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Biosystems
Diversity
ISSN 2519-8513 (Print) ISSN 2520-2529 (Online) Biosyst. Divers., 2020, 28(3), 298-319 doi: 10.15421/012039
Received 10.06.2020 Received in revisedform
07.08.2020 Accepted 09.082020
M G. Kholodny Institute ofBotany, NAS of Ukraine, Tereshchenkivska st., 2, Kyiv, 01004, Ukraine. Tel.: +38-099-079-01-08 E-mail: ddub@ukr.net
Syntaxonomy and ecological differentiation of the pioneer vegetation of Ukraine. 2. Helichryso-Crucianelletea maritimae, Festucetea vaginatae, Koelerio-Corynephoretea canescentis classes
D. V. Dubyna, T. P. Dziuba, S. M. Iemelianova, P. A. Tymoshenko
M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kyiv, Ukraine
Dubyna, D. V., Dziuba, T. P., Iemelianova, S. M., & Tymoshenko, P. A (2020). Syntaxonomy and ecological differentiation of the pioneer vegetation of Ukraine. 2. Helichryso-Crucianelletea maritimae, Festucetea vaginatae, Koelerio-Corynephoretea canescentis classes. Biosystems Diversity, 28(3), 298-319. doi:10.15421/012039
Pioneer psammophytic vegetation is usually developed on wind-drift sandy substrates such as arenas, spits, beaches, river terraces, and this vegetation occupies significant areas in all three natural zones of Ukraine. The Koelerio-Corynephoretea canescentis class was represented by 13 associations, 3 alliances and 1 order, Festucetea vaginatae class by 22 associations, 2 alliances and 1 order, Helichryso-Crucianelletea maritimae by 10 associations, 4 alliances and 1 order. The results of cluster analysis and synoptic tables of the classes are presented. 9 alliances are briefly described. Leading factors of territorial and ecological differentiation are identified. It was found that the territorial distribution of plant communities is influenced by the character of ecotope mesorelief, soil composition and humus horizon thickness, as well as the degree of eolian processes development. The main factors of their ecological differentiation are soil acidity, salt regime and ombroregime. Based on the results of DCA-ordination of syntaxa within certain vegetation classes, it was found that their distribution is influenced by factors that correlate with the environment-specific conditions. It has emerged that an ecological differentiation of syntaxa within Festucetea vaginatae is determined by the integrated effect of gradients, and soil salinity is leading among them. Temperature regime and climate continentality are leading factors in the distribution of syntaxa within the Koelerio-Corynephoretea canescentis class. The gradients of ombroregime and soil humidity have a significant impact. The distribution of communities of the Helichryso-Crucianelletea maritimae class in the ecological space is determined mainly by factors of variability of damping, ombroregime and climate continentality. The author's syntaxonomic concept assumes the independence of the studied classes: Koelerio-Corynephoretea canescentis, Helichryso-Crucianelletea maritimae and Festucetea vaginatae, considering that the leading factors of community differentiation of the Festucetea vaginatae and Koelerio-Corynephoretea canescentis classes are the origin (genesis) of sandy substrates, as well as soil acidity. Phytosociological analysis of a large number of relevés of coastal littoral vegetation also provides support for independence of the Helichryso-Crucianelletea maritimae and Ammophiletea classes different floristically and ecologically. A review of the psammophytic vegetation of Ukraine will determine the place of the selected syntaxonomic units in the pan-European system.
Keywords: psammophytic vegetation, classification, new-formed ecotope, ecological gradients, cluster analysis, DCA-ordination.
Corynephoretea canescentis - "Klasse Koelerio-Corynephoretea Klika in Klika et Novak 1941 - Sandtrockenrasen und Felsgrusfluren von der submeridionalen bis zur borealen Zone" (Dengler, 2004), Helichryso-Crucianelletea maritimae - "Classification of European and Mediterranean coastal dune vegetation" (Marceno et al., 2018). However, discussions about syntaxonomic content and the structure ofhigher classification units of psammophytic vegetation are still ongoing.
The main areas of arenas in Ukraine are located in Polissia, and the smaller one in the forest-steppe zone. The largest continuous sand massifs in the steppe zone are located in the lower valley of the Dnieper River. This territory is called "Oleshkivsky Sands", and it occupies about 200 thousand hectares. Large areas are occupied by "Pridonetski Sands" (80 thousand ha) and sand areas on coasts, spits and islands of the Black and Azov Sea, Lower Danube and the Southern Bug River.
The macrorelief of sand massifs is mostly flat with small differences in elevation. However, the greatest importance in the distribution of plant communities belongs to the mesorelief of the sands, which is usually characterized by an alternation of high, up to 3-5 m, rounded (hills) or elongated dunes and fairly deep depressions. The topography and vegetation of wind-drift sands are mutually determining factors, since the absence of vegetation contributes to the dune formation, and their subsequent overgrowth promotes the formation of sand dunes, which over time are somewhat leveled in the hilly ones (Gordienko, 1969).
Introduction
Pioneer psammophytic vegetation developed on wind-drift sandy substrates such as arenas, spits, beaches, terraces, etc., performs extremely important functions: biological, ecological, regulatory, recreational, and many others. It stabilizes loose scattered soils, creates preconditions for the development of vegetation in subsequent successional stages. The geocomplexes on which such vegetation occurs are sites of biological diversity formation and conservation. They are habitats for many plant and animal species, ways for dispersing their genetic material, and also serve as migration routes by which alien species spread actively. New-formed arena ecotopes are the centers of formation of endemic floristic complexes; they are characterized by generic endemism.
Syntaxonomy of psammophytic vegetation in Europe has been developed in sufficient detail. Classification schemes and characteristics were presented for the territories of the Czech Republic (Chytry, 2007), Slovakia (Valachovic et al., 1995), Germany (Berg et al., 2004), Poland (Matuszkiewicz, 2008), France (Bardat et al., 2004), Romania (Sanda et al., 2008), Italy (Biondi et al., 2014), Spain and Portugal (Rivas-Martinez et al., 2001), Bulgaria (Tzonev et al., 2009), Hungary (Borhidi, 2003), and other countries. An overview of syntaxonomic units of the highest and middle levels in Europe is presented in generalized works: "Vegetation of Europe..." (Mucina et al., 2016), for the class Koelerio-
Pioneer psammophytic vegetation is represented in Ukraine by three classes: Festucetea vaginalae, Helichryso-Crucianelletea maritimae and Koelerio-Corynephoretea canescentis. The syntaxonomy of vegetation of Festucetea vaginatae and Koelerio-Corynephoretea canescentis classes in Ukraine has been the subject of studies conducted by many authors. J. Vicherek on the territory of the lower and middle Dnieper River described Festucion beckeri alliance with 4 associations and, in fact, made an attempt to validate Festucetea vaginatae class, first identified by R. von Soo in 1968 (Vicherek, 1972). Secali-Stipetum borysthenicae association and two alliances: Verbascion pinnatifidi and Cynodonto-Teucrion polii were described for the coastal dune area in the Crimean Peninsula (Korzhenevsky, 1986; Korzhenevsky & Klyukin, 1990). Arena landscapes of the "Biruchiy Island" spit in the Azov Sea were studied by Dubyna et al. (1995). On the territory of the spit, they identified 8 lower-rank syntaxa belonging to Festucetea vaginatae class, and two syntaxa attributed by the authors to Chenopodietea class. Three associations were described as new for science. Psammophytic vegetation of the Bilosaraiska Spit and the sea coast near Mariupol city was studied by Androsova & Solomakha (1996). Didukh & Korotchenko (1996) recorded a Centaureo borysthenicae-Festucetum beckeri association in the southern part of the forest-steppe zone of Ukraine, near the mouth of the Vorskla River and on islands of the Dniprodzerzhinsky and Pechenezhsky reservoirs. Umanets & Solomakha (1999a, 1999b) conducted their research in the 1990s in the Central Black Sea region. In the Black Sea Biosphere Reserve, the vegetation syntaxonomy included without limitation psammophytic coenoses of Festucetea vaginatae class, within which the authors distinguished the Medicagini tenderiensis-Seselietalia tenderiensi order containing two alliances: Medicagini tenderiensis-Seselion tenderiensi and Melico chrysolepi-Ephedrion distachyae. On the territory of the Danube Biosphere Reserve communities of19 associations of the Festucetea vaginatae class were described (Dubyna et al., 1996, 2003). 11 associations of this class were identified on the territory of the National Nature Park "Dzharylhatskyi" (Dubyna & Dziuba 2005a; Davydova, 2019). In the Pryazovia spits: Obitochna, Berdianska, Kryva, Bezymenna, Shyrokinska, Bilosaraiska, Stepanivska and Fedotova, Tyshchenko (2006) recorded the distribution of communities of 9 associations belonging to the Festucetea vaginatae class; while three associations were newly identified in the course of surveys conducted by the author. In the estuarine region of the Dnieper River, Chynkina (2003) identified communities of one association. In addition to the southern regions, coenoses of the class were also observed in the Middle Dnieper River region, on islands of the Dnieper River within Kyiv city (Tsukanova, 2005; Aleshkina, 2011), as well as on the territory of the Kaniv Nature Reserve (Shevchyk & Solomakha, 1996; Shevchyk et al., 1996) where the authors identified a Artemisio dniproicae-Salicion acutifoliae alliance. Its syntaxonomic status is debatable, since such communities involving shrubs are more often classified as Salicetea purpureae Moor 1958 (Mucina et al., 2016).
Phytocoenoses of sandy steppes were also observed in the northern steppe and forest-steppe zone of Ukraine, on the left bank of the Dnieper River (Bairak, 1998; Gomlya, 2005; Dziuba et al., 2010; Chusova, 2019); there the phytocoenoses occupy much smaller areas and are not characterized by coenotic diversity.
The syntaxonomic structure of the class has also been considered in generalized regional publications (Tymoshenko, 1999; Korzhenevsky et al., 2003; Dubyna et al., 2004; Solomakha, 2008; Dubyna et al., 2009).
Within the scope of final surveys on psammophytic vegetation in the Black Sea region, D. Dubyna and T. Dziuba in the "Prodrome of the vegetation of Ukraine" identified a new alliance Artemisio arenariae-Festucion beckeri in the Festucetea vaginatae class and isolated from one the class Helichryso-Crucianelletea maritimae, in which Ephedro distachyae-Medicaginetalia romanicae order was allocated for the Ukraine territory together with the alliance Ephedro distachyae-Medicaginion romanicae (Dubyna et al., 2019a). Unfortunately, the attempt of Korzhenevsky & Kvitnitskaya (2014) to validate the alliances Verbascion pinnatifidi Korzhenevsky et Klyukin 1990 and Cynodonto-Teucrion polii Korzhenevsky et Klyukin 1990, as well as the vegetation associations of aeolian landforms in the Crimea, were not completely successful since expressis verbis "typus" should have been used to indi-
cate the type of syntaxon name, according to article 5 of the ICPN (Weber et al., 2000).
Coenoses of the Koelerio-Corynephoretea canescentis class were first studied in Ukraine by Vicherek (1972) using the Braun-Blanquet approach on riverside sands of the Middle Dnieper River, where he described the Veronico dillenii-Corynephoretum association with its two subassociations. Later, coenoses belonging to this class (within the boundaries of Sedo-Scleranthetea and Festucetea vaginatae) were described for the Kaniv Nature Reserve area (Shevchyk & Solomakha, 1996; Shevchyk et al., 1996). Didukh & Korotchenko (1996) noted the distribution of plant communities belonging to the same association in the southern part of the left-bank forest-steppe zone of Ukraine, in the lower valley of the Vorskla River, in the upper valley of the Berestova River, and on islands of the Dniprodzerzhinsky and Pechenizhsky reservoirs. On left bank of the Dnieper River, coenoses belonging to this class were recorded by Bayrak (1998) and Galchenko (2006). V. Shevchyk and V. Solomakha, and later V. Shevchyk with O. Polishko described the communities of Thymo pallasiani-Centauretum sumensis and Cladonietum associations as part of the vegetation on the Dnieper River pine-forest terrace which grows along the bank of the Kaniv Reservoir (Shevchyk et al., 1996; Shevchyk & Polishko, 2000; Polishko, 2001, 2005). On the Zhytomyr Polissia area, Yakushenko (2004) allocated the Artemisio campestris-Dianthetum borbasii association. Plant communities of the two associations were identified in area of the planned Korostyshivsky National Nature Park (Orlov & Yakushenko, 2005). Coenoses of the Helichryso-Jasionetum association were described on the territory of the Polissky Nature Reserve (Vorobyov et al., 1998), and of Thymetum pulegioido-serpylli association were identified in the Cheremsky Nature Reserve (Konishchuk, 2006). Senchylo (2010) analyzed mainly coenoses of the Koelerio-Corynephoretea canescentis class; the scientist performed phytocoenotic studies on 7 following plant associations in the Dnieper River floodplain area within the forest-steppe zone: Artemisio dniproicae-Sedetum sexangularis, Centaureo borysthenicae-Festucetum beckeri, Veronico dillenii-Secaletum sylvestri, Chamaecytiso ruthenici-Festucetum beckeri, Thymo pallasiani-Centauretum sumensis, Sedo sexangulare-Festucetum beckeri, Diantho borbasii-Agrostietum syreistschikovii. Among the syntaxonomic diversity of vegetation in the Pyryatinsky National Natural Park, Kovalenko (2016) identified communities of Veronico dillenii-Secalietum sylvestris, Linario odorae-Agropyretum dasyanthi and Chamaecytiso ruthenicae-Festucetum beckeri associations, which were assigned to the Festucetea vaginatae class. In Ukrainian Roztochia, Soroka (2008) identified Spergulo vernalis-Corynephoretum and Diantho-Armerietum associations.
Often plant communities of this class were described within Sedo-Scleranthetea and, contrariwise, phytocoenoses associated with areas of poor skeletal soils and crystalline outcrops were sensu lato assigned to Koelerio-Corynephoretea canescentis class. Also, the structures of Festucetea vaginatae and Koelerio-Corynephoretea canescentis classes were sometimes intermixed (Shevchyk et al., 1996; Gaiova, 2015; Kovalenko, 2016). Regional publications summarized the study of the class in Ukraine (Onyshchenko, 2006; Solomakha, 2008; Dubyna et al., 2019a).
One of the generalizing works on classified syntaxa of xerophytic psammophytic vegetation in the forest and forest-steppe zones of the plain area of Ukraine was Kuzemko (2009) which includes syntaxonomic processing of 391 relevés. On the basis of cluster analysis and comparison of synoptic tables, the author identified 12 association-level syntaxa within Koelerio-Corynephoretea canescentis class (incl. Sedo-Scleranthetea and Festucetea vaginatae) and two associations within Molinio-Arrhenathe-retea class. Phytosociological materials from Ukraine consisting of 179 relevés were included in the large-scale analysis of coastal dune vegetation in the Baltic-Atlantic, Mediterranean and Black Sea regions (Marceno et al., 2018). The authors analyzed a large database (11,769 vegetation plots) and created the first formal classification of European coastal dune vegetation, accompanied by an expert system. This allowed them to identify 18 middle-rank syntaxa (alliances) which belong to the classes Ammophiletea, Honckenyo-Elymetea arenarii and Koelerio-Corynephoretea canescentis, and providing a critical revision of the concept ofthe class Ammophiletea used in EuroVegChecklist.
The issues on syntaxonomic structure and independence of Koelerio-Corynephoretea canescentis, Festucetea vaginatae, and Helichryso-Cru-cianelletea maritimae classes are currently discussed. When they survey the areas where continental psammophytic steppes occupy significant areas and differ in phytocoenotic diversity, Ukrainian and Russian phytocoenologists are insistent on the independence of these classes (Demina, 2009, 2015; Dubyna et al., 2019a). Western European researchers more often adhere to a broad understanding of vegetation classes and suggest combining, in particular, Koelerio-Corynephoretea canescentis and Festucetea vaginatae (Mucina et al., 2016), as well as Ammophiletea and Helichryso-Crucianelletea maritimae (Marceno et al., 2018). Phytocoenotic materials of the Sedo-Scleranthetea class very similar to Koelerio-Corynephoretea canescentis (Kuzemko, 2009; Kuzemko et al., 2014; Mucina et al., 2016) are subject to generalized analysis.
The analysis of significant databases of phytosociological relevés allows one to solve many issues on construction and effectiveness of vegetation classification (Lengyel et al., 2018; Willner et al., 2019; Bondareva et al., 2019; Landucci et al., 2020), biodiversity definition (Sabatini et al., 2018), biogeographic distribution of vegetation and its ecological affinity (Chytry et al., 2019), and others. Currently, we have collected more than 1700 phytosociological relevés of psammophytic vegetation in Ukraine which includes more than 1000 relevés belonging to the classes Festucetea vaginatae, Helichryso-Crucianelletea maritimae, Koelerio-Corynephoretea canescentis, syntaxonomic processing of which, in our opinion, will make a certain contribution to studying of psammophytic vegetation in Europe and supplement its syntaxonomy.
In recent years, the ecological component of phytocoenoses has become increasingly used as an object of study and in classification proving (Çoban & Willner, 2019; Zhou et al., 2019; Willner et al., 2019), including for the study of vegetation dynamics (Cao et al., 2019), identifying the main ecological factors of plant communities' differentiation (Kuzemko et al., 2016; Korolyuk et al., 2018), assessing the impact of environment on the floristic richness in phytocoenoses (Jansons et al., 2016; Yousaf et al., 2016; Slezak et al., 2017). The method developed by Didukh (2012) for determining the main eco-factors of the distribution of plant communities by 12 gradients completes phytosociological studies of vegetation and allows us to determine the ecosystem state by indicators of their biotic components.
The purpose of the paper was to generalize the accumulated phytocoenotic materials and existing data on syntaxonomy of the pioneer psammophytic vegetation in Ukraine, to clarify its current state and to develop a syntaxonomic structure, to establish the leading factors of ecological differentiation of phytocoenoses on the basis of ordination analysis, as well as to discuss controversial issues related to the syntaxonomic content of the classes Festucetea vaginatae, Koelerio-Corynephoretea canescentis, Helichryso-Crucianelletea maritimae.
