classification of the class molinio-arrhenatheretea in the

Classification of the class Molinio-Arrhenatheretea in the forest and forest-steppe zones of Ukraine

Anna A. Kuzemko*

Abstract

Aims: This paper reviews the classification of mesic grassland vegetation (phytosociological class Molinio-Ar-rhenatheretea) in Ukraine, and integrates the units recorded into the common European syntaxonomical sys-tem of the Molinio-Arrhenatheretea class. It also proposes solutions to a number of issues that cause conflict between the classical Central European concept of the Molinio-Arrhenatheretea class and its traditional syntax-onomy in the former USSR. Location: Forest and forest steppe zones of Ukraine. Methods: I analysed 2,105 relevés originally assigned to the Molinio-Arrhenatheretea class using the European Expert System and K-means clustering. The units were evaluated for quality and internal homogeneity using Sharpness index and Average Whittaker beta-diversity. I determined the diagnostic species of the vegetation units using calculations of their fidelity based on a phi-coefficient. The environmental assessment of the units follows the Didukh indi-cator values. Results: I interpreted the resulting vegetation units as alliances of the Braun-Blanquet system (Agrostion vinealis, Arrhenatherion elatioris, Cynosurion cristati, Deschampsion cespitosae, Molinion caeruleae, Potentillion anserinae and Calthion palustris) based on a complex of diagnostic species. I also analysed the dis-tribution of the communities of the identified alliances in the study area, revealing their ecological features. Conclusions: The use of modern phytosociological methods in this study to analyse geobotanical data col-lected over a long period and a large area covering the whole range of the Molinio-Arrhenatheretea class in the country, clarified a number of controversial issues previously related to the lack of coordination between the Central and Eastern European phytosociology.

Keywords: Arrhenatheretalia; Europe; expert system; Galietalia veri; grassland; meadow; Molinietalia; syntax-onomy; vegetation database.

Nomenclature: Flora Europaea (Tutin et al. 1968–1993) for vascular plants; Mucina et al. (in press) for higher syntaxa.

Abbreviations: DCA = Detrended Correspondence Analysis; DIV = Didukh indicator values; EVA = Euro-pean Vegetation Archive (http://euroveg.org/eva-database); GIVD = Global Index of Vegetation-Plot Data-bases (http://www.givd.info); ICPN = International Code of Phytosociological Nomenclature; PAM = Parti-tioning around medoids.

Submitted: 22 September 2015; first decision: 15 December 2015; accepted: 7 August 2016

Co-ordinating Editor: Monika Janišová

© 2016 Gebrüder Borntraeger, 70176 Stuttgart, GermanyDOI: 10.1127/phyto/2016/0083

www.borntraeger-cramer.de0340-269X/2016/0083 $ 7.20

Phytocoenologia Vol. 46 (2016), Issue 3, 241–256 Special Issue Palaearctic grasslandsStuttgart, November 2016 Research Paper

*Author’s address: National Dendrological Park “Sofiyivka”, National Academy of Sciences of Ukraine, 12a Kyivska St., Uman’, 20300, Ukraine; [email protected].

Introduction

Mesic grasslands are one of the main types of natural and semi-natural habitats in the forest and forest-steppe zones of Ukraine and Europe. On the one hand, this ve-getation type is very diverse, but on the other, a large number of its syntaxa are widespread, often across the whole European continent. The vast majority of these

syntaxa are found in Ukraine, which, as shown by recent studies (Jiménez-Alfaro et al. 2014) is the second of the Mediterranean countries in the number of classes and al-liances it supports. Due to its importance as pasture and meadow, the state of the vegetation is largely determined by socio-economic factors, in particular, the level of de-velopment of animal husbandry and conservation of tra-ditional land use. Therefore, changes in Ukrainian soci-

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242 Anna A. Kuzemko

ety over the past decades have significantly affected the structure and functioning of meadow vegetation. As a result, there is a need to inventory the meadow vegeta-tion of Ukraine to record its current state as well as to compare it with meadow vegetation in other European countries. A number of works providing overviews of European grassland syntaxonomy, including meadow vegetation, have been published in recent decades (Mucina et al. 1993; Dierschke 1995; Päzolt & Jansen 2004; Chytrý et al. 2007; Janišová 2007; Rūsiņa 2007; Hegedüšová Vantarová & Škodová 2014, etc.). In Ukraine, such recent approaches to phytosociological re-search are little known so far, which significantly compli-cates the integration of information on the plant commu-nity diversity of this country in the global and European system.

For a long time, vegetation in Ukraine was studied fol-lowing the Soviet tradition in geobotany. In the 1980’s, the Braun-Blanquet approach was introduced in Ukraine and only the floodplain meadow vegetation of the Mo-linio-Arrhenatheretea Tx. 1937 class was chosen to test this method in the Ukrainian context. This choice was dictated by the frequent presence of several dominant species in meadow communities (so-called poly-domi-nance), which complicates the use of the Soviet approach for their classification (Mirkin & Shelyag-Sosonko 1979; Shelyag-Sosonko et al. 1989).

It is possible to distinguish three stages in the history of the phytosociological study of mesic grasslands of Ukraine: During the first stage (1980–1983) the flood-plain meadows of some rivers of Ukraine were examined with the aim to collect geobotanic data following the Braun-Blanquet methodology. Data from the valleys of the Dnieper and Dniester rivers were then analysed based on the Braun-Blanquet approach in which environmen-tally specific groups of species were identified. Based on this, the ecological and topological types of vegetation with the ranks of associations, subassociations and vari-ants were obtained in which the associations were com-bined into conventional units of higher rank. Phytocoe-noses were given without complete tables and nomen-clatural types (Shelyag-Sosonko et al. 1980, 1981, 1982; Solomakha 1981а, 1981b, 1982; Shelyag-Sosonko & Solomakha 1981; Sipailova & Shelyag-Sosonko 1981; Sypailova & Shelyag-Sosonko 1982а, 1982b). In the sec-ond stage (1983–1990), the data were systematized and classification schemes were proposed as a complete hier-archy of syntaxonomical units for each studied region (floodplain of the Dnieper, Dniester, Desna, Vorskla riv-ers, small rivers of Ukrainian Polesie) (Shelyag-Sosonko et al. 1985, 1986, 1987) and named following the ICPN (Barkmann et al. 1976). This stage is characterized by the description of new syntaxa including the order Poo-Agrostietalia vinealis Shelyag et al. 1985, the alliances Agrostion vinealis Sypailova et al. 1985 and Festucion pratensis Sypailova et al. 1985, and many associations

(Sypailova et al. 1985; Shelyag-Sosonko et al. 1985, 1987). At the same time, the lack of information on syntaxon-omy of Central Europe was a serious limitation. Thus, some of the European syntaxa were re-described by Ukrainian phytosociologists, which caused a number of syntaxonomical problems that have not yet been re-solved. In the third stage (from 1990 to present) regional classification schemes were systematically developed in order to create the full list of syntaxa for different regions and for Ukrainian vegetation in total, including meadows (Sypailova & Shelyag-Sosonko 1996; Shevchyk et al. 1996; Vorobyov et al. 1997; Senchylo et al. 1997, 1998; Bairak 1998; Kuzemko 1999, 2011a, 2011b; Goncharenko 2000a, 2000b, 2003; Fitsailo 2003; Solomakha et al. 2004; Gomlya 2005; Orlov & Yakushenko 2005; Chorney et al. 2005; Klimuk et al. 2006; Gal’chenko 2006; Onyshchenko 2006; Tertyshnyi 2006; Soroka 2008; Tokaryuk et al. 2009; Solomakha & Chornei 2011). At this stage, foreign literature became more widely available and Ukrainian scientists were able to compare syntaxa described in the Ukraine with the European units. However, this positive phenomenon had somewhat negative consequences, be-cause some of the identified units were included in Euro-pean syntaxa without considering their diagnosis, and often only based on the name-giving taxa.

The accumulation of a large number of relevés and the need for its critical analysis and synthesis induced Ukrainian scientists to produce a four-volume “Vegeta-tion of Ukraine” (Malynovs’kyi & Krichfalushyi 2000; Dubyna 2006; Dubyna et al. 2007; Kuzemko 2009b). This was mainly based on an inductive approach, which meant that the analysis considered relevés from all over Ukraine, which were processed by phytocoenotic table transformation using the FICEN software package (Kos-man et al. 1991). The 4th volume of this edition was pre-pared by the author of this paper and devoted to vegeta-tion of the Molinio-Arrhenatheretea class (Kuzemko 2009b). At the same time, the studies of vegetation in Ukraine on the basis of the Braun-Blanquet approach were summarized in the review of Solomakha (2008).

The structure of the Molinio-Arrhenatheretea class dif-fers considerably in the monographs of Solomakha (2008) and Kuzemko (2009b), especially in the part that concerns xero-mesophytic meadows, since Solomakha distin-guishes two orders which include this type of meadow communities. He defines the Poo-Agrostietalia vinealis Shelyag, V. Solomakha et Sipailova 1985 order as xero-mesophilous and meso-xerophilous communities of the elevated areas of central and riverine parts of floodplains in the Ukrainian plains on turf, sod-meadow and cherno-zem-meadow soils. He defines the Galietalia veri Mirk et Naumova 1986 order as communities of fringes and meadows on medium rich soils with moderately variable atmospheric moisture, and floodplain meadows on slightly developed sandy soil that are moistened by pre-cipitation in the Forest-Steppe zone. However, Kuzemko


Classification of the class Molinio-Arrhenatheretea 243

& Dziuba (2002) consider these two orders to be synony-mous and include all “dry” communities of the Molinio-Arrhenatheretea class in the Galietalia veri (Kuzemko 2009b) order, which they define as xero-mesophilous and meso-xerophylous communities of steppic meadows on turf, sod-meadow and chernozem-meadow sandy and loamy soils. Two alliances Trifolio-Brizion elatioris Didukh & Kuzemko 2009 and Helictotricho-Bistortion officinalis Didukh & Kuzemko 2009 that are given in this monograph were described from Crimea (Didukh & Kuzemko 2009), while the rest represent the vegetation of the plain part of Ukraine and the Carpathians.

