The article presents information about new syntaxa of anthropogenic vegetation in the Kursk Region. We have been conducting research on this type of vegetation since 2003. As a result, a syntaxonomy ...of synanthropic vegetation of Kursk was elaborated (Arepieva, 2015) and a number of papers were published, describing the diversity of phytocenoses of anthropogenic ecotopes of the region. In this paper, 24 associations and 1 derivative community from 4 classes of anthropogenic vegetation are characterized. 210 complete relevés made by the author in 2003–2022 on the territory of Kursk and the regional centers of the Kursk Region were used in the work. Some information about the natural conditions of the region is given in Table 1. Classification is carried out according to Braun-Blanquet approach (Westhoff, Maarel, 1978). The data were treated by IBIS 7.2 software package (Zverev, 2007). The identification of new syntaxa was carried out in accordance with the “International Code of Phytosociological Nomenclature. 4th edition” (Theurillat et al., 2021).The names of the higher syntaxa follow «Vegetation of Europe…» (Mucina et al., 2016). Synoptic tables include only species with constancy above I. Cluster analysis of relevés by the Ward binding method and DCA ordination based on the complete matrix “species composition × relevé” were carried out by PAST 2.17 software package (Hammer et al., 2001). Ecological conditions were assessed with the use of average values by H. Ellenberg et al. (1992) ecological scales and processed by IBIS software (Zverev, 2007). In the class Sisymbrietea, the ass. Hordeetum murini Libbert 1932 was identified (Table 2, Fig. 1). It belongs to the alliance Sisymbrion officinalis and the order Sisymbrietalia sophiae. These communities are found in courtyards, on ruderalized lawns, along sidewalks and highways. Ass. Setarietum verticillatae Méndez 1984 (Table 2, Fig. 2) was assigned to the class Digitario sanguinalis–Eragrostietea minoris, the order Eragrostietalia, the alliance Spergulo arvensis–Erodion cicutariae. These stands grow on ruderalized lawns, along sidewalks, on abandoned flower beds, near buildings. In the class Artemisietea vulgaris and the order Onopordetalia acanthii, 4 associations were identified, they occur near railway embankments and highways, in wastelands. Two associations (Carduo acanthoidis–Onopordetum acanthii Soó ex Jarolímek et al. 1997 and Carduetum acanthoidis Felföldy 1942) were assigned to the alliance Onopordion acanthii (Table 3, Figures 3 and 4). Two associations (Artemisio vulgaris–Echinopsietum sphaerocephali Eliáš 1979 and Artemisio–Oenotheretum rubricaulis Passarge 1977) were assigned to the alliance Dauco-Melilotion (Table 4, Fig. 5). The order Agropyretalia intermedio-repentis and the alliance Convolvulo arvensis–Agropyrion repentis unite communities of the late successional stages. They include 4 associations (Table 5, Fig. 6): Saponario officinalis–Petasitetum spurii Passarge 1964, Anisantho–Artemisietum austriacae Kostylev 1985, Elytrigio repentis–Cirsietum arvensis Prunier et Guenat in Prunier et al. 2018, Dactylido glomeratae–Lupinetum polyphyllis Bulokhov et al. 2020. These communities are common in wastelands, along roads. A derivative community Phalacroloma annuum we also assigned to the class Artemisietea vulgaris (Table 5, Fig. 7). The class Epilobietea angustifolii includes 4 orders and 4 alliances. 4 associations were identified in the order Arctio lappae–Artemisietalia vulgaris and the alliance Arction lappae (Table 6 and 7, Figures 8 and 9): Hyoscyamo nigri–Conietum maculati Slavnić 1951, Helianthetum tuberosi (Moor 1958) Oberdorfer 1967, Geranio sibirici–Arctietum tomentosi Bulokhov et al. 2020 and a new ass. Geranio sibirici–Festucetum giganteae ass. nov. hoc loco. Holotypus: Table 7, relevé 25, Kursk Region, Kursk, courtyard of the house No. 20 on Stepan Razin Street (51.743497° N, 36.190895° E), 22.08.2020, author – L. A. Arepieva. Diagnostic species: Festuca gigantea and Geranium sibiricum. It unites stands with a predominance of Festuca gigantea and Geranium sibiricum. They are common in shaded, infrequently disturbed habitats near the walls of buildings, in parks and squares. There are 4 associations in the order Circaeo lutetianae–Stachyetalia sylvaticae and the alliance Aegopodion podagrariae (Table 10, Fig. 12): Symphyto officinalis–Anthriscetum sylvestris Passarge 1975, Arctio tomentosi–Rumicetum obtusifolii Passarge 1959, Chaerophylletum bulbosi Tüxen 1937, Urtico dioicae–Rubetum caesii Golovanov 2017. They are often found in shaded areas (near the walls of buildings, in gardens and parks). In the order Galio-Alliarietalia and the alliance Geo urbani–Alliarion petiolatae, 3 associations were identified (Table 12, Figure 13): Geo urbani–Chelidonietum majoris Jarolímek et al. 1997, Parthenocissetum insertae Bulokhov et al. 2020, Torilidetum japonicae Lohmeyer ex Görs et Müller 1969. They are common in open and shaded disturbed habitats. Three associations were assigned to the order Convolvuletalia sepium and the alliance Senecionion fluviatilis (Table 13): Calystegio–Angelicetum archangelicae Passarge 1959, Calystegio sepium–Epilobietum hirsuti Hilbig et al. 1972 and Urtico dioicae–Echinocystietum lobatae Bulokhov et Kharin 2008. The latter association includes 2 variants: typica and Arctium tomentosum. Communities of the variant Arctium tomentosum occur in less humid and eutrophic habitats. Whereas other associations are common in wetter habitats.
The article analyzes the monograph by V. G. Storozhenko, V. V. Chebotareva, P. A. Chebotarev «Degradation and regeneration of oak forests in the zone of forest-steppe (results of experimental ...studies)». The research of an author team devoted to the Tellerman oak forest growing system is examined in detail.