Materials and methods
The work is based on the materials of field studies carried out by the authors during 1984-2018 according to the Braun-Blanquet approach to floristic classification (Braun-Blanquet, 1964; Westhoff& van der Maarel, 1973). The standard size of plots was 4 x 4 m. Sometimes, in particular on elongated coastal zones and dunes, the plots have size 1 x 4 or 2 x 5 m. At the same time, requirements for structure uniformity of vegetation cover were met. Phytosociological analysis also included the relevés presented in above-mentioned publications of J. Vicherek, O. Tyshchenko, Y. Didukh, I. Korotchenko, O. Senchylo, O. Umanets, I. Solomakha, V. Solomakha, A. Androsova, O. Orlova, D. Iakushenko, V. Shevchyk, O. Polishko, O. Bayrak, N. Galchenko, M Soroka, A. Davydova, L. Gomlya, T. Chinkina, V. Korzhenevsky and A. Klyukin, and others. The database also included unpublished relevés of psammophytic vegetation from manuscripts and dissertations, kindly provided to us by A. Kuzemko, O. Kovalenko, O. Chusova, O. Senchylo, V. Kolomiychuk, T. Fitsailo, I. Goncharenko, V. Konishchuk, G. Tsukanova, Y. Gayova, D. Iakushenko, V. Konogray, I. Khomyak. The size of plots was not always specified in the relevés, but we used all the available ones.
The materials were ordered by development of a database of geobotanical relevés in TURBOVEG 2.79 (Hennekens & Schaminée, 2001). Interpretation of the phytosociological material was carried out in several steps. At the initial one, the entire database of geobotanical relevés of
halophytic, psammophytic and littoral vegetation in Ukraine (7388 relevés) was processed and divided into smaller groups based on their floristic differences using the method of two-way indicator species analysis (TWINSPAN), in particular its modified algorithm (Hill, 1979; Tichy, 2002; Rolecek et al., 2009) implemented in the JUICE 7.0 software package. "Pseudospecies" cut level was 0%, 5%, 15% and 30%. The Whittaker's beta was chosen as cluster heterogeneity measure (Whittaker, 1978). Further, the clusters were selected which corresponded to the classes of psammophytic vegetation, based on diagnostic species characteristics.
Since a significant number of relevés included only vascular plants, or when mosses and lichens were identidied only up to their genus, authors excluded mosses, lichens and algae species from processing in order to "align" the relevés. These species were added again at the final stages of compiling the phytocoenotic tables. Cluster analysis was carried out using the PC-ORD program after the rejection of a certain part of relevés related to Ammophiletea Br.-Bl. et Tx. ex Westhoff et al. 1946, Salicetea purpureae Moor 1958 (Artemisio dniproicae-Salicetum acutifoliae Shevchyk et Solomakha 1996), Pyrolo-Pinetea sylvestris Korneck 1974, Sedo-Scleranthetea Br.-Bl. 1955, relevés of ruderal vegetation, relevés attributed by the authors to communities, and also not containing dominant species at all. The Sörensen coefficient (Sörensen, 1948) was chosen as the similarity measure and grouping was performed using the "flexible" beta method at -0.25. This resulted in finding smaller groups approximately corresponding to the association's rank. Relevés of Festucetea vaginatae class were grouped according to Ward's method (Ward, 1963). Identification of diagnostic species in syntaxa was carried out in accordance with the fidelity index - the Phi coefficient (Willner et al., 2009), the threshold values of which were assumed at the level of 25. All relevé groups were standardized to equal size, and non-essential fidelity values were removed based on the Fisher exact test.
The taxonomic nomenclature was given by "Flora Europaea" (Tutin et al., 1964-1980). Some of the species (taxonomically problematic, narrow, considered differently by various authors) we have combined into aggregates. Specifically, Medicago sativa ssp. falcata agg. combined the species M kotovii Wissjul., M romanica Prodan, M. tenderiensis Opperman ex Klokov and actually M. falcata L. Crepis tectorum agg. includes C. tectorum L. and C. ramosissima D'urv. Artemisia campestris agg. combines A. campestris L., A marschalliana Spreng. and A. dniproica Klokov.
Identification of the obtained phytocoenoses was carried out on the basis of their diagnostic species and floristic composition by comparison with foreign and Ukrainian publications (Demina, 2009, 2015; Mucina et al., 2016; Bulokhov, 2019; Dubyna et al., 2019a).
The method of DCA-ordination (Hill & Gauch, 1980; ter Braak & Smilauer, 2015) of the R-project software (Venables & Smith, 2008) through the JUICE software package was used to determine the features of ecological differentiation of communities. The calculation of environmental parameters was carried out according to phytoindication scale developed by Didukh (2011), which allows ordination analysis to be performed on 12 factors: soil humidity (Hd), variability of damping (fH), soil aeration (Ae), soil nitrogen content (Nt), soil acidity (Rc), salt regime (Sl), carbonate content (Ca), temperature regime (Tm), ombroregime (Om), climate continentality (Kn), cryoregime (Cr) and light intensity (Lc).
Results
Cluster analysis of psammophytic vegetation database includes 1785 relevés and allowed several classes to be identified (Fig. 1). In particular, clusters 1-4 corresponded to the class Salicetea purpureae (Artemisio scopariae-Tamaricion ramosissimae), 5-12 - Ammophiletea, 13-15 -Cakiletea maritimae, 16-20 and 32-35 - Helichryso-Crucianelletea maritimae, 21-31 - Koelerio-Corynephoretea canescentis, 36-41 and 49-70 - Festucetea vaginatae, 42-48 - Stellarietea mediae (ruderal vegetation on sandy soils).
At the next stage of processing, after exclusion of relevés related to Ammophiletea, Salicetea purpureae, Pyrolo-Pinetea sylvestris, relevés of disturbed habitats, as well as those that do not contain dominant species, phytocoenons with the studied classes remained: Helichryso-Crucianelle-tea maritimae, Koelerio-Corynephoretea canescentis and Festucetea vaginatae - 955 relevés (Fig. 2). Then we performed cluster analysis
separately per classes. It should be noted that a significant number of sample sites within the studied classes were identified by us as agglo-merative groups representing the initial stages of sand overgrowth, or as
rankless (basal/derrivate) communities (a total of 258 relevés), which were not included by the authors at this stage in the classification scheme.
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Fig. 1. Hierarchical cluster classification of relevés ofpsammophytic vegetation ofUkraine (1785 relevés) (PC-ORD)
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Based on processing ofphytocoenotic tables and cluster analysis results, a refined classification scheme of psammophytic vegetation of Ukraine was obtained.
Classification scheme ofpsammophytic vegetation ofUkraine Koelerio-Corynephoretea canescentis Klika in Klika et Novak 1941 Corynephoretalia canescentis Klika 1934 Corynephorion canescentis Klika 1931
Veronico dillenii-Corynephoretum Passarge 1960 Corniculario aculeatae-Corynephoretum canescentis Steffen 1931 (incl Helichryso-Jasionetum sensu Vorobyov et al. 1998non Libbert 1940) Armerion elongatae Potsch 1962
Diantho deltoidis-Armerietum elongatae Krausch 1959 Koelerion glaucae Volk 1931
Veronico dillenii-Secaletum sylvestris Shevchyk et Solomakha 1996 Chamaecytiso ruthenicae-Festucetum beckeri Shevchyk et al. 1996 Jasiono montanae-Thymetum serpyli Bulokhov 2019 (Thymetum pulegioido-serpylli Konishchuk2003 nom. inval (syntax. syn)) Jasiono montanae-Festucetum ovinae Klika 1941 CorynephoroSilenetum tataricae Libbert 1931 Artemisio campestris-Dianthetum borbasii Yakushenko 2004 Artemisio dmprocae-Sedetum sexangularis Shevchyk et Solomakha in Shevchyk et al. 1996 (incl. Sedo sexangulare-Festucetum Solomakha, Shevchyk et Senchylo 1997 (art. 10), Poetum bulbosae Shevchyk et Polishko 2000 (syntax. syn.))
Diantho borbasii-Agrostietum syreistschikovii Vicherek 1972 Thymo angustifoliiFestucetum beckeri Vicherek 1972 Centaureo borysthenicae-Festucetum beckeri Vicherek 1972 Festucetea vaginatae Soó exVicherek 1972 Festucetalia vaginatae Soó 1957
Artemisio arenariae-Festucion beckeri Dubyna et Dziuba in Dubyna et al. 2019 Aperetum maritimae Popescu, Sanda et Doltu 1980 (incl. Apero maritimi-Chry^pogonetum grylli Davydova 2019 p. p. (syntax. syn.) Dauco gutШ-Chrysapoganetum grylli Popescu, Sanda et Doltu 1980 (incl. Apero maritimi-Chrysapogonetum grylli Davydova 2019 (syntax. syn.))
Carici cokhncae-Hobschœfetum vulgaris Sorbu et al. 1995
Salici rasmarinifoIiae-HoIaschaenetum vulgaris Mititelu et al. 1973 (incl.
Galio ruthenici-Salicetum rosmarinifoliae I. Solomakha, Vorobyov et
Moysienko 2015 nom. inval. (syntax. syn.))
Allio guttati-Festucetum rupicolae Umanets et Solomakha 1999
Heliotropo dolosi-Brometum japonici Dubyna, Neuhâuslovà et Shelyag-
Sosonko 1995
Trago-Anthemietum ruthenicae Pitjcaru-Soroceanu et al. 1963 Plantaginetum arenariae (Buia et al. 1960) Popescu et Sanda 1987 Linario adoraeAgropyretum dasyanthi Vicherek 1972 Cynodonla^Medicaginelum minimae Popescu et Sanda 1975 Secdi-Cymcbrtetum dactyli Dubyna, Neuhâuslovà et Shelyag-Sosonko 1995
Secali-Stipetum borysthenicae Korzhenevsky ex Dubyna, Neuhâuslovà et Shelyag-Sosonko 1995
Secaletum sylvestris Popescu et Sanda 1973 Secali sylvestri-Brometum tectorum Hagtai 1940 Centaureo odessanae-Caricetum colchicae Tyschenko 1999 (incl Secali sylvestri-Caricetum colchicae Davydova 2019 (syntax. syn)) Centaureo odessanae-Stipetum capillatae Dubyna, Neuhauslová et Shelyag-Sosonko 1995
Poo bulbosae-Caricetum colchicae Dubyna, Neuhauslová et Shelyag-Sosonko 1995
Achilleo setacei-Festucetum beckeri Chusova 2019 nom inval (art. 3b) Centaureo gerberi-Chamaecyisetum borysthenici Demina2009 Festucion beckeri Vicherek 1972
Festucetum beckeri Ad. Oprea 1998 Centaureo odessanae-Festucetum beckeri Vicherek 1972 Centaureo brevicipiti-Festucetum beckeri Vicherek 1972 Helichryso-Crucianelletea maritimae Géhu et al in Sissingh 1974
Ephedro dstachyae-Medcaginetalia romanicae Dubyna et Dziuba in Dubyna et al 2019 (incl. Medicagfni tenderiensis-Seselietalia tenderiensi Umanets et I. Solomakha 1999 (syntax. syn.))
Ephedro distachyae-Medicaginion romanicae Dubyna et Dziuba in Dubyna et al. 2019
Anisantho tectorum-Medicaginetum kotovii Tyschenko 1996 Anisantho tectorum-Helichrysetum arenariae Tyschenko 1999 Ephedro-Caricetum colchicae (Prodan 1939) Sanda et Popescu 1973 Scabiosion ucranicae Sanda et al. 1980
Scabioso ucranicae-Caricetum ligericae (Simon 1960) Krausch 1965 Secali sylvestri-Alyssetum borzaeani (Borza 1931) Morariu 1959 Medicagini tenderiensis-Seselion tenderiensi Umanets et I. Solomakha 1999 (incl. Melico chrysolepi-Ephedrion distachyae Umanets et I. Solomakha 1999 (syntax. syn.))
Medicagini tenderiensis-Seselietum tenderiensi Umanets et I. Solomakha 1999 (incl. Sileno subconicae-Galietum tenderiensi Umanets et I. Solomakha 1999 (syntax. syn.); Elytrigio elongatae-Galietum tenderiensi Umanets et I. Solomakha 1999 (syntax. syn.); Consolido paniculatae-Anchusetum gmelini Umanets et I. Solomakha 1999 (syntax. syn.)) Cynodonto-Teucrion polii (Korzhenevsky et Klyukin 1990) Korzhenevsky et Kvitnytskaya 2014 nom. inval. (art. 5) (incl. Verbascion pinmatifidi Korzhenevsky et Klyukin 1990 (syntax. syn.))
Cynodonto-Ajugetum chiae (Korzhenevsky et Klyukin 1990) Korzhenevsky et Kvitnytskaya 2014 nom. inval. (art. 5) Carici liparicarpo-Centaureetum adpressae (Korzhenevsky et Klyukin 1990) Korzhenevsky et Kvitnytskaya 2014 nom. inval. (art. 5)
Table 1
Synoptic table ofthe class Koelerio-Corynephoretea canescentis
Astragale borysthenici-Ephedretum distachyae (Korzhenevsky et Klyukin 1990) Korzhenevsky et Kvitnytskaya 2014 nom. inval. (art. 5) Leymo-Verbascetum pinnatifidi (Korzhenevsky et Klyukin 1990) Korzhenevsky et Kvitnytskaya 2014 nom. inval. (art. 5)
Koelerio-Corynephoretea canescentis class includes pioneer communities on fluvioglacial and riverine sandy, shallow, weakly mobile soils, mostly dry and poor in nutrients, having acidic reaction. According to our results, the class in Ukraine include 1 order, 3 alliances and 13 associations (Fig. 3, Table 1).
