The syntaxonomic diversity of forest meadows at the Ob-Inya watershed was studied within the Novosibirsk Region. The territory is a part of the Sokur elevation, which approaches the Salair ridge in ...the south-east. Common is the erosion of the landscape reflected in dense network of river valleys and draws. The vegetation is transitional between subtaiga and forest steppe (Reverdatto, 1931; Vandakurova, 1957; Kuminova, 1973; etc.). Forest meadows (the order Carici macrourae–Crepidetalia sibiricae Ermakov et al. 1999 of the class Molinio-Arrhenatheretea R. Tx. 1937) occur on the edges and glades of grassy aspen and birch forests and are an element of vegetation of both forest steppe zone and the subtaiga subzone. Their set of species and syntaxonomic diversity reflect the soil and climatic conditions of the location and may serve as a criteria for defining the zonal and subzonal borderlines.
The aim of the study was to reveal and to characterize the syntaxonomic diversity of the forest meadows of the Ob-Inya watershed.
The studies were conducted at the territories of the Novosibirsk, Moshkovo, and Bolotnoye districts. 114 reléves served as a basis for the study.
Forest meadows are represented by 2 associations, 2 subassociations, 2 variants and 2 communities from 2 suballiances of the alliance Crepidion sibiricae Mirkin ex Ermakov et al. 1999 (Table 1).
Communities of the ass. Crepidetum sibiricae Dymina ex Ermakov et al. 1999 are most common, and occur everywhere in the studied area (Table 2). The association is represented by two subassociations differing by the degree of the moisture content in the habitats. Communities of the subass. typicum occur in depressed highly moistered landscapes primarily in the axial part of the Sokur elevation and in the north-eastern regions of the Ob-Inya watershed. A new subass. Crepidetum sibiricae fragarietosum viridis subass. nov. uniting more xerophytic communities which are uniformly dispersed over the territory in well-drained flat and convex parts of the watershed and on the gentle slopes of the draws. Character for these is participation of xeromesophytes from the diagnostic combination of dry forest meadows of the suballiance Aconito barbati–Vicenion unijugaeKorolyuk et al. 2016. The nomenclature type of the subassociation (holotypus): Table 2, rel. 17 (field number Z20-021), Novosibirsk region, Bolotnoye district, near the Kozlovka village, 55.86934° N, 84.15000° E, 31.08.2020, the author E. G. Zibzeev.
Under the increased grazing pressure on the communities of the subass. C. s. fragarietosum viridis, pasture meadows of the var. Poa angustifolia, mostly dominating by Poa angustifolia,were formed near the settlements of the south-western part of the Sokur elevation. There are some meadow-forest plants of the alliance Crepidion sibiricae of the order Carici macrourae–Crepidetalia sibiricae in such communities.
A new forest meadow ass. Galatello biflorae–Artemisietum macranthae ass. nov. (Table 3) unites dry forest meadows occurring on the upper parts of the convex slopes of the draws of sunlit exposures in the axial part of the Sokur elevation The communities developed under conditions ecologically closed to these of the subass. Crepidetum sibiricaefragarietosum viridis meadows have more xerophytic composition: the tall-forb species are absent; the abundance and continuity of the meadow mesophytes is lower, while the proportion of meadow-steppe elements is higher. The nomenclature type (holotypus): Table 3, rel. 1 (field number mr20-033), Novosibirsk region, Moshkovo district, the area near the Verkh-Balta village, 55.32588° N, 83.76177° E, 30.08.2020, the author M. P. Tishchenko.
This association may be considered as subtaiga variant of the dry forest meadows at the Ob-Inya watershed (the right-bank region of the Ob River). For the species composition and environmental occurrence, communities are similar to the these of the ass. Aegopodio podagrariae–Brachypodietum pinnati Tishchenko et al. 2015 from the subtaiga of the left-bank region of the Ob River. Xeromesophytes as well the hygrophilous species growing in the subtaiga forests are common in their coenofloras. The communities of the ass. Galatello biflorae–Brachypodietum pinnati Korolyuk 1998 on the left-bank region of the Ob River and of the ass. Filipendulo vulgaris–Brachypodietum pinnati Makunina et al. 2010 (Korolyuk, Kipriyanova, 1998; Makunina, 2016) in the right-bank region, occur in similar forest-steppe habitats.
In the elevated sites of the central part of the Ob-Inya watershed and in the right-bank region of the Inya River, dry forest meadows of community Iris ruthenica are described (Table 4, rel. 1–6). Character for these is participation of Iris ruthenica and Filipendula vulgaris, rare in the studied area.
The community Carex praecox (Table 4, rel. 7–10) includes the forest meadows of the driest habitats, where some steppe species are found. The communities occur in the eastern part of the Ob-Inya watershed on the steep southern and south-western slopes into the valley of the Iksa River, the tributary of the Ob River, in their upper part.
To summarize, the forest meadows of the region have more mesophytic composition compared to the forest-steppe communities: no xerophytes and many meadow-steppe species (Peucedanum morisonii, Ligularia glauca, Adonis vernalis, Campanula altaica, Dianthus superbus et al.) common in forest meadows in the forest-steppe zone. The Ob-Inya watershed forest meadow syntaxonomy is rather peculiar, compare to the forest-steppe zone.
Mesophilous broad-leaved forests predominating in south-western part of Russia were included in alliance Aceri campestris–Quercion roboris Bulokhov et Solomeshch in Bulokhov et Semenishchenkov 2015, ...association Fraxino excelsioris–Quercetum roboris Bulokhov et Solomeshch 2003 and three geographic subassociations. The subassociation F. e.–Q. r. typicum Bulokhov et Solomeshch 2003 in Bulokhov et Semenishchenkov 2015 unites communities occurring mainly in the broad-leaved forests zone of the Central Russian Upland. The subassociation F. e.–Q. r. stellarietosum nemori Semenishchenkov et al. 2015 occurs in the northern part of the association range in the zone of broad-leaved forests and close to the southeastern border of the hemiboreal subzone. Mesophilous broad-leaved forests occurring in the forest-steppe zone of the Central Russian Upland were included in the subass. F. e.–Q. r. crataegetosum curvisepalae Semenishchenkov in Bulokhov et Semenishchenkov 2015. The geographical distribution of the syntaxa is related to the climate continentality gradient — from the subass F. e.–Q. r. crataegetosum curvisepalae (the largest continentality indices) to the subass. F. e.–Q. r. stellarietosum nemori (the smallest continentality indices).