Along with the species composition and richness of species, an important component of the communities structure is the ratio of groups of species that perform similar functions, the so-called ...functional diversity (Ackerly and Cornwell, 2007; Diaz et al., 2007; Vasilevich, 2016). Functional diversity is assessed by a set of species properties, one of the commonly used species traits is the life form of species. Life forms (LF) reflect the adaptation of plants to macro- and microecological habitat conditions and characterize the structure of communities. When classifying communities, a taxonomic only approach may have limitations (Mucina, 1997), but despite this, the use of species traits to assess community type and structure is not sufficient. The main questions of this paper: 1) How do the spectra of life forms (biomorphological spectra) look like in different types of forest communities? 2) Are differences in species composition reflected in the biomorphological spectra of forest communities? And how are differences in life form spectra exhibited for different categories of the classification system? 3) Are secondary forests different from native forests by their biomorphological spectra? The study of the structure of syntaxa is based on the forest communities of the Moscow Region (Morozova et al., 2017, 2021, 2022 a, b; Chernenkova et al., 2018), located in the center of the Russian Plain. According to remote sensing data, the forests cover 51 % of region area (Chernenkova et al., 2020). Classification of forests was performed by the Braun-Blanquet approach. The forests of the Moscow Region belong to 10 associations of 4 classes of forest vegetation: Vaccinio-Piceetea Br.-Bl. in Br.-Bl. et al. 1939, Carpino-Fagetea Jakucs ex Passarge 1968, Alnetea glutinosae Br.-Bl. et Tx. ex Westhoff et al. 1946, Alno–Populetea albae P. Fukarek et Fabijanic 1968. More than a third part of the forests are secondary, they represent the facies of zonal associations, which they replaced. Taking into account the facies of secondary forests, the analysis was performed for 15 syntaxa. The classification of LF was carried out according to the Serebryakov system with additions (Zhmylev et al., 2017). The variety of LF of vascular plants was summarized in 13 large groups according to the state of skeletal axes, type of root system, and life longivity (Table 1). The shares of LF groups were calculated as weighted by coverage. To compile a “complete” portrait of communities, we included the proportion of bryophytes in the spectrum. Differences between syntaxa in terms of LF shares were analyzed using analysis of variance (ANOVA). A clear connection was revealed between the number of LF and the number of species of vascular plants in the communities: the more species, the more LF. For logarithmic values, it has a linear form (Fig. 1), however, to assert or refute the universality of the direct relationship between species and functional diversity, it is necessary to expand the testing units from different vegetation classes. For the considered syntaxa, the proportions of LF groups in communities differ significantly (F = 21.971, p = 0.00001). Basically, differences appear between syntaxa of different classes, i.e. within the same class, syntaxa according to the corresponding LF may not be differentiated (Table 2, Figs. 2–4). Formulas for the highest classification units have been compiled according to the predominant LF groups. Thus, in LF spectra of the Carpino-Fagetea class, trees LF (Tr) predominate; second and third places are occupied by long-rhizome herbaceous perennials (LR) and shrubs (Shrub), followed by short rhizome herbaceous perennials (SR) (Fig. 3). In general, the spectrum of syntaxa of this class can be represented as Tr–LR–Shrub or или Tr–LR–Shrub. The Carpino-Fagetea syntaxa are also characterized by the highest proportion of taproots (S), an increased proportion of bulbs/stem tubers (Bu) and root sprouting (R) species among herbaceous perennials in at least one of the syntaxa of the class. According to the biomorphological spectra, secondary forests of the Moscow Region differ slightly from the communities of native forests, which emphasizes their relatively good state and the possibility of transition to native forests in the course of further succession. In the Vaccinio-Piceetea, trees predominate, followed by mosses and dwarf shrubs (SShr) or shrubs. The generalized formula for the spectrum of mesophytic syntaxa of this class is Tr–Mosses–SShr/Shrub (Fig. 4). A significant proportion of tufted LF species (Tu) is recorded in this class compared to other forest classes. However, the biomorphological spectra of different communities of this class differ greatly, which reflects the modern view on the syntaxa system. In contrast to the mesophytic communities of the boreal forest class, the spectra of associations of the order Vaccinio uliginosi–Pinetalia sylvestris Passarge 1968are dominated by mosses, followed by either shrubs (ass. Vaccinio uliginosi–Pinetum de Kleist 1929) or trees LF (ass. Vaccinio uliginosi–Betuletum Libb. 1933). Structure formula of the syntaxon of sphagnum pine forests is Mosses–SShr–Tr. This indicates a clear transitional character of boggy pine communities and the possibility of placing the association both in the class of boreal forests and in the class of open sphagnum bogs of the class Oxycocco-Sphagnetea Br.-Bl. et Tx. ex Westhoff et al. 1946 (Table 4). However, according to the results of comparison of biomorphological spectra between boggy pine forests and forested open bogs (Fig. 6), sphagnum pine forests of the Moscow Region are assigned to ass. Vaccinio uliginosi–Pinetum of the class Vaccinio-Piceetea. The spectrum formula of associations included in the class of black alder forests Alnetea glutinosae is Tr–LR–SR, with mosses on the fourth place (Fig. 4). A similar biomorphological spectrum is for a syntaxon from the class Alno–Populetea albae, but in these communities, shrubs are on the fourth place, and among the all considered forest syntaxa of the Region, the proportion of annuals/biennials (An/Bi) is the highest here. The revealed patterns can be used to characterize communities, they adequately reflect the species richness, structure of communities and emphasize the ecological differences of syntaxa and can be an additional argument for placing the syntaxon in the system of higher units.
