 1 1 . . . . + + . . . . . III . . . + . Achillea clypeolata Sm. 1 + . 1 . . . . + . . . . . . . . . . . I 2a 2a . 1 . Seseli tortuosum L. . . . . . . . . . . 1 . + 1 . . . . . . I 1 + + . . Eryngium campestre L. . . . . . . . + . + . . . . . . . . . . I . . + . . Sanguisorba minor Scop. . . . . . . . . . . . . . . . . . . . . . . . . . Erysimum diffusum Ehrh. 1 . . . . . . . . . . . . . . . . . . . I . . . . . Centaurea rhenana Boreau . . . . . . . 1 . . . . . . . . . . + . I . . . . . Galium octonarium (Klokov) Soo . . . . . . . . . . . . . . . . . . . . . . . . . Linaria genistifolia (L.) Miller. . 1 1 1 . . . . . . . . . . . . . . . . I . . . . . Sternbergia colchiciflora Waldst.et Kit. . . . . . . . . . . . . . . . . . . . . . . + . . Hyacinthella leucophaea (Steven .ex Kunth.) Schur. . . . . . . . . . . . . . . . . . . . . . . + . . Adonis volgensis DC. . . . . . . . . . . . . . . . . . . . . . . . . . Allium rotundum L. . . . . . . . . . . . . . . . . . . . . . . . . . Artemisia austriaca Jacq. . . . . . . . . . . . . . . . . . . . . . . . . . Astragalus onobrychis L. subsp. chlorocarpus . . . . . . . . . . . . . . . . . . . . . . . . . Carlina vulgaris L. . . . . . . . . . . . . . . . . . . . . . . . . . Filipendula vulgaris Moench. . . . . . . . . . . . . . . . . . . . . . . . . . Galium verum L. . . . . . . . . . . . . . . . . . . . . . . . . . Hieracium echioides Lumn. . . . . . . . . . . . . . . . . . . . . . . . . . Medicago falcata L. subsp. falcata . . . . . . . . . . . . . . . . . . . . . . . . . Minuartia setacea (Thuill.) Hayek subsp. setacea . . . . . . . . . . . . . . . . . . . . . . . . . Plantago media L. . . . . . . . . . . . . . . . . . . . . . . . . . Poa angustifolia L. . . . . . . . . . . . . . . . . . . . . . . . . . Salvia nemorosa L. . . . . . . . . . . . . . . . . . . . . . . . . . Inula salicina L. . . . . . . . . . . . . . . . . . . 1 . I . . . . . Orobanche purpurea Jacq. . . + + . + + . . . . . . . . . . . . . I . + . . . Silene bupleuroides Chater et Walters . . . . . . . . . . . . . . . . . . . . r . . . . 18 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia 26272829 30313233343536 3738394041424344454647484950515253 54555657585960616263 .... I....... ................. ......... . . . . .IIIIV . 433332aIV. . 3 .2a2b2b . 332a2a2b 3433IV332a2b2a2a2a 3 . 3VIV . . . . 12a2a 11. .IV. . . .2a111. . .+.1. . .II2b2a2a2a2a2a2a2a2a 1VIV . . . . . . +2a+12aIV. . .2a2a 12a11 . .2a112a.2aIV. . . . . . . . . . III .... I++11++2aV...++.1.r+2a..++++III+..+......IIII .... ..2a2a .+1III..+..2a...........I2a2b2a2a2a2a 1.+.IVIII . . . . I.2a2a .2m2m+ IV. . . . .+.2a. . . .2a+. . .II+. . . . . . . .2mIII .... ..2a....I................. .......... I .... 1.++..1III..+...1..........I.......... I .... ....... .............+..+I.......... I . . . .IIII. .12b+2a2a IV2a. . . .2a12a. . .2a. .2a. .II2a2a ++1+12a12aVIV 2a...IIIIII ....... 2a1.2a..11...2a..132aIII.1........III . . . . . I 1 12a2b2b2b2a V2a2b2b2b 12a32b+2a2b 32b32a2b2a V2a2a2b2a 32b3 .2b2a VV . . . . .I.+. . . . .I.2b.3. .2a+. . . . .1. . .II1+. . . . . . . .III .....I..32b2b 33IV........2m....2a...I.......... II .....I....... .........2b.2a..+..I.......... I ..++III....... ................. .......... ..... ....