The associations Aceri campestris–Tilietum cordatae Zaugolnova et Braslavskaya 2003 (nom. inv.) from the Kaluga and Tula Regions (Zaugolnova, Braslavskaya, 2003) and Aceri campestris–Quercetum roboris (nom. inv.) (var. Acer tataricum) described from the Voronezh Region (Starodubtseva, Khanina, 2009) were included in the association Fraxino excelsioris–Quercetum roboris after comparative syntxonomical analysis.
On the basis of a comparative analysis, diagnostic combinations of subassociations were compiled, consisting of species with constancy above 20 % and values of the statistical φ-coefficient for selections above 20: F. e.–Q. r. stellarietosum nemori — Carex sylvatica (5059.9), Equisetum hyemale (2842.3), Galeobdolon luteum (10087.4), Matteucia struthiopteris (4257.7), Ranunculus cassubicus (5857.9), Stellaria nemorum (3349.6); F. e.–Q. r. crataegetosum curvisepalae — Acer tataricum (5362.0), Crataegus rhipidophylla (5964.0), Poa nemoralis (2736.9), Pyrus pyraster (3840.8), Scilla siberica (4558.4), Viola odorata (3440.5).
Analysis of the differentiaal table of mesophilous broad-leaved and spruce-broad-leaved forests syntaxa made it possible to conclude that associations previously (Onyshchenko, 2009) assigned in Ukraine to alliance Scillo sibericae–Quercion roboris have a high floristic similarity with the syntaxa of Aceri campestris–Quercion roboris from the South-West of Russia, especially in the forest-steppe part of its range. Selections of three subassociations of the association Mercurialo perennis–Quercetum roboris from Ukraine floristically close to the syntaxa of alliance Querco roboris–Tilion cordatae from the South-West of Russia. In the dendrogram of cluster analysis, the compared syntaxa are grouped into two blocks corresponding to the rank of alliances: Aceri campestris–Quercion roboris and Querco roboris–Tilion cordatae. Based on the review, the authors believe that the «Hierarchical floristic classification system…» (Mucina et al., 2016) may be amended in terms of recognizing the syntaxonomical independence of the alliance Aceri campestris–Quercion roboris and its geographical interpretation.
Derivative communities disturbed by humans in the place of mesophylous broad-leaved forests are classified as anthropogenic variants and non-rank «communities» named after the dominant species: Betula pendula, Populus tremula, Tilia cordata. Forests with a poor floristic composition, in which the diagnostic species of association and subassociation drop out of the coenoflora are classified as inops variants. They are characterized by a noticeable decrease in the average floristic saturation in comparison with typical communities. Derivative forests with a predominance of Betula pendula have a higher species richness in comparison with aspen or linden forests, as well as typical phytocoenoses of the association.
The psammophytic communities on sandy accumulative coasts of the Black and Azov Seas were studied in the Krasnodar Territory (Russia) in 2004, 2006 and 2009, when 1610 relevés were made. Of these, ...203 relevés were previously classified and published. In this paper based on 23 relevés two new associations and two new subassociations of the class Ammophiletea Br.-Bl. et Tx. ex Westhoff et al. 1946 and one rankless transitional community (Fig. 1) are described according to Braun-Blanquet approach. The abundance of plants estimated in the field as a percentage of the projective cover was converted to scale points: 5 — > 50 %, 4 — 26–50 %, 3 — 16–25 %, 2 — 6–15 %, 1 — 1–5 %, + — < 1 %.
Clustering of the relevés (phytocenon isolation) was carried out by flexible beta linkage (β = –0.25) based on the Sørensen coefficient in PC-ORD 5.0, available through the JUICE 7.1 software package (Tichý, Jason, 2006). As a result four phytocenons were established. Their species composition was compared with the species lists of the lower 676 coastal syntaxa of the Azov-Black Sea region, taken from literature and stored in authors’ syntaxon database (GIVD ID EU-RU-005) based on the TURBOVEG program (Hennekens, Schaminée, 2001). An initial assessment of the similarity of littoral syntaxa and phytocenons was performed using several methods available in the JUICE 7.1 software package. In all cases bryophytes and lichens as well as vascular plant species with frequency less than 20 % in any community were excluded from the analysis. All these excluded species are present in Tables 1 and 2. To classify the established phytocenons, their species composition was compared with similar protologues of the lower syntaxa (Table 1).
The vascular plant taxa names are by Tutin et al., (2001). In naming the taxa below, a broad understanding of species (s. l.), their aggregation (agg.), or the combination of several species (with “+”) are used: Artemisia campestris+A. tschernieviana, Cakile maritima s. l. (C. maritima, C. maritima subsp. euxina), Centaurea arenaria s. l.(C. arenaria, C. arenaria subsp. odessana), Leymus racemosus s. l. (L. racemosus, L. racemosus subsp. sabulosus); S. kali aggr. (Salsola kali, S. kali subsp. ruthenica or S. kali subsp. tragus) based on P. Uotila (2011) and S. L. Mosyakin (2017); Xanthium strumarium s. l. (X. strumarium, X. strumarium subsp. italicum, X. strumarium subsp. strumarium × subsp. italicum). Few bryophyte and lichen taxa are given with their authors. The names of the new syntaxa are formed according to the ICPN rules (Theurillat et al., 2021).
The terrain in the study area is flat. The natural banks are represented by abrasion and accumulative types. The latter is often in the form of sandy or sandy-shelly spits. There are low-lying near-mouth and delta accumulative banks at the mouths of large rivers. Widespread are solonchaks both not vegetated or with halophytic communities. The climate is temperate with continental features. The month mean temperatures and sums of precipitation amounts for the ten-year period (2001–2010) are reflected in the climate diagrams (Fig. 2).
The closest in species composition to the established phytocenons were the lower syntaxa (among the protologues) described in Russia, Romania and Ukraine (Table 1), attributed by their authors to the alliance Elymion gigantei Morariu 1957 (class Ammophiletea Br.-Bl. et Tx. ex Westhoff et al. 1946). However, the identified species differences did not allow us to assign the phytocenons (Table 1) to these syntaxonomical units. All phytocenons are classified as new syntaxa (Table 2). The geographical location of the lower syntaxa presented in Table 1 see on Fig. 3.