Thus, two recently published classification schemes of the Molinio-Arrhenatheretea class in Ukraine signifi-cantly differ and are not consistent with the syntaxon-omy of the class accepted in other European countries. This indicates a need for further critical revision of the vegetation class using modern research methods.

In this regard, the aims of this paper are:i) to carry out a syntaxonomical analysis of available

data on mesic and wet grasslands of the Molinio-Ar-rhenatheretea class to alliance level using modern nu-merical phytosociological methods

ii) to analyze the geographical distribution of the units obtained in the forest and forest-steppe zones of Ukraine and to determine the features of their eco-logical differentiation;

iii) to integrate the units obtained in the common Euro-pean syntaxonomical system of the Molinio-Ar-rhenatheretea class, and

iv) to resolve a number of critical issues that create con-tradiction between the classical Central European concept of the Molinio-Arrhenatheretea class and its traditional syntaxonomy in the former USSR.

Study area

The study area covers the forest and forest-steppe zones of the Ukrainian plains. In general, I follow the bounda-ries of the territory in accordance with the latest version of geobotanical zonation of Ukraine (Didukh & Shelyag-Sosonko 2003) (Fig. 1). Overall, the study area covers about 54% of Ukraine (~326 000 km²).

Much of the territory belongs to the Polissian (maxi-mum 316 m a.s.l.) and Dnieper (236 m) lowlands. The right bank of the Dnieper River is dominated by upland areas, namely the Podolian (471 m), Volyn (361 m) and Dnieper (323 m) uplands (Marinich et al. 1985; Kryzhova & Kulyk 2007), in which different rocks (granite, lime-stone, chalk, sandstone, etc.) are often exposed on the surface. This causes considerable diversity of steppe and petrophyte grassland vegetation. However, the river val-leys in the upland regions are often narrow with very small floodplains, and little space for mesic and wet grass-lands. In contrast, lowland regions are characterized by

extensive floodplains with high diversity of mesic, damp and wet grasslands.

In the Polissian lowland area, the zonal soils are sod-podzols. In big lowlands with relatively frequently high water tables, the soils are gleying in the lower horizons and the landscape is dominated by bog-podzolic soils. In depressions in continental meadows, sod-gley soils are formed with varying degrees of gleying, depending on the depression depth. The periphery of peatlands usually contains peat-bog soils. The transition from Polissya (forest zone) to forest-steppe is associated with the ap-pearance of carbonate loess sediments that change from non-carbonate glacial deposits. In the forest-steppe, the zonal soils are light grey and grey forest soils as well as typical chernozems. Floodplains have mainly meadow and meadow-marsh soils (Vernander 1986).

The large size of the study area (about 500 km from north to south and more than 1,100 km from west to east) means there is a wide variety of climatic conditions, which obviously have a great influence on the grassland formations, including meadow vegetation. The average annual temperature increases slightly from north to south. It is the lowest in the northeast (6–7°C), and the highest in the southwest (9–10°C). Due to the air circula-tion patterns, precipitation in Ukraine decreases from the north and northwest towards the south and southeast. In the cold period (November-March), precipitation throughout most of the territory is 175–200 mm. In the warm season (April-October), the spatial distribution of precipitation is similar to the annual distribution (Lipins’kyi et al. 2003). The highest annual rainfall occurs in the western part of the study area (450–475 mm) and the lowest along the boundaries of the forest-steppe and steppe zone (325–350 mm). Thus, during the warm sea-son precipitation gradually decreases in the direction from northwest to southeast. The plain part of Ukraine is located within two climatic regions, with the study area being assigned to the Atlantic-Continental regional cli-mate (Lipins’kyi et al. 2003).

Methods

Data preparation

The study is based on the vegetation relevés collected from the literature and from unpublished sources stored in the Ukrainian Grassland Database (GIVD code EU-UA-001) (Kuzemko 2012b), which is also included in the European Vegetation Archive (EVA) (Chytrý et al. 2016).

Only those relevés previously assigned to the Molinio-Arrhenatheretea class were selected, resulting in a dataset with 2,105 relevés and 1,047 species. The plot size of the relevés ranged from 16 to 100 m2, which is considered ac-ceptable for grassland vegetation. The floristic data were


244 Anna A. Kuzemko

further analysis in order to obtain a more homogeneous dataset. I also excluded all species identified only to the genus level (except the Alchemilla genus). The dataset was stratified following a Heterogeneity-Constrained Resampling (Lengyel et al. 2011) to the relevés from one cell of the geographical grid of 1.25 minutes of longitude × 0.75 minutes of latitude (approximately 1.5 × 1.4 km). The relevés were then sorted in order to maximize beta-diversity (according to the Bray-Curtis Index) in the range from 1 to 5 relevés per stratum. As a result of this stratification 1,197 relevés with 681 species were obtained and used for further analysis.

Data analysis

All analyses were performed in JUICE 7.0 (Tichý 2002). In particular, I tested divisive methods of clustering: TWINSPAN algorithm (Hill 1979), modified TWIN-SPAN (Roleček et al. 2009) measuring of heterogeneity

revised and corrected according to Flora Europaea (Tutin et al. 1968–1993). There are some mismatches in nomen-clature used in Ukrainian and European taxonomy. In some cases, I used species aggregates as presented in the Supplement S1 in order to avoid introducing error into the identification of vegetation units based on floristic composition.

For more accurate assignment of relevés to the Mo-linio-Arrhenatheretea class, I carried out additional ana-lyses using the European Expert System (Mucina et al., in press) in the JUICE program (Tichý 2002), which allows raw automatic classification of plant community data us-ing some selected target species and thus assigning them to a class of vegetation. During this analysis, 414 relevés were re-assigned to other classes of vegetation. A total number of 1,691 relevés that according to the expert sys-tem were correctly classified in the class Molinio-Ar-rhenatheretea class were finally selected. Since not all relevés contained information about cryptogams or tree and shrub layers, all of these species were removed before

Fig. 1. Forest and forest-steppe zones of Ukraine (Didukh & Shelyag-Sosonko 2003). 1-4 the borders of the units of geobot-anical zonation of Ukraine – zones (1), provinces (2), subprovinces (3) and districts (4); A-B – European broad-leaves forests zone: Central European province of broad-leaved forest (A), Eastern European province of mixed and broad-leaved forest (B), C – Forest-Steppe zone, Forest-Steppe province of the oak forests, steppe meadow and meadow steppes.



with Sørensen, Simpson, Jaccard dissimilarity and Whit-taker beta-diversity, and also agglomerative methods based on cluster analysis in PC-ord software (McCune & Mefford 2006). I used different combinations of distance measures and group linkage method, with and without logarithmic transformation of the data, and also cluster algorithms of non-supervised K-means (Legendre & Legendre 1998) and PAM (Kaufman & Rousseuw 1990) with number of starts 10 and number of related plots for definition of starting centroid three or five. To determine the optimal number of clusters I used functions “crisp-ness of classification” (Botta-Dukát et al. 2005) the re-sults of which are presented in Supplement S2.

The units obtained in the analysis have been evaluated for quality using Sharpness index (Chytrý & Tichý 2003) and for internal homogeneity by calculating Average Whittaker beta-diversity. When selecting the optimal method of classification, an advantage was given to the method that provided the receiving units with: 1) higher values of Sharpness index, 2) lower values of Average Whittaker beta-diversity, and 3) lower values of standard deviation for both of these indexes (Supplement S3). However, the final choice of the best method of classifi-cation was based on expert knowledge of the author.

Diagnostic species of the vegetation units were deter-mined using calculations of their fidelity based on a phi-coefficient (Tichý & Chytrý 2006). Diagnostic species were considered as those with a phi-coefficient higher than 0.2, and highly diagnostic species were those with phi > 0.4 (bold in the text). When calculating this index I used the following options: 1) calculation of fidelity based on the presence / absence of species; 2) standardi-zation of all relevés to groups of equal size; 3) exclusion of non-essential values of fidelity based on Fisher’s exact test at p < 0.01.

The identification of the obtained syntaxa was carried out on the basis of analysis of their diagnostic species us-ing the author’s own expert assessment and the literature (Päzolt & Jansen 2004; Hájková et al. 2007, 2014; Rūsiņa 2007; Kącki et al. 2013). The most useful sources were those using numerical methods of vegetation classifica-tion, an inductive approach and a clear, ecologically grounded concept of syntaxa.

I used the Didukh indicator values (DIV; Didukh 2011) for the environmental assessment of the identified units including 10 factors: soil water regime (Hd), soil acidity (Rc), total salt regime (Sl), carbonate content in soil (Ca), nitrogen content in soil (Nt), thermoregime (Tm), continentality of climate (Kn), humidity of climate (Om), cryo-climate (Cr) and light (Lc). The values of the environmental factors were determined in the program JUICE by using the tool “Indicator values”. To identify the patterns of syntaxa arrangement in multidimensional space of environmental factors I used a method of DCA-ordination with R-project, integrated into JUICE with DIV as environmental vectors.