+..I................. .......... I +...III....... ................. ......... . .... IV....... ..............11+I.......... I 1...IIIIII ....... ................. ......... . .... III....... .1...............I.......... I .... III2a1.2a2a ..III.1..11........2a1.II.......... II .2a.+IIIIII ....... ++............+11II....+.....II .+++IIIII . . . . .+.I. . .+. .+. .2m.2a. .+.+II. . . .++++. .IIII .... II....... ................. ......... . . . . .III2a2a +1+1.V+. . .1+.+.2a. .+2b+11III11.+. . .+1.IIIIV . . . .IIII+. . . .2a1III2b12b112a2a 12a2b2a 12a32a2a2a V+12a+2a2a +2a.2aVV ....IIII...11..II2a2a 11...+......12b.III.......... II . . . .II1+111+2aV++.++11++112a1+.2a.V+1+. . . .+. .IIIV . . . . . .+112a.III++2a2a 12a.+2a. . . . .2a.2aIII11111+.+. .IVIV .+++III2a.2a12a.+IV.11++1.1. .1. .1. . .III+2a.111.1.+IVIII .... I...+1..II.............1...I....++..++IIII .... ....... ......+.+.11.2a+..II1....2a....III ...+II.....+.I................. ........1.II ....II....... .....1+..........I.......... I ....II++..+..III......+....+..+..I.......... I .... ...2b1..II................. ........1.II .... ......+I................. .......... I .... 2a.2a..2a2a III................. .......... I .... +1+++.+V................. .......... I .... ....... ................+I.......... I .... .1..1..II..+1....+.....+..II.......... I .... +......I..............1..I.......... I .... .....2a.I................. ..+.......II .... +..12a.2aIII......1.........2aI.......... I .... +2m .....II......2m..........I.......... I .... +......I................. .......... I .... 1.2b1...III....+............I...1......II .... +......I................. .......... I .... I....... ................. ......... . ....II....... ................. ......... . ....II....... ................. ......... . 19 Hacquetia 5/1 • 2006, 5–23 Number of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Other taxa Tanacetum millefolium (L.) Tzvel. 1 1 1 3 . . . . . . . 1 . + + 2b . . . . II . . 1 2a . Adonis flammea Jacq. 2m . . . . . . . . . . . . . . . . . + . I + + . . . Valerianella pumila (L.) DC. . . . . 2m . . . . . . . . . . . . . . . I 2m + + . . Onosma echioides L. . . . 2a . 2a + 1 1 . . . + . 1 . . . + . II . . 2a . . Reseda luteola L. . 1 . . . 1 . 1 + . . . . . 1 . . . + . II . . . . . Crepis sancta (L.) Babcock . . . . . . . . . . . . . . . . . . . . + + + . . Bromus racemosus L. . . . . . . . . . . . . . . . . . . . . . . . . . Crataegus monogyna Jacq. . . . . . . . 1 . . . . . . . . . . . . I . . + . . Euphorbia helioscopia L. + . . . . . . . . . . . . . . . . . . . I . . . . . Trigonella gladiata Steven ex Bieb. . . . . . . . . . . . . . . . . . . . . + + + . . Thlaspi arvense L. . . + + . . . . . . . . . . . . . . . . I + + . + . Ziziphora capitata L. . . . 2m . . . . . . . . . . r . . . . . I . r r . . Leontodon hispidus L.subsp. hyspidus . . . . . . . . + . . . . . . . . . 1 . I . . . . . Vicia peregrina L. . . . . 2a . . . . . . . . . . . . . . . I . . + . . Cichorium inthybus L. . . . . . . . + + . . . . . . . . . . . I . . . . . Orobanche amethystea Thuill. . . . r . . . . . . . . . . . . . . . . I . . . . . Xeranthemum annuum L. . . . . . . . . . . . . . . . . . . . . . . . . . Ornithogalum gussonii kochii Parl. . . . . . . . . . . . . . . . . . . . . . . . . . Senecio vernalis Waldst. et Kit. . . + . . . . . . . . . . . . . . . . . I . . . . . Achillea millefolium L. . . . . . . . . . . . . . . . . 