Fig. 3. Cluster analysis of phytosociological data ofKoelerio-Corynephoretea canescentis class: clusters: 1 - Veronico dillenii-Secaletum sylvestris association; 2 - Chamaecytiso ruthenicae-Festucetun beckeri', 3 - Diantho deltoidis-Armerietum elongatae; 4 - Jasiono montanae-Thymetum serpylli; 5 -Jasiono montanae-Festucetum ovinae; 6 - Corynephoro-Silenetum tataricae; 7 - Artemisio campestris-Dianthetum borbasii; 8 - Artemisio dniproicae-Sedetum
sexangularis, 9 - Veronico dillenii-Secaletum sylvestris (sensu Kovalenko); 10 - Diantho borbasii-Agrostietum syreistschikovii; 11 -Thymo angustifolii-Festucetum beckeri; 12 - Centaureo borysthenicae-Festucetum beckeri; 13 - Veronico dillenii-Corynephoretum; 14 - Corniculario aculeatae-Corynephoretum canescentis
No. of syntaxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Number of relevés 14 24 7 7 38 6 8 44 12 9 25 29 15 83
Aristolochia clematitis Tragopogon ucrainicus Vicia tetrasperma Cardaminopsis arenosa Salix acutifolia Galium ruthenicum Peucedanum oreoselinum Verbascum densiflorum Pulsatilla pratensis Centaurea sumensis Jurinea ewersmanii Artemisia vulgaris Poa angustifolia Arabis glabra Veronica spicata s. incana Stellaria graminea Viola tricolor s. matutina Jurinea cyanoides s. tenuiloba Silene nutans
Sedum telephium s. maximum Vincetoxicum hirundinaria Thesium ebracteatum Luzula pallescens Filipendula vulgaris Stachys recta
Centaurea arenaria s. majorovii Artemisia austriaca Phleum phleoides Euphorbia esula s. tommasiniana Scorzonera purpurea Sempervivum ruthenicum Dianthus deltoides
42.4
38.7 32.3
25.8 25.8 25.8
15.3
64.1
63.7
59.8
56.3 52.6
52.4
50.8 44.3 43.6
39.6
37.7
34.2 33.6
32.5
30.9 27.9 27.9 27.9 27.9 27.9 27.9 27.9 27.9 27.9
27.8
19.3
19.3
96.0
No. of syntaxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Number of relevés 14 24 7 7 38 6 8 44 12 9 25 29 15 83
Armeria maritima s. elongata Deschampsia cespitosa Poa pratensis Dactylis glomerata Anthoxanthum odoratum Agrostis capillaris Elymus repens Medcago lupulina Thuidium abietinum Cerastium arvense Lolium perenne Equisetum arvense Veronica chamaedrys Knautia arvensis Rumex thyrsiflorus Ranunculus acris Alyssum alyssoides Vicia hirsuta Festuca pratensis Lotus corniculatus Fragaria viridis Senecio sylvaticus Briza media Bromus commutatus Scirpus sylvaticus Alchemilla vallesiaca Plantago major Trisetum sibiricum Potentilla reptans Prunella vulgaris Leucanthemum vulgare Rubus caesius Centaureajacea Plantago media Hylocomium splendens Agrimonia eupatoria Agrimonia procera Poa annua Daucus carota Clinopodium vulgare Thymus pulegioides Silene armeria Galeopsis ladanum Anthemis cotula Silene vulgaris Sedum telephium s. ruprechtii Trifolium alpestre Ononis arvensis Potentilla collina
Dianthus arenarius s. pseudoserotinus
Jovibarba globifera s. globifera
Polypodium vulgare
Centaurea rhenana s. pseudomaculosa
Potentilla cinerea
Chondrillajuncea
Centaurea phrygia
Artemisia scoparia
Jasione montana
Pteridium aquilinum
Achillea millefolium
Anthericum ramosum
Oenothera rubricaulis
Allium oleraceum
Ajuga reptans
Scabiosa ochroleuca
Centaurea rhenana
Silene tatarica
Verbascum lychnitis
Medicago sativa s. falcata
Euphorbia esula
Koeleria macrantha
Viola arvensis
Silene latifolia s. alba
Senecio jacobaea
Leontodon hispidus
Erigeron annuus
Echium vulgare
Genista tinctoria
Poa compressa
Potentilla argentea
83.6 74.4
74.4
63.1 62.1 61.9 54.0 52.0 52.0 52.0 52.0 48.6 40.8 40.6
39.5
36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6 36.6
33.2 25.0
11.6
16.8
11.9
19.7
21.4
16.2
23.4 15.9
53.1 -
52.0 -
52.0 -
49.7 -
44.3 -
37.7 -
36.6 -
36.6 -
36.6 -
36.6 -
36.6 -
36.6 -
33.6 -
32.6 -
30.6 11.4
- 63.5
- 57.6
- 45.1
- 35.5
- 32.7
- 30.0
- 28.1
- 27.2
- 27.2
20.7
17.8 18.8
20.4 16.8
9.7
13.4
23.1
100.0
98.1 80.8
71.2
69.4 62.8
56.3
49.5 46.9 44.7 43.7 43.1 34.5 33.1
14.8
24.2
85.0 54.6
15.1
18.6
4.5
12.4
21.2
No. of syntaxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Number of relevés 14 24 7 7 38 6 8 44 12 9 25 29 15 83
Verbascum nigrum ----- - 48.6 - - -----
Melampyrum nemorosum ----- - 48.6 - - -----
Oenothera biennis 15.5 ----- 45.5 - - -----
Quercus robur ----- - 42.9 - - - - - - 7.2
Calamagrostis epigejos - 16.2 - - - - 40.3 13.3 - -----
Hieracium pilosella - - 23.8 - - - 35.8 - - - - - - 17.0
Solidago virgaurea ----- - 34.9 - - - - - - 18.1
Elytrigia intermedia ----- - 34.2 - - -----
Polygonatum odoratum ----- - 34.2 - - -----
Luzula pilosa ----- - 34.2 - - -----
Lepidium campestre ----- - 32.5 - - -----
Acinos arvensis ----- - 30.6 - - -----
Bromus inermis - - - - - - - 68.6 - -----
Tanacetum vulgare - - - - - - - 62.4 - -----
Veronica spicata - - - - - - - 51.6 - -----
Achillea cartilaginea - - - - - - - 48.6 - -----
Eryngium planum - - - - - - - 46.3 - -----
Gratiola officinalis - - - - - - - 46.3 - -----
Veronica praecox - - - - - - - 43.9 - -----
Atriplexprostrata - - - - - - - 38.7 - -----
Veronica verna - - - - - - - 36.9 - -----
Allium angulosum - - - - - - - 32.6 - -----
Scirpus holoschoenus - - - - - - - 29.1 - -----
Sedum argutum - - - - - - - 26.3 - -----
Bromus squarrosus - - - - - - - 26.1 - -----
Bromus hordeaceus - - - - - - - - 97.8 -----
Spergularia rubra - - - - - - - - 97.5 -----
Arenaria serpyllifolia v. stepicola - - - - - - - - 91.0 -----
Spergula arvensis - - - - - - - - 90.5 -----
Polygonum aviculare - - - - - - - - 80.9 -----
Buglossoides arvensis s. arvensis - - - - - - - - 72.3 -----
Carex praecox - - - - - - - - 54.4 -----
Vicia cracca - 14.4 - - - - - - 46.9 -----
Linaria vulgaris - 13.4 - - - - - - 44.7 -----
Thesium procumbens - 20.2 - - - - - - 28.3 -----
Taraxacum officinale - - - - - - - 12.6 28.0 -----
Hieracium umbellatum - 6.5 - - - - - - - 84.3 - - - -
Agrostis vinealis - - - - - - - 16.6 - 70.1 - - - -
Crepis tectorum agg. - - - - - - - - - 52.7 - - - -
Herniaria polygama - - - - - - - - 24.2 44.7 22.7 12.8 - -
Linaria genistifolia - 12.6 - - - - - - - 29.9 18.8 18.0 - -
Thymus serpyllum s. serpyllum - - - - - - - - - - 93.4 -
Dianthus arenarius s. pseudosquarrosus - -- -- - - -- - 69.5 21.4
Jurinea cyanoides ---- - - - -- - 69.4 21.6
Sedum telephium s. telephium ---- - - - -- - 56.8 -
Androsace septentrionalis ---- - - - -- - 47.6 -
Hierochoe stepporum ---- - - - -- - 43.4 -
Astragalus arenarius ---- - - - -- - 43.4 -
Sempervivum zeleborii ---- - - - -- - 27.3 -
Thymus pallasianus ---- - - - -- -- 91.9
Minuartia viscosa ---- - - - -- -- 84.2
Agropyron cristatum s. sabulosum - -- -- - - ----75.4
Achillea micrantha ---- - - - -- -- 75.4
Polygonum arenarium s. arenarium - -- -- - - -- -- 75.4
Alyssum tortuosum ---- - - - -- -- 75.4
Astragalus varius ---- - - - -- -- 75.4
Tragopogon brevirostris s. brevirostris - -- -- - - ----73.0
Syrenia cana ---- - - - -- -- 72.7
Dianthus platyodon ---- - - - -- -- 70.6
Cladonia subcariosa ---- - - - -- -- 70.6
Asperula graveolens ---- - - - -- -- 70.6
Senecio borysthenicus ---- - - - -- -- 69.9
Xanthopaemelia somloensis ---- - - - ----68.1
Anchusa gmelinii ---- - - - ----68.1
Dianthus squarrosus ---- - - - -- -- 65.6
Stipa borysthenica ---- - - - -- -- 64.9
Agropyron dasyanthum ---- - - - -- -- 62.9
Asperula leiograveolens ---- - - - -- -- 57.3
Scabiosa argentea ---- - - - -- -- 57.3
Syrenia montana ---- - - - -- -- 57.3
Seseli arenarium ---- - - - -- -- 54.3
Minuartia setacea ---- - - - -- -- 51.1
Allium guttatum ---- - - - -- -- 47.8
Gypsophila paniculata ---- - - - -- - 18.3 44.8
Scorzonera ensifolia ---- - - - -- -- 44.2
Pulsatilla patens ---- - - - -- -- 43.7
Jurinea paczoskiana ---- - - - -- -- 40.3
Carex ligerica ----- - - 12.9 - - 16.1 39.0
Jurinea polyclonos ---- - - - -- -- 36.0
Artemisia tschernieviana - - - - - - - - - - - 31.1
No. of syntaxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Number of relevés 14 24 7 7 38 6 8 44 12 9 25 29 15 83
Diploschistes scruposus - -
Cladonia pocillum - -
Cladonia degenerans - -
Cladonia mitis - -
Hypochoeris radicata - -
Cladonia rangiferina - -
Scleranthus perennis - -
Arenaria serpyllifolia - -
Dicranum scoparium - -
Logfia minima - -
Secale sylvestre 55.0 -
Anthemis ruthenica 40.9 -
Lepidium densiflorum 31.3 -
Asparagus officinalis 27.4 -
Chamaecytisus ruthenicus - 47.9
Centaurea arenaria s. borysthenica - 45.1
Galium verum - 38.9
Rumex acetosa - 25.4
Euphorbia cyparissias - -
Pinus sylvestris - -
Festuca ovina - -
Trifolium arvense - -
Sedum sexangulare - -
Berteroa incana - -
Carex hirta - -
Plantago lanceolata - -
Artemisia absinthium - -
Pinus sylvestris - -
Genista tinctoria - -
Sedum telephium - -
Scleranthus annuus - -
Sedum acre s. acre - -
Gypsophila muralis - -
Rumex acetosella s. acetosella v. tenuifolius - -
Silene conica - -
Polytrichum piliferum - -
Tortula ruralis - -
Coelocaulon aculeatum - -
Euphorbia seguierana - -
Alyssum minutum - -
Hieracium echioides - -
Silene borysthenica - -
Cerastium semidecandrum - 19.6
Racomitrium canescens - -Helichrysum arenarium
31.1
17.9
63.4 50.2
38.4
30.0
28.1
26.5 26.1 26.0 26.0
- - 12.2
29.2 - 29.7
- 41.7 -
33.8 49.6
- 32.0
- 8.7
24.2
30.1 30.8
52.5 47.1
45.6
33.5
30.6 28.9 28.6
20.8
23.3
42.1
35.4
47.2 60.2
32.7 34.5
8.8
31.4 50.9
10.8
31.9
42.0 77.2 36.6
16.5
7.4
61.6
44.0
26.0 15.9
48.2
25.7
49.4
50.4 70.7 62.7 47.1 46.3
40.5
3.8
7.8
7.0
28.3
13.0 24.5
15.2 35.8
58.8
56.1
49.3 49.3
37.0
36.9 36.5
29.1 25.9
25.9
35.2
32.3
54.1
50.4 61.1
48.4
43.5 43.3 45.5
40.1
13.3 20.7
11.9
35.8
Corynephorus canescens - - - - - - - - - - 23.0 - 48.8 48.0
Festuca beckeri 19.3 35.8 - - - - - - - - 46.2 46.2 - -
Achillea collina - 6.4 31.3 - - 49.9 25.4 - - - - - - -
Thymus serpyllum - - - 54.2 - 31.4 28.5 - - - - - - 4.7
Sedum acre - - - 32.2 7.7 - 25.8 28.5 15.8 - - - - -
Dianthus borbasii - - - - - - 34.0 21.1 - 34.0 29.5 - - -
Erophila verna - - - - - - - - - 45.4 40.4 45.4 12.2 -
Ceratodon purpureus - - - - - - 19.3 - - 41.9 41.9 31.5 21.8 -
Myosotis stricta - - - - - - - - - 33.8 48.1 45.9 13.9 -
Bassia laniflora - - - - - - - - - 32.9 45.0 56.2 - -
Cladina arbuscula - - - - - - - - - 29.2 44.5 36.7 - -
Tragopogon brevirostris s. podolicus - - - - - - - - - 27.3 50.8 49.5 - -
Cladonia rangiformis - - - - - - - - - - 48.3 51.9 33.3 -
Artemisia campestris agg. - - - - - 25.3 25.3 24.0 - 25.3 23.1 25.3 - -
Poa bulbosa - - - - - - - - 37.9 43.0 28.4 38.8 - -
Koeleria glauca - - - - 9.7 - - - 30.2 30.2 30.2 30.2 - -
Cladonia foliacea - - - - - - - - - 41.5 40.9 48.6 27.2 -
Veronica dillenii 12.6 - - - - - - - 32.6 32.6 28.1 30.7 32.6 -
Conyza canadensis Cladonia coniocraea Digitaria sanguinalis Polytrichum juniperinum Nardus stricta
Campanula patula s. abietina Verbascum phlomoides Calluna vulgaris Cladonia mediterranea Setaria viridis Cladonia uncialis Agrostis canina Cladonia species Logfia arvensis Herniaria glabra Erigeron acer Brachythecium albicans
23.8 23.7 21.2 21.2 21.2 21.2 18.3 18.3 18.3 18.3 18.3 17.0
13.7
12.8 10.6 9.0 8.3
No. of syntaxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Number of relevés 14 24 7 7 38 6 8 44 12 9 25 29 15 83
Festuca rubra Hypericum perforatum Bromus tectorum Bromus secalinus Calamagrostis arundinacea Draba nemorosa Fallopia convolvulus Rhinanthus minor Lychnis viscaria Carexpediformis s. rhizodes Carex ericetorum Pinus sylvestris Juncus compressas Campanula patula Chenopodium rubrum Glechoma hederacea Dichanthium ischaemum Cerastium fontanum s. vulgare Teucrium chamaedrys Spergula morisonii Leontodon autumnalis Festuca valesiaca Descurainia sophia Fragaria vesca_
8.3 5.6
24.8
20.6 20.6 20.6 20.6 20.6
15.1
22.6
22.1 17.9
22.1 18.4 22.1 22.1 22.1 22.1 21.1
24.9
9.1
22.1
17.9
24.0 24.0
Notes: numbers mark syntaxa: 1 - Veronico dillenii-Secaletum sylvestris, 2 - Chamaecytiso ruthenicae-Festucetum beckeri; 3 - Diantho deltoidis-Armerietum elongatae, 4 - Jasiono montanae-Thymetum serpylli; 5 - Jasiono montanae-Festucetum ovinae; 6 - Corynephoro-Silenetum tataricae; 7 - Artemisio campestris-Dianthetum borbasii; 8 - Artemisio dniproicae-Sedetum sexangularis; 9 - Veronico dillenii-Secaletum sylvestris (sensu Kovalenko); 10 - Diantho borbasii-Agrostietum syreistschikovii; 11 - Thymo angustifolii-Festucetum beckeri; 12 - Centaureo borysthenicae-Festucetum beckeri; 13 - Veronico dillenii-Corynephoretum; 14 - Corniculario aculeatae-Corynephoretum canescentis.
The alliance Corynephorion canescentis combines coenoses in the structure of which mosses and especially lichens play a significant role, and which develop on weakly mobile sandy soils on fluvioglacial deposits, mainly in the Ukrainian Polissia The Koelerion gfauoae alliance includes multi-species communities growing on neutral sandy soils which developed mainly on alluvial deposits. They are common in the Ukrainian Polissia and, less often, in the forest-steppe on the pine-forest terraces of rivers and natural levees. Their territorial differentiation is determined by the reliefand soil reaction.
According to the results of ordination analysis of communities in the class Koelerio-Corynephoretea canescentis, it was found that temperature regime and climate continentality play a leading role in their ecological differentiation (Fig. 4). These parameters are most important for coenoses within Corynephorion canescentis. Also, the distribution of communities is significantly affected by the gradients of ombroregime and soil humidity. Communities of Corynephoro-Silenetum tatarioae, Thymo angustifolii-Festucetum
Table 2
Synoptic table of the Festucetea vaginatae class
beckeri, Jasiono montanae-Thymetum serpylli and Jasiono montanae-Festucetum ovinae associations are more "sensitive" to changes in these parameters. Communities of the last two syntaxa will also respond to concentration of available nitrogen forms in soil substrate. Variability of damping (fH) can be considered the leading ecological factor for the coenoses Diantho borbasii-Agrostietum syreistschikovii, Veronico dillenii-Secaletum sylvestris and Chamaecytiso ruthenicae-Festucetun beckeri. Communities of the Diantho deltoidis-Armerietum elongatae association were described on the territory of the Ukrainian Roztochia on specific moraine deposits, and their distribution in the ecological space determines content of carbonates and other salts in the soil. Diantho deltoidis-Armerietum elongatae, Corynephoro-Silenetum tataricae, Artemisio dniproicae-Sedetum sexangularis and Thymo angustifolii-Festucetum beckeri communities differ significantly in their ecological amplitude (Fig. 4).