Ass. Eryngio maritimi–Leymetum sabulosi ass. nov: Table 2, rel. 1–13, Fig. 4; holotypus: Table 2, rel. 6, Krasnodar Territory, Anapa sandbar opposite the settlement Blagoveshchenskaya (coast of the Black Sea), fixed dunes in the distance from the sea, 25.08.2004. Diagnostic taxa (d. t.): Artemisia campestris+A. tschernieviana, Eryngium maritimum, Leymus racemosus s. l. The association includes psammophytic herb–dwarf-semishrub communities on fixed and mobile sandy substrates. Subass. Eryngio maritimi–Leymetum sabulositypicum subass. nov.: Table 2, rel. 1–7; holotypus: Table 2, rel. 6, Fig. 5. The diagnostic attributes of the typical subassociation communities are the same as of the association. Plant communities of the subassociation inhabit both mobile and weakly fixed sandy sediments of the accumulative coasts of the Black and Azov Seas. Subass. Eryngio maritimi–Leymetum sabulosi crambetosum maritimae subass. nov.: Table 2, rel. 8–13, Fig. 6; holotypus: Table 2, rel. 10, Krasnodar Territory, 4 km ENE from the settlement Golubitskaya, coast of the Temryuk Bay (Azov Sea), low dune (1.5–2 m high), 04.09.2004, D. t. of the subassociation: Crambe maritima, Lactuca tatarica. Communities occur on fixed sands on accumulative banks.
Community Conium maculatum–Centaurea arenaria: Table 2, rel. 14–18. D. t.: Astragalus onobrychis, Centaurea arenaria s. l., Conium maculatum, Galium humifusum, Torilis arvensis. The communities of this rankless transitional unit occur on flat or depressed terrain areas on the Temryuk Bay coast.
Ass. Cakilo euxinae–Glycyrrhizetum glabrae ass. nov.: Table 2, rel. 19–23, Fig. 7; holotypus: Table 2, rel. 19, Krasnodar Territory, Bugazsk Spit, opposite Veselovka village, the coast between the Black Sea and Bugaz Liman, slope of sandy sediment (north-east exposure, 1–2° slope) toward the estuary, 19.08.2004. D. t.: Cakile maritima s. l., Glycyrrhiza glabra, Carex ligerica, Salsola kali aggr. Communities are distributed on the southern part of the Taman Peninsula. They are found on tops of the avandunes, as well as away from them on the elevated parts of the marine spits.
Relevés of communities (94 in total) with shrub willows — Salix glauca, S. lanata, S. phylicifolia, and tree-like willows — S. viminalis, S. dasyclados were made in the Bolshezemelskaya tundra (11 ...sites, Fig. 1). Six new associations and one community type are described in 2 alliances — Polemonio acutiflorum–Salicion glaucae Lavrinenko et Lavrinenko 2021 and Galio borealis–Salicion viminalis all. nov. (Tables 1–4).
The alliance Polemonio acutiflorum–Salicion glaucae unites herb and herb-moss willow scrubs (predominantly with Salix glauca, S. lanata) in lakeside depressions, runoff troughs, the slopes of hills on watersheds, slopes of bedrock terraces, above-floodplain terraces and occasionally flooded floodplains in river valleys. The range of this alliance is the plain tundra in the East European and Siberian sectors of the Arctic. Associations Polemonio acutiflorum–Salicetum lanatae Zanokha ex Lavrinenko et Lavrinenko 2021, Triseto sibirici–Salicetum glaucae Lavrinenko et Lavrinenko 2021 and Climacio dendroidis–Salicetum lanatae Lavrinenko et Lavrinenko 2021 were earlier described on Taymyr Peninsula, Kolguev and Vaygach Islands (Lavrinenko, Lavrinenko, 2021). New associations Chamaepericlymeno suecici–Salicetum glaucae, Geo rivalis–Salicetum glaucae, Hylocomio splendentis–Salicetum glaucae are described in the Bolshezemelskaya tundra (Table 1). Some communities of the associations Dicrano majoris–Salicetum lanatae Khitun in Telyatnikov et al. 2021 and Calliergono cordifolii–Salicetum lanatae Khitun in Telyatnikov et al. 2021, described on the Tazovskiy and Gydanskiy Peninsulas (Telyatnikov et al., 2021a and b) can be attributed to this alliance. However, the decisions for both these associations need to be revision (see Table 5).
The characteristic species of the alliance Polemonio acutiflorum–Salicion glaucae were confirmed and clarifed. The phi-coefficient values for them are in the range 84.7–42.3 (Table 6).
A new order of the same name is proposed for the willow scrubs of this alliance.
Order Polemonio acutiflorum–Salicetalia glaucae ord. nov.
Herb and herb-moss willow scrubs with Salix glauca and S. lanata in the plain tundra in the East European and Siberian sectors of the Arctic.
Nomenclature type (holotypus): alliance Polemonio acutiflorum–Salicion glaucae Lavrinenko et Lavrinenko 2021 (Lavrinenko, Lavrinenko, 2021: 99).
Differentiating species combination: Salix glauca, S. lanata; Bistorta vivipara, Petasites frigidus, Poa arctica, Polemonium acutiflorum, Ranunculus propinquus, Rubus chamaemorus, Valeriana capitata; Hylocomium splendens.
Constant taxa: Achillea millefolium, Cardamine pratensis subsp. angustifolia, Chrysosplenium alternifolium subsp. sibiricum, Equisetum arvense s. l., Poa pratensis s. l., Rubus arcticus, Veratrum lobelianum, Viola biflora; Bryum pseudotriquetrum, Rhizomnium pseudopunctatum, Sanionia uncinata.
Habitats. Intrazonal biotopes on watersheds (lakeside depressions, runoff troughs, hillsides), slopes of high river terraces, above-floodplain terraces and rarely flooded parts of floodplains in river and stream valleys.
Willow scrubs from tree-like (Salix dasyclados, S. viminalis) and shrub (Salix lanata, S. phylicifolia) species, occupying the near-channel part of the floodplains of large and small rivers, channels and oxbows and experiencing regular flooding, are described in the southern tundra, northern forest-tundra and in the Pechora River delta. For such willow scrubs we have described a new alliance Galio borealis–Salicion viminalis in order Salicetalia purpureae Moor 1958 and class Salicetea purpureae Moor 1958.
Alliance Galio borealis–Salicion viminalis all. nov.