Kedrovaya Pad Reserve is located in the southwestern part of Primorye Territory, Russian Far East. This nature reserve was established more than 100 years ago aiming to preserve unique stands of ...virgin south Manchurian mixed forest with Abies holophylla, Pinus koraiensis and numerous broadleaf species. The structure of vegetation was studied thoroughly, however, classification was performed with the emphasis on dominant species and geographical characteristics of the location (Vasilyev et al., 1984), which can be rather useful for forestry but usually does not reflect all peculiarities of described community. Proposed here classification based on Braun-Blanquet approach provides another glance on vegetation structure and allows to consider results within the frames of the existing syntaxonomic system for Northeast Asia. We analyzed 109 relevés, which we collected during the years 2008–2018 (Fig. 1). Cover of each species was estimated in percent. We applied unsupervised classification — modified TWINSPAN (Roleček, 2009) via JUICE 7.0 (Tichý, 2002) to receive seven clusters (Fig. 2, Table 1). Clusters experienced expert revision and were turned into seven groups. These groups were interpreted as associations or subassociations (Table 2). Diagnostic species were identified basing on calculated constancy and phi coefficient (Chytrý et al., 2002). The studied forest vegetation of Kedrovaya Pad Reserve is presented by six associations and three subassociations. One of the seven groups incorporates three relevés, which describe deciduous communities on freshly formed alluvial soils. However, these relevés differ significantly between each other and obtained group doesn’t have any syntaxonomic meaning. Spodiopogono sibirici–Quercetum mongolicae ass. nov. hoc loco Holotypus: relevé 18 in Table 4. Diagnostic species: Artemisia keiskeana, Polygonatum humile, Spodiopogon sibiricus. Communities of this association prefer steep and moderate south or west slopes with mild water deficiency. Trees form one or two strata. Quercus mongolica dominates the canopy, often accompanied by Tilia amurensis. Second tree stratum, if presented, formed by Acer pseudosieboldianum and, again, Tilia amurensis. Typical for the shrub layer is Lespedeza bicolor. Most noticeable species of the herb layer are Artemisia keiskeana, Carex siderosticta & C. nanella, Melampyrum roseum, Spodiopogon sibiricus and Calamagrostis brachytricha. Viti amurensis–Quercetum mongolicae ass. nov. hoc loco Holotypus: relevé 10 in Table 5. Diagnostic species: Chloranthus japonicus, Geranium eriostemon, Heracleum dissectum, Vitis amurensis. Important feature of the habitats for communities of this association is sufficient amount of moisture and this can be met in the lower parts of the slopes, as well as on the top of the ridges (450–500 m a. s. l.), where sea fogs bring additional humidity. Trees form two strata. Quercus mongolica, Tilia amurensis and Fraxinus rhynchophylla form the upper one. Acer mono is ordinary in the second stratum. Shrub layer is dominated by Corylus mandshurica, other species are Rubus crataegifolius, Philadelphus tenuifolius and Weigela praecox. Vines — Vitis amurensis, Schisandra chinensis and Actinidia kolomikta are also common. In the herb layer there are no strong dominants, most characteristic are Carex lanceolata & C. siderosticta, Artemisia stolonifera and several Vicia species. Rhododendro mucronulati–Betuletum schmidtii ass. nov. hoc loco Holotypus: relevé 9 in Table 6. Diagnostic species: Betula schmidtii, Carex nanella, Rhododendron mucronulatum. Communities of this association occupy moderate slopes or tops of the ridges, facing north-east or north-west; soils are rocky and dry. Quercus mongolica and Betula schmidtii form the highest stratum. Acer pseudosieboldianum presents second stratum. Rhododendron mucronulatum dominates in shrub layer. Carex nanella, C. siderosticta and young shoots of vine Schisandra chinensis prevail in the herb layer. Matteuccio struthiopteridis–Juglandetum mandshuricae ass. nov. hoc loco Holotypus: relevé 13 in Table 7. Diagnostic species: Brachybotris paridiformis, Cardamine leucantha, Carex pallida, Filipendula palmata, Hylomecon vernalis, Juglans mandshurica, Matteuccia struthiopteris, Oxalis obtriangulata, Phlomoides maximowiczii, Sanicula rubriflora, Stellaria bungeana, Valeriana fauriei. Communities of this association can be met within river terraces, which are most close to the floodplain. Amount of moisture is sufficient or excess. Upper trees belong to the vast number of species; most common are Tilia amurensis, Fraxinus mandshurica, Juglans mandshurica, Pinus koraiensis, Abies holophylla. Second stratum is also rich in species, with constant Acer mono. The latter one along with Carpinus cordata forms the third stratum. In the shrub layer common are Eleutherococcus senticosus, Viburnum sargentii and several Lonicera species. Among numerous species typical for the herb layer are diagnostic ones. Two subassociations of this association are distinguished. M. s.–J. m. typicum subass. nov. hoc loco occupies the same habitat as specified for association and has the same set of diagnostic species. M. s.–J. m. chrysosplenietosum villosi subass. nov. hoc loco prefers more wet habitats with poorly drained soils. From the aforementioned subassociation it differs by the composition of the herb layer. Holotypus: relevé 6 in Table 7. Diagnostic species: Chrysosplenium pseudofaurie, C. villosum, Cornopteris crenulatoserrulata, Geum aleppicum, Pseudostellaria japonica. Polysticho tripteron–Pinetum koraiensis typhicum Gumarova et al. ex Krestov et al. ex Krestov et al. 2023 Releves belonging to this syntaxon are presented in Table 8. Communities belonging to this subassociation occupy north facing gentle or moderate slopes. The tree layer consists of three strata. The upper one is formed by Abies holophylla up to 50 m high. Besides fir, Tilia amurensis, Quercus mongolica, Pinus koraiensis and Kalopanax septemlobus are common. The second stratum is built by Abies holophylla and Tilia amurensis, accompanied by Acer mono. Acer pseudosieboldianum and Carpinus cordata form the lowest tree stratum. In shrub layer Philadelphus tenuifolius dominates. Species of the herb layer can be divided into two groups: constant with low abundance — Phryma asiatica, Maianthemum dilatatum, Thalictrum filamentosum и T.tuberiferum, Carex siderosticta; and ferns Dryopteris crassirhizoma & D. expansa, Osmundastrum asiaticum with the cover 5–50 % each. Vines Schisandra chinensis, Actinidia arguta & A. kolomikta are well pronounced. Weigelo praecocis–Aceretum ukurunduensis ass. nov. hoc loco Holotypus: relevé 4 in Table 9. Diagnostic species: Euonymus maximowicziana, Gymnocarpium dryopteris, Weigela praecox. Communities of this association were observed only on the slopes of Mt. Uglovaya higher than 600 m a. s. l. The tree layer consists of two or three strata. Abies nephrolepis, Tilia amurensis and birches — Betula costata or B. lanata form the highest one. Lower strata are built by Acer ukurunduense, A. pseudosieboldianum & A. tegmentosum. For the shrub layer common are Weigela praecox, Philadelphus tenuifolius, Deutzia amurensis. In the herb layer Carex siderosticta is abundant, other noticeable group is ferns and vine Actinidia kolomikta.