No. of syntaxa 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 26 27
Number of relevés 10 17 20 13 18 4 9 43 9 10 14 5 10 7 5 12 10 13 21 8 47 16 10 10 10 92 47
Juncus bufonius 51 21
Trifolium fragiferum 44
Aster tripolium 44
Sonchus palustris 44
Carex extensa 44
Barbarea stricta 44
Odontites verna s. serotina 41 18
Medcago lupulina 41 15 -
Limonium bellidifolium 38
Plantago major 37
Agrostis gigantea s. maeotica 34
Phragmites australis 34 19 - 18
Rumex acetosa 31
Inula britannica 31
Elytrigia intermedia 31
Aster sedfolius s. dracunculoides 31
Xanthium strumarium 29
Cirsium ukranicum 28
Bromus squarrosus 28 16 - - 14 - ----25--- ------- 14 - - - - -
Chrysopogon gryllus 22 85
Picris hieracioides s. hieracioides - 36
Asparagus litoralis - 35
Convolvulus lineatus - 34
Orchis mono s. picta - 34
Plantago maritima 21 33
Schoenus nigricans - 32
Rumex acetosella - 31 18 12
306 Biosyst. Divers., 2020, 28(3)
No. of syntaxa 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 26 27
Number of relevés 10 17 20 13 18 4 9 43 9 10 14 5 10 7 5 12 10 13 21 8 47 16 10 10 10 92 47
Orchis coriophora _ 31 _______________________
Lactuca tatarica __ 35 _______ 19 ______________
Falbpia dumetorum ---73 - -- -- -- -- -- -- -- -- -- --Solidago virgaurea ---41 - -- -- -- -- -- -- -- -- -- --Rhamnus catharticus ---39 - -- -- -- -- -- -- -- -- -- --Tanacetum vulgare ---39 - -- -- -- -- -- -- -- -- -- --Verbascum orientale ---27 - -- -- -- -- -- -- -- -- -- --Verbascum chaixii s. austriacum ---27 - -- -- -- -- -- -- -- -- -- -Viola lavrenkoana ---27---------------------
Veronica spicata s. barrelieri - - - 27 - - - - - - - - - - - - - - - - - - - - -Jurinea longifolia ---27 - -- -- -- -- -- -- -- -- -- -Viola hymettia ---27 - -- -- -- -- -- -- -- -- -- -Cuscuta monogyna ----40---- - 20 - -- -- -- -- -- -- -
Vincetoxcumfuscatum ----40--------------------
Festuca rubra ----38 - -- -- -- -- -- -- -- -- -- -
Stipa borysthenica ---- 36 - -- -- 20 - -- -- -- -- 22 - 22 - 18
Medicago minima -----90 17 - -- -- -- -- -- -- -- -- -
Ajuga chamaepitys s. chia -----80) - -- -- -- -- -- -- -- -- --
Potentilla humifusa -----70 - -- -- -- -- -- -- -- -- --
Carduus uncinatus - - - - - 63- - - - 8- - - - - - - - 18 - 12- - -
Potentilla astracanica ----- 62 - -- -- -- -- -- -- 18 - 23---
Silene conica s. subconica - - - - 9 52 - 9 - - 10 - - - - - - - - 15 - - - - -
Cruciata pedemontana -----49 - -- -- -- -- -- -- -- -- --
Aegilops lorentii -----49 - -- -- -- -- -- -- -- -- --
Allium scorodoprasum s. rotundum - -- --49 - -- -- -- -- -- -- -- -- --
Hordeum murinum 15 - -- -41 - -- -- -- -- -- -- -- -- --
Linum austriacum s. euxinum - - - - - 40 - - - - - - - - - - - - - 19 - - - - -
Asperula tenella -----36-------------------
Crepis foetida s. rhoeadifolia - - - - - 33 - - - - - - - - - - - - - - - 15 - - -
Elymus elongatus 12 - -- --52 - -- -- -- -- -- -- -- -- -
Consolida regalis - - - - - - 46 - - - - - - - - 24 - - - - - - - - -
Artemisia santonica ----14 -41 - --24 - -- -- -- -- - 12---
Cuscuta europaea ------31------------------
Asparagus maritimus - - - - - - 30 - - - 25 - - - - - - - - 21 7 - - - -
Plantago cornuti ------30------------------
Camelina rumelica - - - - - - 28 - - - 17 - - - - - - - - - - - - - -Gypsophila perfoliata 23 - 14 - -- 27 6 - -- -- -- -- -- 19 ----Vicia cracca --------41----------------
Alyssum desertorum - - - - - - - 15 41 - - - - - - - - - - - - - - - -
Centaurea biebersteinii s. biebersteinii - -- -- -- - 33 - -- -- -- -- -- -- -- -
Allium scorodoprasum s. waldsteinii - -- -- -- - 33 - -- -- -- -- -- -- -- -
Cannabis sativa v. spontanea ------- - 33 - -- -- -- -- -- -- -- -
Saponaria officinalis --------33----------------
Tragopogon ucrainicus - - - - - - - - 30 - - - - - - - - - - - - - - - -Centaurea arenaria s. borysthenica - -- -- -- - 28 - -- -- -- - 19 - -- -- -Festuca rupicola - - - - - - - - - 93 - - - - - - - - - - - - - - -Daucus carota -------- - 59 - -- -- -- -- -- -- --
Agrostis gigantea -------- - 52 - -- -- -- -- -- -- --
Phleum phleoides ---------46---------------
Thalictrum simplex ---------44---------------
Potentilla recta - - - - - - - - - 43 - - - - - - - - - - - - - - -
Bromus inermis ---------42---------------
Vicia tetrasperma - - - - - - - - - 42 - - - - - - - - - - - - - - -Asparagus officinalis s. officinalis - -- -- -- -- 38 - -- -- -- -- -- - 24 -Apera spica-venti ------- 22 - 33 --------------Linaria vulgaris - - - - - - - - - 31 - - - - - - - 23 - - - - - - -Festuca pratensis ---------31---------------
Potentilla species - - - - - - - - - 31 - - - - - - - - - - - - - - -
Teucrium chamaedrys ---------31---------------
Astragalus arenarius - - - - - - - - - 31 - - - - - - - - - - - - - - -
Centaurea triniifolia -------- - 31 - -- -- -- -- -- -- --
Centaurium erythraea ---------31---------------
Erigeron acer - - - - - - - - - 31 - - - - - - - - - - - - - - -
Lavatera thuringiaca ---------31---------------
Carex melanostachya - - - - - - - - - 31 - - - - - - - - - - - - - - -
Trifolium campestre -------- - 28 - -- -- -- -- -- -- --
Coelocaulon species ---------28---------------
Heliotropium dolosum - - - - - - - - - - 97 - - - - - - - - - - - - - -
Lepidium campestre --------- - 88 - -- -- -- -- -- -- -
Bromus japonicus ----- - 19 - --77 - -- -- -- -- -- -- -
Aeluropus littoralis --------- - 75 - -- -- -- -- -- -- -
Lithospermum arvense ----------72--------------
Diplotaxis muralis - - - - - - - - - - 64 - - - - - - - - - - - - - -
Sisymbrium altissimum --------- - 59 - -- -- -- - 21-----
Allium pervestitum - - - - - - - - - - 42 - - - - - - - - - - 21 - - -
Orobanche cernua ----------37--------------
Artemisia pontica - - - - - - - - - - 37 - - - - - - - - - - - - - -
Elymus uralensis s. viridiglumis - - 17 21 17 - -- -- 37 - -- -- -- -- -- -- -
Melilotus alba --------- - 28 - -- -- -- -- - 12---
Asparagus tenuifolius - - - - - - - - - - 26 - - - - - - - - - - - - - -
No. of syntaxa 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 26 27
Number of relevés 10 17 20 13 18 4 9 43 9 10 14 5 10 7 5 12 10 13 21 8 47 16 10 10 10 92 47
Polygonum arenarium s. pulchellum — — — — — — — — — — 26 — — — — — — — — — — — — — — — —
Halimione verrucifera --------- - 26 - -- -- -- -- -- -- -- -
Thesium linophyllon --------- - 26 - -- -- -- -- -- -- -- -
Tragus racemosus ---------- - 72 - -- -- -- - 10 - -- -- -
Lotus tenuis ---------- - 66 - -- -- -- -- -- - 20 --
Eragrostis pilosa ---------- - 48 - - 22 - -- -- -- -- -- -
Corispermum ucrainicum ---- 10 - 24 - -- - 45 - -- -- -- -- -- -- --
Thlaspi arvense ---------- - 34 - -- -- -- -- -- -- --
Corispermum nitidum ----------- - 99 - -- -- -- -- -- -- -
Linaria odora ----------- - 99 - -- -- -- -- -- -- -
Tribulus terrestris --------- - 24 - 77 - -- -- -- -- -- -- -
Linaria dulcis -------------93-------------
Jurinea ewersmanii ------------ - 75 - -- -- -- -- -- --
Digitaria sanguinalis ------------ - 72 - -- -- -- -- -- -Silene borysthenica -------------60----- - 12 - - 20 20 --
Jurinea cyanoides s. tenuiloba ------------ - 59 - -- -- -- -- -- -Ambrosia artemisiifolia ------------ - 55 - -- 20 - -- -- -- -Lactuca serriola ---16 - -- -- -- -- 51 - -- -- -- -- -- --Calamagrostis epigejos --7 24 - -- -- 14 - -- 38 - -- 20 - -- -- -- --Sisymbrium orientale ------- - 16 - -- --69 - -- -- -- -- -- -
Anchusa officinalis --------------58------------
Achillea collina - - - - - - - - - - - - - - 52 - - - - - - - - - - - -
Sisymbrium polymorphum -11 9----- - 14 - -- -50 - -- -- -- -- -- -
/nula salicina -16 - -- -- -- -- -- - 29 - -- -- -- -- -- -
Ephedra distachya --- - 10 - -- -- 11 - -- - 54 - -- 24-24 18 ----
Dianthus capitatus ---------------28-----------
Dianthus bessarabicus - - - - - - - - - - - - - - - 28 - - - - - - - - - - -
Onobrychis arenaria -------------- - 28 - -- -- -- -- --
Marrubium pestalozzae -------------- - 28 - -- -- -- -- --
Sinapis alba ---------------28-----------
Echinops ritro - - - - - - - - - - - - - - - 28 - - - - 17 - - - - - 14
Thymus dimorphus - - 22 - 16 - -- -- 22 - -- - 27 - -- 24 - -- -- --
Achillea setacea ----------------89----------
Potentilla argentea - - - - - - - - - - - - - - - - 78 24 - - - - - - - - -
Hieracium pilosella ----------------76----------
Poa angustifolia - - - - - - - - - 21 - - - - - - 73 - - - - - - - - - -
Trifolium arvense --------------- - 72 - -- 4- -- -- -
Cerastium fontanum s. vulgare --------------- - 70 - -- -- -- -- -
Taraxacum officinale --------------- - 57 - -- -- -- -- -
Artemisia austriaca ---------- - 21 - -- - 56 --- - 20 -----
Jurinea consanguínea s. arachnoidea
Sedum acre ----------------46----------
Poa annua - - - - - - - - - - - - - - - - 44 - - - - - - - - - -
Galium album ----------------44----------
Thesium arvense - - - - - - - - - - 15 - - - - - 41 - - - 7 - - - - - -
Stellaria graminea ----------------39----------
Herniaria glabra - - - - - - - - - - - - - - - - 36 17 - - - - - - - - -
Sanguisorba officinalis --------------- - 31 - -- -- -- -- -
Convolvulus arvensis ----------------28----------
Jurinea polyclonos - - - - - - - - - - - - - - - - - 73 - - - - - - - - -
Achillea nobilis ------------ - 22 - - 14 66---------
Centaurea arenaria s. majorovii - - - - - - - - - - - - - - - - - 61 10 - - - - - - - -
Asparagus officinalis ---------------- - 58 - -- -- -- --
Jasione montana ---------------- - 56 - -8------
Tragopogon ruthenicus s. tanaiticus - - - - - - - - - - - - - - - - - 47 - - - - - - - - -
Agrostis vnealis -----------------47---------
Gypsophila paniculata - - - - - - - - - - - - - - - 14 - 42 - - 10 - - - - - -
Reseda lutea ---------------- - 39 - -- -- -- --
Verbascum thapsus ---------------- - 38 - -- -- -- --
Salix acutifolia ---------------- - 37 - -- -- -- --
Potentilla cinerea ---------------- - 29 - -- -- -- -11
Geranium robertianum - - - - - - - - - - - - - - - - - 27 - - - - - - - - -
Aster amellus -----------------27---------
Pulsatilla vulgaris - - - - - - - - - - - - - - - - - 27 - - - - - - - - -Astragalus sulcatus ---------------- - 27 - -- -- -- -Viola ambigua -----------------27---------
Potentilla thyrsiflora - - - - - - - - - - - - - - - - - 27 - - - - - - - - -
Silene chlorantha ---------------- - 27 - -- -- -- --
Equisetum arvense ---------------- - 27 - -- -- -- --
Elymus hispidus s. barbulatus ---------------- - 27 - -- -- -- -Melissa officinalis -----------------27---------
Poa compressa - - - - - - - - - - - - - - - - - 27 - - - - - - - - -
Delphinium cuneatum ---------------- - 27 - -- -- -- --
Melilotus officinalis ---------------- - 25 - -- -- -- --
Leymus racemosus s. sabulosus - -- -- -- -- -- -- -- -- - 39 - -18-----
Elymus farctus s. bessarabicus ----------------- - 37 - -- -- -- -
Eryngium maritimum ----------------- - 36 - -- -- -7-
Astrodaucus littoralis ----------------- - 30 - -- -- -- -
Festuca arundinacea s. orientalis - -19 - -- -- -- -- -- -- -- 28 - -- -- -- -
54
No. of syntaxa 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 26 27
Number of relevés 10 17 20 13 18 4 9 43 9 10 14 5 10 7 5 12 10 13 21 8 47 16 10 10 10 92 47
Stipa capillata ___________________ 85 ______
Coronilla varia ___________________ 59 ______
Dianthus pseudarmeria — — — — 17 — — — — — 16 — — — — — — — — 55 — — — — — —
Alliumflavum s. tauricum — — — — 19 — — — — — 11— — — — — — — — 48 — — — — — —
Trigonella monspeliaca ----17 - -- --14 - -- -- -- - 42 - -- -- -
Descurainia sophia ------------------ - 39 - -- -- -
Siiene chersonensis ------------------ - 38 - -- -- -
Buglossoides arvensis ------------------ - 35 - -- -- -
Afcea rugosa -------------------35------
Cladonia species - - - - - - - - - - - - - - - 21 - - - 34 - - - - - -
Gagea arvensis ------------------ - 28 - -- -- -
Gagea dubia ------------------ - 28 - -- -- -
Saxífraga tridactylites ------------------ - 28 - -- -- -
Salvia aethiopis -------------------28------
Agropyron cimmericum - - - - - - - - - - - - - - - - - - - 27 - - - - - -
Cirsium alatum ------------------ - 27 - -- -- -
Cirsium vulgare 20 - -- -- -- -- -- -- -- -- - 25 - -- -- -
Ceratodon purpureus --------------------50-----
Cladonia convoluta - - - - - - - - - - - - - - - - - - - - 45 - - - - -
Allium paniculatum ------------------- - 43-----
Neofuscelia species ------------------- - 41-----
Scorzonera ensifolia ------------------- - 33 - -- -12
Achillea micrantha ----------- - 18 - -- - 16 - - 27 - - 18 23 -
Rumex confertus -------------------- - 43----
Crepis sancta -------------------- - 43----
Artemisia absinthium -------------------- - 43----
Papaver dubium -------------------- - 43----
Cichorium intybus 23 - -- -- -- -- -- -- -- 23 --- - 41 ----
Lepidium latifolium -------------------- - 38----
Koeleria macrantha ---------------------35----
Chenopodium opulifolium - - - - - - - - - - - - - - - - - - - - - 35 - - - -
Falcaria vulgaris --------------- - 21--- - 34----
Lamium amplexicaule -------------------- - 33----
Echium vulgare --------------- - 15 - - 20 - 26----
Bassia prostrata -------8------------ - 25----
Elymus farctus --------------------- - 83---
Artemisia tschernieviana --------- - 24 - -- -- -- -- -- 76 ---
Alyssum borzaeanum ----------------------74---
Asperula setulosa - - - - 7 - - - - - 22 - - - - - - - - 23 - - 70 - - -
Sifene exaltata --------------------6-62---
Orobanche coerulescens --------------------- - 54---
Onosma arenaria ------------------- - 12-49 - --
Leymus racemosus ----------------------44---
Linaria genistifolia - - - - - - - - - - - - - - - - - - - - 18 23 27 17 - -
Tnufa salicina s. aspera -------- - 21 - -- -- -- -- -- -- - 80-
Rumex acetosella s. acetosella v. 7
tenuifolius
Vicia hirsuta ----------------------- - 54--
Centaurium pulchellum ------------------------46--
Veronica spicata ------------------------42--
Verbascum phoeniceum - - - - - - - - - - - - - - - - - - - - - - - - 42 - -
Echinops ritro s. ruthenicus ------------------- - 17 - - 18 40--
Scabiosa ochroleuca - - - - - - - - - - - - - - - - - - - - 9 - - 22 31 - -
Thalictrum minus ----------------------- - 31--
Euphorbia cyparissias ----21 - -- -- -- -- -- -- -- -- -- -- 40
Centaurea rhenana - - - - - - - - - - - - - - - - - - - - - - - - - - 29
Chamaecytisus ruthenicus --------------------------27
Apera spica-venti s. maritima 58910- - 42 - -- -- -- -- -- -- -- -- -- -Lotas corniculatus 35 - -- -- -- -- -- -- -- 35 - -- -- -- -- -Matricaria perforata 30 - -- -- -- -- -- -- -- -- -- - 28 -----
Milium vernale 21 31 -35 - -- -- -- -- -- -- -- -- -- -- --
Scirpus holoschoenus - 14 32 25 - -- -- 14 16 - -- -- -- - 15 - -- - 32 --
Salix rosmarinifolia - - - 63 - - - - - - - - - - - - - - - - - - - 63 - - -
Galium ruthenicum ---41 - -- --38 - -- -- -- -- -- -- - 21--
Conyza canadensis - - - 29 - - - - - - - - - 47 - - - - - - - - - - - - -
20 55
Arabidopsis toxophylla - - - - 38 - - - - - 29
Bassia sedoides ----32 - -- --33
Arenaria serpyllifolia - - - - 26 - - - - - - - - - - - - - - 41 - - - - -
Thlaspi perfoliatum ---- 26 - -- -- 49 - -- -- -- -- -- -- -
Anthemis ruthenica - - - - 7 40 - - - - - 56 - - - - - - - - - - - - -Limonium meyeri ----- 34 - -- -- -- -- -- -- -- 39 --Arenaria serpyllifolia s. leptoclados - -- -- 30 - -- -- -- -- -- -- - 14 21 - - 52 Marrubium peregrinum - - - - - 30 - - - - 61 - - - - - - - - 21 - - - - -Pea bulbosa ----- 29 - -- -- -- -- - 20 19 - -- -- - 29
Artemisia taurica - - - - - - 56 - - - 27 - - - - - - - - - - - - - -
Senecio vernalis ----- - 34 - -- 50 - -- -- -- -- -- -- -
Galium humifusum - - - - - - 34 - - - 20 - - - - - - - 11 - - 39 - - -
Xanthopaemelia camschadalis - - 18 - 10 - 28 - - - 51- - - - 19- - - 17- - - - -
Anthemis arvensis ------ - 30 32 - -- -- -- -- -- -- -- -
Linum perenne - - - - - - - - - 45 - - - - - - - - - - - - 33 - -
No. of syntaxa 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 26 27
Number of relevés 10 17 20 13 18 4 9 43 9 10 14 5 10 7 5 12 10 13 21 8 47 16 10 10 10 92 47
Picris hieracioides -------- - 25 49 - -- -- -- -- -- 18 --
Eragrostis minor --------- - 63 - -- -- -- - 27 -----
Alyssum hirsutum - - 13 - 15 - 24 - -- 38 - -- -- -- - 36 - 18 - -Silene spergulifolia --------- - 31 - -- -- -- -- - 27---
Syrenia cana --------- - 29 - -- -- -- -- 12 - --37
Crambe maritima --------- - 27 - -- -- -- 17 - - 30 ---
Lappula squarrosa ----------- 60 - 27 - -- -- -- -- --
Lithospermum officinale ----- - 16 - -- -33 - -51 - -- -- -- -- -
Achillea aspkrifolia - 10 - - 18 - -- -- 13 28 - -- -- -- 26 -----
Seseli tortuosum --- - 19 - -- -- - 28 - -- 17 - -9 29 -----
Polygonum arenarium s. arenarium - -- -- -- -- -- - 73 - -- -- -- -- 39 --
Agropyron dasyanthum ----------- - 45 - -- -- -- 30 - - 18 24
Anchusa gmelinii ------------40------ - 24 - -- 31
Syrenia montana ----------- - 32 - -- -- -- 22 - 38 - 15
Polygonum arenarium - - - - - - - - - - - - - 70 - - - - - - 40 - - - -
Veronica verna ------------ - 58 - - 57 - -- -- -- -
Centaurea apiculata s. adpressa - - - - - - - - - - - - - - 37 - - - - - - - 44 - -
Medicago sativa s.falcata --- - 16 - -- -- 20 - -- 21 33 - -- 29 - - 16 --
Gypsophila muralis --------------- - 43 41 - -- -- --
Eryngium campestre --------------- - 34 - - 36 -23---
Berteroa incana --------------- - 31 30 - -- 23---
Chamaecytisus borysthenicus - - - - 16 - - - - - - - - - - - - - 32 - - 14 - - 65 - - -
Hieracium echioides - - ------ - - - - - - - - - 28 - - 20 - - 21 29 - -
Centaurea margaritacea s. breviceps - - - ------ - - - - - - - - - - - - 60 - - 32 25 - -
Jurinea albicaulis s. laxa - - ------ - - - - - - - - - - - - 47 - - 42 - - -
Alyssum tortuosum - - ------ - - - - - - - - - - - - 43 - - - 40 - -
Minuartia viscosa - - ------ - - - - - - - - - - - - 39 - - 61 20 - -
Thymus pallasianus - - ------ - - - - - - - - - - - - 35 - - 60 - - 8
Cladina arbuscula - - ------ - - - - - - - - - - - - 34 - - - 72 - -
Thymus borysthenicus - - 31 - 36 25
Helichrysum arenarium - - 23 17 - - 25 - - 27 23 8 13
Seseli arenarium - - 12 - - 57 12 44 - -
Silene conica - - ------ - - - - - - - - - - - - 13 - - 46 - 36 - -
Herniaria polygama - - ------ - - - - - - - - - - - - 23 - - 38 28 - -
Cladonia subcariosa - - 21 - 36 61 - -
Cerastium semidecandrum - - ------ - - - - - 15 - - 13 - - - 10 - 24 35 35 - -
Goniolimon graminifolium - - 21 - 31 31 - -
Scabiosa argentea - - - - 17 - - - - - - - - - - - - - - - - 25 - - 31 42 - -
Astragalus varius - - - - - 12 - - - - - 9 - - - - - 16 - - - 6 - - 27 45 - -
Dianthus platyodon - - ------ - - - - - - - - - - - - 22 - - 26 26 12 -
Cerastium pumilum s. glutinosum - 35 16 - - 58 - -- - 26 - -- -- -- -Plantage lanceolata -----46--- - 26 - -- -- 46---
Bassia hirsuta ------40-- - 53 - -- - 29 ----
Cynanchum acutum -- 21 - -- 30 - -- 29 - -- - 21 - -- 28
Teucrium polium ---- 20 - 29 - -- 44--- - 15 - -- 34
Bromus tecterum ------ 27 5 18 - 7 19 27 27 - - 11 - - -
Plantage arenaria ----------- - 49 49 49 ----
Erophila verna --------------- - 37--
Myosotis stricta --------------- - 33--
Centaurea arenaria s. odessana - -- -15 - -- -- -- -- -- -- 19 44 Bassia laniflora ------------------Veronica dillenii ------------------Tragopogon brevirostris s. brevirostris - -- -- -- -- -- -- - 23----
Agropyron cristatum s. sabulosum - -- -- -- -- -- -- -- -- -Senecio borysthenicus ------------25------
Cladonia foliacea - - - - - - - - - - - - - - - - - - -
Xanthopaemelia somloensis -------------------
Alyssum minutum - - - - - - - - - - - - - - - - - - -
Cladonia rangiformis - - - - - - - - - - - - - - - - - - -
Coelocaulon aculeatum - - - - - - - - - - - - - - - - - - -
Centaurea diffusa - - 5 - - 45 33 - - - 30 - - - - 14 29 - -
Cynodon dacylon 10 13 - - 14 31 31 - - 10 25 - - - 14 13 - - - 26
Artemisia campestris agg. - - - - - - - - 28 - - - - - - - - - -
Tortula ruralis -----18 10 - --29 - -- -11 - --18
Asperula graveolens - - - - - - - - - - - - 31 - - - - - -
Koeleria glauca_- - - -- -- -- -- -- 27 - -- 20 - - 26 - 18 27 27 - 27
Aofes: only plant species having diagnostic value were listed; due to the table's widlh restriction, phi values were listed to integer, numbers mark syntaxa: 1 - Aperetum maritimae; 2 -Dauco guttati-Chrysapogonetum grylli; 3 - Caric' celchicae-Heloschoenetum vulgaris, 4 - Salici rosmarinifoliae-Holoschoenetum vulgaris; 5 - Secali-Stipetum borysthenicae; 6 -Cynodonto-Medicaginetum minimae; 7 - Secali-Cynadantetum dactyli; 8 - Secaletum sykestris, 9 - Saca/Ï sylvestri-Brometum tectorum; 10 -Alio guttati-Festucetum rupicolae; 11 -Heliatrapia dohsi-Bmmetum japonici; 12 - Trago-Anthemietum ruthemcae; 13 - Linario odorae-Agropyretum dasyanth;14 - Linario adorae-Agrapyretum dasyanth (sensu Kovalenko); 15 - Plantaginetum arenariae; 16 - Poo butbosae-Caricetum colchicae, 17 - Achilleo setacei-Festucetum beckeri; 18 - Centaureo gerberii-Chamaecytisetum borysthenici; 19 - Centaureo odessanae-Caricetum colchicae; 20 - Centaureo odessanae-Stipetum capillatae; 21 - Centaureo brevicipiti-Festucetum beckeri var. typica; 22 -Centaureo odessanae-Festucetum beckeri (sensu Kolomiychuk et al.); 23 - Centaureo adessanae-FestuceUm beckeri (Vicheiek relevés); 24 - Centaureo brevicipiti-Festucetum beckeri salicetosum rosmarinifoliae; 25 - Centaureo brevicipiti-Festucetum beckeri inuletosum sabuletorum; 26 - Festucetum beckeri var. typica; 27 - Festucetum beckeri var. Koeleria glauca.