Tall grass-herb willow scrubs (Salix dasyclados, S. lanata, S. phylicifolia, S. viminalis) in riparian habitats in the southern tundra, forest-tundra and northern taiga in the European North of Russia.
Nomenclature type (holotypus): ass. Senecioni nemorensis–Saliсetum viminalis ass. nov. (Table 2, rel. 19–25; Table 4, syntaxon 19).
Characteristic species: Salix viminalis and Veronica longifolia (the same species of class Salicetea purpureae and order Salicetalia purpureae), Aconitum septentrionale, Adoxa moschatellina, Calamagrostis purpurea s. l., Filipendula ulmaria, Galium boreale, Lamium album, Myosotis palustris, Ranunculus repens and Vicia cracca (phi-coefficient varies from 77.7 to 47.6) (Table 6). Apparently, Salix dasyclados, with which one syntaxon has been described, can also be attributed to these.
Constant species: Equisetum arvense s. l. (often dominant or codominant), Poa pratensis s. l., Veratrum lobelianum, Viola biflora.
Habitats. Near the riverbed and central parts of the floodplain of large and small rivers, alluvial soils under conditions of regular flooding.
Those willow scrubs are classified as 3 new associations and one community type: Vicio sepium–Salicetum lanatae, Senecioni nemorensis–Saliсetum viminalis, Bromopsio inermis–Salicetum viminalis, Calamagrostis purpurea–Salix dasyclados (Table 2).
Floristically close communities are described in the floodplain of the Pechora River in the northern taiga (Shushpannikova, 1996), so we expanded the Galio borealis–Salicion viminalis range to this subzone. Further south the alliance Salicion triandrae T. Müller et Görs 1958 from the same order and class is widespread. Communities of Salicion triandrae occupy riparian habitats in the middle and southern taiga subzones (Taran, 1999; Vasilevich, 2009) and in the nemoral zone of Europe (Neuhäuslová et al., 2013; Mucina et al., 2016).
The results of DCA-ordination confirm the correctness of the alliances Polemonio acutiflorum–Salicion glaucae and Galio borealis–Salicion viminalis establishment (Fig. 14).
Ass. Carici stantis–Salicetum phylicifoliae ass. nov. (Table 3) unites willow scrubs (Salix glauca, S. lanata и S. phylicifolia) swamp-herb (Comarum palustre, Epilobium palustre, Equisetum fluviatile, Eriophorum angustifolium, Menyanthes trifoliata)–sedge (Carex aquatilis subsp. stans). If focusing on the composition of the lower layers, the place of association will undoubtedly be in the class Scheuchzerio palustris–Caricetea nigrae Tx. 1937, since most of the listed species are characteristic for it or for the order Caricetalia nigrae Koch 1926. However, the presence of rather tall and highly closed shrub layer, which determines the structure and physiognomy of communities, calls into question their attribution to mire. For the time being, we leave such communities outside the higher units.
Differences in the floristic composition of the syntaxa in the East European tundras, in the mountainous Fennoscandia (Salicion phylicifoliae Dierssen 1992) and Chukotka (Aulacomnio turgidi–Salicion glaucae Sinelnikova 2001 (2009) and Saussureo oxyodontae–Salicion Sekretareva 2001 all. prov.) are significant (Table 4), which is clearly demonstrated be the results of cluster analysis (Fig. 15). Syntaxa of another northern alliance — Geranio albiflorum–Salicion Sekretareva 2011, described in Polar Ural, are joined in one cluster with the willow scrubs of the alliance Galio borealis–Salicion viminalis (Fig. 15).
We have not assigned to any class the new order Polemonio acutiflorum–Salicetalia glaucae ord. nov., as opposed to Salicetalia glauco-lanatae Bœuf et al. ex Mucina et Daniёls in Mucina et al. 2016 (subarctic and boreal herbrich willow scrubs and birch krummholz of the Holarctic), placed in the class Betulo carpaticae–Alnetea viridis Rejmánek ex Bœuf, Theurillat, Willner, Mucina et Simler in Bœuf et al. 2014 (subalpine and subarctic herb-rich alder and willow scrubs and krummholz of the Alps, the Carpathians, the Balkans, the Caucasus, Northern Europe and Greenland). We believe that for the willow scrubs of the tundra zone a revision of all the accumulated data and description of a new class are required.
Being one of the most characteristic types of plant communities found within settlements, the synanthropic communities of the Republic of Bashkortostan (RB) have been studied since the end of the XX ...century (Ishbirdin et al., 1988; Ishbirdinа, Anischenko, 1989; Sachapov et al., 1990; Anischenko, 1995; Ryabova, 1996, etc.). At the beginning of the XXI century geobotanical research was actively continued, while significant attention was paid to phytocenoses located within the towns and villages of the republic (Govorov, 2004; Edrenkina, 2005; Yamalov et al., 2008; Yamalov, Suyundukova, 2008; Usmanova et al., 2013, 2014). Urban vegetation is most fully presented in the series of publications on the classification of vegetation of towns of the Southern industrial zone (Golovanov, Abramova, 2012, 2013; Golovanov et al., 2017). The given article presents results of the long-term (2005-2018) research of vegetation of the towns of Bashkir Pre-Urals, as well as rural settlements of Trans-Urals. 63 complete relevés of new communities for RB included in the geobotanical base of synantropic vegetation of the Southern Urals were used. The geobotanical database was implemented using the TURBOVEG software (Hennekens, Schaminée, 2001). The area of relevé depended on the community size and homogeneity. Classification is carried out according to Braun-Blanquet method using the JUICE software (Tichý, 2002). 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 are given by “Vegetation of Europe...” (Mucina et al., 2016). The ecological conditions were assessed with use of average values of E. Landolt (1977) ecological scales and processed by IBIS software (Zverev, 2007). Average values are calculated on the following scales: humidification (M), acidity (R), soil richness with mineral nutrition elements (N), humus (H) content, mechanical composition and soil structure (D), illumination (L). At the beginning of the 2010s prodrome of synanthropic communities of RB, included 7 classes, 12 orders, 17 alliances and 61 associations (Yamalov et al., 2012). Currently, the synthaxonomy of the synanthropic vegetation of the study area requires adjustment in accordance with the modern state of European synthaxonomy. Special geobotanical studies of recent years have allowed expanding the synthaxonomic scheme of synanthropic vegetation of the region. Earlier, data have already been published on new associations for the region of the classes Artemisietea vulgaris and Galio-Urticetea (Golovanov, 2017). This article extends our research. This article considers the synanthropic vegetation of two synanthropic classes — Sisymbrietea and Digitario sanguinalis–Eragrostietea minoris. These