The Polar-Alpine Botanical Garden-Institute named after N. A. Avrorin (PABGI) Kola Research Centre RAS, Russian Botanical Society, and the Russian geobotany have suffered an irreparable loss. On ...April 6, 2024 after a serious illness the outstanding geobotanist, a well-known specialist in the field of studying flora and vegetation of the Far North, a leading research scientist of the PABGI, PhD Natalia Evgenievna Koroleva has left us. Natalia Evgenievna was born in 1963 in Chernovskoe village, Kirov Region, in the family of agronomist and school teacher. After school graduation, in 1981 she entered the Moscow State University, where she specialized in the Department of Geobotany under the supervision by Prof. T. A. Rabotnov and Prof. A. P. Tyrtikov. After the University, in 1986 she was firstly enrolled in the Laboratory of Biocenology and afterwards in the Laboratory of Flora and Plant Resources of the PABGI. From 1986 to 1990, she passed the postgraduate probation in the Komarov Botanical Institute RAS (St. Petersburg) under the supervision by Prof. B. N. Norin. In 1991, she defended her PhD thesis «Vegetation classification of the mountain tundra belt of Khibiny» and received the degree of candidate of biology (PhD). Throughout her scientific life, Natalia Evgenievna was associated with PABGI having gone from probationer to the senior researcher in the Laboratory of Flora and Plant Resources. For many years, she was a Head of Murmansk Branch of the Russian Botanical Society (RBO). She organized and led a lot of geobotanical expeditions to the Murmansk Region and Spitsbergen Archipelago. The geography of her field researches was highly diverse. She brilliantly knew not only the vegetation of the Kola Peninsula, but as well other regions of Russian Arctic, Norway, Finland and Spitsbergen. The study and conservation of species and community diversity of Arctic and Subarctic vegetation, namely the plant cover of the Kola Peninsula and Spitsbergen Archipelago and revealing its transformation under the man impact, was in the focus by Natalia Koroleva scientific interests. She made a great contribution to the study of the Murmansk Region flora and vegetation, the regional Red Book conducting, plant cover monitoring and Nature conservation. In the Natalia Koroleva person, Russian vegetation science has lost an outstanding scientist, the specialist in phytocoenology, vegetation classification, the study of biodiversity and Nature conservation. She elaborated the vegetation classification method based upon the analysis of the uniformity of plant species distribution with similar ecology in particular plant community groups. Using this method, Natalia Evgenievna and her colleagues developed a detailed classification of plant communities and habitat types of zonal and mountain tundra and highland heaths of the Kola Peninsula and Spitsbergen that has resulted in numerous publications. She had published about 150 scientific papers and 10 monographs, books and guides to the Nature reserved territories of Khibiny Mountains, Lovozero Tundra, Spitsbergen and many others that put a huge impact in the development of vegetation science and Nature conservation in the Murmansk Region. She was an editor of many scientific proceedings and collective monographs. Natalia Evgenievna greatly contributed to the training of young scientists: For ten years (2001–2011), she gave lectures at the Apatity Branch of Moscow State Technical University in the ecology, biodiversity, and vegetation classification. Some Master’s and PhD theses were prepared under her supervision. Natalia Evgenievna devoted a lot of time and efforts to the work in RBO. For about 25 years she was a Head of Murmansk Branch of RBO and a member of its Council, took part in the Organizing Committees of numerous conferences, symposiums and workshops held by RBO. The memory of Natalia Evgenievna Koroleva — an outstanding geobotanist, teacher, environmental activist and popularizer of science — will always remain in her numerous scientific works, and in the hearts of students and colleagues.
Dono-Tsimlyansk sand massif is one of the largest in the South-East of the European part of Russia. It represents a complex of river terraces of different ages within the ancient continental delta ...between the Tsimla and Aksenets rivers. A cluster analysis of 140 relevés was carried out using the Ward method and Czekanowski-Dice-Sørensen similarity coefficient (Fig. 4). The dataset was divided into two large groups. The first one (A) represents communities, which are formed mainly by obligate psammophytes (Table 1), many of species are used in the diagnosis of the class Festucetea vaginatae Soó ex Vicherek 1972. The second group (B) is characterized by high importance values of xerophytes and mesoxerophytes, typical for steppes and xeric meadows of the class Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947. Analysis at the level of 6 clusters shows that clusters 1-4 are poorly differentiated in terms of species importance values (Table 2). Clusters 5 and 6 are clearly distinguished by groups of dominant plants. Cluster 5 represents shrub communities with mesoxerophites: Scirpoides holoschoenus, Poa angustifolia, Galium ruthenicum and Calamagrostis epigeios. Cluster 6 unites forb-bunchgrass steppes, usually dominated by Festuca rupicola, they correspond to the Festuco-Brometea. Most of the communities belong to the class Festucetea vaginatae and the order Festucetalia vaginatae. Diagnostic species of class and order (D.s.): Artemisia campestris, A. marschalliana, Astragalus varius, Chondrilla graminea, C. juncea, Euphorbia seguieriana, Festuca beckeri, Gypsophila paniculata, Kochia laniflora, Koeleria