The class Festucetea vagfnatae combines communities of sandy the Azov Sea regions. These coenoses are less common on riverine sands. steppes and sandy meadows. In Ukraine, these coenoses are common in the According to our study, in Ukraine, the class includes one order, two allianc-steppe zone on the lower Dnieper River arenas, in the Black Sea region and es and 22 associations. Cluster analysis suggests that Artemisio arenariae-
28 - - - 56
20 - - 51 51
- 40 38 - -
33 - - 49 35
28 - - 57 57
28 - - 38 44
28 - - 26 73
27 - - 37 50
21 - 47 47 47
13 - 46 31 39
15 - 29 50 43
15 - 28 44 39
23 - 25 57 51
28 - - 28 28 :
! 11 10 29 29 29
30 - - 48 31
26 - 18 27 27
Festucion beckeri alliance is comprised of 4-5 sub-alliances, but it is not still possible to identity clear diagnostic blocks (Fig. 5, Table 2). The leading factors of territorial differentiation of phytocoenoses also determining their coenotic diversity are the mesorelief patterns (elements of dunes, interdunal depressions, flat areas in arenas), soil type, its humus content and density, as well as the degree of aeolian processes influence. The hyperspace of ecological conditions from humid (deep depressions) to semi-desert (dune tops) contributes to their significant coenotic richness and diversity. The alliance Festucion beckeri includes coenoses occurring on sandy areas of low hilly coastal and riverine dunes which consist of non-carbonate soils. The alliance Artemisio arenariae-Festucion beckeri includes steppe plant communities occurring on sandy and sandy-shell soils of the Black Sea and the Azov Sea coastal areas.
According to the results obtained from ordination analysis ofFestucetea vagfnatae associations it was found that the ecological differentiation of syntaxa within the class is determined by the complex influence of ecologi-
Table 3
Synoptic table ofHelichryso-Crucianelletea maritimae class
cal factors, and most of plant communities develop under similar environmental conditions (Fig. 6). The general leading factor is the soil salinity regime. Moisture, content of carbonates in the soil and its aeration are of crucial importance to the communities existing in Allio guttati-Festucetum rupicolae and Achilleo setacei-Festucetum beckeri associations. The distribution of Aperetum maritimae, Dauco guttati-Chrysopogonetum grylli and Carici colchicae-Holoschoenetum vulgaris coenoses in the ecological space is determined mainly by content of available nitrogen forms in the soil. Coenoses in the Secaletum sylvestris association are characterized by the widest ecological amplitude. Their placement is determined by the variability of damping and temperature conditions.
Coenoses belonging to Helichryso-Crucianelletea maritimae class occupy specific ecotopes in stabilized overgrown (grey) dunes and overgrown uprush berm crest areas on the coasts of the Black Sea and Azov Sea. In Ukraine, the class includes 1 order, 4 alliances and 10 associations (Fig. 7, Table 3).
No. of syntaxa 1 2 3 4 5 6 7 8 9 10
Number of relevés 13 10 7 10 24 14 10 14 17 9
Onopordum tauricum 60.0 ---------
Matricaria perforata 59.2 ---------
Poa angustifolia 55.1 ---------
Cynanchum acutum 54.5 ---------
Seseli tortuosum 50.8 ---------
Falcaria vulgaris 46.9 ---------
Cerastium gracile 46.3 ---------
Alliumflavum s. tauricum 46.3 ---------
Berteroa incana 45.0 ---------
Camelina microcarpa 41.3 ---------
Bromus squarrosus 40.7 ---------
Bassia laniflora 39.5 ---------
Allium scorodoprasum s. waldsteinii 37.5 ---------
Phragmites australis 37.5 ---------
Galium humifusum 35.3 ---------
Crambe maritima 32.8 ---------
Alyssum desertorum 32.4 ---------
Odontites verna s. serotina 29.5 ---------
Elymus repens 28.8 ---------
Helichrysum arenarium Veronica verna
Arenaria serpyllifolia s. leptoclados Silene spergulfolia Conyza canadensis Scabiosa argentea Lithospermum officinale Centaurea margaritacea s. breviceps Medcago minima Gypsophila paniculata Asperula setulosa Seseli arenarium Artemisia tschernieviana Elymus farctus Cerastium semidecandrum Silene conica Alyssum minutum Syrenia montana Onosma arenaria
Polygonum arenarium s. arenarium Orobanche coerulescens Koeleria glauca
Asparagus officinalis s. officinalis
Festuca beckeri
Linum perenne
Picris hieracioides
Centaurea arenaria s. odessana
Echinops sphaerocephalus
Galium ruthenicum
Centaurea arenaria s. borysthenica
Glycyrrhiza glabra
Consolida regalis
Seseli campestre
Asperula graveolens
Galium tenderiense
Melica transsilvanica s. klokovii
Limonium meyeri
Anchusa gmelinii
Elymus elongatus
Eleocharis palustris
Cuscuta monogyna
51.1 42.9 40.8 35.0 26.3
86.2 57.5
51.4 49.1
35.5
20.0
16.8
21.2
22.3
20.2
15.0
14.2
16.9
14.5
39.1 33.8 33.8 27.5 27.5
87.7 78.6
74.3 63.5 56.2
51.4 51.4 44.4 44.4
No. of syntaxa 1 2 3 4 5 6 7 8 9 10
Number of relevés 13 10 7 10 24 14 10 14 17 9
Linaria dulcis
Papaver dubium s. laevigatum Consolida paniculata Artemisia scoparia Scabiosa ochroleuca Senecio vernalis Bromus tectorum Apera spica-venti s. maritima Aster oleifolius Echium vulgare Anthemis cotula Limonium latifolium Dianthus marschallii Goniolimon tataricum Helianthemum salicifolium Viola kitaibeliana Salvia aethiopis Scorzonera cana Peganum harmala Petrorhagia prolifera Centaurea orientalis Capsella bursa-pastoris Satureja rumelica Potentilla astracanica Valerianella pumila Asperula supina
Euphorbia esula s. tommasiniana Carex liparocarpos Linaria sabulosa Thymus pannonicus Clypeolajonthlaspi Medcago praecox Trigonella monspeliaca Eragrostis minor Erysimum diffusum Arenaria serpyllifolia Stipa capillata Alcea rugosa Sisymbrium orientale Convolvulus lineatus Erodium cicutarium Setaria viridis Thesium arvense Leontodon hispidus Lappula barbata Achillea leptophylla Psilurus incurvus Minuartia hybrida Hieracium umbellatum Tribulus terrestris Artemisia taurica Minuartia glomerata Erophila verna s. praecox Cerastium pumilum s. glutinosum Plantago lanceolata Silene conica s. subconica Papaver rhoeas Heliotropium ellipticum Cerastium fontanum s. vulgare Artemisia austriaca Artemisia campestris agg. Centaurea diffusa Salvia scabiosifolia Centaurea sterilis Apera spica-venti Crataegus laciniata Rosa canina Adonis aestivalis Melica ciliata s. monticola Koeleria macrantha Festuca rupicola Arum elongatum Anthriscus cerefolium Sambucus nigra
Bromus hordeaceus s. hordeaceus Alyssum calycocarpum Trinia crithmifolia Geum urbanum Valerianella coronata Dactylis glomerata Galium aparine Milium vernale
Agropyron cristatum s. ponticum Thalictrum minus
24.2
39.8 36.7
36.3 36.1 36.1 31.5
30.9
26.4
21.3
15.5
24.0
18.8
91.9 91.9
90.1 85.4
84.4
83.2
78.6
77.3
69.7
66.5
63.4 63.1
62.6
61.8 60.1 60.1
59.0 58.9 52.7 51.7 50.9
49.1
48.5
48.5 45.9 44.4 44.4 42.7 41.9
41.6
39.7
38.0
36.1 36.1 34.7 33.0 28.9
24.7
17.0
23.9
20.8
24.8
20.7
66.7 62.2 57.4 52.2 52.2 52.2 46.6 46.6
46.6 40.2 40.2 40.2 40.2 40.2 40.2 40.2 40.2 40.2 40.2 36.9
32.7 32.7
24.6
21.1
No. of syntaxa 1 2 3 4 5 6 7 8 9 10
Number of relevés 13 10 7 10 24 14 10 14 17 9
Dactylis glomerata s. hispanica - - - - - - - - 32.7 -
Buglossoides arvensis - - - - - - - - 32.5 -
Lactuca tatarica - - - - - - - - 31.9 -
Carex ligerica - - - 23.4 - - - - 30.1 -
Agropyron cimmericum - - - - - - - - 27.1 -
Potentilla species - - - - - - - - - 76.3
Linaria genistifolia s. genistifolia - - - - - - - - - 64.7
Asperula tenella - - - - - - - - - 64.7
Onosma tinctoria - - - - - - - - - 60.1
Cerastium pumilum - - - - - - - - - 60.1
Medicago lupulina - - - - - - - - - 55.7
Cichorium intybus - - - - - - - - - 49.8
Elymus farctus s. bessarabicus - - - - - - - - - 49.8
Holosteum umbellatum - - - - - - - - - 45.2
Crepis foetida s. rhoeadifolia - - - - - - - 24.5 - 43.7
Hieracium echioides - - - - - - - - 19.0 42.8
Astrodaucus littoralis - - - - - - - - - 30.1
Coronilla varia - - - - - - - 18.3 - 26.3
Artemisia santonicum 57.2 - - - - 25.6 - - - -
Silene exaltata 53.3 - - 54.0 - - - - - -
Medicago sativa s. falcata 33.5 - - - - - - 23.9 - | 26.0
Gypsophila perfoliata 30.8 25.8 - - - - - - - -
Potentilla argentea 30.8 25.8 - - - - - - - -
Linaria genistifolia 29.5 - - 43.2 - - - - - -
Leymus racemosus s. sabulosus - 47.0 - - 32.9 - - - - -
Eryngium maritimum - 32.6 - - - - - - - 48.2
Melilotus alba - 28.7 - - - - - - - 56.0
Syrenia cana - - 50.6 - - 27.3 - - - -
Anthemis ruthenica - - 41.8 - - - 38.3 - - -
Festuca valesiaca - - 39.7 - - - - 33.0 | - -
Alyssum borzaeanum - - - 64.4 - 58.6 - - - -
Ephedra distachya - - - - 34.3 34.3 - - - -
Centaurea apiculata s. adpressa - - - 24.8 - 30.8 - 30.8 - -
Ajuga chamaepitys s. chia - - - - - - 71.8 46.9 - -
Marrubium peregrinum - - - - - - 53.9 32.6 - -
Cynodon dactylon - - - - - - 47.1 47.1 - -
Linum austriacum s. euxinum - - - - - - 42.3 50.0 - -
Rumex tuberosus - - - - - - 39.5 - 33.7 -
Echinops ritro - - - - - - 37.3 - 46.6 23.6
Eryngium campestre - - - - - - 36.9 41.1 - -
Carduus uncinatus - - - - - - 35.5 58.8 - 24.3
Potentilla taurica - - - - - - 27.8 66.8 - -
Silene densiflora - - - - - - 26.2 50.4 - -
Sideritis montana s. montana - - - - - - - 60.8 - 57.5
Thymus dimorphus - - - - - - - 53.4 - 59.0
Teucrium polium - - - - - - - 52.8 - 28.2
Silene syreistschikowii - - - - - - - 47.5 - 39.2
Elymus uralensis s. viridiglumis - - - - - - - 45.3 20.0 26.9
Stipa borysthenica - - - 21.9 - - - 37.6 34.4 -
Allium guttatum - - - - - - - 33.0 - 27.8
Alyssum tortuosum - - - - - - - 29.5 - 44.4
Plantago arenaria - - - - - - - 26.9 - 43.2
Verbascum pinnatifidum - - - - - - - 23.3 46.8 55.7
Jurinea albicaulis s. laxa - - - - - - - - 39.9 57.3
Leymus racemosus - - - - - - - - 33.3 65.8
Astragalus onobrychis 26.1 - - - - - - - 33.8 41.9
Alyssum turkestanicum - - - - - - - 54.9 41.7 29.9
Alyssum hirsutum - - - - - - - 44.8 37.3 25.9
Chondrillajuncea - - - - - - - 33.6 40.5 33.1
Astragalus varius - - - - - - 23.7 26.8 31.5 36.8
Secale sylvestre - - - - - - - - 24.4 20.9
Euphorbia seguierana - - - 19.6 - - - 23.4 - -
Astragalus cicer - - - - 23.0 - - - - -
Melica ciliata - - - - - - - - 23.8 -
Xanthium strumarium s. italicum - - - - - - - - 21.9 -
Poa bulbosa - - - - - - - 15.7 13.5 -
Notes: numbers mark syntaxa: 1 - Anisantho tectorum-Medicaginetum kotovii; 2 - Anisantho tectorum-Helichrysetum arenariae, 3 - Scabioso ucranicae-Caricetum ligericae; 4 - Secali sylvestri-Alyssetum borzaeani; 5 - EphedroCaricetum colchicae; 6 - Medicagini tenderiensis-Seselietum tenderiensi; 7 - Cynodonto-Ajugetum chiae, 8 - Carici liparicarpoCentaureetum adpressae; 9 -Astragalo borysthenici-Ephedretum distachyae; 10 - Leymo-Verbascetum pinnatifidi.