classes are first given for the territory of the RB. The article contains their regional diagnostic species. Previously, they were a part of the wider class — Stellarietea mediae, combining annual weed and ruderal vegetation. For the class Sisymbrietea, three synthaxa are given: the associations Descurainietum sophiae and Poo annuae–Stellarietum mediae and the subass. Malvetum pusillae urticetosum urentis. The last two synthaxa are new. They unite nitrophilic communities of low-growth annuals along the house walls, fences, shoulders and other disturbed habitats both in rural settlements and in towns. Association Descurainietum sophiae unites communities found in various disturbed habitats: landfills, deposits, as well as near buildings. They occur mainly within the small settlements of the Bashkir Trans-Urals, less often in the Bashkir Cis-Urals. A comparison of the coenoflora of communities described in territory of RB and in other regions of Europe (Table 2) showed that the composition of diagnostic species groups is quite similar. The greatest similarity is with the communities located in the Czech Republic. Association Malvetum pusillae, subass M. p. urticetosum urentis (Table 3). holotypus: Table 3, relevé 2, the Republic of Bashkortostan, Khaibullinsky district, Stepnoe, along the wall of the building, 51.81988° N, 58.15081° E, 14.06.2005, author — G. Ya. Suyundukova. We compared communities dominated by Urtica urens described in territory of RB with the closest associations of the Malvion neglectae alliance of Eastern Europe (Table 4). Significant differences are visible both in the almost complete absence of main diagnostic species, and in the features of coenoflora. The cenoses described in the Czech Republic are characterized by the high constancy of Malva neglecta, completely absent in the Southern Urals, as well as the significant presence of species resistant to trampling. This indicates the different nature of environmental conditions in the compared syntaxa. Comparison with other syntaxa of the Malvion neglectae alliance in RB (Table 6) showed that the floristic composition and ecology of the Urtica urens dominated communities and the ass. Malvetum pusillae cenoses are very similar. Thus, we consider the specific subass. M. p. urticetosum urentis as part of the above association. Its physiognomy is characterized by the dominance of Urtica urens. Basically, communities are found within the rural settlements of the Bashkir Trans-Urals in the steppe and forest-steppe zones. Association Poo annuae–Stellarietum mediae (табл. 3; Fig. 1). holotypus: Table 5, relevé 1, Republic of Bashkortostan, town district Salavat, boulevard Salavat Yulaev, d. 17, house plot, 10.08.2008, 53.34177° N, 55.93038° E, author — Ya. M. Golovanov. Communities dominated by Stellaria media in Eurasia were described as a rule within the framework of weed vegetation. The closest in ecology are the cenoses of the Euphorbia peplus–Stellaria media community described within the alliance Malvion neglectae in Poland (Ratynska et al., 2010). In the ruderal communities described by us, species of the alliance Malvion neglectae and other classes of synanthropic vegetation occur with high constancy. Association communities are found near the building walls of shadow expositions, along the edges of abandoned palisades in the towns of the Bashkir Cis-Urals. Compare to other syntaxa of the Malvion neglectae alliance in the territory of RB (Table 6), the ass. Poo annuae–Stellarietum mediae occupies the most humidified habitats on slightly less rich substrates. At the same time, such communities have the richest coenoflora due to the presence of species of the classes Molinio-Arrhenatheretea and Epilobietea angustifolii. Communities of subassociation Malvetum pusillae urticetosum urentis are similar in coenofloras and occupy similar habitats. The ass. Eragrostio–Amaranthetum albi within the class Digitario sanguinalis–Eragrostietea minoris is firstly identified. It combines specific synanthropic communities of railway embankments with predominance of annual species. Associations with the dominance of Eragrostis minor of the Digitario sanguinalis–Eragrostietea minoris class in different regions were compared (Table 8, Fig. 2). Significant similarity of the cenoses described in the republic with the communities of the ass. Eragrostio–Amaranthetum albi located in Kursk was revealed. Differences are probably due to the nature of the use of railway embankments within stations, as well as the accidental drift of certain species.
The dominance of xeropetrophytic plant communities with original species composition along with high phytocenotic diversity is the first to determine the distinctness of the South Ural low-mountain ...steppe province within the Ural Mountains. As the zonal steppe ecosystems are totally destroyed and transformed, petrophytic cenoses represent the predominant type of natural vegetation. They are of high conservation importance due to preserving unique biodiversity with a large number of rare and endangered species, relics and endemics (Ryabinina, 2003; Yamalov et al., 2019).
Stony steppes in the Southern Urals have been studied for more than 100 years. Initiated in the 1980s, the syntaxonomical research covers now only the limited territories, mainly within the protected areas (Unikalnye ..., 2014; Prirodnye ..., 2018). The recent studies are focused on the relationship between the petrophytic vegetation and macroecological gradients in the context of geographical zonation (Zolotareva et al., 2019; Korolyuk et al., 2020).
Our study presents the results of the analysis of 265 relevés of petrophytic communities of the steppe zone, carried out by the authors on the territory of the Gaysky, Kuvandyksky, Saraktashsky and Belyaevsky administrative districts of the Orenburg region, as well as the Khaibullinsky district of the Bashkortostan Republic in 2014–2018 (Fig. 1).
Six phytocoenons were identified using the TWINSPAN algorithm followed by manual sorting. They represent the main diversity of petrophytic vegetation in the studied region and differ in the groups of species identified by formal criteria (Korolyuk et al., 2016; Zolotareva et al., 2019). We conducted the cluster analysis to compare these with the petrophytic associations of the Urals: 26 associations from the classes Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 and Anabasietea cretaceae Golovanov et al. 2021 (Fig. 2, 3). Five communities were assigned to the steppe vegetation, and their position in the syntaxonomical system was determined. Communities 1–3 regarded as typical steppes have similar floristic composition and dominant plants; community 4 is closer to the rich forb-bunchgrass steppes; community 5 and the ass. Anthemido trotzkianae–Thymetum guberlinensis have transitional features to desert steppes.