1 . . . I . . . . . Astragalus hamosus L. . . . . . . . . . . . . . . . . . . . . . . . 1 . Avena eriantha Durieu . . . . . . . . . . . . . . . . . . . . . . 3 . . Alyssum minutum Schleicht. ex DC . . . . . . . . . . . . . . + . . . . . I . . . . . Gypsophilla glomerata Pallas ex Bieb. . . . . . . . . . . . . . . . . . . . . 2a 1 . . . Bupleurum rotundifolium L. . . + . . . . . . . . . . . . . . . . . I . r . . . Euphorbia agraria Bieb. . . . . + . . . . . . . . . . . . . . . I . . . . . Viola kitaibeliana Schultes . . . . + . . . . . . . . . . . . . . . I . . . . . Haplophyllum suaveolens (DC.) G. Donf. . . . 2a . . . . . . . . . . . . . . . . I . . . . . Rosa canina L. . . . . . . . 1 . . . . . . . . . . . . I . . . . . Erodium ciconium (L.) L’Her. . . . . . . . . . . . . . . . . . . . . . . . . . Asparagus verticillatus L. . . . . . . + . . . . . . . . . . . . . I . . . . . Bromus tectorum L. . . . . . . . . . . . . . . . . . . . + I . . . . . Linum tauricum Willd. subsp. tauricum . . . 1 . 2a . 2a 1 1 . + . . . 1 . . 2a . II . . . . . Jasminum fruticans L. . . . 2a . 1 . . . . . . . . . 1 1 + . . II . . . . . Galium flavescens Borbas 2a 1 . . . . . . . . . . . . . . . . . . I . . . . . Paliurus spina-christi Miller . . 1 . . . . + . . . . . . . . . . . . I . . . . . Prunus mahaleb L. . . + . . . . . . . . . . . . . . . . . I . . . . . Asparagus maritimus (L.) Miller . . . 1 . . . . . . . . . . . . . . . . I . . . . . Coronilla emerus L. subsp. emeroides . . . 1 . . . . . . . . . . . . . . . . I . . . . . Cotinus cogyggria Scop. . . . . 1 2a . . . . . . . . . . . . . . I . . . . . Fraxinus ornus L. . . . . . 1 . . 1 . . . . . . . . . . . I . . . . . Colutea arborescens L. . . . . . . . . . + . . . . . . . . . . I . . . . . Allium saxatile Bieb. . . . . . . . . . . . 1 . . . . . . . + I . . . . . Genista sessilifolia DC. subsp. trifoliata . . . . . . . . . . . . . . 2a . . . . . I . . . . . Tamarix tetrandra Pallas ex Bieb. . . . . . . . . . . . . . . . . . 1 . . I . . . . . Carduus pycnocephalus L. . . . . . . . . . . . . . . . . . . . . . + + . . Papaver dubium L. . . . . . . . . . . . . . . . . . . . . + . . . . Medicago disciformis DC. . . . . . . . . . . . . . . . . . . . . . . 2m . . Acinos suaveolens (Sibth. et Sm.) G. Don . . . . . . . . . . . . . . . . . . . . . . . . . Cladonia rangiformis Hoffm. . . . . . . . . . . . . . . . . . . . . . . . . . Euphorbia nicaeensis All. subsp. candilathri . . . . . . . . . . . . . . . . . . . . . . . . . Minuartia hybrida (Vill.) Schischkin . . . . . . . . . . . . . . . . . . . . . . . . . Sedum sartorianum Boiss. subsp. ponticum . . . . . . . . . . . . . . . . . . . . . . . . . Euphorbia myrsinites L. . . . . . . . . . . . . . . . . . . . . . . . . . Asphodeline lutea (L.) Reichenb. . . . . . . . . . . . . . . . . . . . . . . . . . Marrubium peregrinum L. . . . . . . . . . . . . . . . . . . . . . . . . . Carduus acanthoides L. . . . . . . . . . . . . . . . . . . . . . . . . . Ajuga chamaepytis (L.) Schreber . . . . . . . . . . . . . . . . . . . . . . . . . 20 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia 26272829 30313233343536 3738394041424344454647484950515253 54555657585960616263 . . .