Ca -1" X 8 / / ) /
Lc -i fH \ "^Ab \
Fig. 4. Results of ordination analysis of syntaxa ofthe class Koelerio-Corynephoretea canescentis; numbers mark syntaxa: 1 - Veronico dillenii-Secaletum sylvestris, 2 - Chamaecytiso ruthenicae-Festucetum
beckeri; 3 - Diantho deltoidis-Armerietum elongatae; 4 -Jasiono montanae-Thymetum serpylli; 5 - Jasiono montanae-Festucetum ovinae;
6 - Corynephoro-Silenetum tataricae; 7 - Artemisio campestris-Dianthetum borbasii; 8 - Artemisio dniproicae-Sedetum sexangularis; 9 -Vetonico dillenii-Secaletum sylvestris (sensu Kovalenko); 10 - Diantho
borbasii-Agrostietum syreistschikovii; 11 - Thymo angustifolii-Festucetum beckeri; 12 - Centaureo borysthenicae-Festucetum beckeri; 13 - Veronico dillenii-Corynephoretum; 14 - Corniculario aculeatae-Corynephoretum canescentis, here and further the scale of ecological factors developed by Didukh (2011) was used: Hd - soil moisture; Nt -content of available nitrogen forms in the soil; Ca - content ofcarbonates; Rc - soil acidity; Sl - salt regime; fH - soil moisture variability; Ae - soil
aeration; Lc - light intensity; Kn - climate continentality; Om -ombroregime; Tm - temperature regime; Cr - cryoregime; DCA1, DCA2 - ordination axes
Fig. 5. Cluster analysis of phytosociological data of the Festucetea vaginatae class; clusters: 1 - Aperetum maritimae; 2 - Dauco guttati-Chrysopogonetum grylli; 3 - Carici colchicae-Holoschoenetum vulgaris, 4 - Salici rosmarinifoliae-Holoschoenetum vulgaris, 5 - Secali-Stipetum borysthenicae; 6 - Cynodonto-Medicaginetum minimae; 7 - Secali-Cynodontetum dactyli; 8 - Secaletum sylvestris, 9 - Secali sylvestri-Brometum tectorum; 10 - Allio guttati-Festucetum rupicolae; 11 -Heliotropio dolosi-Brometum japonici; 12 - Trago-Anthemietum ruthenicae; 13 - Linario odorae-Agropyretum dasyanthi; 14 - Linario odorae-Agropyretum dasyanthi (sensu Kovalenko); 15 - Plantaginetum arenariae; 16 - Poo bulbosae-Caricetum colchicae; 17 -Achilleo setacei-Festucetum beckeri; 18 - Centaureo gerberii-Chamaecytisetum borysthenici; 19 - Centaureo odessanae-Caricetum colchicae; 20 -Centaureo odessanae-Stipetum capillatae; 21 - Centaureo brevicipiti-Festucetum beckeri var. typica; 22 - Centaureo odessanae-Festucetum beckeri (sensu Kolomiychuk); 23 - Centaureo odessanae-Festucetum beckeri (Vicherek relevés); 24 - Centaureo brevicipiti-Festucetum beckeri
salicetosum rosmarinifoliae; 25 - Centaureo brevicipiti-Festucetum beckeri inuletosum sabuletorum; 26 - Festucetum beckeri var. typica; 27 - Festucetum beckeri var. Koeleria glauca
The Ephedra distachyae-Medicaginion romanicae alliance includes plant communities of stabilized coastal dunes and uprush berm crest areas covered with dense sand-shell soils in the Black Sea and Azov Sea. The Medicagini tenderiensis-Seselion tenderiensi alliance is presented by coenoses of mesophytic sand steppes on accumulative sand-shell
macroforms in the northwest coast of the Black Sea. The Scabiosion ucranicae alliance is presented by coenoses of stabilized coastal dunes in the Western Black Sea Region. The Cynodonto-Teucrion polii alliance includes plant communities on young dunes, as well as on uprush limit parts of berm crest areas growing on washed sandy and shelly soils, enriched with detritus, on the Azov-Black Sea coast of Crimea.
L
Kn Tm //Xj ^^^tUCr ;, . .JO. si
/4 j J Nt
Om Hd V10\
Fig. 6. Results ofordination analysis obtained for syntaxa ofFestucetea vaginatae class; numbers mark syntaxa: 1 -Aperetum maritimae, 2 - Dauco guttati-Chrysopogonetum grylli; 3 - Carici colchicae-Holoschoenetum vulgaris, 4 - Salici rosmarinifoliae-Holoschoenetum vulgaris, 5 - Secali-Stipetum borysthenicae; 6 - Cynodonto-Medicaginetum minimae; 7 - Secali-Cynodontetum dactyli; 8 - Secaletum
sylvestris, 9 - Secali sylvestri-Brometum tectorum; 10 - Allio guttati-Festucetum rupicolae; 11 - Heliotropio dolosi-Brometum japonici; 12 -Trago-Anthemietum ruthenicae; 13 - Linario odorae-Agropyretum dasyanthi (Vicherek relevés); 14 - Linario odorae-Agropyretum dasyanthi (sensu Kovalenko); 15 -Plantaginetum arenariae; 16 -Poo bulbosae-Caricetum colchicae; 17 - Achilleo setacei-Festucetum beckeri; 18 -Centaureo gerberii-Chamaecytisetum borysthenici; 19 - Centaureo odessanae-Caricetum colchicae; 20 - Centaureo odessanae-Stipetum capillatae; 21 - Centaureo brevicipiti-Festucetum beckeri var. typica; 22 -Centaureo odessanae-Festucetum beckeri (sensu Kolomiychuk et al.); 23 -Centaureo odessanae-Festucetum beckeri (Vicherek relevés); 24 -Centaureo brevicipiti-Festucetum beckeri salicetosum rosmarinifoliae; 25 - Centaureo brevicipiti-Festucetum beckeri inuletosum sabuletorum; 26 - Festucetum beckeri var. typica; 27 - Festucetum beckeri var. Koeleria glauca
Fig. 7. Cluster analysis of phytosociological data ofHelichryso-Crucianelletea maritimae class: 1 - Anisantho tectorum-Medicaginetum kotovii; 2 - Anisantho tectorum-Helichrysetum arenariae; 3 - Scabioso ucranicae-Caricetum ligericae; 4 - Secali sylvestri-Alyssetum borzaeani; 5 - Ephedro-Caricetum colchicae; 6 - Medicagini tenderiensis-Seselietum tenderiensi; 7 - Cynodonto-Ajugetum chiae; 8 - Carici liparicarpo-Centaureetum adpressae; 9 - Astragalo borysthenici-Ephedretum distachyae; 10 - Leymo-Verbascetum jrirmatifidi
The leading factors of territorial differentiation of these communities are the landforms and their relative age, groundwater level, substrate density and type (sandy, shelly), as well as the degree of dune geomorphostructures stabilization depending on the activity of the sea and aeolian processes.
-2 0 2 4 6
DCA1
Fig. 8. Results obtained from ordination analysis of associations in the class Helichryso-Crucianelletea maritimae: 1 - Anisantho tectorum-Medicaginetum kotovii; 2 - Anisantho tectorum-Helichrysetum arenariae;
3 - Scabioso ucranicae-Caricetum ligericae; 4 - Secali sylvestri-Alyssetum borzaeani; 5 - Ephedro-Caricetum colchicae; 6 - Medicagini tenderiensis-Seselietum tenderiensi; 7 - Cynodonto-Ajugetum chiae; 8 -Carici liparicarpo-Centaureetum adpressae; 9 - jAstmgalo borysthenici-Ephedretum distachyae; 10 - Leymo-Verbascetum pinnatifidi
Om DCA3 Ca
Fig. 9. Results of ordination analysis obtained for the classes Festucetea vaginatae, Koelerio-Corynephoretea canescentis and Helichryso-Crucianelletea maritimae by alliances: numbers mark alliances: 1 - Koelerion glaucae; 2 - Armerion elongatae; 3 - Corynephorion canescentis; 4 - Artemisio arenariae-Festucion beckeri; 5 - Festucion
beckeri; 6 - Ephedro distachyae-Medicaginion romanicae; 7 -Scabiosion ucranicae ; 8 - Medicagini tenderiensis-Seselion tenderiensi;
9 - Cynodonto-Teucrion polii
Within the ecological space, distribution of communities is mainly determined by the variability of damping, ombroregime and climate continentality factors (Fig. 8). They have a greater influence on the coenoses in Anisantho tectorum-Helichrysetum arenariae, Scabioso ucranicae-Caricetum ligericae and Secali sylvestri-Alyssetum borzaeani associations. In addition to these factors, Anisantho tectorum-Medicaginetum kotovii, Ephedro-Caricetum colchicae and Medicagini tenderiensis-Seselietum tenderiensi communities also depend on the degree of soil aeration, soil humidity and the content of various salts. Anisantho tectorum-Helichrysetum arenariae, Scabioso ucranicae-Caricetum ligericae and Ephedro-Caricetum colchicae syntaxa have greater stenotopy, while coenoses of Cynodonto-Teucrion polii alliance developing in the Crimea demonstrate stenotopy by most ecological parameters.
Discussion
Studies on the vegetation cover of natural zones in Eurasia are important for understanding the processes of their development and functioning at the continental scale (Erdos et al., 2018). Therefore, a critical analysis of the accumulated phytocoenological materials against the background of modern knowledge is important. At the present time, the syntaxonomic content of the classes Festucetea vaginatae and Koelerio-Corynephoretea canescentis is still a debatable issue, which is facilitated by the transitional position of their syntaxa and the belonging of certain diagnostic species of the highest classification ranks to the same genus, in particular, coenose-forming species Festuca psammophila, F. polesica, F. vaginata, and F. beckeri. As already noted, a number of European authors classified phytocoenoses of overgrown sands as belonging to the order Festucetalia vaginatae Soo 1957 in the class Koelerio-Corynephoretea canescentis (Dring et al., 2002; Borhidi, 2003; Biondi et al., 2014), when others classified them as entering into the class Festuco-Brometea (Rodwell et al., 2002; Tzonev et al., 2009). Complementary to Ukraine, scientists from the Czech Republic (Chytry, 2007), Romania (Sanda et al., 2008) and Russia (Demina, 2009, 2011; Ermakov, 2012) recognized the class independence, where syntaxonomic diversity of sandy steppes is higher than that of Central Europe. According to the authors, the leading factors of differentiation of communities in Festucetea vaginatae and Koelerio-Corynephoretea canescentis classes are the origin (genesis) of sandy substrates, as well as a reaction of soil solution. In the habitats of coenoses of the class Festucetea vaginatae, sandy soils were developed by marine and river sediments and have an alkaline reaction. Communities of Koelerio-Corynephoretea canescentis class were developed mainly on fluvioglacial sands having acidic reaction. This was confirmed by the ordination analysis results of alliances in these classes (Fig. 9). Climatic conditions oftheir distribution are also important in the steppe (Festucetea vaginatae) and forest and forest-steppe zones (Koelerio-Corynephoretea canescentis), respectively.
Ordination analysis of psammophytic vegetation alliances allowed us to determine the main factors of ecological differentiation of the communities (Fig. 9). They are primarily soil acidity (Rc) and salt regime (Sl). Most of the alliances are located along the DCA1 axis, i.e. along these vectors. According to soil pH gradient, coenoses of the classes Koelerio-Corynephoretea canescentis and Helichryso-Crucianelletea maritimae occupy the opposite positions. Communities of Festucetea vaginatae are located in the center, occupying an intermediate position and overlapping most of the Helichryso-Crucianelletea maritimae syntaxa, which indicates that they occur under similar environmental conditions. However, as the graph shows, one of the main parameters for the latter class is the content of carbonates (Ca) and nitrogen (Nt) in the soil. It is reasonable to assume an influence of ombroregime (Om) on distribution of Koelerio-Corynephoretea canescentis communities associated with forest and forest-steppe zones.
Syntaxa ratio in the classes Koelerio-Corynephoretea canescentis, Sedo-Scleranthetea (vegetation of rock outcrops) and Festucetea vaginatae also remains debatable. Individual European authors recognize the class Koelerio-Corynephoretea canescentis in a broad sense, combining all orders into a single class (Borhidi, 2003). Most researchers combine Koelerio-Corynephoretea canescentis and Sedo-Scleranthetea into one (Rodwell et al., 2002; Dengler, 2004; Chytry, 2007; Sanda et al., 2008; Tzonev et al., 2009). In the latest review of European vegetation, Sedo-Scleranthetea was assigned to a separate class, and Festucetea vaginatae was included in Koelerio-Corynephoretea canescentis as an order (Mucina et al., 2016). Slovak (Valachovic et al., 1995; Jarolimek & Sibik, 2008), Italian (Biondi et al., 2014), Spanish and Portuguese (Rivas-Martinez et al., 2001), Russian (Ermakov, 2012; Dulepova et al., 2018), as well as the majority of Ukrainian phytocoenologists maintain the position of the three classes being independent.
The syntaxonomic position of psammophytic communities with the participation of the shrub Salix rosmarinifolia (association Salici rosmarinifoliae-Holoschoenetum vulgaris Mititelu et al. 1973 and subassociation Centaureo brevicepsis-Festucetum beckeri salicetosum rosmarinifoliae Vicherek 1972) has to be clarified in the future. This is due to the fact that such coenoses were classified as Salicetea arenariae
Weber 1999 on the Atlantic coast area of Western Europe (Mucina et al., 2016), but in Ukraine some authors classified it to the class Nerio-Tamaricetea Br.-Bl. et O. de Bolos 1958, the order Salicetalia arenariae Preising et Weber 1997, and to the provisionally allocated alliance Scirpoido holoschoeni-Salicion rosmarinifoliae I. Solomakha, Vorobyov et Moysienko 2015 prov. (Solomakha et al., 2015), and others classified it to the class Festucetea vaginatae (Dubyna et al., 2003, 2019a).
It should be noted that the position of Jasiono montanae-Festucetum ovinae Klika 1941 within Koelerion glaucae is obviously somewhat ambiguous; since there is clearly a transitional position of its communities to meso-xerophytic vegetation on shell skeletal silicate soils of the Sedo-Scleranthetea class. Czech scientists attributed this association to the alliance Hyperico perforati-Scleranthion perennis Moravec 1967 within the Sedo-Scleranthetea class (Sadlo & Chytry, 2007), but in Ukraine they are described on sandy habitats having a weak acidic reaction of soil solution; so, the authors previously assigned these communities to the Koelerio-Corynephoretea canescentis class. CoenoHora composition and syntaxo-nomic position of the association Festuco psammophilae-Koelerietum glaucae sensu Gal'chenko 2006, non Klika 1931 remain unclear, since the diagnostic species given by the author was Festuca valesiaca (Galchenko, 2006), and the floristic composition differs significantly from the protolog. Further conducting of phytosociological studies of communities with the participation of such psammophytic coenose-forming species as Festuca psammophila and F. polesica are required, in particular, on the territory of the Ukrainian Polesie, which in this regard has not been sufficiently studied. Presumably, phytocoenoses of Festuco psammophilae-Koelerietum glaucae Klika 1931, Spergulo-Festucetum psammophilae Passarge 1960, Diantho arenarii-Festucetum polesicae R. Tx. 1937 and other associations should be distributed on the territory of Ukraine.
Probably, further research on communities in the little-known class Pyrolo-Pinetea sylvestris in Ukraine will allow us to determine the syntaxonomic position of associations Thymo pallasiani-Centauretum sumensis Shevchyk et Solomakha in Shevchyk, Solomakha et Voytuk 1996 and Cladonietum Shevchyk et Polishko 2000 nom. inval. (art. 3b) originally identified by the authors as part of Koelerion glaucae alliance (Shevchyk et al., 1996; Shevchyk & Polishko, 2000). Previously, according to results obtained from the analysis of psammophytic vegetation, the associations in their floral composition were closer to native dry pine forests on sandy soils developed on the sandy terraces of large rivers (Koelerio glaucae-Pinion sylvestris alliance Ermakov 1999, Pyrolo-Pinetea sylvestris class).
Class Helichryso-Crucianelletea maritimae has been allocated recently on the territory of Ukraine (Dubyna et al., 2019a). Previously, its communities were considered within the class Festucetea vaginatae. There are currently two orders were allocated in Western Europe: Artemisio-Koelerietalia Sissingh 1974 (sandy meadows and shrubs on the rich stabilized grey dunes on the coasts of the Atlantic Ocean and of the Northern, the Ligurian and the Adriatic seas) and Crucianelletalia maritimae Sissingh 1974 (Mediterranean, Cantabro-Francoatlantic dwarf shrubs and Xerothermic meadows on stabilized coastal grey dunes). Their plant communities differ significantly from those of the Black Sea region; because of this a separate order was proposed in the class for the territory of Ukraine: Ephedro distachyae-Medicaginetalia romanicae (Dubyna et al., 2019a). Although some researchers see no rationale for separation of this class and consider it as a synonym ofAmmophiletea (Marceno et al., 2018), the results obtained allow us to maintain the position of its ecological and floral isolation. Despite the fact that both of these high-rank syntaxa include several common species (Eryngium marilimum, Leymus racemosus ssp. sabulosus, Ephedra distachya), the class Helichryso-Crucianelletea maritimae is characterized by the presence of a block of diagnostic species that distinguish it from the complex of diagnostic species Ammophiletea and Festucetea vaginatae, at least on the territory of Ukraine. These species are the following: Artemisia tschernieviana, Asparagus maritimus, Astragalus onobrychis, Centaurea apiculata ssp. adpressa, Erodium cicutarium, Medicago romanica (~ M sativa ssp. falcata), Melica transsilvanica ssp. Uokovii, Scabiosa argentea, Tamarix gracilis, Teucrium polium, Trachomitum sarmatiense, Verbascum pinnatifidum. Ecologically stable dune communities also are distinct in that they do not experience periodic effects of sea waves, since they are protected by the berm crest, and their habitats have morphological differences. Most often, these are gently sloping uprush parts ofberm crest
and, to some extent, elevated sand deposits (dunes) anchored mostly by perennial vegetation. According to our data, communities of Helichryso-Crucianelletea maritimae class were even closer to Festucetea vaginatae than to Ammophiletea.