Results

Classification

The 414 relevés originally assigned to the Molinio-Ar-rhenatheretea class that were not confirmed by the Euro-pean Expert System were re-assigned to the classes Phragmito-Magno-Caricetea Klika in Klika et Novák 1941 (123 relevés), Artemisietea vulgaris Lohmeyer et al. ex von Rochow 1951 (104 relevés), Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 (80 relevés), Koelerio-Coryne-phoretea Klika in Klika et Novák 1941 (79 relevés), Scheuchzerio-Caricetea fuscae Tx. 1937 and Stellarietea mediae Tx. et al. ex von Rochow 1951 (7 relevés each), Nardetea strictae Oberd. 1949 (six relevés), Galio-Urtice-tea Passarge ex Kopecký 1969, Bidentetea Tx. et al. ex von Rochow 1951 and Trifolio-Geranietea sanguinei T. Müller 1962 (two relevés each), Polygono-Poetea annuae Rivas-Mart. 1975 and Salicetea purpureae Moor 1958 (one relevé each).

I accept that differentiation at the alliance level reflects the results with the number of clusters from six to 10 that depends on the method chosen. When using the cluster analysis, available in PC-ord, with different distance measures and group linkage methods, the units obtained are characterized by a quite low values of the Sharpness Index and high values of the Whittaker beta-diversity. This indicates that they are less suitable for analysis of the data set compared to other methods (Supplement S3). The best results for sharpness and homogeneity of the units were obtained using the modified TWINSPAN al-gorithm. However, when using this method the mean values of the Sharpness Index and Whittaker beta-diver-sity were characterized by very high values of the stand-ard deviation, which is explained by the uneven number of relevés in clusters, which ranged from three to 590. When clustering methods using K-means and PAM were used (with number of starts being 10 and number of re-lated plots for definition of starting centroid being five), the results were approximately similar, but with K-means clustering the units are characterized by lower figures of Whittaker beta-diversity, i.e. they are internally more homogeneous. Thus, the best results were obtained by the clustering method of K-means and seven clusters, which allowed the best interpretation of the results (Ta-ble 1).

The resulting vegetation units were interpreted as alli-ances of the Braun-Blanquet system based on complex of diagnostic species (Supplement S4).

Cluster 1 was identified as the Agrostion vinealis alliance. Diagnostic species: Achillea millefolium subsp. millefo-lium, Agrostis vinealis, Allium angulosum, Bromopsis iner-mis, Carex praecox, Centaurea jacea subsp. jacea, Cicho-rium intybus, Crepis tectorum, Dianthus borbasii subsp. borbasii, Elytrigia repens subsp. repens, Eryngium planum,


246 Anna A. Kuzemko

Euphorbia virgata, Festuca valesiaca subsp. valesiaca, Fili-pendula vulgaris, Galium verum subsp. verum, Koeleria delavignei, Lotus corniculatus subsp. corniculatus, Poa an-gustifolia, Potentilla argentea, Rumex acetosella subsp. acetosella, Rumex thyrsiflorus, Stellaria graminea, Vicia tetrasperma.

It includes relevés of communities that are distributed mainly in the northeastern part of Ukraine, on the left bank of the Dnieper River (Fig. 2a). In the right-bank of the Dnieper River (Right-Bank Ukraine) they occur only in the Dnieper basin. This syntaxon includes psamo-xero-mesophilous communities common in riverine part of floodplains. The communities are usually used as pas-ture although they have low productivity. Dominants in this vegetation are representatives of the steppe and psammophytic communities Agrostis vinealis, Poa an-gustifolia and Koeleria delavignei. The complex of diag-nostic species is well defined, which is also confirmed by high values of the Sharpness Index (see Table 1) and also includes a significant proportion of representatives of the afore-mentioned types of vegetation. The species with the highest constancy in this alliance are Plantago lanceo-lata, Festuca pratensis, Alopecurus pratensis and Trifolium pratense.

Cluster 2 was identified as the alliance Arrhenatherion elatioris Luquet 1926. Diagnostic species: Achillea millefolium subsp. millefo-lium, Anthoxanthum odoratum, Bromus hordeaceus subsp. hordeaceus, Carex hirta, Carum carvi, Cerastium fontanum subsp. fontanum, Dactylis glomerata subsp. glomerata, Daucus carota subsp. carota, Erigeron annuus subsp. annuus, Galium mollugo, Medicago lupulina, Oe-nothera rubricaulis, Pimpinella saxifraga, Plantago lanceolata, Trifolium dubium, Veronica chamaedrys subsp. chamaedrys.

It includes communities distributed mainly in the northern part of the study area within the forest zone, but in the forest-steppe they are rare (Fig. 2b). The syn-taxon includes communities of mesic floodplains and up-land meadows, mainly distributed in flat, sometimes de-pressed areas of riverbed parts of floodplain and slightly

elevated areas in the central part of floodplains. This alli-ance is characterized by the highest internal heterogene-ity as the value of Whittaker beta-diversity is greater than that of the other alliances (see Table 1). Dominant species of the communities are typical of mesic grasslands Fes-tuca rubra, F. pratensis, Poa pratensis. The same species as well as Ranunculus acris, and Trifolium pratense have the highest constancy.

Cluster 3 includes relevés assigned to the Cynosurion cristati Tx. 1947 alliance. Diagnostic species: Festuca pratensis, Leontodon autum-nalis, Lolium perenne, Poa pratensis subsp. pratensis, Ra-nunculus acris subsp. acris, Taraxacum officinale, Trifo-lium pratense subsp. pratense, Trifolium repens subsp. repens.

Communities of this alliance are quite evenly distrib-uted throughout the study area, while in the north-west-ern regions they are somewhat less common (see Fig. 2c). The alliance includes communities of floodplain, upland or lowland mesic grasslands confined to the plain, rarely slightly raised or depressed areas of the central and ter-race part of floodplain. Outside floodplains they occur on the flat-depressed areas of the watershed, in the bot-toms of the shallow flowing valleys and other depres-sions. The vegetation of this alliance is formed only un-der fairly intensive grazing or combined use. The alliance is characterized by the lowest value of the Sharpness In-dex (see Table 1). The communities are dominated by Festuca pratensis, as well as Lolium perenne, Poa pratensis and Trifolium repens, which are the main cenosis-form-ing species in mesic grasslands in Ukraine and have wide ecological amplitudes as well as being typical pasture spe-cies resistant to trampling. This complex of diagnostic species is generally consistent with the Central European version, with the exceptions of the name-giving taxon Cynosurus cristatus and Bellis perennis, because these species are at the southern and eastern limits of their con-tinuous distribution in the study area. The highest con-stancy in the relevés is reached by Achillea millefolium, Potentilla anserina, Ranunculus repens.

Table 1. Characteristics of the distinguished vegetation units.

Cluster N Syntaxa Number of relevés Sharpness Index Whittaker beta-diversity Index

1 Agrostion vinealis 158 32.40 13.78

2 Arrhenatherion elatioris 207 20.20 18.86

3 Cynosurion cristati 253 11.39 16.53

4 Deschampsion cespitosae 120 29.03 10.73

5 Molinion caeruleae 91 38.79 12.38

6 Potentillion anserinae 165 20.99 17.81

7 Calthion palustris 203 21.60 17.06



Agrostion vinealis Arrhenatherion elatioris

Cynosurion cristati Deschampsion cespitosae

Molinion caeruleae Potentillion anserinae

Fig. 2. Distribution of the alliances of the Molinio-Ar-rhenatheretea class in the forest and forest-steppe zones of Ukraine.


248 Anna A. Kuzemko

Cluster 4 was identified as the alliance Deschampsion cespitosae Horvatić 1930, and was distributed mainly in the north-eastern part of the study area in the forest zone, sometimes in the forest-steppe, but mainly on the Left-Bank part of Ukraine (Fig. 2d). Diagnostic species: Achillea pyrenaica, Agrostis canina subsp. canina, Agrostis gigantea subsp. gigantea, Allium angulosum, Alopecurus pratensis, Beckmannia eruci-formis, Carex vulpina, Cnidium dubium, Gratiola offic-inalis, Inula britannica, Juncus atratus, Lysimachia num-mularia, Lythrum virgatum, Myosotis cespitosa, Poa palustris, Ranunculus flammula, Ranunculus repens, Ve-ronica longifolia.

This alliance includes communities of wet floodplain meadows, which are confined to depressions of different parts of floodplain, often in riverine and central parts, en-riched by alluvial deposits. The formation of these com-munities is determined by flooding intensity, which is typical for floodplains in the lower reaches of large rivers. The land use is mainly hay mowing or mowing and graz-ing at moderate intensity. The alliance is characterized by a moderately high value of the Sharpness Index and at the same time it has the lowest value of the Whittaker beta-diversity (see Table 1). The communities are dominated by the tall grasses Alopecurus pratensis, Poa palustris and Agrostis gigantea. Potentilla anserina and Mentha arven-sis have the highest constancy.