+IIII331.2a2b2a V32a2a2b2b2b2b2a2b .2a+. . .2a+IV+2b2a 1. .2a. . .IIIIV . ++ .IIIII +2a2a2m2m2m+ V2m.2m. +2m+2m2m+2m+2m2m . ++V. . . .2m. . ++ . IIIV . . + +IIIII 2m. . . +2a2a III2a2m . . . .2m2m2a . +2a2a2m+ 2a+IV. . . . + . . + . . IIII . . . .III+. . . .2b2a III1.11.2a. . .12a1. .2a2a 1III1.2a. . . . . . .IIII .... II....... ................. ...+......II . + . .IIII2m2m2a + .2m2a V2a2a2a2a2a2b .2a2a . . .2m2a . . .III. . .2m2m . .2m. . IIIII .+. .II.+1+1.1IV. . .+2a. . . . .+. . .+. .II1. . .11. . . .IIII . . . .II+1+1.2a.IV. . . . .1. . . .1.1. . . .I+. .+. . . . .2bIIII .... I..+...+II.2a1.+1+1+........III........1.III .2a. .IIII. . .2a+2m2a III. . .+. .+2a. .++. . .+.II. . . .+. . . . .III . . . .III. .2m+ +2m+ IV2a.+++2m . . . . .2a2m+ . . .III. . . . . . . . . . II ....III....... ........r........I.......... I .... I....... .........2a..1....I1.....1...II . . . .II. . . . .2a.I. . . . . .+.1. . . . . . . .I. . . . . . . . .+II .... I....... ......+......1...I.+........II ..+.II....... .....+...........I.1........II .+.+III....... 1.11.........2a++.II.......... I .+..II...1.++III...........1....+I.......... I .... I+1.....II......1+.........I.......... I .... I3312a1..IV........+........I.......... I .+..III......+I.......+.........I.......... I ....II....... ....3..3.........I.......... I .... I......2aI.......2m.........I.......... I ....III....... ............+....I.......... I ....II....... ......+..........I.......... I .... I....... ...........2a.....I.......... I .... I....... ......2m..........I.......... I .... I..2a.1..II................. .......... I .... I++.....II................. .......... I .+.1III....... ................. .........+II .... I....... ................. ........+.II .... I....... ................. .........2aII .... II....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .+..II....... ................. ......... . .... I....... ................. ......... . .... I....... ................. ......... . .+.+IIII....... ................. ......... . ....II....... ................. ......... . ....II....... ................. ......... . .+..II....... ................. ......... . . . . . . .+. .2m.II. . .+. .+2a+2a2b2b 3. . . .III2a2m2m2a .+. . .2aIIIIII . . . . .+12a2a2a 1V+.12a1.+.2a.2a. . . .11III. .2a1++. . .2aIIIIII .... ....... ......+...2m+ .....I.2m2m+ ..+...IIII .... ....2m..I.....2a.2a..2a.2a2a ..2mII.++...2m...IIII .... ....... .........2b2a2b .2a...II..++..2a...IIII .... ....... ++.+1.+2a....+..+.III+..+......III .... 2a+.2a1+1V....+.1......2a...I....+..+..III .... ++1....III.....+...+.......I..+...+...III .... .2a1.2b+1IV..........+...1+.I.......+..III 21 Hacquetia 5/1 • 2006, 5–23 Number of relevé 1 2 3 4 5 6 7 8 9 1011121314151617181920 2122232425 Erodium cicutarium (L.) L’Her. . . . . . . . . . . . . . . . . . . . . . . . . . Aegilops geniculata Rothm. . . . . . . . . . . . . . . . . . . . . . . . . . Ruta graveolens L. . . . . . . . . . . . . . . . . . . . . . . . . . Tortella flavovirens (Bruch) Broth. . . . . . . . . . . . . . . . . . . . . . . . . . Thymus callieri Borbas ex Velen. subsp. urumovii . . . . . . . . . . . . . . . . . . . . . . . . . Centaurea marshalliana Sprengel . . . . . . . . . . . . . . . . . . . . . . . . . Erophila verna (L.) Chevall . . . . . . . . . . . . . . . . . . . . . . . . . Sherardia arvensis L. . . . . . . . . . . . . . . . . . . . . . . . . . Filago vulgaris Lam. . . . . . . . . . . . . . . . . . . . . . . . . . Cladonia foliacea (Huds.) Schaer. . . . . . . . . . . . . . . . . . . . . . . . . . Muscari racemosum DC. . . . . . . . . . . . . . . . . . . . . . . . . . Thymus pannonicus All. . . . . . . . . . . . . . . . . . . . . . . . . . Arabis recta Vill. . . . . . . . . . . . . . . . . . . . . . . . . . Bromus mollis L. . . . . . . . . . . . . . . . . . . . . . . . . . Carpinus orientalis Miller . . . . . . . . . . . . . . . . . . . . . . . . . Nonea pulla (L.) DC. . . . . . . . . . . . . . . . . . . . . . . . . . Silene conica L subsp. conica . . . . . . . . . . . . . . . . . . . . . . . . . Arenaria serpilifolia L. . . . . . . . . . . . . . . . . . . . . . . . . . Tortulla ruralis (Hedw.) Crome . . . . . . . . . . . . . . . . . . . . . . . . . Sideritis montana L. . . . . . . . . . . . . . . . . . . . . . . . . . Carthamus lanatus L. . . . . . . . . . . . . . . . . . . . . . . . . . Papaver hybridum L. . . . . . . . . . . . . . . . . . . . . . . . . . Platago lanceolata L. . . . . . . . . . . . . . . . . . . . . . . . . . Ajuga genevensis L. . . . . . . . . . . . . . . . . . . . . . . . . . Anthemis tinctoria L. . . . . . . . . . . . . . . . . . . . . . . . . . Centaurea napulifera Rochel subsp. thirkei . . . . . . . . . . . . . . . . . . . . . . . . . Coronilla scorpioides (L.) C. Koch. . . . . . . . . . . . . . . . . . . . . . . . . . Lamium amplexicaulis L. . . . . . . . . . . . . . . . . . . . . . . . . . Ranunculus oxyspermus Bieb. . . . . . . . . . . . . . . . . . . . . . . . . . Veronica prostrata L. . . . . . . . . . . . . . . . . . . . . . . . . . Centaurea caliacrae Prodan . . . . . . . . . . . . . . . . . . . . . . . . . Androsace maxima L. . . . . . . . . . . . . . . . . . . . . . . . . . Bellevalia ciliata (Cyr.) Ness . . . . . . . . . . . . . . . . . . . . . . . . . Dactylis glomerata L. . . . . . . . . . . . . . . . . . . . . . . . . . Geranium pusillum L. . . . . . . . . . . . . . . . . . . . . . . . . . Stachys recta L. subsp. subcrenata . . . . . . . . . . . . . . . . . . . . . . . . . Saxifraga tridactilites L. . . . . . . . . . . . . . . . . . . . . . . . . . Taraxacum erytrospermum Andrz. ex Besser . . . . . . . . . . . . . . . . . . . . . . . . . Anthemis ruthenica Bieb. . . . . . . . . . . . . . . . . . . . . . . . . . Digitalis lanata Ehrh. . . . . . . . . . . . . . . . . . . . . . . . . . Localities: 1 – Eastern from the town of Balchik; 2, 3, 5 and 6 – Cape “Ikantulukö; 4, 7, 15, 16, 17 and 18 – north from “Tuzlata” Resort; 8, 9, 10, 11, 12, 13, 14 and 19 – Topola village; 20 – “Bolata dere”; 21, 22, 23, 24, 25, 26, 27, 28 and 29 – “Chirakman” locality; 30 and 31 – “Kayryaka”, Bozhuretz village; 32, 33, 34, 35 and 36 – “Gyorensko dere”, Vidno village; 37 and 38 – “Kaliakra” Reserve, the Castle; 39, 40, 41, 42, 43, 44, 54, 55, 56, 57, 58, 59, 60, 61 and 62 – “Kaliakra” Reserve, the road to the “Bolata dere”; 45 and 50 – Kamen bryag village, north of “Yaylata” locality; 46, 47, 48, 49, 51, 52 and 53 – game-breeding farm “Kushlata”, north of the “Bolata” Inlet; 63 – Taukliman, “Roussalka” Resort. 