We identified the new alliance Elytrigion pruiniferae all. nov. D. s.: Allium tulipifolium, Alyssum turkestanicum, Artemisia salsoloides, Centaurea turgaica, Elytrigia pruinifera, Ferula tatarica, Linaria uralensis, Poa bulbosa s. l., Polycnemum arvense, Spiraea hypericifolia. Holotypus — the association Elytrigietum pruiniferae Lebedeva ass. nov. Alliance represents xeropetrophytic vegetation of the steppe zone of the South Urals. Its area covers the watersheds of the Sakmara, Ural and Ilek rivers. Its communities were also found on the right bank of the Sakmara river (the Guberlya and Kuragan basins), as well as in the lower part of the Bolshoy Ik river basin. The communities occupy convex slopes and tops of ridges with immature gravelly soils. In the southern part of the alliance distribution area, they are confined to slopes of various shapes and exposures, while in the northern part mainly to convex southern slopes or to the top of the ridges. The alliance represents xerophytic part of the order Helictotricho-Stipetalia. It replaces the more mesophytic alliance Helictotricho desertori–Orostachyion spinosae on the moisture gradient (Korolyuk, 2017). The alliance includes 3 associations and 1 community type.
Ass. Elytrigietum pruiniferae Lebedeva ass. nov. (Table 4), holotypus — rel. 1 (field no. 15-086): Orenburg region, Kuvandyk district, 1.5 km southwest of Yalnair village, near-summit convex part of a rocky slope, 51.28635°N, 57.81782°E, 10.06.2015, author — A. Yu. Korolyuk. D. s.: Alyssum turkestanicum, Centaurea turgaica, Elytrigia pruinifera, Linaria uralensis, Poa bulbosa s. l., Polycnemum arvense, Spiraea hypericifolia, Thymus guberlinensis, Tulipa scythica. The association unites petrophytic steppes in hilly lands and low-mountains of the Saraktash, Kuvandyk and Gaysky districts of Orenburg region and on the Turatka mountain in Khaibullinsky district of Bashkiria. The cenoses are confined to convex slopes and tops of ridges. They are developed on gravelly soils on outcrops of basic and metamorphic rocks, less often on sedimentary rocks. There are 4 variants within the association.
Ass. Thymo guberlinensis–Galatelletum villosae Yamalov ass. nov. (Table 5), holotypus — rel. 2 (field no. 16-015): Orenburg region, Kuvandyk district, 19 km SW of Zaluzhye village, 51.18268°N, 56.91858°E, author — A. Yu. Korolyuk. D. s.: Astragalus tenuifolius, Ephedra distachya, Galatella villosa, Hedysarum argyrophyllum, Meniocus linifolius, Scorzonera stricta, Stipa lessingiana. The association represents xerophytic stony steppes. They occur in all regions confined mainly to the southern slopes and tops of ridges with fine gravelly soils on sedimentary rocks. There are 4 variants within the association.
Ass. Stipo zalesskii–Centauretum turgaicae ass. nov. (Table 6, rel. 7–22), holotypus — rel. 10 (field no. 17-122): Orenburg region, Gaysky district, east of the Kazachya Guberlya village, slightly convex slope of the ridge in the near-top part, 51.14663°N, 58.03227°E, 06.18.2017, author — A. Yu. Korolyuk. D. s.: Amygdalus nana, Campanula sibirica, Carex pediformis, Cephalaria uralensis, Erysimum canescens, Helictotrichon desertorum, Pulsatilla patens, Stipa pulcherrima, S. zalesskii. The association was described in the southeastern part of the Guberlinsky Uplands (Gaysky District). The communities occupy habitats typical for petrophytic steppes — convex slopes and tops of ridges on outcrops of basic and metamorphic rocks.
The associations Elytrigio pruiniferae–Stipetum sareptanae Golovanov ass. nov. and Anthemido trotzkianae–Thymetum guberlinensis Golovanov et al., 2021 were described in the study area as well.
Ass. Elytrigio pruiniferae–Stipetum sareptanae ass. nov. (Table 7), holotypus — rel. 10 (field no. GY18-043): Orenburg region, Kuvandyk district, 4 km SE Novyi village, Suyunduksai Balka, 51.02067°N, 57.33993°E, author — Ya. M. Golovanov. D. s.: Artemisia lerchiana, Astragalus tenuifolius, Atraphaxis frutescens, Ephedra distachya, Gypsophila rupestris, Hedysarum argyrophyllum, H. razoumovianum, Meniocus linifolius, Sterigmostemum tomentosum. The association is found mainly to the south of the latitudinal section of the Ural River within the Guberlinsky Uplands (Kuvandyk District), as well as of the hilly lands of the South Urals (Saraktashsky District). The communities are confined to slopes of different exposure and steepness on outcrops of sedimentary rocks. The plants of dry and desert steppes, as well as those preferring chalk outcrops, are among the diagnostic species. This brings the association together with desert-steppe communities of the order Agropiretalia Korolyuk et Laktionov 2021, class Artemisietea lerchianae Golub 1994 (Korolyuk, Laktionov, 2021). Nevertheless, the composition and structure of cenoses confirm the assignment of the association to the class Festuco-Brometea. There are 4 variants within the association.
The diversity of xeropetrophytic vegetation of the South Urals is much higher than that described in the article. The involvement of data from the adjacent regions of Kazakhstan will facilitate the further development of its syntaxonomical system.
The tendency to gradual reduction of crop is observed now in the North-West of the Russian Federation. From 2010 for 2021 the total area under crops in four regions (Leningrad, Novgorod, Pskov, ...Vologda) has decreased from 1150.7 to 907.5 thousand hectares. Studies of specific structure of weed communities of the North-West of the Russian Federation started in the first decade of the XX century with inspections of crops by A. I. Maltsev in the St. Petersburg region (Maltsev, 1908, 1909). Since the beginning of the XXI the study of weed plants were focused at the All-Russian Institute of Plant Protectionwhere monitoring of crops of the North-West of the Russian Federation (Mysnik, 2012; Luneva, Mysnik, 2016, 2017, Luneva et al., 2009) has became the priority direction. In spite of previous wide floristic researches, the diversity of weed communities of this region is still not established. The first results of the floristic classification of weed vegetation in the North-East of the Russian Federation are presented.