Cluster 5 includes relevés assigned to the Molinion caeru-leae Koch 1926 alliance. Diagnostic species: Salix cinerea, Salix rosmarinifolia; Anthoxanthum odoratum, Briza media, Carex davalli-ana, Carex flava subsp. flava, Carex pallescens, Carex panicea, Epipactis palustris, Filipendula ulmaria, Galium uliginosum, Genista tinctoria, Gentiana pneumonanthe, Holcus lanatus, Linum catharticum subsp. catharticum, Luzula multiflora subsp. multiflora, Lysimachia vulgaris, Molinia caerulea, Parnassia palustris, Polygala amarella, Potentilla erecta, Sanguisorba officinalis, Schoenus fer-rugineus, Selinum carvifolia, Serratula tinctoria subsp. tinctoria, Stachys officinalis subsp. officinalis, Succisa pratensis, Trifolium montanum subsp. montanum, Vale-riana simplicifolia.

Communities of this cluster are distributed mainly in the northern regions of Ukraine within the forest zone, their largest concentration is observed in the west and northeast of the study area (Fig. 3e). The alliance includes communities of floodplain, upland and lowland moder-ately wet meadows in shallow depressions of the terraced area, with less in the central part of floodplain. In addition these commuinities occur outside floodplains in depres-sions on the river terraces or watersheds, often in place of partially reclaimed terrace marshes and in flat depressions with low amplitude of moisture fluctuations. They also occur in areas with low to moderate grazing intensity, sometimes disturbed by peat extraction, or in overgrown

former peat extraction sites. The land use is mowing and grazing with moderate intensity. The alliance exceeds all other alliances in the value of the Sharpness Index, and yet it has a very low Whittaker beta-diversity value (see Table 1). The dominant species is Molinia caerulea, and Poten-tilla anserina, Deschampsia cespitosa and Ranunculus acris have the highest constancy values.

Cluster 6 was identified as the alliance Potentillion an-serinae Tx. 1947, including relevés throughout the study area with an exception of the northwestern regions, and also the south forest-steppe of the Right-Bank of the Dnieper River (Fig. 2f). Diagnostic species: Agrostis stolonifera subsp. stoloni-fera, Alopecurus geniculatus, Bidens tripartitus subsp. tripartitus, Eleocharis palustris subsp. palustris, Glyceria fluitans, Mentha arvensis, Persicaria maculosa, Plantago major subsp. major, Potentilla anserina subsp. anserina, Rorippa amphibia, Schoenoplectus tabernaemontani, Trifolium fragiferum subsp. fragiferum, Triglochin palustris.

The alliance combines the vegetation of floodplain and lowland meadows, common in flat and shallow depres-sions of the terrace part (in floodplains of large and me-dium rivers) and the central part (floodplains of small riv-ers). They often occur in the floodplains affected by drainage amelioration. Outside floodplains, the commu-nities are confined to depressed areas of river terraces, watershed depressions, bottoms of ravines and gullies, drained valley marshes. They are formed in place of damp and wet meadows under conditions of intense grazing, which leads to the deterioration of the air and water re-gime of soil, often accompanied by the formation of characteristic hubble microrelief. The dominant species is Agrostis stolonifera which is well adapted to grazing. The grazing tolerant species Ranunculus repens and Trifolium repens also have highest constancy.

Cluster 7 is represented by communities assigned to the alliance Calthion palustris Tx. 1937 and evenly distrib-uted over the study area (Fig. 2g). Diagnostic species: Caltha palustris, Calystegia sepium, Carex acutiformis, Cirsium rivulare, Equisetum palustre, Galium palustre, Iris pseudacorus, Juncus effusus, Juncus inflexus, Lythrum salicaria, Phalaroides arundinacea, Scirpus sylvaticus, Symphytum officinale subsp. officinale, Valeriana officinalis.

It includes the most humid communities of lowland and floodplain wet and damp meadows of the Molinio-Arrhenatheretea class. These communities are confined to eutrophic and meso-eutrophic habitats in shallow flat depressions (slightly flowing or not flowing), along the edges of floodplain ponds and marshes in terraces and central parts of floodplains, as well as outside floodplains in the depressions of river terraces and watersheds, on the margins of terrace marshes. The landuse regime is mow-



ing or combined mowing and grazing, rarely grazing or abandoned. This alliance, like the previous one, has aver-age values of the Sharpness Index and a relatively high internal heterogeneity (see Table 1). The communities are dominated mostly by Scirpus sylvaticus and Carex cespi-

tosa. The species with the highest constancy are Ranun-culus repens and Potentilla anserina.

The proposed synopsis of the studied class in the present study is the following:

Molinio-Arrhenatheretea Tx. 1937 Galietalia veri Mirkin et Naumova 1986 (syn. Poo-Agrostietalia vinealis Shelyag-Sosonko et al. 1985 (nom. inval.))

Agrostion vinealis Sypailova et al. 1985Arrhenatheretalia Pawłowski et al. 1928

Arrhenatherion elatioris Luquet 1926 (syn. Festucion pratensis Sipailova et al. 1985)Cynosurion cristati Tx. 1947

Molinietalia Koch 1926 (incl. Deschampsietalia caespitosae Horvatić 1958, Potentillo-Polygonetalia Tx. 1947)Deschampsion cespitosae Horvatić 1930 (syn. Alopecurion pratensis Passarge 1964)Molinion caeruleae Koch 1926 Potentillion anserinae Tx. 1947 Calthion palustris Tx. 1937 (incl. Filipendulion ulmariae Segal 1966, Veronico longifoliae-Lysimachion vulgaris (Passarge 1977) Bal.-Tul. 1981)

Ordination

According to the results of the DCA-ordination (Fig. 3) the first ordination axis correlates strongly with moisture vector, along which the identified alliances are arranged from the driest Agrostion vinealis to the wettest Calthion palustris. The vector of carbonate content in the soil is also close to the first axis and is directed toward the op-posite of moisture vector. Thus, the right part of the or-dination diagram contains the xero-mesophilous alliance Agrostion vinealis and mesophilous alliances Arrhenathe-rion and Cynosurion cristati. The left-hand section con-tains the alliances of wet meadows Potentillion anserinae and Calthion palustris, and an intermediate position in the center of the diagram is occupied by the fresh meadow alliances Deschampsion and Molinion. The second ordi-nation axis is strongly correlated with the vector of soil salt regime and less strongly with the vectors of soil acid-ity, continentality and thermoregime. According to this distribution, the lower area of the diagram contains the alliances Potentillion anserinae, Cynosurion cristati and less Deschampsion, because their communities undergo intensive, mostly pasture use. The upper area contains the alliances Molinion, Arrhenatherion and less Calthion, which are used less intensively and primarily as hayfields. The Agrostion vinealis alliance occupies an intermediate position with its centroid set right on the second ordina-tion axis. Other factors play a smaller role in differentiat-ing the syntaxa. Thus, the vector of nitrogen content in the soil separates the Potentillion anserinae alliance, which is linked to the management regime mentioned above. The Molinion and Arrhenatherion alliances are separated by a factor of cryoregime, the Molinion and Calthion by a factor of ombroregime due to the distribu-

tion of majority of these alliances communities in the north of the study area within the humid zone.

Discussion

Using the European Expert System resulted in about 20% of the relevés previously assigned to the Molinio-Arrhenatheretea class being excluded from the analysis, and assigned to 12 other classes. At the same time, 93% of these relevés were assimilated to just four classes. It is therefore logical that a large number of relevés of wet meadows, previously included mainly to the Calthion al-liance were classified by the expert system within Phrag-mito-Magno-Caricetea class, since the boundaries be-tween classes are rather blurred. The other three classes, Artemisietea vulgaris, Festuco-Brometea and Koelerio-Corynephoretea included mainly the relevés previously contained within the Galietalia veri order. The order originally assigned to the Trifolion montani alliance came into Festuco-Brometea, those ordered to the Agrostion vinealis came in the Koelerio-Corynephoretea and trans-formed communities of both alliances came in the Arte-misietea vulgaris. This situation once again proved to be a debatable syntaxonomy of the “dry wing” of the Mo-linio-Arrhenatheretea class. This problem has been dis-cussed repeatedly in the literature (Kuzemko & Dziuba 2002; Kuzemko 2009a, 2009b, 2012a; Kuzemko et al. 2014). These communities are almost absent in Europe, with the results that they have no analogues among clas-sical Central European syntaxonomic units. There is also no consensus on the syntaxonomy of such communities among phytocenologists from Eastern Europe. Most re-searchers include them in the various alliances of the


250 Anna A. Kuzemko

Galietalia veri Mirkin et Naumova 1986 order (Kuzemko 2009b; Bulokhov 2001; Ermakov 2012; Yamalov et al. 2012; etc.). Recent publications have suggested distribut-ing these communities between classes Koelerio-Coryne-phoretea, Festuco-Brometea and to a lesser extent Mo-linio-Arrhenatheretea and Trifolio-Geranietea (Kuzemko et al. 2014). In the present analysis, the use of the Euro-pean Expert System actually led to some of the relevés previously considered as a part of the the Galietalia veri order being excluded from the Molinio-Arrhenatheretea class and referred to the three abovementioned classes of dry grasslands. However, many of the relevés still remain in the Molinio-Arrhenatheretea class. These relevés form a separate cluster, identified as Agrostion vinealis alliance with a distinct diagnostic block of species, as evidenced by the high value of the Sharpness Index (the value of this index for this alliance is second only to that of the Molin-ion caeruleae, which characterizes it as a “good” unit) and no signs of internal differentiation, sufficient for identification within a cluster of several units of the alli-ance rank, as evidenced by the low Whittaker beta-diver-

sity Index. Thus, the Agrostion vinealis occupies an inter-mediate position between the Molinio-Arrhenatheretea class and other classes of dry grasslands. In the present paper, an affiliation of the syntaxa to the Molinio-Ar-rhenatheretea class was confirmed by the analysis using the European Expert System, but nevertheless the alli-ance was clearly separated by a set of diagnostic species. However, it was somewhat surprising that, based on the results, it was not possible to distinguish another xe-romesophilous alliance, Trifolion montani Naumova 1986, which was distinguished earlier using other meth-ods (Kuzemko 2009a, 2009b, 2012a). The differentiation of these alliances was not observed in the variants of ana-lysis with larger numbers of clusters with the conse-quence that they only made the interpretation of the units at the level of alliances more difficult. Obviously, the communities that were previously considered as a part of this alliance, were assigned by the expert systems to the Festuco-Brometea class.