22 Rossen tzonev, veska Roussakova, MaRius DiMitRov: tHe WesteRn-Pontic stePPe vegetation in BulgaRia 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 57 58 59 60 61 62 63 . . . . + . + . . . 2m III . . + . . 2m + . . . . . . 2m . + . II . . . . . . . . . . II . . . . . . . 2m . + . II . . . . . . + + 3 + . . . . . + . II . . . . . . . . . . II . . . . . . . . . . . . . . . . . . . . 1 1 . 3 . . . . I . . . 1 . . 2a . 3 . II I . . . . . . . . . . . . . . . . . 2m . . . . . . . . . . I 2a + 2a . . . . . . . II I . . . . . . 2a 2a . . . II . . . . . 2a . . . . . . . 2b . . . I 2m . . . . . . . 2m . I I . . . . . . . . . . . . . . . . + . . . . . . . . . . + I + . . . . . . . . 1 I I . . . . . . 2m . . . + II . . . . . . . . . . . . . . . . . . + . . . . . + . . I I . . . . . . 2a . . . . I . . . . + . . 1 3 . . . . . . . . I . . . . . . . . . + I I . . . . . . . + . . + II . . . . . 1 . . . . . . . . . 1 . I . . . . + . . . . . I I . . . . . . . 2a + 2a . III . . . . . . 2a . . . . . . . . . . I . . . . . . . . . . I . . . . . + . 1 . 1 1 III . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . + 2a . . . . 2a III . . . . . 2a . . . . . . . . . . . I . . . . . . . . . . I . . . . . . . . . 2m . I . . . . . . + . . . . . . . . . . I . . . . . . . . . . I . . . . . . + . . . . I . . . . . . . . . . . . . . . . + I . . . . . . . . . . I . . . . . . . . . 2a . I . . . . . . . . . . . . . . . . + I . . . . . . . . . . I . . . . 1 . . . . . . I . . . . . . . . . . . . . + . . . I . . . . . . . . . . I . . . . . . + . . . . I . . . . . . . + . . . . . . . . . I . . . . . . . . . . I . . . . . . 2a + + . . III . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . 2a 2a 2a . III . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . + + + . III . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . 1 1 . . . II . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . 1 + . . . II . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . + . . 1 . II . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . 2a . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . . . 2a . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . 2a . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . . . + . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . + . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . 1 . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . 1 . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . I . . . . . . . . . . . . . . . . . . . . + 2a . . . . . . I . . . . . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . + . I . . . . . . . . . . I . . . . . . . . . . . . . r . . . . . . . . . . . . . . I . . . . . . . . . . I . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . I . . . . . . . . . . I . . . . . . . . . . . . . . . . . . 2a . . . . . . . . . I . . . . . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . . . . . + I . . . . . . . . . . I . . . . . . . . . . . . . . . . . . . . . . . + . . . . I . . . . . . + . . . I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 I I 2 HACQUETIA 5/1 • 2006, 25–36 HAloPHIlE VEgETATIon oF THE sloVEnIAn sEACoAsT: THERO-SALICORNIETEA AnD SPARTINETEA MARITIMAE Mitja kALIgARIČ*,** et Sonja ŠkoRnIk* Abstract Halophile vegetation of the Slovenian sedimentary seacoast have been sampled with the standard