The dataset is based on 278 relevés made by N. N. Luneva, I. N. Nadtochiy, E. V. Filippova, T. D. Sokolova, Yu. V. Eroshina for field seasons in 1999–2916. The research area covered Leningrad, Novgorod, Pskov and Vologda regions. Data processing was carried out according floristic classification approach (Westhoff, Maarel, 1978), using TWINSPAN algoritm in JUICE software packages (Tichy, 2002). The new syntaxa are named in accordance with the rules of the International Code of Phytosociological Nomenclature (Theurillat et al., 2021). DCA-ordination using the CANOCO 4.5 software package (Ter Braak, Smilauer, 2002) was produced to identify patterns of environmental differentiation. The habitat moisture and soil richness-salinity scales status of communities (Ramenskiy et al., 1956) was calculated and basic agroclimatic parameters were identified to characterize the ecological variability of habitats.
The communities were classified within the order of Aperetalia spica-venti J. Tx. et Tx. in Malato-Beliz et al. 1960 of the class Papaveretea rhoeadis S. Brulo et al. 2001 and were assigned to the alliance Scleranthion annui (Kruseman et Vlieger 1939) Sissingh in Westhoff et al. 1946), which unites the most mesophytic weed communities distributed mainly on gray forest, sod-podzolic, podzolic and other types of soils of the forest zone. The alliance diagnostic species are Stellaria media, Tripleurospermum inodorum, Capsella bursa-pastoris, Lepidotheca suaveolens, Thlaspi arvense, Centaurea cyanus. Three species from the diagnostic groups of ruderal vegetation classes are also active in their cenoflora — Elytrigia repens, Viola arvensis, Taraxacum officinale.
From the diagnostic group of the Papaveretea rhoeadis class and the order Aperetalia spica-venti with high constancy, species are found Cirsium setosum, Chenopodium album, Sonchus arvensis, less often are Fallopia convolvulus. The group of species of the alliance Scleranthion annui includes Stellaria media, Tripleurospermum inodorum, Capsella bursa-pastoris, Lepidotheca suaveolens, Thlaspi arvense, Centaurea cyanus. Unlike the weed communities of other regions of the Russian Federation (Khasanova et al., 2021 Yamalov et al., 2021a, b), apophytes are not significantly presented in cenophlora, but such ruderal species as Elytrigia repens, Viola arvensis, Taraxacum officinale are active.
The identified associations and communities are different geographically, in terms of crop affinity and the system of agricultural machinery, the type of soils and heat conditions. The most widespread are the communities of the ass. Tussilagi farfarae–Centauretum cyani, which occur in the Leningrad, Novgorod, Pskov and Vologda regions. The associations Galeopsetum bifidae (Novgorod, Pskov and Vologda regions) and Lamio purpurei–Persicarietum lapathifolii (Leningrad, Novgorod and Pskov regions) are of slightly smaller range.
Syntaxa are differentiated according to the level of habitat heat conditions from the higher (ass. Echinochloo crusgalli–Galietum aparinis, ass. Lamio purpurei–Persicarietum lapathifolii) to the lower (community Fumaria officinalis) ones.
Communities of the associations Echinochloo crusgalli–Galietum aparinis and Lamio purpurei-Persicarietum lapathifolii are formed on swamp, peat and podzolic-swamp soils. The community Polygonum aviculare and the ass. Mentho arvensi–Sonchetum arvensis are formed on weakly and medium-podzolic, as well as sod-podzolic soils. The other syntaxa are confined to sod-carbonate soils.
According to the spectrum of crop, syntaxa were distributed as follows: associations Echinochloo crusgalli–Galietum aparinis, Mentho arvensis–Sonchetum arvensis, Lamio purpurei–Persicarietum lapathifolii and the community Polygonum aviculare occur mainly on fields with vegetable crops (potatoes, carrots, cabbage, beets, etc.); Tussilagi farfarae–Centauretum cyani, Galeopsetum bifidae, the communities of Fumaria officinalis and Achillea millefolium — on grain crops (barley, oats, rye, triticale and wheat).
The main factors of differentiation of the studied communities are the type of soil, cultivated crop and humidification.
The article provides a survey of grass psammophylous vegetation syntaxa in the Southern Nechernozemye of Russia and validation of syntaxa in accordance with the requirements of the International Code ...of Phytosociological Nomenclature (Theurillat et al., 2021); questions of syntaxonomy and diagnosis of units of different levels of the syntaxonomical hierarchy are discussed.
The data for the analysis was the base obtained in 2018–2021, including 312 relevés of psammophylous grass vegetation in the Southern Nechernozemye of Russia (Bryansk, Kaluga, south-west of Moscow, north-west of Oryol, Smolensk Regions). Most of the relevés were made on the watershed of two large river systems: the Dnieper (the Sozh river basin) and the Volga (the Oka river basin) while some ones are in the northwestern part of the Smolensk region (Demidovsky district, Smolenskoye Poozerye National Park), which belongs to the Zapadnaya Dvina basin. Natural psammophytic habitats in this region are widespread on outwash plains, sandy river terraces dominated by pine forests of the alliance Dicrano–Pinion sylvestris (Libb. 1933) W. Mat. 1962 nom. conserv. propos. Open sands are formed in the place of such forests after clear felling of pine with the destruction of the living land cover. In addition, psammophylous grass vegetation forms on non-flooded or short-flooded sandy ridges in river floodplains, on plowed sands that were opened during construction, along sand pits, on clearings under power lines, along old abandoned roads on sand embankments and sandy roads fallow lands and pastures with sandy and sandy loam soils (Kupreev et al., 2020).
The psammophylous grass vegetation of the studied region is represented by 12 associations comprising 4 alliances and 2 orders of the class Koelerio–Corynephoretea canescentis Klika in Klika et Novák 1941, which unites dry grasslands on sandy soils and on rocky outcrops of the temperate to boreal zones of Europe, the North Atlantic islands and Greenland (Mucina et al., 2016). On the basis of comparative analysis for alliances and orders we compiled regional combinations of diagnostic species with constancy in coenoflora above 20 % and values of the statistical φ-coefficient above 20 (at p <0.01): Armerion elongatae (Armeria maritima, Astragalus arenarius, Dianthus fisheri, Jovibarba globifera, Jurinea cyanoides, Koeleria glauca), Corynephorion canesentis (Corynephorus canescens), Hyperico perforati–Scleranthion perennis (Berteroa incana, Festuca ovina, Helichrysum arenarium, Hypericum perforatum, Jasione montana, Pilosella officinarum, Rumex acetosella, Scleranthus perennis, Trifolium arvense). The following species diagnose the orders of psammophylous vegetation: Corynephoretalia canescentis (Chamaecytisus ruthenicus, Cladonia arbuscula, Corynephorus canescens, Jurinea cyanoides, Koeleria glauca, Sedum acre) and Trifolio arvensis–Festucetalia ovinae (Abietinella abietina, Elytrigia repens, Hieracium umbellatum, Jasione montana, Festuca ovina, F. rubra, Galium mollugo, Pilosella officinarum, Plantago lanceolata, Poa angustifolia, Trifolium arvense, Viola tricolor). Diagnostic combinations of alliances partly duplicate those of orders, and some species are the diagnostic for two alliances, which does not contradict the logic of syntaxonomy.