Another problem was the differentiation of the “dry wing” of the Molinio-Arrhenatheretea class at the level of

Fig. 3. DCA-ordination of the dataset with centroids of particular clusters. Number of the centroids corresponds to the num-bers of clusters in the Table 1. Ecological vectors: Hd – soil water regime, Rc – soil acidity, Sl – total salt regime, Ca – carbon-ate content in soil, Nt – nitrogen content in soil, Tm – thermoregime, Om – humidity of climate (ombroregime), Kn – continen-tality of climate, Cr – cryoregime and Lc – light; alliances 1 – Agrostion vinealis, 2 – Arrhenatherion elatioris, 3 – Cynosurion cristati, 4 – Deschampsion caespitosae, 5 – Molinion caeruleae, 6 – Potentillion anserinae, 7 – Calthion palustris. Eigenvalues: 1st axis (DCA1) 0.4278, 2nd axis (DCA2) 0.2547.



orders. In the original publication, the Agrostion vinealis alliance was included to the Arrhenatheretalia order (Sypailova et al. 1985), and later assigned to the newly described order Poo-Agrostietalia vinealis (Shelyag-So-sonko et al. 1985). Only one recently published synthesis on the vegetation of Europe (Rodwell et al. 2002) places the Agrostion vinealis alliance within the Poo-Agrostieta-lia vinealis order, which in this publication is, however, not included into Molinio-Arrhenatheretea class, but into the Festuco-Brometea class. Moreover, in that publi-cation it is stated, probably erroneously, that communi-ties of the Agrostion vinealis alliance are formed on leached chernozem-like soils, whereas in fact they are confined to light sod and meadow-sod soils. Earlier it was proved that the Poo-Agrostietalia vinealis and Gali-etalia veri orders are synonymous (Kuzemko & Dzyuba 2002; Kuzemko 2009b), the first of them is not valid ac-cording to the current edition of ICPN (Weber et al. 2000). The assignment of the Agrostion vinealis alliance to the Arrhenatheretalia order is not appropriate because, unlike typical mesophytic communities that are tradi-tionally assigned to this order, it includes xero-mesophy-lous communities. In addition, the author adheres to the class concept Molinio-Arrhenatheretea, which provides differentiation in the “dry” (Galietalia veri order), “mes-ophytic” (Arrhenatheretalia order) and “wet” (Molini-etalia order) part, and not just on mesophytic and wet parts as in the traditional Central-European syntaxon-omy.

The syntaxonomy of mesic meadows is less controver-sial, but the status of the Festucion pratensis alliance and its relationship with the Arrhenatherion alliance never-theless remains unclear. These two alliances have been traditionally considered as vicariants (Sypailova et al. 1985; Kuzemko 2009b; Birzniece et al. 2011). However, the results of the analysis did not show sufficient separa-tion of the alliance. The reason for this can be seen in the features of the original diagnosis of the alliance, which includes species with wide ecological amplitude such as Festuca pratensis, F. rubra, Poa pratensis, Phleum prat-ense, Trifolium pratense. All these species are present in the Arrhenatheretalia order communities actually within the entire range from the Iberian Peninsula (Rodriquez-Rojo et al. 2014) to Bashkortostan (Yamalov 2005). The name Festucion pratensis obviously needs to be removed in accordance with Article 24 of the ICPN (Weber et al. 2000) and classified as synonym of the Arrhenatherion elatioris alliance, since the association Festucetum praten-sis Soó 1938 that was chosen as nomenclature type of the Festucion pratensis alliance (Sypailova et al. 1985) belongs to the Arrhenatherion elatioris alliance in the original publication (Soó 1939). When comparing the complex of diagnostic species of this alliance in Ukraine with the di-agnostic species of the alliance in Central Europe, par-ticularly Czech Republic (Hájková et al. 2007), Poland (Kącki et al. 2013), Slovakia (Hájková et al. 2014) it is

clear that many of the species that are diagnostic for the alliance in Central Europe (Arrhenatherum elatius, Cam-panula patula, Leucanthemum vulgare, Knautia arvensis, etc.), have a relatively high fidelity in this cluster but not enough to consider them as diagnostic in Ukraine. These differences between Ukrainian and Central European communities resulted in the description of the Festucion pratensis alliance. However, the present analysis did re-vealed no differentiation within this vegetation unit that justifies considering these two alliances separately. The cluster that includes relevés assigned to the Arrhenathe-rion alliance has the greatest internal heterogeneity com-pared with other units (see Table 1), which indicates the need to confirm the results obtained by the analysis using data from wider area. It is notable that most of the relevés of transformed pastures previously identified within the Festucion pratensis alliance were included in the present analysis in the Cynosurion cristati alliance. This alliance has the lowest number of diagnostic species, and the low-est Sharpness Index among all analyzed alliances of the class. This is understandable when considering the exces-sive transformation of plant communities due to grazing that leads to a leveling of environmental conditions and loss of specialist species, to be replaced by generalists.

Taking into account the largest number of described alliances, the syntaxonomy of the “wet” part of the Mo-linio-Arrhenatheretea class, is difficult, however, as all of them were described in Central Europe, they are there-fore less controversial compared to previous units. Some inconsistency is observed on the status of certain alliances of damp and wet meadows. Hence, for the countries of Central Europe the Deschampsion alliance is traditional (Päzolt & Jansen 2004; Hájková et al. 2007, 2014), and its diagnosis almost overlaps with the alliance Alopecurion pratensis, which is traditional for East European phyto-cenology (Bulokhov 2001; Grigor’ev et al. 2002; Kuzemko 2009b; Yamalov et al. 2012 etc.). Most Euro-pean authors consider the Alopecurion pratensis alliance together with Agrostion albae Soó 1941, Cnidion venosi Bal.-Tul. 1965 and Veronico longifoliae-Lysimachion vulgaris (Passarge 1977) Bal.-Tul. 1981 alliances as syn-onymous with the Deschampsion alliance (Botta-Dukát et al. 2005; Hájková et al. 2007). However, this rule has many exceptions, e.g. in the last survey of Polish vegeta-tion (Kącki et al. 2013) such communities are considered in the Cnidion venosi alliance. Instead, in the “Prodromus of higher vegetation units of Russia” (Ermakov 2012), the syntaxonomical status of the Deschampsion alliance is consistent with the Central European status, and the Alo-pecurion pratensis alliance is considered a synonym. Thus, according to the results of the present analysis, these communities from Ukraine should also be included in the Deschampsion alliance, especially as this solution has already been established for Central Europe using statistical methods (Botta-Dukát et al. 2005). However, in many sources from Eastern Europe the Deschampsion


252 Anna A. Kuzemko

alliance is named, but with a description that is much more consistent with the Central European Potentillion anserinae alliance. Despite the fact that the communities of this alliance have not been previously mentioned for the territory of Ukraine, as far as the author knows, there is reason to suggest that these communities of wet mead-ows with pasture use should be regarded as part of the Potentillion anserinae alliance.

One of the most clearly defined alliances of meadow vegetation is Molinion caeruleae, which in the present analysis was characterized by the highest Sharpness In-dex and the highest internal homogeneity. However, this alliance proved also to have the lowest number of relevés. The reason for this could be that a necessary condition for the formation of the communities of the Molinion al-liance is late mowing of herbage for the collection of litter for animal bedding (Poschlod et al. 2009). However, this practice is now very rare in Ukraine and only found in some western regions. In the rest of the area these com-munities are rapidly reducing in area due to increasing land use intensity, especially by grazing, or are otherwise abandoned and overgrown with trees and shrubs. An-other issue not yet sufficiently investigated in the syntax-onomy of wet meadows, is the feasibility of differentia-tion of their “wettest” wing. In some sources, especially from Poland, a proportion of such communities with a predominance of tall grasses was assigned to the alliance Filipendulion ulmariae Segal 1966 (Matuszkiewicz 2008; Kuzemko 2009b) or in the Veronico longifoliae-Lysima-chion vulgaris alliance (Kącki et al. 2013). However, the results of the present analysis did not demonstrate such differentiation within this unit, so that the wettest com-munities of the Molinio-Arrhenatheretea class were placed within the Calthion alliance (sensu lato).

There are also certain disagreements regarding the as-signment of alliances of damp and wet meadows to the orders. In most of the analyzed sources, all wet meadows belong to the same order Molinietalia. However, some publications separately considerd the Deschampsietalia order (Sypailova & Shelyag-Sosonko 1996; Päzolt & Jansen 2004), while others included the Potentillo-Polygonetalia Tx. 1947 order (Ermakov 2012; Kącki et al. 2013). As already mentioned, the concept that I adopted, namely that the Molinio-Arrhenatheretea class contains three orders that are differentiated by the degree of mois-ture, does not provide differentiation of orders by man-agement or transformation, which is the main criterion for differentiation at the alliance level.