glauca, K. sabuletorum, Pilosella echioides, Secale sylvestre, Silene borysthenica, Stipa borysthenica. We identified the new alliance Agropyrion tanaiticum all. nov. D. s.: Agropyron tanaiticum, Chamaecytisus borysthenicus, Dianthus polymorphus, Holosteum umbellatum, Scabiosa ucranica, Senecio borysthenicus, Seseli tortuosum, Thymus pallasianus, Tragopogon tanaiticus, Veronica verna. Holotypus — the association Scabioso ucranicae–Stipetum borysthenicae ass. nov. The alliance unites sandy steppes with the main range in the lower Don basin. Ass. Scabioso ucranicae–Stipetum borysthenicae ass. nov. (Table 4, rel. 1–21), holotypus — Table 4, rel. 5 (field no. 22-214): Rostov Region, Tsimlyansky district, south of the Nizhengnutov village, Tsimlyansky nature reserve, gentle slope of sandy mound, 47.87754°N, 42.45692°E, 06.08.2022, author — A. Yu. Korolyuk. D. s.: Agropyron lavrenkoanum, Artemisia austriaca, Astragalus varius, Centaurea adpressa, C. arenaria, Eryngium campestre, Galium ruthenicum, Scabiosa ucranica, Seseli tortuosum, Stipa borysthenica. The association unites communities of sandy steppes, occupying leveled, well-fixed sands. It dominates the territory of the Tsimlyansky Reserve, where an anthropogenic pressure is limited. The coenoses are rich, on average 37 species per relevé, with an average cover of 40 %. The main dominant is Stipa borysthenica, its cover can reach 25–35 %. Euphorbia seguieriana and Artemisia marschalliana are constantly co-dominating. These three species form the upper layer 40–60 cm high. Agropyron lavrenkoanum, Festuca beckeri, Poa bulbosa and Potentilla arenaria are recorded in some stands with a cover of more than 10%. A moss layer is usually developed, composed of Syntrichia ruralis with a cover of up to 30 %. The association is synonymous with the subassociation Secali–Stipetum borysthenicae jurinetosum cyanoidis Demina 2009 described from Dono-Tsimlyansk sand massif. Association Secali–Stipetum borysthenicae Korzh. 1987 ex Dubyna et al. 1995 is distributed on the coastal dunes of the Black Sea and Azov; it does not occur on continental sands. Ass. Centaureo gerberi–Chamaecytisetum borysthenicae Demina 2009 (Table 5, оп. 1–36). D. s.: Agropyron tanaiticum, Centaurea gerberi, Chamaecytisus borysthenicus, Dianthus polymorphus, Thymus pallasianus, Tragopogon tanaiticus. The association represents the communities on moderately fixed sands. They form large contours, occupying slopes and tops of sandy hills, bottoms of deflation areas. The total cover varies from 15 to 60 %. Moss layer of Syntrichia ruralisis usually well developed, in some stands covering almost the entire surface. The species richness varies from 16 to 33, averaging 22 species per relevé. The set of dominant species is not constant, Artemisia arenaria, A. marschalliana, Chamaecytisus borysthenicus, Euphorbia seguieriana, Festuca beckeri, Helichrysum arenarium, Jurinea polyclonos, Koeleria sabuletorum, Secale sylvestre, Silene borysthenica are noted with a cover of more than 10 %. As part of the association, we distinguish 3 variants: typica (Table 5, rel. 1–17), Secale sylvestre (Table 5, rel. 18–24), Jurinea polyclonos (Table 5, rel. 25–36). Ass. Secalo sylvestris–Salicetum rosmarinifoliae ass. nov. (Table 4, rel. 22–30), holotypus — Table 4, rel. 28 (field no. 22-256): Rostov Region, Tsimlyansky district, south of the Nizhengnutov village, Tsimlyansky nature reserve, flat depression, 47.87348°N, 42.49095°E, 12.08.2022, author — A. Yu. Korolyuk. D. s.: Salix rosmarinifolia (дом.), Spiraea hypericifolia (дом.), Calamagrostis epigeios, Galium ruthenicum, Inula salicina, Poa angustifolia, P. bulbosa, Scirpoides holoschoenus, Secale sylvestre. The association represents shrub communities with the participation of psammophytes, occupying flat depressions in leveled and hilly sands. The coexistence in the same coenoses of Salix rosmarinifolia, common in wetlands, and xerophytic plants is typical for sandy landscapes. A characteristic feature of communities is a shrub layer composed of Salix rosmarinifolia and Spiraea hypericifolia with total cover from 10 to 35 % at a height of 80–120 cm. Among forbs the most common dominant is Scirpoides holoschoenus, less often Secale sylvestre, Galium ruthenicum, Poa bulbosa, Anisantha tectorum dominate. The moss layer with up to 15–30 % coverage was formed by Syntrichia ruralis. The class Festuco-Brometea, the order Festucetalia valesiacae and alliance Festucion valesiacae represent steppes. The suballiance Artemisio campestris–Stipenion capillatae unites grasslands on sandy soils. Ass. Scabioso ucranicae–Festucetum rupicolae ass. nov. (Table 4, rel. 31–41), holotypus — Table 4, rel. 38 (field no. 22-269): Rostov Region, Tsimlyansky district, south of the Nizhengnutov village, Tsimlyansky nature reserve, flat depression, 47.94400°N, 42.46205°E, 13.08.2022, author — A. Yu. Korolyuk. D. s.: Allium sphaerocephalon, Apera spica-venti, Centaurea adpressa, Festuca rupicola, Linaria genistifolia, Scabiosa ucranica, Stipa borysthenica, Vicia villosa. The association represents forb-bunchgrass sandy steppes. They are found along fixed sands, where they occupy periphery of flat, often extensive depressions, usually near forests and along the edge of wetlands. The coenoses are rich, with an average of 44 species per relevé and total cover from 50 to 70 %. The main dominant is Festuca rupicola, whose cover can reach 40–45 %. It forms the main layer about 30 cm high. Artemisia austriaca, Stipa borysthenica, Salvia tesquicola and Galium ruthenicum are permanent co-dominants. Agropyron lavrenkoanum, Bromopsis inermis, Carex colchica, C. supina, Euphorbia seguieriana, Poa bulbosa, Vicia villosa, Securigera varia occasionally have a high abundance.