Braun- Blanquet procedure. All the 140 collected relevés have been classified using the Syn-TAx 2000 software. The resulting dendrogram separated 5 well-defined clusters, characterized by different dominant species. Two well separated clusters have been further elaborated in this study. The floristically-poor association Suaedo maritimae-Salicornietum patulae Brullo et Funari ex géhu et géhu Franck 1984 tend to form monodominate stands with Salicornia europaea s.l. on mudflat hypersaline stands. The association Suaedo maritimae-Bassietum hirsutae Br.-Bl. 1928, with high abundance of Suaeda maritima occupy smaller surfaces on drier stands. Both were classified within halophile annual swards of the class Thero-Salicornietea. Spartina maritima-dominated perennial halophyte saltmarshes are represented with association Limonio-Spartinetum maritimae (Pignatti 1966) Beeft. et géhu 197 (class Spartinetea maritimae), which colonizes muddy islets, perturbed by high tide and sea turbulence and supports brackish water, which should be rich in nutrients. Key words: phytosociology, halophile vegetation, classification, north Adriatic, Thero-Salicornietea, Spartinetea maritimae. Izvleček S standardno Braun-Blanquetovo metodo smo vzorčili halofitno vegetacijo na sedimentni morski obali v Sloveniji. Vseh 140 zbranih popisov je bilo klasificiranih s pomočjo programa Syn-TAx 2000. nastali dendrogram je ločil 5 dobro definiranih šopov, okarakteriziranih z različnimi dominantnimi vrstami. Dva dobro ločena šopa sta bila nadalje obdelana v tej študiji. Floristično revna asociacija Suaedo maritimae-Salicornietum patulae Brullo et Funari ex géhu et géhu Franck 1984 teži h graditvi enovrstnih sestojev z vrsto Salicornia europaea s. l. na muljastih hipersalinih rastiščih. Asociacija Suaedo maritimae-Bassietum hirsutae Br.-Bl. 1928 z visoko abundanco vrste Suaeda maritima zaseda manjše površine na bolj suhih rastiščih. obe smo uvrstili v razred halofitnih enoletnic Thero-Salicornietea. obmorska močvirja z dominantno vrsto Spartina maritima predstavljajo asociacijo Limonio-Spartinetum maritimae (Pignatti 1966) Beeft. et géhu 197 (razred Spartinetea maritimae), ki kolonizira muljaste otočke, izpostavljene plimovanju in valovanju, in dobro prenašajo brakično vodo, ki mora biti bogata s hranili. Ključne besede: fitocenologija, halofitna vegetacija, klasifikacija, Severni Jadran, Thero-Salicornietea, Spartinetea maritimae. 1. InTRoDUCTIon substrate is almost perfectly matching with the ter ritory of Slovenia, only a part, in the Muggia/Milje Slovenia has only 46 km of seacoast, which is under peninsula and bay are lying predominantly in Italy. the pressure of urbanization, tourism and industry Flysch substrate results in a dense hydrological sys(port of koper). Its peculiarity is the geological tem above ground due to its impenetrable prop- substrate. With some exceptions, it consistes of cal-erties. Three streams/rivers have their mouths in careous sandstone – Eocene flysch substrate. This the gulf of Trieste. Alluvial deposits on the mouths * Department of Biology, University of Maribor, SI-2000 Maribor, koroška 160, Slovenia, mitja.kaligaric@uni-mb.si, sonja.skornik@uni-mb.si ** ZRS koper, Institute for biodiversity studies, garibaldijeva 1, SI–6000 koper, Slovenia 25