Acknowledgements

The author is grateful to Jürgen Dengler for an idea of the paper and inspiring its preparation, as well as for Monika Janišová and Iveta Škodová for the discussion of controver-sial syntaxonomic issues and improving the skills of the au-

thor in the JUICE program during the research stay in Slo-vakia with the support of IAVS travel grant. Also, the author is deeply grateful to Sergei Yamalov and two anonymous referees, as well as Monika Janišová as the Co-ordinating Editor and Pavel Krestov as the Receiving Editor for valua-ble comments on the manuscript. Also, many thanks to Laura Sutcliffe and Andrew N. Gillison for linguistic editing of the text.

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Supplement S1: A list of species aggregates.Supplement S2: A results of the “crispness of classification” analysis.Supplement S3: A results of the Sharpness Index and Whittaker beta-diversity Index calculation for the results of different methods of cluster analysis and differen number of clusters (mean ± SD).Supplement S4: Synoptic table.

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Supporting Information to the paper Kuzemko, A.A. Classification of the class Molinio-

Arrhenatheretea in the forest and forest-steppe zones of Ukraine

Supplement S1. A results of the Sharpness Index and Whittaker beta-diversity Index calculation for

the results of different methods of cluster analysis and differen number of clusters (mean ± SD)

Method Sharpness Whittaker beta-

of cluster analysis Index diversity index

Sorensen/Flexible beta -25/non-transf 6 clusters 13.73±11.870 15.83±3.118

Sorensen/Flexible beta -25/non-transf 10 clusters 11.59±8.493 12.81±2.287

Sorensen/Flexible beta -25/log 6 clusters 13.64±9.752 15.49±5.059

Rel Sorensen/Flexible beta -25/non-transf 8 clusters 12.73±12.685 13.82±4.577

Rel Sorensen/Flexible beta -25/log 6 clusters 12.21±10.941 15.97±3.811

Rel Sorensen/Flexible beta -25/log 8 clusters 12.73±8.313 14.24±2.897

Euclidean/Ward/non-transf 7 clusters 13.45±15.326 14.57±4.977

Euclidean/Ward/non-transf 12 clusters 10.45±10.255 11.59±4.725

Euclidean/Ward/log 7 clusters 12.72±13.860 14.93±4.858

Euclidean/Ward/log 10 clusters 13.06±11.831 12.59±4.880

TWINSPAN mod Sorensen 9 clusters 21.53±17.707 11.44±6.610

TWINSPAN mod Sorensen 12 clusters 20.71±13.653 10.38±5.563

TWINSPAN mod Simpson 10 clusters 22.37±16.797 10.55±6.697

TWINSPAN mod Jaccard 9 clusters 21.53±17.707 11.44±6.610

TWINSPAN mod Jaccard 12 clusters 20.71±13.653 10.38±5.563

TWINSPAN mod Whittaker 10 clusters 13.20±12.796 12.52±2.939

K-means 6 clusters 18.78±4.866 16.36±2.879K-means 6 clusters 18.78±4.866 16.36±2.879

K-means 7 clusters 17.24±7.120 15.31±3.036

K-means 8 clusters 15.40±9.159 14.57±2.555

K-means 9 clusters 16.53±9.263 13.83±2.455

K-means 10 clusters 14.25±8.337 13.33±2.885

PAM 6 clusters 19.46±8.167 17.82±2.896

PAM 7 clusters 17.29±7.589 16.82±3.042

PAM 8 clusters 14.60±8.984 15.84±2.915

PAM 9 clusters 15.90±8.305 15.18±3.084

PAM 10 clusters 15.33±8.370 14.47±2.582

Supporting Information to the paper Kuzemko, A.A. Classification of the class Molinio-Arrhenatheretea in the

forest and forest-steppe zones of Ukraine

Supplement S4 Synoptic table with modified fidelity phi coefficient and percentage frequency (7 columns) - Fisher's exact test > 3 Group No. 1 2 3 4 5 6 7 No. of releves 158 207 253 120 91 165 203 Agrostis vinealis 54.8 47 --- 6 --- 2 --- 8 --- . --- . --- . Poa angustifolia 53.2 51 --- 10 --- 2 --- 7 --- 2 --- 2 --- 1 Koeleria delavignei 50.1 48 --- 4 --- 1 --- 8 --- 13 --- . --- . Potentilla argentea 48.0 45 14.9 21 --- 2 --- 1 --- . --- 1 --- . Carex praecox 46.6 45 --- 5 --- 1 9.7 18 --- 2 --- 1 --- . Galium verum subsp. verum 44.2 51 6.3 19 --- 4 --- 8 --- 12 --- 1 --- 1 Stellaria graminea 41.2 44 --- 16 --- 8 --- 8 --- 4 --- 1 --- 1 Filipendula vulgaris 34.7 24 --- 4 --- 2 --- 1 --- 5 --- . --- 1 Festuca valesiaca subsp. valesiaca 31.7 17 --- 6 --- . --- . --- . --- . --- . Rumex thyrsiflorus 31.2 34 --- 5 --- 5 17.9 24 --- 3 --- 1 --- 2 Dianthus borbasii subsp. borbasii 30.7 13 --- 1 --- . --- . --- 1 --- . --- . Vicia tetrasperma 30.4 20 --- 7 --- 1 --- 2 --- . --- . --- 1 Lotus corniculatus subsp. corniculatus 29.7 56 --- 30 --- 30 --- 13 --- 27 --- 8 --- 5 Centaurea jacea subsp. jacea 27.1 49 --- 26 --- 17 --- 12 11.5 33 --- 7 --- 8 Elytrigia repens subsp. repens 22.5 38 --- 22 --- 17 --- 25 --- . --- 15 --- 4 Cichorium intybus 22.1 16 12.9 11 --- 3 --- 2 --- . --- . --- . Eryngium planum 21.9 9 --- 3 --- . --- . --- 1 --- . --- . Rumex acetosella subsp. acetosella 21.6 26 11.3 18 --- 8 --- 7 --- 5 --- 4 --- 2 Bromopsis inermis 20.6 9 --- 1 --- 1 --- 3 --- . --- 1 --- . Crepis tectorum 20.5 10 --- 1 --- 1 --- 2 --- 1 --- . --- 1 Euphorbia virgata 20.1 13 --- 3 --- 2 --- 7 --- 1 --- . --- 1 Dactylis glomerata subsp. glomerata --- 6 37.4 32 --- 8 --- . --- 4 --- 1 --- 3 Veronica chamaedrys subsp. chamaedrys --- 3 33.2 27 --- 8 --- 1 --- 3 --- 1 --- 3 Plantago lanceolata 17.3 55 31.0 71 12.8 50 --- 12 --- 37 --- 8 --- 10 Cerastium fontanum subsp. fontanum --- 20 29.0 42 8.6 24 --- 4 --- 8 --- 10 --- 4 Carex hirta --- 16 25.7 47 --- 30 --- 8 --- 7 --- 22 --- 20 Daucus carota subsp. carota --- 4 25.3 21 --- 9 --- 1 --- 5 --- 1 --- 2 Bromus hordeaceus subsp. hordeaceus --- 6 21.9 15 --- 5 --- . --- 1 --- 2 --- 1 Trifolium dubium --- 1 21.8 10 --- 2 --- 1 --- . --- 1 --- 1 Medicago lupulina --- 23 21.8 37 16.6 32 --- 7 --- 4 --- 12 --- 3 Carum carvi --- 4 20.8 15 --- 7 --- 1 --- . --- 2 --- 2 Galium mollugo --- 1 20.8 21 --- 11 --- 3 --- 7 --- 1 --- 8 Pimpinella saxifraga --- . 20.4 7 --- 1 --- . --- 1 --- . --- . Oenothera rubricaulis --- . 20.4 5 --- . --- . --- . --- . --- . Erigeron annuus subsp. annuus --- 5 20.4 15 --- 3 --- 3 --- 4 --- . --- 1 Taraxacum officinale --- 31 --- 23 40.0 64 --- 8 --- 5 --- 22 --- 6 Trifolium repens subsp. repens --- 20 --- 20 39.9 67 --- 17 --- 5 15.7 41 --- 2 Trifolium pratense subsp. pratense 14.1 46 10.0 42 30.3 64 --- 12 --- 19 --- 22 --- 8 Poa pratensis subsp. pratensis --- 38 --- 39 27.0 59 --- 16 --- 20 --- 13 --- 17 Festuca pratensis --- 47 10.5 48 25.2 66 --- 25 --- 30 --- 15 --- 22 Ranunculus acris subsp. acris --- 43 --- 41 23.6 68 --- 17 --- 45 --- 22 --- 40 Lolium perenne --- . --- 6 22.9 13 --- . --- . --- 3 --- 1 Leontodon autumnalis --- 20 --- 17 22.1 43 12.8 34 --- 15 --- 18 --- 1 Achillea pyrenaica --- 5 --- . --- 5 51.9 47 --- 2 --- 6 --- 1 Poa palustris --- 8 --- 1 --- 14 45.5 68 --- 13 --- 22 --- 26 Carex vulpina --- 6 --- 2 --- 5 43.6 46 --- 4 --- 5 --- 13 Beckmannia eruciformis --- 1 --- . --- 1 42.3 26 --- . --- 2 --- 1 Gratiola officinalis 4.0 13 --- 1 --- 1 40.4 39 --- 10 --- 3 --- 1 Lythrum virgatum --- 3 --- 1 --- 1 33.8 22 --- . --- 4 --- 2 Alopecurus pratensis 19.8 46 --- 9 --- 22 32.8 60 --- 8 --- 7 --- 24 Ranunculus flammula --- 1 --- 1 --- 11 28.6 30 --- 5 --- 11 --- 7 Lysimachia nummularia --- 15 --- 4 --- 15 28.5 42 --- 3 --- 13 --- 21 Juncus atratus 9.1 13 --- 1 --- 4 27.3 25 --- 4 --- 4 --- . Agrostis gigantea subsp. gigantea --- 25 --- 13 7.8 26 25.9 43 --- 2 --- 8 --- 12 Cnidium dubium --- 4 --- 1 --- 3 24.4 24 19.5 21 --- 1 --- 2 Inula britannica --- 16 --- 2 --- 11 24.3 32 --- 4 7.4 19 --- 3 Agrostis canina subsp. canina --- 2 --- . --- 3 21.8 18 10.9 12 --- 4 --- 1 Ranunculus repens --- 20 --- 17 12.2 58 21.7 69 --- 12 17.1 64 14.6 61 Myosotis cespitosa --- 2 --- . --- . 21.3 9 --- 1 --- . --- 1 Veronica longifolia --- 4 --- 1 --- 2 20.7 18 --- 8 --- 3 --- 5 Potentilla erecta --- 9 --- 3 --- 6 --- 8 72.4 80 --- 1 --- 2 Molinia caerulea --- 1 --- 1 --- 1 --- 2 68.5 57 --- 1 --- 2 Succisa pratensis --- . --- 1 --- 1 --- . 44.7 26 --- 1 --- 1 Carex panicea --- 8 --- 9 --- 14 --- 2 43.4 54 --- 12 --- 8 Briza media --- 2 8.3 14 --- 2 --- . 37.5 33 --- 2 --- 3 Carex flava subsp. flava --- 1 --- 1 --- 5 --- 2 35.4 30 --- 6 --- 6 Galium uliginosum --- . --- 1 --- 5 --- 2 31.4 24 --- 2 --- 7 Schoenus ferrugineus --- . --- . --- . --- . 30.9 11 --- . --- . Sanguisorba officinalis --- 3 --- 4 --- 1 --- 2 30.8 22 --- . --- 5 Linum catharticum subsp. catharticum --- 1 --- . --- 2 --- . 30.6 16 --- 3 --- 1 Salix rosmarinifolia --- . --- 1 --- . --- 1 29.6 12 --- . --- 1 Carex davalliana --- . --- . --- 1 --- . 28.8 12 --- 1 --- 1 Serratula tinctoria subsp. tinctoria --- . --- 1 --- . --- . 26.7 9 --- . --- . Valeriana simplicifolia --- . --- . --- 1 --- . 26.4 11 --- . --- 2 Gentiana pneumonanthe --- 1 --- . --- . --- 1 25.8 10 --- 1 --- . Holcus lanatus --- 1 10.9 15 --- 4 --- . 24.9 24 --- 1 --- 10 Lysimachia vulgaris --- 1 --- 3 --- 6 7.6 22 24.3 37 --- 7 18.3 32 Luzula multiflora subsp. multiflora --- 4 --- 4 --- 1 --- 1 24.2 14 --- 1 --- . Stachys officinalis subsp. officinalis --- 1 --- 1 --- . --- . 23.6 8 --- . --- . Parnassia palustris --- . --- . --- 1 --- . 22.6 10 --- 3 --- 1 Genista tinctoria --- 3 --- 1 --- . --- 1 22.6 10 --- . --- . Polygala amarella --- . --- 1 --- 1 --- . 22.4 8 --- . --- 1 Selinum carvifolia --- . --- . --- 1 --- 1 22.3 10 --- . --- 3 Carex pallescens --- 1 9.2 7 --- . --- . 22.1 12 --- . --- 1