There is a lot of studies concerning steppe vegetation in the Transbaikalia region, though detailed floristic classification has been developed recently (Korolyuk, 2017, 2019). However, some of the ...specific types of steppe communities are still out of that classification so far. In particular, meadow steppe communities on permafrost soils at the northern part of the forest-steppe subzone on the Vitimskoye Plateau (Transbaikalia) still are not classified. The paper aims to design the syntaxonomic position and brief characteristics of this particular vegetation. Rather similar communities are known as cryoxerophytic steppes from more northern regions (Yurtsev, 1978). Taking into account that studied area is slightly less cold we call Tranbaikalian meadow steppe communities on permafrost soils as hemicryomesoxerophytic steppe. Our studies have been carried out within the Eravninskaya depression (Fig. 1) in the southwest of the Vitimskoye Plateau (Transbaikalia). The depression of hollow-concave shape with flat relief extends for about 80 km to the northeast, while the width is about 63.5 km, and elevation is about 950 m a. s. l. The climate is sharply continental, with frosty snowless winters and short and warm summers. The permafrost is common throughout the depression. The mean annual precipitation is 250–400 mm, with 60 % of the total amount falls in July–August. The main types of soils are represented by permafrost chernozem and turf soil (Badmayev et al., 2006). Meadow steppe in combination with the Gmelin larch and white birch coppices form the forest-steppe landscape. Field studies have been conducted in 2009 and 2018, and 26 relevés (of 100 m2) of steppe communities were collected. The Braun-Blanquet approach for the classification of vegetation has been applied, and the nomenclature of syntaxa follows to ICPN (Theurillat et al., 2021). Steppe vegetation of the south-west of Vitimskoye Plateau belongs to the class of Cleistogenetea squarrosae Mirkin et al. ex Korotkov et al. 1991, order Helictotrichetalia schelliani Hilbig 2000, and alliance Helictotrichion schelliani Hilbig 2000. Habitats of hemicryomesoxerophytic steppe communities differ from those of suballiances described within the Helictotrichion schelliani in relation to position across landforms and the soil profile pattern. Differences are noticeable contrast to habitats of the Thymenion baicalensis Korolyuk 2017 communities, and are less pronounced compare to those of the Stipenion baicalensis Korolyuk 2017. The special peculiarity of the floristic composition of hemicryomesoxerophytic steppes is imparted by the presence of Kobresia filifolia, K. myosuroides and Thalictrum alpinum, indicating the cryophilic habitats. Moreover, there is a set of montane-steppe species also evidencing the same environment situation. Based on the floristic composition (Table) and ecological specificity we describe a new suballiance of steppe vegetation, containing 2 new associations. The suballiance Leontopodio conglobati–Caricenion pediformis Anenkhonov suball. nov. Nomenclatural type of the suballiance (holotypus) — ass. Kobresio filifoliae–Festucetum lenensis Anenkhonov et Naidanov. Diagnostic species (d. s.): Bistorta alopecuroides, Bupleurum sibiricum, Festuca lenensis, Gentiana decumbens, Leontopodium conglobatum, Lychnis sibirica, Rumex acetosa, Salix kochiana, Tephroseris integrifolia, Thalictrum alpinum. The suballiance represents hemicryomesoxerophytic steppe communities confined to the permafrost soils in the cold and sharply continental climate in the northern part of the forest-steppe subzone of the Transbaikalia region. The suballiance communities occupy flatland habitats and produce the forest-steppe landscape in combination with larch (Larix gmelinii) or birch (Betula platyphylla) coppices. Less often these communities are confined to the southern slopes when forest dominates in the landscape. The floristic composition of the hemicryomesoxerophytic steppes is significantly different from that of the cryophytic steppes described in other regions. Particularly, there are sufficient differences comparing to cryoxerophytic steppe of the Altai mountains, the subalpine montane steppe and kobrezian communities of Mongolia, and meadow-steppe communities in northern areas of Yakutia and West Chukotka. All of this support establishing the new syntaxon to integrate the hemicryomesoxerophytic steppe of Vitimskoye Plateau into the syntaxonomy of steppe vegetation. Defining the rank of the Transbaikalian hemicryomesoxerophytic steppes was based on the specificity of habitats, especially confinement to permafrost soils. Such criterion clearly separates these communities from steppes of the alliance Stipenion baicalensis, which are developed on seasonally frozen soils. The suballiance includes 2 associations: (1) Kobresio filifoliae–Festucetum lenensis — relatively more cryophytic; (2) Orostachyo malacophyllae–Festucetum lenensis — relatively more thermophytic. Ass. Kobresio filifoliae–Festucetum lenensis Anenkhonov et Naidanov ass. nov. (Table, rel. 1–15, Fig. 2). Nomenclatural type (holotypus hoc loco): Table, rel. 4 (field number nb18-23), Republic of Buryatia, Eravninskiy district, 43 km NE Sosnovo-Ozerskoye village, 52.82825° N, 111.93925° E, Varioherbosae-Sedge-Kobresia meadow steppe, 10.07.2018. Authors — B. B. Naidanov, O. A. Anenkhonov. D. s.: Artemisia rupestris, Equisetum arvense, Galatella dahurica, Heteropappus altaicus, Ixeridium gramineum, Kobresia filifolia, K. myosuroides, Potentilla anserina, P. arenosa. The association represents hemicryomesoxerophytic steppe communities on meadow-chernozem permafrost soils. The habitats are relatively more cryophytic (comparing to the 2nd association). Communities usually occupy open flatlands or slightly concaved surfaces of various configurations, and also cover shady (the most often northerly and easterly) sites close to coppices. Ass. Orostachyo malacophyllae–Festucetum lenensis Anenkhonov et Naidanov ass. nov. (Table, rel. 16–26; Fig. 3). Nomenclatural type (holotypus): Table 1, rel. 20 (field number nb18-22), Republic of Buryatia, Eravninskiy district, 36 km NE Sosnovo-Ozerskoye village, 3.3 km SSE Khorga settlement, 52.78897° N, 111.84138° E, Varioherbosae-Fesque-Pasqueflower meadow steppe, 10.07.2018. Authors — B. B. Naidanov, O. A. Anenkhonov. D. s.: Aconogonon angustifolium, Achillea asiatica, Campanula glomerata, Dianthus versicolor, Orostachys malacophylla, Schizonepeta multifida, Silene jeniseensis, Veronica incana. This association, like a previous one, represents hemicryomesoxerophytic steppe communities, however, it is relatively more thermo- and xerophytic. Communities occur on slightly elevated flat surfaces, hillocks, sites on well–warmed forest edges – most often from the southern and western sides of coppices, and on southern and south-western slopes of hills. To resume, hemicryomesoxerophytic steppe communities of Eravninskaya depression on the south-west of Vitimskoye Plateau represent a peculiar type of steppe vegetation, originated due to special habitats on the permafrost soils at the northern part of the main area of steppe vegetation in Transbaikalia region. These steppe communities belong to the new suballiance (with two new associations) of steppe vegetation.