Analysis of the comparative table showed that all syntaxa of psammophylous vegetation taken for analysis can be fully assigned to the class Koelerio–Corynephoretea canescentis in accordance with its current diagnosis (Mucina et al., 2016). According to the ratio of the diagnostic species of the classes Koelerio–Corynephoretea canescentis and Sedo-Scleranthetea, it is not possible to divide syntaxa into two groups. In general, in the Southern Nechernozemye of Russia, typical natural habitats characteristic of the Central European communities of the class Sedo-Scleranthetea are virtually absent. They are imitated by anthropogenic ecotopes with washed out or trampled, usually heavily disturbed, rubble substrates. Probably, it is possible to classify some types of grass communities that form here on the slopes of railways and roads, on overgrown stony dumps, strongly compacted as a result of trampling or the passage of vehicles, to the class Sedo-Scleranthetea. However, the assigning of these communities to this class is debatable.
A significant part of the studied phytocoenoses was assigned to 7 non-rank units — «communities» within the class Koelerio–Corynephoretea canescentis. These are, firstly, pioneer and, as a rule, sparse or floristically incomplete psammophylous communities at the initial stages of succession with the participation of widespread grass psammophilic oligotrophs. Secondly, these are monodominant phytocoenoses formed after anthropogenic disturbance of psammophylous vegetation, for example, on fallow lands and clearings in pine forests on sandy and sandy loam soils; after fires, trampling. The names of these syntaxa are given according to the dominant plant species.
A nearly circumpolar hypoarctic species Stuckenia subretusa (Hagstr.) Holub is a rare species that grows in Yakuyia in lakes and reservoirs of deltaic systems of the large rivers (Lena, Kolyma, Yana, ...Indigirka, Anabar) mainly north of 68° N(Bobrov, Mochalova, 2014, 2017; Egorova, 2016; Opredelitel’…, 2020). The species is listed for Taymyr Peninsula as the most common in the plain part of the southern tundra and in the forest tundra (Pospelova, Pospelov, 2007) and as rarer in the typical tundra (Polozova, Tikhomirov, 1971). In general, this is a rather rare species. Besides the Taymyr Peninsula and Yakutia it occurs in the Bolshezemelskaya tundra, in the mouth of the Yenisey River, in Chukotka Peninsula and Alaska (Yurtsev et al., 2010; Bobrov et al., 2021).
Stuckenia subretusa is a critical taxon associated by some authors (Kaplan, 2008; Konspekt…, 2012) with S. vaginata (Turcz.) Holub. However, both morphological and molecular genetic differences between these taxa were shown later (Volkova et al., 2017). It was also shown that S. subretusa is found in the lower reaches and estuarine areas of the large rivers in Asia, mainly north of 68° N, in Europe — of 67°, while the northern limit of the distribution of S. vaginata in Yakutia — 64°, in Europe — 66° (Bobrov, Mochalova, 2014), therefore these species are geographically separated.
In July 2021, during a geobotanical survey of the islands of the southern part of the Lena River delta, thickets of S. subretusa were found on Sasyl-Ary Isl. (Fig. 1) in a shallow flow (channel) of 2 km total length, 130–150 m width and from 4–5 cm to 1.5 m depth with a very sluggish current. The bottom substrate is silty sands. Permafrost is 2–3 m depth. Thickets of S. subretusa were found only at 4–30 cm water depths. The thickets are monospecies with projective cover from 35 to 80 % (Fig. 2). Single specimens of Tephroseris palustris (L.) Rchb. were met rarely, at 4–5 cm water depths. Part of the thickets was located on wet sand along the shallows of the channel. S. subretusa was not found on neighboring islands.
Five geobotanical relevés were made within the flow according to the generally accepted methodology with georeferencing using 12-channel GPS in the WGS-84 coordinate system on sample plots of 10×10 m. The classification was carried out on the principles of the ecological-floristic approach (Westhoff, van der Maarel, 1973). Estimation of the species projective cover in Table is given according to the Brown-Blanquet scale (Becking, 1957): r — single; + — less than 1 %; 1 — 1–5 %; 2 — 6–25 %; 3 — 26–50 %; 4 — 51–75 %; 5 — 76–100 %. The syntaxon name is given in accordance with the International Code of Phytosociological Nomenclature (Theurillat et al., 2021).
The described communities belong to the class Potamogetonetea Klika in Klika et Novák 1941, the order Potamogetonetalia Koch 1926, and the alliance Potamogetonion Libbert 1931. We consider these as a part of a new association.
The ass. Stuckenietum subretusae ass. nov. (Table). Nomenclature type (holotypus) — relevé No. L21-108а (Table, relevé 3). Republic of Sakha (Yakutia), Bulunsky district, Sasyl-Ary Isl., shallow channel in the Lena River delta, in water (N 72.35989°, E 126.41968°), 07/20/2021. N. N. Lashchinskiy is an author. Diagnostic species of the association is Stuckenia subretusa (= Potamogeton subretusus).
The association includes monospecies communities of Stuckenia subretusa, sometimes with a single presence of other species. It is possible that communities of this species are floristically richer in southerner regions. These are reliably found in other parts of the Lena River delta (Nikolin et al., 2017), on Taimyr Peninsula (Krasnoyarsk Territory) in the Khatanga River (Pospelova, Pospelov, 2007), as well as in the Komi Republic (Chemeris, Bobrov, 2020), where their productivity is from 142 to 409 g/m2 of air-dry matter.
In general, the association distribution area apparently coincides with the species range (northern regions of the Komi Republic, Krasnoyarsk Territory, Yakutia, Chukotka Autonomous District (Russia) and Alaska (USA).