Worldwide, coastal dunes and other psammophytic habitats are recognized as highly vulnerable and dynamic ecosystems (Rannow & Neubert, 2014; Acosta & Ercole, 2015; Sarmati et al., 2019); in Ukraine and other countries they are systematically over-used. Inadequate land management contributes to the fact that these habitats are threatened with extinction, especially in the Mediterranean region (Sabatini et al., 2018). Frequent invasions of alien and atypical species into the structure of communities have been recorded; it resulted in their transformation and degradation (Dubyna et al., 2019b). All of this indicates the need for urgent actions to protect and preserve them.
Conclusion
Pioneer psammophytic vegetation of Ukraine is represented by 45 associations, 9 alliances and 3 orders belonging to 3 classes (Koelerio-Corynephoretea canescentis, Helichryso-Crucianelletea maritimae and Festucetea vaginatae). The main factors of territorial and ecological differentiation of pioneer psammophytic vegetation in Ukraine were identified. It was found that the territorial distribution of communities of Koelerio-Corynephoretea canescentis, Festucetea vaginatae and Helichryso-Crucianelletea maritimae were mainly influenced by ecotope mesorelief pattern, soil composition and the thickness of humic horizon, as well as the degree of aeolian processes development. The main factors of their ecological differentiation are soil acidity, salt content and ombroregime. Based on the results of DCA-ordination of syntaxa within certain vegetation classes, it was found that their distribution is influenced by factors that correlate with the environment-specific conditions. It was found that an ecological differentiation of syntaxa within Festucetea vaginatae is determined by the integrated effect of gradients, and soil salinity can be distinguished among these. The main factors in syntaxa distribution within the Koelerio-Corynephoretea canescentis class were temperature regime and climate continentality, as well as the gradients of ombroregime and soil humidity. The distribution of communities of Helichryso-Crucianelletea maritimae class in the ecological space is determined mainly by factors of variability of damping, ombroregime and climate continentality. Coenoses of Secaletum sylvestris, Diantho deltoidis-Armerietum elongatae, Corynephoro-Silenetum tataricae, Artemisio dniproicae-Sedetum sexangularis, Thymo angustifolii-Festucetum beckeri, Anisantho tectorum-Helichrysetum arenariae, Scabioso ucranicae-Caricetum ligericae and Ephedro-Caricetum colchicae associations were characterized by the widest ecological amplitude.
The authors adhere to the position of independence of the studied classes: Koelerio-Corynephoretea canescentis, Helichryso-Crucianelletea maritimae and Festucetea vaginatae, considering that the main factors of differentiation of communities in the classes Festucetea vaginatae and Koelerio-Corynephoretea canescentis were the origin (genesis) of sandy substrates, as well as the reaction of soil solution. Phytosociological analysis of a large number of coastal littoral vegetation relevés also provide support for independence of Helichryso-Crucianelletea maritimae and Ammophiletea classes based on their floristic and ecological differences.
The conducted research will allow the place of selected syntaxonomic units of the pioneer psammophytic vegetation of Ukraine to be determined in the pan-European system. The results of the work will be suitable for development of the Nature Reserve network and Eco-network of Ukraine, maintaining Prodrome of the vegetation of Ukraine, Prodrome of the vegetation of Europe, preparing the next volume of the publication "Vegetation of Ukraine" and "Green Data Book ofUkraine", for further study of successional and adaptation processes in extreme environmental conditions of pioneer habitats, establishing patterns of regenerative geosystems' development and finding out the ways to manage them. The ability of plant communities to indicate the environment state (phytoindication of environmental conditions), which in recent years has become an urgent sociological necessity, will allow the environmental situation in the regions to be monitored, as well help in as determining trends in their further changes. The authors gratefully acknowledge A. Kuzemko, O. Kovalenko, O. Chusova, O. Senchylo, V. Kolomiychuk, T. Fitsailo, I. Goncharenko, V. Konischuk, G.
Tsukanova, Y. Gaiova, D. Iakushenko, V. Konogray, for the kindly provided relevés
of psammophyte vegetation from their dissertation manuscripts.
References
Acosta, A., & Ercole, S. (Eds.). (2015). Gli habitat delle coste sabbiose italiane: Ecologia e problematiche di conservazione [Italian coastal dune habitats: Ecology and conservation issues]. ISPRA, Serie Rapporti (in Italian).
Aleshkina, U. M. (2011). Rastitelnye soobshchestva zelenoy zony g Kieva [Plant communities of the green zone of Kiev]. In: Vserossiyskaya nauchnaya konferentsiya "Otechestvennaya geobotanika: Osnovnye vekhi i perspektivy". Boston-Spektr, Sankt-Petersburg. Pp. 7-10 (in Russian).
Androsova, A. Y., & Solomakha, T. D. (1996). Psamofilna roslynnist' Bilosarayskoi kosy i morskoho uzberezhzhya poblyzu m. Mariupolya [Psammophilous vegetation of the Bilosarai spit and the sea coast near Mariupol]. Ukrainian Phytosociological Collection, Series A, 1, 41-49 (in Ukrainian).
Bairak, O. M. (1998). Florystychna klasyfikatsiya roslynnoho pokryvu Livoberezhnoho Prydniprov'ya [Floristic classification of vegetation of the Left Bank Dnieper]. Ukrainian Botanical Journal, 55(2), 139-145 (in Ukrainian).
Bardat, J., Bioret, F., Botineau, M., Boullet, V., Delpech, R., Genu, J.-M., Haury, J., Lacoste, A., Rameau, J.-C., Royer, J.-M., Roux, G., & Touffet, J. M. (2004). Prodrome des végétations de France [The prodrome of French vegetation]. Muséum National d'Histoire Naturelle, Paris (in French).
Berg, C., Dengler, J., Abdank, A., & Iermann, M. (Eds.). (2004). Die Pflanzengesellschaften Mecklenburg-Vorpommerns und ihre Gefährdung [The plant communities of Mecklenburg-Vorpommern and their vulnerability]. Wiessdorn Verlag, Jena (in German).
Biondi, E., Blasi, C., Allegrezza, M., Anzellotti, I., Azzella, M. M., Carli, E., Casavecchia, S., Copiz, R., Del Vico, E., Facioni, L., Galdenzi, D., Gasparri, R., Lasen, C., Pesaresi, S., Poldini, L., Sburlino, G., Taffetani, F., Vagge, I., & Zivkovic, L. (2014). Plant communities of Italy: The vegetation prodrome. Plant Biosystems, 148(4), 728-814.
Bondareva, V. V., Nikolaychuk, L. F., & Golub, V. B. (2019). Kserofitnye rastitelnye soobshchestva doliny Nizhney Volgi [Xerophytic plant communities ofthe Lower Volga valley]. Phytodiversity of Eastern Europe, 13(1), 4-29 (in Russian).
Borhidi, A. (2003). Magyarorszag növcnytärsuMsai [Plant associations in Hungary]. Akadémiai Kiadö, Budapest (in Hungarian).
Braun-Blanquet, J. (1964). Pflanzensoziologie - Grundzüge der Vegetationskunde [Phytosociology - Broad Vegetation Science]. 3rd ed. Springer, Wien (in German).
Bulokhov, A. D. (2019). Soobshchestva klassa Sedo-Scleranhetea Br.-Bl. 1955 v poyme reki Desny [Communities of the class Sedo-Scleranhetea Br.-Bl. 1955 in floodplain of the Desna River]. Diversity of Plant World В, 3(3), 56-66 (in Russian).
Cao, G., Tsuchiya, K., Zhu, W., & Okuro, T. (2019). Vegetation dynamics of abandoned paddy fields and surrounding wetlands in the lower Tumen River Basin, Norlheast China. PeerJ, 7, 1-17.
Chusova, O. O. (2019). Roslynnist' ta biotopy baseynu r. Krasna [Vegetation and biotopes ofthe Krasna river basin]. Phytosociocentre, Kyiv (in Ukrainian).
Chynkina, T. B. (2003). Syntaksonomiya i antropohenna dynamika roslynnosti hyr-lovoji oblasti Dnipra [Syntaxonomy and anthropogenic dynamics of vegetation of the mouth of the Dnieper]. Publishing Kherson National University, Kyiv (in Ukrainian).
Chytry, M. (Ed.). (2007). Vegetace Ceské republiky. 1. Travinna a kerickova vegetace [Vegetation of the Czech Republic. 1. Grassland and Heathland Vegetation]. Academia, Praha (in Czech).
Chytry, M., Chiarucci, A., Pärtel, M., Pillar, V. D., Bakker, J. P., Mucina, L., Peet, R. K., & White P. S. (2019). Progress in vegetation science: Trends over the past three decades and new horizons. Journal of Vegetation Science, 31(1), 1-4.
Çoban, S., & Willner, W. (2019). Numerical classification ofthe forest vegetation in the Western Euxine Region of Turkey. Phytocoenologia, 49 (1), 71-106.
Davydova, А О. (2019). Syntaksonomiya roslynnosti Natsionalnoho Pryrodnoho Parku "Dzharylhatskyi". Klas Festucetea vaginatae [Syntaxonomy of vegetation of national nature park "Dzharylhatskyi". The class Festucetea vaginatae]. Biology and Ecology, 5(1), 34-43 (in Ukrainian).
Demina, O. N. (2009). Soobshchestva klassa Festucetea vaginatae Soö em. Vicherek 1972 na territorii Tsimlyanskikh peskov v Rostovskoy oblasti [Communities of the class Festucetea vaginatae Soö em. Vicherek 1972 on the territory of Tsimlyansk sands in the Rostov region]. In: Materialy Moskovskogo Tsentra Russkogo Geograficheskogo Obshchestva. Biogeografiya, Moscow, 15, 27-38 (in Russian).
Demina, O. N. (2015). Klassifikatsiya rastitelnosti stepey basseyna Dona [Classification of the steppe vegetation of the Don basin]. Publisher of the Southern Federal University, Rostov na Donu (in Russian).
Dengler, J. (2004). Klasse Koelerio-Corynephoretea Klika in Klika et Novak 1941 -Sandtrockenrasen und Felsgrusfluren von der submeridionalen bis zur borealen Zone [Class Koelerio-Corynephoretea Klika in Klika et Novak 1941 - dry sand
turf and rocky meadows from the submeridional to the boreal zone]. In: Berg, C., Dengler, J., Abdank, A., & lermann, M. (Eds.). Die Pflanzengesellschaften Mecklenburg-Vorpommerns und ihre Gefährdung. Wiessdorn Verlag, Jena. Pp. 301-326 (in German).
Didukh, Y. P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Phytosociocentre, Kyiv.
Didukh, Y. P. (2012). Osnovy bioindykatsii [Fundamentals of bioindication]. Naukova Dumka, Kyiv (in Ukrainian).
Didukh, Y. P., & Korotchenko, I. A. (1996). Stepova roslynnist' pivdennoi chastyny Livoberezhnoho Lisostepu Ukrainy. 1: Klasy Festucetea vaginatae ta Helianthemo-Thymetea [Steppe vegetation of the southern part of the Left-Bank Forest-Steppe of Ukraine. 1: Classes Festucetea vaginatae and Helianthemo-Thymetea]. Ukrainian Phytosociological Collection, Series A, 2, 56-63 (in Ukrainian).
Dring, J., Hoda, P., Mersinllari, M., Mullaj, A., Pignatti, S., & Rodwell, J. (2002). Plant communities of Albania - a preliminary overview. Annali di Botanica, 2, 7-30.
Dubyna, D. V., Neuhäuslova, Z., & Shelyag-Sosonko, Y. R. (1995). Vegetation of the "Birjucij Island" Spit in Azov Sea. Sand Steppe Vegetation. Folia Geobotanica et Phytotaxonomica, 30, 1-31.
Dubyna, D. V., Dziuba, T. P., Zhmud, O. I., Tymoshenko, P. A., & Shelyag-Soson-ko, Y. R. (1996). Syntaksonomiya roslynnosti Zhebryyanskoho prymorskoho pasma (Odeska obl.). I. Pisky [Syntaxonomy of vegetation of the Zhebriyansky coastal ridge (Odessa region). I. Sands]. Ukrainian Phytosociological Collection, Series A, 2, 44-56 (in Ukrainian).
Dubyna, D. V., Shelyag-Sosonko, Y. R., Zhmud, O. I., Dvoretskiy, T. V., Dziuba, T. P., & Tymoshenko, P. A. (2003). Dunayskyi biosfernyi zapovidnyk. Roslyn-nyi svit [Dunaisky Biosphere Reserve. Plant Kingdom]. Phytosociocentre, Kyiv (in Ukrainian).
Dubyna, D. V., Neuhäuslova, Z., Dziuba, T. P., & Shelyag-Sosonko, Y. R. (2004). Klasyfikatsiya ta prodromus syntaksonomichnoi riznomanitnosti vodoym, perezvolozhenykh terytoriy ta aren Pivnichnoho Prychornomor'ya [Classification and Prodrome of vegetation of reservoirs, overmoisturized territories and aren of the Nothern part of the Black Sea region]. Phytosociocentre, Kyiv (in Ukrainian).
Dubyna, D. V., & Dziuba, T. P. (2005). Fitotsenotychna riznomanitnist' ostrova Dzharylhach (Khersonska oblast) [Phytocenotic diversity of Dzharylgach island (Kherson region)]. Ukrainian Botanical Journal, 62(2), 255-269 (in Ukrainian).
Dubyna, D. V., Tymoshenko, P. A., & Dvoretskyi, T. V. (2009). Ekoloho-florys-tychni osoblyvosti uhrupovan klasu Festucetea vaginatae v Ukraini ta zavdannya yikh okhorony [Ecological and floristic features of groups of the class Festucetea vaginatae in Ukraine and tasks of their protection]. Chornomorski Botanical Journal, 5(4), 491-501 (in Ukrainian).
Dubyna, D. V., Dziuba, T. P., Iemelianova, S. М., Bagrikova, N. О., Borysova, О. V., Borsukevych, L. M., Vynokurov, D. S., Gapon, S. V., Gapon, Y. V., Davydov, D. A, Dvoretsky, T. V., Didukh, Y. P., Zhmud, О. I., Kozyr, М. S., Konyschuk, V. V., Kuzemko, А А., Paskevych, N. А., Ryff, L. Е., Solomakha, V. A, Felbaba-Klushyna, L. М., Fitsailo, Т. V., Chorna, G. A, Chorney, I. I., Shelyag-Sosonko, Y. R., & Iakushenko, D. М. (2019a). Prodromus roslynnosti Ukrainy [Prodrome of the vegetation of Ukraine]. Naukova Dumka, Kyiv (in Ukrainian).
Dubyna, D. V., Iemelianova, S. M., Dvoretzkiy, T. V., Dziuba, T. P., & Tymoshenko, P. A. (2019b). Adventization of coenofloras of the classes of pioneer vegetation in Ukraine. Ukrainian Botanical Journal, 76(6), 499-510 (in Ukrainian).
Dulepova, N. A, Korolyuk, А Y., Yamalov, S. M., Lebedeva, M. V., & Golovanov, Y. M. (2018). Rastitelnost peschanykh stepey Orenburgskoy oblasti [Sandy steppe vegetation in Orenburg Region]. Vegetation of Russia, 33, 53-65 (in Russian).
Dziuba, T. P., Solomakha, T. D., & Tymoshenko, P. A. (2010). Roslynni uhrupovannya [Vegetation communities]. In: Pereyaslav-Khmelnytskiy. Nature: Vegetation. Critical inventory annotated notes of flora and vegetation: Vascular plants, bryophytes, lichens, and algae. FOP Majdanchenko l. S., Korsun-Shevchenkivsky, 86-95 (in Ukrainian).
Erdös, L., Ambarli, D., Anenkhonov, O. A., Batori, Z., Cserhalmi, D., Kiss, M., Kröel-Dulay, G., Liu, H., Magnes, M., Molnar, Z., Naqinezhad, A., Semenishchenkov, Y. A., Tölgyesi, C., & Török, P. (2018). The edge of two worlds: A new review and synthesis on Eurasian forest-steppes. Applied Vegetation Science, 21, 345-362.
Ermakov, N. B. (2012). Prodromus vysshikh edinits rastitelnosti Rossii [Prodrome of higher units of vegetation in Russia]. In: Mirkin, B. M., & Naumova, L. G. (Eds.). Sovremennoe sostoyanie osnovnykh kontseptsiy nauki o rastitelnosti [The current state of the basic concepts of vegetation science]. Gilem, Ufa. Pp. 377-483 (in Russian).
Gaiova, J. Y. (2015). Psamofitna roslynnist' Cherkasko-Chyhyrynskoho heobotani-chnoho rayonu [The Psamofit Vegetation of Cherkasy-Chigirin Geobotanic Region]. Naukovyi Visnyk NLTU Ukrainy, 25(9), 131-138 (in Ukrainian).
Gomlya, L. M. (2005). Roslynnist' dolyny richky Khorol [Vegetation of the Khorol River valley]. Ukrainian Phytosociological Collection, Series A, 22, 2-187 (in Ukrainian).
Gordienko, I. I. (1969). Oleshskie peski i biogeotsenoticheskie svyazi v protsesse ikh zarastaniya [Oleshski Sands and biogeocenotic connections in the process of their overgrowth]. Naukova Dumka, Kyiv (in Ukrainian).
Galchenko, N. P. (2006). Rehionalnyi landshaitnyi park "Kremenchutski plavni". Roslynnyi svit [Kremenchuk Plavni Regional Landscape Park. Plant Kingdom]. Phytosociocentre, Kyiv (in Ukrainian).