In the framework of the project “Classification of Vegetation of the Russian Federation” (Plugatar et al., 2020; Kipriyanova et al., 2021), a revision of the modern coenotic diversity of Russia, ...including aquatic vegetation, is being carried out. It turned out that the types of some plant communities have not yet been described, including the association with the dominance of the Ranunculus mongolicus (Krylov) Serg.
Ranunculus mongolicus was first reported in the Flora of West Siberia (Flora..., 1931) by P. N. Krylov as a subspecies of the water buttercup R. aquatilis ssp. mongolicus Kryl. (Krylov, 1931), L. P. Sergievskaya raised the taxon rank to species (Sergievskaya, 1964). In the modern monographic treatment of aquatic buttercups, this taxon is given at the species rank (Wiegleb et al., 2017). It indicates that Ranunculus mongolicus is habitually similar to R. aquatilis, although smaller in all parts. Lamellar leaves are deeply dissected, peduncles are long, all parts of the plant are densely hairy. In arid regions, it behaves like an annual (Wiegleb et al., 2017).
In the course of expedition work in 2020 in the Ulagan district of the Republic of Altai, five geobotanical descriptions of communities dominated by the Mongolian buttercup (see Table) were performed, and it was decided to describe a new association. The classification was carried out on the principles of the ecological-floristic approach of J. Braun-Blanquet in accordance with the Code of Phytosociological Nomenclature (Theurillat et al., 2021).
Ass. Ranunculetum mongolici ass. nov. unites the communities dominated by the Mongolian buttercup (Fig. 1, 2). The diagnostic species is Ranunculus mongolicus (= Batrachium mongolicum (Krylov) Krecz.). Nomenclature type of association (holotypus hoc loco) — relevé No 5 in the Table. Republic of Altai, Ulagan district, vicinity of the mouth of the Chulyshman River, the oxbow lake of the Chulyshman River, 21.08.2020, 51.33130 N 87.73750 E. Author — L. M. Kipriyanova.
We attributed the new association to the alliance Batrachion aquatilis Passarge 1964 of the order Callitricho hamulatae–Ranunculetalia aquatilis Passarge ex Theurillat in Theurillat et al. 2015 of the class Potamogetonetea Klika in Klika et Novák 1941.
The range of the community apparently coincides with the range of the species. In Asia, R. mongolicus is found in Siberia, the Russian Far East, Kyrgyzstan, Afghanistan, Mongolia, China, and Nepal (Wiegleb et al., 2017). It is found along the west coast of North America where it is confused with R. aquatilis (as R. aquatilis var. aquatilis and R. aquatilis var. hispidulus); likewise, most South American “R. aquatilis” refers to R. mongolicus. (Wiegleb et al., 2017). In Eastern Siberia, R. mongolicus is rare and does not form communities (Chepinoga, 2015).
In addition to the Mongolian buttercup communities from Ulagan district, there are publicly available photographs of cenoses with R. mongolicus taken by A. I. Pyak in the Kosh-Agach district of the Republic of Altai (valley of the Kyzylchin (Buguzun) River, 49.992579° N, 89.088472° E, 15.07.2007, https://www.inaturalist.org/observations/99636689) (see Fig. 3). There are also photos of Yu. O. Kopylov-Guskov with thickets of Mongolian buttercup from the Republic of Tuva (Mongun-Taiginskiy district, 50.180458° N, 90.136025° E, 15.07.2016, https://www.inaturalist.org/observations/65270151) (see Fig. 4), such as A. L. Ebel photo from the Krasnoyarsk Territory (Kuraginskiy district, near the village of Kuragino, muddy bank of a channel of the Tuba River. 18.07.2018, 53.85932° N, 92.60829° E, https://www.plantarium.ru/page/image/id/719518.html) (see Fig. 5) In addition, R. mongolicus forms communities on the Taimyr Peninsula (Krasnoyarsk Territory, Khatanga River valley, Malaya Balakhnya oxbow lake, 22.08.2001, http://byrranga.ru/ranunculaceae/batrachium_aquatile/index.htm) (see Fig. 6).
Thus, at the moment it is possible to speak about the communities of the Mongolian buttercup from the Republic of Altai, the Republic of Tuva and the Krasnoyarsk Territory.
In the Republic of Altai, association communities are confined to flooded ecotopes, mainly with mineral (clay and sandy) silty substrates and depths from 20 to 60 cm in oligomesotrophic and mesotrophic cold-water reservoirs in the low-mountain and high-mountain belts of Altai, which makes it possible to attribute the association to the alliance Potamogetonion graminei Westhoff et Den Held 1969 combining the vegetation of rooted macrophytes of nutrient-poor (oligo-mesotrophic, sometimes dystrophic) shallow-water fresh water bodies of mountainous regions (Westhoff and Den Held, 1969; Mucina et al., 2016). However, the features of the ecology of communities throughout all the distribution area are not known yet, so for the time being we leave it in the alliance Batrachion aquatilis of the order Callitricho hamulatae–Ranunculetalia aquatilis.
Open sands are usual elements of subarid and arid landscapes in Eurasia. The unique psammophytic flora makes an important contribution to the regions’ biological diversity. From the syntaxonomical ...point of view, psammophytic vegetation of south-east European part of Russia still poorly explored. The Sarykum sand massif is a unique object, due to its location in piedmonts of the Caucasus at a considerable distance from vast sands of Terek-Kum and Caspian lowlands (Fig. 1, 2).
Our study presents the results of the analysis of 66 relevés. The data set was stored and processed in IBIS 7.2 software. As a result of performing hierarchical cluster analysis using Ward’s method, dendrogram was constructed reflecting the similarity of the relevés. Sokal/Sneath coefficient No. 1 which considers positive and negative species concurrences was used. Cluster analysis made it possible to reveal general differentiation patterns in psammophytic vegetation (Fig. 3). As a result, the relevés set was divided into two clusters differing in 28 species (Table 1). Cluster A represents poor of species and sparse communities on not fixed sands — in average 9 species per relevé and 18 % of the plants cover. Cluster B includes coenoses of fixed sands, with an average richness of 23 species and cover of 27 %.