Trifolium montanum subsp. montanum 6.6 4 --- 1 --- . --- . 22.1 10 --- . --- . Epipactis palustris --- . --- . --- . --- . 21.8 5 --- . --- . Salix cinerea --- . --- 1 --- . --- 2 21.6 14 --- 5 5.1 6 Filipendula ulmaria --- 4 --- 4 --- 10 7.3 22 20.1 34 --- 4 17.9 32 Agrostis stolonifera subsp. stolonifera --- 1 --- 2 --- 24 --- 14 --- 5 58.9 75 --- 8 Trifolium fragiferum subsp. fragiferum --- 3 --- 1 11.4 13 --- . --- . 32.3 25 --- . Bidens tripartitus subsp. tripartitus --- 1 --- . --- 3 --- 2 --- . 31.8 19 --- 3 Alopecurus geniculatus --- 1 --- 1 --- 4 --- 2 --- . 30.1 19 --- 1 Glyceria fluitans --- 1 --- 1 --- 2 --- 8 --- . 28.5 21 --- 4 Potentilla anserina subsp. anserina --- 18 --- 27 19.5 74 --- 58 --- 41 26.9 83 --- 51 Plantago major subsp. major --- 15 --- 11 14.7 27 --- 4 --- 3 26.6 38 --- 4 Triglochin palustris --- . --- . --- 4 --- . --- . 23.6 11 --- 1 Persicaria maculosa --- 1 --- 1 --- 2 --- 3 --- 2 22.8 14 --- 2 Eleocharis palustris subsp. palustris --- 2 --- 1 --- 4 9.2 12 --- 3 22.3 19 --- 2 Mentha arvensis --- 6 --- 7 --- 23 17.7 45 --- 27 21.5 49 --- 24 Schoenoplectus tabernaemontani --- . --- . --- . --- . --- . 20.7 6 --- 1 Rorippa amphibia --- . --- . --- . --- 6 --- . 20.1 12 --- 4 Symphytum officinale subsp. officinale --- 2 --- 2 --- 4 --- 12 --- 1 --- 4 38.6 35 Caltha palustris --- 1 --- 1 --- 9 --- 11 --- 5 10.4 22 38.2 46 Scirpus sylvaticus --- . --- . --- 4 --- 1 --- 2 --- 6 35.1 25 Lythrum salicaria --- 2 --- 3 --- 6 --- 7 12.9 26 --- 21 29.5 41 Phalaroides arundinacea --- 1 --- 1 --- 2 --- 11 --- 1 --- 3 27.9 22 Equisetum palustre --- 1 --- 2 --- 8 --- 2 --- 10 15.0 22 26.8 32 Juncus effusus --- 1 --- 6 --- 3 --- 7 --- 8 --- 7 26.7 26 Carex acutiformis --- . --- . --- 2 --- 1 --- 3 --- 1 26.0 13 Calystegia sepium --- . --- 3 --- 1 --- 2 --- 1 --- 5 25.9 16 Iris pseudacorus --- . --- . --- 4 --- 7 --- 7 --- 7 24.2 21 Cirsium rivulare --- . --- 1 --- 6 --- . 16.3 18 --- 3 23.6 22 Juncus inflexus --- . --- 1 --- 3 --- . --- . --- 5 23.6 13 Valeriana officinalis --- 2 --- 1 --- 2 --- 7 --- 12 --- 1 22.4 20 Galium palustre --- 1 --- 2 --- 6 16.4 29 --- 10 7.9 22 21.9 34 Achillea millefolium subsp. millefolium 30.2 73 41.2 86 --- 46 --- 5 --- 34 --- 10 --- 8 Allium angulosum 22.0 22 --- 1 --- 1 28.8 26 --- 2 --- . --- . Anthoxanthum odoratum --- 3 24.8 29 --- 6 --- . 28.7 32 --- . --- 3 Persicaria amphibia var. amphibia --- . --- . --- 2 --- 2 --- . --- 6 18.8 11 Angelica sylvestris subsp. sylvestris --- 1 --- 2 --- 1 --- . --- 5 --- . 18.5 10 Galium aparine --- 2 --- 2 --- 4 --- 1 --- 1 --- 4 18.3 12 Carex acuta --- 1 --- . --- 9 9.6 18 --- 2 14.2 22 17.2 24 Typha latifolia --- . --- . --- 1 --- . --- . --- 1 17.0 5 Cirsium palustre --- . --- . --- 1 --- . 13.5 8 --- 1 16.6 9 Cirsium oleraceum --- . --- . --- 1 --- . --- 2 --- . 16.3 6 Epilobium palustre --- . --- . --- 2 --- 1 --- 5 11.3 10 16.2 13 Scrophularia umbrosa subsp. umbrosa --- . --- . --- 1 --- . --- . --- . 16.0 3 Glyceria maxima --- . --- . --- 1 --- 8 --- . --- 7 15.1 11 Urtica dioica subsp. dioica --- . --- 5 --- 2 --- 2 --- 1 --- 5 15.1 11 Lycopus europaeus subsp. europaeus --- 1 --- 2 --- 5 --- . 11.6 16 12.3 17 14.8 19 Rumex hydrolapathum --- . --- 1 --- 1 --- 4 --- . --- 4 14.3 8 Lathyrus pratensis subsp. pratensis --- 3 --- 9 --- 8 --- 10 --- 10 --- 2 14.3 18 Phragmites australis --- . --- 1 --- 6 --- 3 15.4 18 --- 9 14.1 17 Lathyrus palustris --- . --- . --- 1 11.2 7 --- . --- 2 13.1 7 Carex rostrata --- . --- . --- 1 --- 1 --- 1 --- 2 13.1 5 Mentha longifolia --- 1 --- 3 --- 4 --- . --- 1 --- 2 13.1 8 Succisella inflexa --- . --- . --- . --- . --- . --- . 13.0 2 Cirsium canum --- 2 --- 2 --- 1 --- 1 --- 1 --- . 13.0 6 Equisetum fluviatile --- . --- . --- 2 --- . --- 3 --- 5 12.6 7 Myosotis palustris --- 1 --- 3 --- 8 8.6 21 --- 12 14.9 26 12.0 24 Geum rivale --- . --- 2 --- 7 --- 1 17.3 15 --- 2 11.8 12 Carex vesicaria --- 1 --- 1 --- 3 --- 6 --- 7 --- 7 11.7 11 Eupatorium cannabinum --- . --- 1 --- 1 --- 1 --- 9 --- 7 11.5 9 Geranium palustre --- 1 --- . --- 1 --- . --- 2 --- 5 11.4 6 Omphalodes scorpioides --- . --- . --- . --- . --- . --- . 11.3 1 Berula erecta --- . --- . --- . --- . --- . --- . 11.3 1 Dactylorhiza incarnata subsp. incarnata --- 3 --- 5 --- 2 --- 2 --- 4 --- . 10.5 8 Achillea salicifolia --- 1 --- 1 --- 2 --- 5 --- . --- 1 10.4 6 Scutellaria galericulata --- 2 --- . --- 2 --- 2 --- 4 --- 4 10.0 7 Mentha aquatica --- . --- 1 --- . --- 1 --- 4 --- 3 9.8 5 Carex nigra --- 3 --- 6 --- 20 --- 3 --- 22 17.5 32 9.8 25 Carex cespitosa --- 1 --- 2 --- 2 --- 8 --- 9 --- 7 9.7 11 Cardamine pratensis subsp. pratensis --- . --- 1 --- 4 --- 3 --- . --- 2 9.6 6 Deschampsia cespitosa subsp. cespitosa --- 9 --- 20 12.8 40 --- 12 14.9 42 --- 21 9.5 36 Stachys palustris --- . --- 1 --- . --- 4 --- 1 --- 1 9.1 4 Sonchus palustris --- . --- 1 --- 1 --- . --- . --- 4 8.9 3 Dactylorhiza majalis subsp. majalis --- 1 --- 1 --- 2 --- 2 --- 3 --- 1 8.3 5 Galium rivulare --- . --- 1 --- . --- . --- 2 --- . 8.3 2 Stellaria palustris --- 2 --- . --- 5 12.0 11 --- 2 --- 4 8.2 9 Juncus conglomeratus --- . --- 1 --- 1 --- 2 18.9 11 --- . 8.0 6 Alisma plantago-aquatica --- . --- 1 --- . --- 6 --- . 11.4 7 6.8 5 Carex caryophyllea --- . --- 1 --- 1 --- . --- . --- . --- . Rubus caesius --- 1 --- 1 --- 1 --- 1 --- 3 --- . --- 2 Carex leporina 14.5 32 --- 21 --- 20 --- 24 --- 16 --- 5 --- 8 Galium album subsp. album --- 1 --- . --- . --- . --- 2 --- . --- . Salvia nemorosa subsp. nemorosa --- . --- 1 --- . --- . --- . --- . --- . Armoracia rusticana --- . --- . --- 1 --- . --- . --- . --- . Vicia tenuifolia subsp. tenuifolia --- . --- 1 --- . --- . --- . --- 1 --- 1 Festuca rupicola --- . --- 1 --- . --- . --- . --- . --- . Euphorbia palustris --- 1 --- . --- . --- 2 --- . --- . --- 2 Carex juncella --- . --- . --- . --- . 16.7 4 --- 1 --- 1 Viola palustris subsp. palustris --- 1 --- . --- . --- . 17.8 4 --- . --- . Carex appropinquata --- . --- . --- 1 --- . 17.8 8 --- 2 --- 3 Plantago cornutii --- 1 --- . --- 1 --- . --- . --- 1 --- . Melampyrum nemorosum subsp. nemorosum --- . --- 1 --- . --- . --- 1 --- . --- . Veratrum lobelianum --- . --- . --- . --- . 15.3 3 --- . --- 1 Allium oleraceum --- 1 --- . --- 1 --- . 16.0 4 --- . --- . Stellaria media --- 1 --- 1 --- 3 --- . --- 1 --- 3 --- 1 Vaccinium oxycoccos --- . --- . --- . --- . --- 1 --- . --- . Carex serotina --- . --- . --- . --- . --- 1 --- . --- . Poa annua --- . --- 1 --- 2 --- . --- . --- . --- 1 Salix aurita --- . --- 1 --- . --- . 17.1 7 --- . --- 3 Achillea pannonica --- . --- . --- . --- . --- 1 --- . --- . Campanula patula subsp. patula --- 3 15.9 12 --- 2 --- . 12.3 10 --- . --- 1 Campanula rotundifolia subsp. rotundifol 14.5 7 --- 3 --- 1 --- . --- 3 --- . --- . Asparagus officinalis subsp. officinalis 14.0 3 --- . --- . --- 1 --- . --- . --- . Carex otrubae --- . --- . 9.4 3 --- . --- . --- 2 --- 1