Hennekens, S. M., & Schaminee, J. H. J. (2001). Turboveg, a comprehensive date base management system for vegetation data. Journal of Vegetation Science, 12, 589-591.
Hill, M. O., & Gauch, H. G. (1980). Detrended correspondence analysis: An improved ordination technique. Vegetatio, 42, 47-58.
Hill, M. O. (1979). Twinspan - a Fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and the attributes. Ithaca, NY.
Jansons, A, Robalte, L., Cakss, R., & Matisons, R. (2016). Long-term effect of whole tree biomass harvesting on ground cover vegetation in a dry Scots pine stand. Silva Fennica, 50(5), 1661.
Jarolimek, I., Sibik, J., Hegedusova, K., Janisova, M., Kliment, J., Kucera, P., Majekova, J., Michalkova, D., Sadlonova, J., Sibikova, I., Skodova, I., Tichy, L., Ujhazy, K., Ujhazyova, M., Uhlifova, J., Valachovic, M., & Zaliberova, M. (2008). Diagnostic, constant and dominant species of the higher vegetation units of Slovakia. Veda, Bratislava.
Konishchuk, V. V. (2006). Otsinka riznomanitnosti ekosystem Cheremskoho Pryrodnoho Zapovidnyka na osnovi kartohrafichnoho modelyuvannya [Estimation of ecosystem diversity of Cheremsky Nature Reserve on the basis of cartographic modeling]. Dia, Kyiv (in Ukrainian).
Korolyuk, A Y., Lebedeva, M. V., Yamalov, S. M., Golovanov, Y. M., Dulepova, N. A., Zolotareva, N. V., & Teptina, A. Y. (2018). The petrophytic steppes of the Urals: Diversity and ecological drivers. The Fourth International scientific conference on ecology and geography of plants and plant communities. KnE Life Sciences. Pp. 88-94.
Korzhenevsky, V. V. (1986). Rastitelnost dyun Kryma [The vegetation of the Crimean dunes]. Collection of Scientific Papers of the State Nikitsky Botanical Garden, 98, 122-133 (in Russian).
Korzhenevsky, V. V., & Klyukin, A. A. (1990). Rastitelnost abrazionnykh i akkumulyativnykh form rel'efa morskikh poberezhiy i ozer Kryma [Vegetation of abrasion and accumulative relief forms of the Crimean sea coasts and lakes]. Moscow, Dep. v VINITI 10.07.1990, No 3822-B90 (in Russian).
Korzhenevsky, V. V., Bagrikova, N. A., Ryff, L. E., & Levon, A. R. (2003). Prodromus rastitelnosti Kryma (20 let na platforme floristicheskoy klassifikatsii) [Prodromus of the Crimean vegetation (20 years on the platform of floristic classification)]. Bulletin of Main Botanical Garden of RAN, 186, 32-51 (in Russian).
Korzhenevsky, V. V., & Kvitnitskaya, A. A. (2014). Sintaksonomiya rastitelnosti eo-lovogo relefa Kryma [Syntaxonomy of vegetation of the Crimean eolian relief]. Works of the State Nikitsky Botanical Gardens, 136, 41-55 (in Russian).
Kovalenko, O. A. (2016). Flora, roslynnist' ta fitosozolohichni aspekty NPP "Pyryatynskyi" [Flora, vegetation and phytososological aspects of NPP "Pyriatynsky"]. Mria, Sumy (in Ukrainian).
Kuzemko, A. (2009). Dry grasslands on sandy soils in the forest and forest-steppe zones of the plains region of Ukraine: Present state of syntaxonomy. Tuexenia, 29, 369-390.
Kuzemko, A., Becker, T., Didukh, Y. P., Arde-lean, I. V., Becker, U., Beldean, M., Dolnik, C., Jeschke, M., Naqinezhad A, Ugurlu, E., Unal, A., Vassilev, K., Vorona, E. I., Yavorska, O. H., & Dengler, J. (2014). Dry grassland vegetation of Central Podolia (Ukraine) - a preliminary overview of its syntaxonomy, ecology and biodiversity. Tuexenia, 34, 391-430.
Kuzemko, A. A., Steinbauer, M. J., Becker, T., Didukh, Y. P., Dolnik, C., Jeschke, M., Naqinezhad, A., Ugurlu, E., Vassilev, K., & Dengler, J. (2016). Patterns and drivers of phytodiversity of steppe grasslands of Central Podolia (Ukraine). Biodiversity and Conservation, 25(12), 2233-2250.
Landucci, F., Sumberova, K., Tichy, L., Hennekens, S., Aunina, L., Bita-Nicolae, C., Borsukevych, L., Bobrov, A., Carni, A., De Bie, E., Golub, V., Hrivnak, R., Iemelianova, S., Jandt, U., Jansen, F., K^cki, Z., Lajer, K., Papastergiadou, E., Silc, U., Sinkeviciene, Z., Stancic, Z., Stepanovic, J., Teteryuk, B., Tzonev, R., Venanzoni, R., Zelnik, I., & Chytry, M. (2020). Classification of the European marsh vegetation (Phragmito-Magnocaricetea) to the association level. Applied Vegetation Science, 23 (2), 297-316.
Lengyel, A., Landucci, F., Mucina, L., Tsakalos, J. L., & Botta-Dukat, Z. (2018). Joint optimization of cluster number and abundance transformation for obtaining effective vegetation classifications. Journal of Vegetation Science, 29, 336-347.
Marceno, C., Guarino, R., Loidi, J., Herrera, M., Isermann, M., Knollova, I., Tichy, L., Tzonev, R. T., Acosta, A. T. R., FitzPatrick, U., Iakushenko, D., Janssen, J. A. M., Jimenez-Alfaro, B., K^cki, Z., Keizer-Sedlakova, I., Kolomiychuk,
V. P., Rodwell, J. S., Schaminée, J. H. J., Silc, U., & Chytry, M. (2018). Classification of European and Mediterranean coastal dune vegetation. Applied Vegetation Science, 21(1), 1-27.
Matuszkiewicz, W. (2008). Pizewodnik do oznaczania zbiorowisk roslinnych Polski [Guide for identification of the plant communities of Poland]. Pañstwowe Wydawnictwo Naukowe, Warszawa (in Polish).
Mucina, L., Bültman, H., Dierssen, K., Theurillat, J.-P., Dengler, J., Carni, A., Sumberová, K., Raus, T., Di Pietro, R., Gavilán Garcia, R., Chytry, M., Iakushenko, D., Schaminée, J. H. J., Bergmeier, E., Santos Guerra, A., Daniels, F. J. A., Ermakov, N., Valachovic, M., Pigantti, S., Rodwell, J. S., Pallas, J., Capelo, J., Weber, H. E., Lysenko, T., Solomeshch, A., Dimopoulos, P., Aguiar, C., Freitag, H., Hennekens, S. M., & Tichy, L. (2016). Vegetation of Europe: Hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science, 19(1), 3-264.
Onyshchenko, V. A. (2006). Florystychna klasyfikatsiya roslynnosti Ukrainskoho Polissya [Floristic classification of vegetation of Ukrainian Polissya]. In: Fitoriznomanittya Ukrainskoho Polissya ta yoho okhorona [Phytodiversity of Ukrainian Polissya and its protection]. Phytosociocentre, Kyiv. Pp. 43-84 (in Ukrainian).
Orlov, O. O., & Yakushenko, D. M. (2005). Roslynnyi pokryv proektovanoho Ko-rostyshivskoho natsionalnoho pryrodnoho parku [Vegetation of the projected Korostyshiv National Nature Park]. Phytosociocentre, Kyiv (in Ukrainian).
Polishko, O. D. (2001). Syntaksonomiya roslynnosti dilyanky borovoi terasy (Prokhorivske lisnytstvo Cherkaskoi oblasti) [Syntaxonomy of pine terrace vegetation (Prochoriv forestry of Cherkasy region)]. Zapovidna Sprava v Ukraini, 7(1), 11-20 (in Ukrainian).
Polishko, O. D. (2005). Syntaksonomiya roslynnosti dilyanky borovoi terasy Dnipra (Chyhyrynske lisnytstvo Cherkaskoi oblasti) [Syntaxonomy of vegetation of the Dnieper pine terrace area (Chyhyryn forestry of Cherkasy region)]. Aktualni Problemy Botaniky ta Ekolohii, 1, 163-176 (in Ukrainian).
Rannow, S., & Neubert, M. (Eds.). (2014). Managing protected areas in Central and Eastern Europe under climate change. Advances in Global Change Research. Vol. 58. Springer Nature.
Rivas-Martínez, S., Fernández-González, F., Loidi, J., Lousa, M., & Penas, A
(2001). Syntaxonomical checklist of vascular plant communities of Spain and Portugal to association level. Itinera Geobotanica, 14, 5-341.
Rodwell, J. S., Schaminee, J. H. J., Mucina, L., Pignatti, S., Dring, J., & Moss, D.
(2002). The diversity of European vegetation. An overview of phytosociological alliances and their relationships to EUNIS habitats. National Centre for Agriculture, Nature Management and Fisheries, Wageningen.
Rolecek, J., Tichy, L., Zeleny, D., & Chytry, M. (2009). Modified Twinspan classification in which the hierarchy respects cluster heterogeneity. Journal of Vegetation Science, 20, 596-602.
Sabatini, F. M., Jiménez-Alfaro, B., Burrascano, S., Lora, A., Chytry, M. (2018). Beta-diversity of Central European forests decreases along an elevational gradient due to the variation in local community assembly processes. Ecography, 41, 1038-1048.
Sádlo, J., & Chytry, M. (2007). Jasiono montanae-Festucetum ovinae Klika 1941. In: Chytry M. (Ed.), Vegetace Ceské republiky. 1. Travinná a kerícková vegetace [Vegetation of the Czech Republic. 1. Grassland and Heathland Vegetation]. Pp. 353-355 (in Czech).
Sanda, V., Ollerer, K., & Burescu, P. (2008). Fitocenozele din Romania. Sintaxonomia, structura, dinamica si evolutie. Ars Docendi, Bucure§ti (in Romanian).
Sarmati, S., Bonari, G., & Angiolini, C. (2019). Conservation status of Mediterranean coastal dune habitats: Anthropogenic disturbance may hamper habitat assignment. Rendiconti Lincei. Scienze Fisiche e Naturali, 30, 623-636.
Senchylo, O. O. (2010). Roslynnist' zaplavy Dnipra v mezhakh Lisostepu [Vegetation ofthe Dnieper floodplain within the Forest-Steppe zone]. Phytosociocentre, Kyiv (in Ukrainian).
Shevchyk, V. L., & Polishko, O. D. (2000). Syntaksonomiya roslynnosti dilyanky borovoi terasy Dnipra (Liplyavske lisnytstvo Cherkaskoi oblasti) [Syntaxonomy of pine terrace vegetation (Liplavo forestry of Cherkasy region)]. Ukrainian Phytosociological Collection, Series A, 16, 67-89 (in Ukrainian).
Shevchyk, V. L., & Solomakha, V. A. (1996). Syntaksonomiya roslynnosti ostroviv Kruhlyk ta Shelestiv Kanivskoho pryrodnoho zapovidnyka [The syntaxonomy of vegetation of Kruglyk and Shelestiv islands in Kaniv Nature Reserve]. Ukrainian Phytosociological Collection, Series A, 1, 12-27 (in Ukrainian).
Shevchyk, V. L., Solomakha, V. A., &Voityuk, Y. O. (1996). Syntaksonomiya ros-lynnosti ta spysok flory Kanivskoho pryrodnoho zapovidnyka [The syntaxono-my of vegetation and list of the flora of Kaniv Nature Reserve]. Ukrainian Phytosociological Collection, Series B, 1(4), 2-120 (in Ukrainian).
Slezák, M., Hrivnák, R., & Machava, J. (2017). Environmental controls of plant species richness and species composition in black alder floodplain forests of central Slovakia. Tuexenia, 37, 79-94.
Solomakha, I. V., Vorobyov, Y. O., & Moysiyenko, I. I. (2015). Roslynnyi pokryv lisiv ta chaharnykiv Pivnichnoho Prychornomor'ya [Vegetation of forests
and shrubs of the Northern Black Sea Coast]. Phytosociocentre, Kyiv (in Ukrainian).
Solomakha, V. A. (2008). Syntaksonomia rosnynnosti Ukrainy. Tretie nablyzhennia [The syntaxonomy of vegetation of the Ukraine. The third approximation]. Phytosociocentre, Kyiv (in Ukrainian).
Sörensen, T. A. (1948). A method of establishing groups of equal amplitude in plant sociology based on similary of species content, and its application to analyses of the vegetation on Danish commons. Kongelige Danske Videnskabernes Selskabs Biologiske Skrifter, 5, 1-34.
Soroka, M. I. (2008). Roslynnist' Ukrainskoho Roztochchya [Vegetation of the Ukrainian Roztocze]. Svit, Lviv (in Ukrainian).
Ter Braak, C. J. F., & Smilauer, P. (2015). Topics in constrained and unconstrained ordination. Plant Ecology, 216(5), 683-696.
Tichy, L. (2002). Juice, software for vegetation classification. Journal of Vegetation Science, 13, 451-453.
Tsukanova, H. O. (2005). Florystychne ta tsenotychne riznomanittya ostroviv Dnipra v mezhakh m. Kyyeva ta yoho okhorona [Floristic and coenotic diversity of the Dnieper islands within the city of Kyiv and its protection]. Kyiv (in Ukrainian).
Tymoshenko, P. A. (1999). Suchasnyi stan klasyfikatsii roslynnosti aren pivdnya Ukrainy [The modern state of classification of vegetation of the arenas of Southern Ukraine]. Ukrainian Phytosociological Collection, Series A, 12-13, 201-205 (in Ukrainian).
Tyshchenko, O. V. (2006). Roslynnist' prymorskykh kis pivnichnoho uzberezhzhya Azovskoho morya [The vegetation of the northern coastal spits of the Sea of Azov]. Phytosociocentre, Kyiv (in Ukrainian).
Tzonev, R. T., Dimitrov, M. A., & Roussakova, V. H. (2009). Syntaxa according to the Braun-Blanquet approach in Bulgaria. Phytologia Balcanica, 15(2), 209-233.
Umanets, O. Y., & Solomakha, I. V. (1999a). Syntaksonomiya roslynnosti Chornomorskoho biosfernoho zapovidnyka II. Ostriv Tendra [Syntaxonomy of vegetation of the Black Sea Biosphere Reserve. П. Tendra Island]. Ukrainian Phytosociological Collection, Series A, 12-13, 63-77 (in Ukrainian).
Umanets, O. Y., & Solomakha, I. V. (1999b). Syntaksonomiya roslynnosti Chornomorskoho biosfernoho zapovidnyka. lll. Dilyanka lvano-Rybalchanska [Syntaxonomy of vegetation of the Black Sea Biosphere Reserve. Ш. Ivano-Rybalchanska site]. Ukrainian Phytosociological Collection, Series A, 14, 84102 (in Ukrainian).
Valachovic, M., Ot'ahel'ova, H., Stanova, V., & Maglocky, S. (Eds.). (1995). Rastlinne spolocenstva Slovenska 1. Pionierska vegetacia [Plant communities of Slovakia 1. Pioneer vegetation]. Veda, Bratislava.
Venables, W. N., & Smith, D. M. (2008). An introduction to R. 2nd edition. Network Theory Ltd.
Vicherek, J. (1972). Die Sandpflanzengesselschaften des unteren und mittleren Dnieprstromgebietes (die Ukraine). Folia Geobotanica et Phytotaxonomica, 7, 9-46.
Vorobyov, Y. O., Balashov, L. S., & Solomakha, V. A. (1997). Syntaksonomiya roslynnosti Polis'kogo pryrodnogo zapovidnyka [The syntaxonomy of vegetation of the Polesie Nature Reserve]. Ukrainian Phytosociological Collection, Series B, 8, 2-128 (in Ukrainian).
Ward, J. H., Jr. (1963). Hierarchical grouping to optimize an objective function. Journal ofthe American Statistical Association, 58, 236-244.
Weber, H. E., Moravec, J., & Theurillat, J.-P. (2000). International code of phytosociological nomenclature. 3rd edition. Journal of Vegetation Science, 11, 739-768.
Westhoff, V., & van der Maarel, E. (1973). The Braun-Blanquet approach. 2-nd ed. In: Whittaker, R. (Ed.). Classification of Plant Communities. Pp. 287-399.
Whittaker, R. H. (1978). Approaches to classifying vegetation on classification of plant communities. 2-nd ed. The Hague, Junk.
Willner, W., Rolecek, J., Korolyuk, A., Dengler, J., Chytry, M., Janisova, M., Lengyel, A., Acic, S., Becker, T., Cuk, M., Demina, O., Jandt, U., K^cki, Z., Kuzemko, A., Kropf, M., Lebedeva, M., Semenishchenkov, Y., Silc, U., Stancic, Z., Staudinger, M., Vassilev K., & Yamalov, S. (2019). Formalized classification of semi-dry grasslands in Central and Eastern Europe. Preslia, 91, 25-49.
Willner, W., Tichy, L., & Chytry, M. (2009). Effects of different fidelity measures and contexts on the determination of diagnostic species. Journal of Vegetation Science, 20, 130-137.
Yakushenko, D. M. (2004). Nova asotsiatsiya psamofilnoi roslynnosti zi skhodu Zhytomyrskoho Polissya [New association of psammophilous vegetation from the east of Zhytomyr Polissya]. Visnyk of Lviv National University. Biology Series, 35, 95-101 (in Ukrainian).
Yousaf, A., Shabbir, R., Jabeen, A., Erum, S., & Ahmad, S. S. (2016). Linkage between herbaceous vegetation and soil characteristics along rawal dam Islamabad. Journal of Soil Science and Plant Nutrition, 16(1), 88-100.
Zhou, L., Shen, H., Chen, L., Li, H., Zhang, P., Zhao, X., Liu, T., Liu, Sh., Xing, A., Hu, H., & Fang, J. (2019). Species richness and composition of shrub-encroached grasslands in relation to environmental factors in northern China. Journal of Plant Ecology, 12(1), 56-66.