The studied communities belong to the class Festucetea vaginatae Soó ex Vicherek 1972, which represents the sandy steppes. It is distributed in the forest-steppe and steppe zones of Eastern and South-Eastern Europe, as well as in the south of Western Siberia and Northern Kazakhstan. The class includes the only order Festucetalia vaginatae Soó 1957. Psammophytic vegetation described from various regions of southern Russia belongs to the alliance Festucion beckeri Vicherek 1972, but none of its diagnostic species was noted in relevés. Comparison of our data with the associations described in the class on the territory of Russia and Ukraine allowed us to identify a group of species differentiating the psammophytic vegetation of the Sarykum massif: Artemisia tschernieviana, Astragalus brachylobus, Asperula diminuta, Centaurea arenaria, Syrenia siliculosa, Thesium maritimum, Tragopogon dasyrhynchus ssp. daghestanicus. These species can become the basis for the diagnosis of a new alliance. Its definition will be possible as a result of comparative analysis of psammophytic vegetation of the southeast of European Russia, primarily of the Caspian and Terek-Kum sands.
Аss. Senecioni schischkiniani–Artemisietum tschernievianae ass. nov. (Table 2, relevés 1–18). Holotypus — relevé 1 in Table 2. (field no. 21-004), Republic of Dagestan, Kumtorkalinsky region, WSW from Korkmaskala village, Sarykum sands, leveled area in the upper part of the dune, 43.00751°N, 47.23290°E, 12.05.2021, author — A. Yu. Korolyuk.
Diagnostic species (D. s.): Artemisia tschernieviana, Senecio schischkinianus, Melilotus polonicus, Tragopogon dasyrhynchus ssp. daghestanicus. Communities are found in the central, most open part of the Sarykum massif. They occupy slopes, as well as plane areas characterized by active sands winding (Fig. 4). Artemisia tschernieviana usually dominates.
Аss. Leymo racemosi–Artemisietum tschernievianae ass. nov. (Tabl. 2, relevés 19–26), Holotypus — relevé 19 in Table 2. (field no. 21-026), Republic of Dagestan, Kumtorkalinsky region, WSW from Korkmaskala village, Sarykum sand massif, gentle northern slope of the dune, 43.01028°N, 47.23391°E, 13.05.2021, author — A. Yu. Korolyuk. D. s.: Artemisia tschernieviana (dom.), Leymus racemosus. The association represents pioneer cenoses that are widespread in the central elevated part of the massif. They are formed on slopes that are actively covered with sand (Fig. 5). The communities are sparse, with a cover of 10 to 25 % and the only dominant is Artemisia tschernieviana.
Аss. Jurineo ciscaucasicae–Stipetum borysthenicae ass. nov. (Table 3, relevés 1–31). Holotypus — relevé 1 in Table 3. (field no. 21-030), Republic of Dagestan, Kumtorkalinsky region, WSW from Korkmaskala village, Sarykum sand massif, gentle slope of small hill in the northern foot of the dune, 43.0112°N, 47.2333°E, 13.05.2021, author — A. Yu. Korolyuk. D. s.: Agropyron fragile, Artemisia tschernieviana, Cephalaria uralensis, Cerastium semidecandrum, Jurinea ciscaucasica, Medicago caerulea, Pleconax conica, Scorzonera biebersteinii, Stipa borysthenica, Syrenia siliculosa, Teucrium orientale, Thesium maritimum. The communities of the association represent the background type of vegetation of the more fixed sands of the Sarykum massif. They develop on plane areas, gentle slopes of hillocks and ridges, along the bottoms of old blowed hollows (Fig. 6, 7). More than other species Cephalaria uralensis dominates, less are Agropyron fragile, Artemisia tschernieviana, Euphorbia seguieriana, Scorzonera biebersteinii, Secale sylvestre, Stipa borysthenica, Teucrium orientale, Ziziphora serpyllacea. As part of the association, we distinguish three variants: var. typica (Table 3, op. 1–16) is widely distributed to the north and northeast of the Sarykum dune, var. artemisia marschalliana (Table 3, op. 17–24) represents the richest communities of sandy steppes, var. inops (Table 3, op. 25–31) was noted only in the southern part of the massif on the sections adjacent to the railway.
Five communities were described at the studied area. Com. Leymus racemosus occupy small patches, usually they form narrow strips between open sand and sagebrush coenoses (Fig. 8). Com. Eremosparton aphyllum was described in the middle part of the massif on the ridge slope (Fig. 9). Com. Calligonum aphyllum was observed on the top of a small sandy hill (Fig. 10). Com. Salix caspica was found on the tops of sand hills and represents large willow clones 2–3 meters high. Com. Imperata cylindrica was registered in the southwestern part of the massif along the slopes and bottom of depression (Fig. 11).
Differentiation of classes Festucetea vaginatae and Artemisietea lerchianae Golub 1994 is an interesting syntaxonomical problem. In the European Vegetation Checklist (Mucina et al., 2016), the class Artemisietea tschernievianae Golub 1994, representing psammophytic vegetation, is assigned to the second class as a synonym. We carried out a formalized analysis of Dagestan communities and syntaxa of three classes described in Astrakhan, Saratov, and Rostov Regions (Fig. 12). Cluster analysis divided the syntaxa into two groups corresponding to the classes Artemisietea lerchianae (A) and Festucetea vaginatae (B). Desert associations of the class Artemisietea lerchianae united together with ass. Artemisietum tschernievianae Golub 1994. This association is the holotype of the class Artemisietea tschernievianae, so the latter one should be recognized as synonym of the class Artemisietea lerchianae, representing deserts and desert steppes (Korolyuk, Laktionov, 2021). The ass. Koelerietum sabuletorum Golub 1994 joined together with other syntaxa of the class Festucetea vaginatae representing sandy steppes and vegetation of fixed sands.
The vegetation of the Sarykum sand massif is represented by original plant communities and numerous plant species that are typical for Middle Asia. In addition, about 20 plant species listed in the regional and federal Red Books grow on the territory of «Sarykum dune» cluster of Dagestan reserve and its protected zone. Despite the protection regime, the dune ecosystems are under human impact, which negatively affects the dune ecosystems.