Cirsium vulgare --- 1 --- 3 --- 4 --- . --- . --- 2 --- 1 Sagina procumbens --- 1 --- . --- . --- . --- . --- . --- . Epilobium tetragonum subsp. tetragonum --- . --- . --- 1 --- . --- . --- 1 --- . Lamium album --- . --- 1 --- 1 --- . --- . --- . --- . Limonium suffruticosum --- 1 --- . --- . --- . --- . --- . --- . Euphrasia rostkoviana subsp. rostkoviana --- 1 --- 1 8.4 4 --- 1 --- 4 --- . --- . Cerinthe minor subsp. minor --- 1 --- . --- . --- . --- . --- . --- . Thesium linophyllon --- 1 --- . --- . --- . --- . --- . --- . Juncus subnodulosus --- . --- . --- . --- . --- 1 --- . --- . Anthyllis vulneraria subsp. polyphylla --- 1 --- 1 --- 1 --- . --- 2 --- . --- . Helichrysum arenarium 13.1 3 --- 1 --- . --- . --- . --- . --- . Carex spicata --- 1 --- 2 --- 4 --- 1 --- 2 --- 1 --- 2 Carex limosa --- . --- . --- . --- . --- . --- . --- 1 Verbascum thapsus subsp. thapsus --- . --- 1 --- 1 --- . --- . --- . --- . Heracleum sibiricum --- 3 --- 4 8.5 6 --- 1 --- 1 --- . --- 3 Ononis arvensis --- 2 --- 2 --- 1 --- . --- 4 --- 1 --- 1 Carex distans --- 1 --- 2 7.1 5 --- . --- 5 --- 2 --- 1 Falcaria vulgaris --- . --- 1 9.7 2 --- . --- . --- . --- . Euphorbia lucida --- . --- . --- . --- . --- . --- 1 --- . Scorzonera cana --- 1 --- . --- . --- . --- . --- 1 --- . Crepis biennis --- 1 --- 1 --- . --- . --- 1 --- . --- . Rorippa austriaca --- 1 --- 1 --- . --- 2 --- . --- 1 --- . Veronica serpyllifolia subsp. serpyllifo --- 1 --- . 9.2 2 --- . --- . --- . --- . Alliaria petiolata --- . --- . --- 1 --- . --- . --- . --- . Galium boreale 14.7 25 --- 6 --- 6 19.2 28 11.4 22 --- 1 --- 2 Cerastium sylvaticum --- . --- . --- . --- . --- . --- . --- 1 Carex hostiana --- . --- 1 --- . --- . --- . --- . --- . Tanacetum vulgare subsp. vulgare 7.2 3 --- 1 --- . --- . 11.8 4 --- . --- 1 Eryngium campestre --- 1 --- . --- . --- . --- . --- . --- . Hieracium umbellatum 14.7 15 --- 3 --- 1 8.8 12 9.5 12 --- 1 --- 1 Leontodon hispidus --- . --- 2 --- 1 --- . 12.1 4 --- 1 --- 1 Myosotis stricta 15.9 5 --- 1 --- . --- 1 --- . --- 1 --- . Tragopogon orientalis --- . --- 1 --- . --- . --- . --- . --- . Trifolium hybridum subsp. hybridum 19.0 24 --- 5 --- 11 --- 12 --- 10 --- 4 --- 4 Phleum pratense subsp. pratense 10.1 26 --- 21 12.0 28 --- 12 --- 11 --- 13 --- 7 Carex canescens --- . --- . 10.9 3 --- . --- . --- 1 --- 1 Bellis perennis --- . --- 5 10.6 6 --- . --- 1 --- 2 --- 1 Medicago falcata subsp. romanica --- 3 12.0 9 8.4 7 --- . --- . --- 4 --- 1 Prunella vulgaris 9.4 37 --- 29 18.3 47 --- 25 --- 29 --- 15 --- 8 Convolvulus arvensis 15.0 9 --- 5 --- 2 --- 3 --- . --- 1 --- 2 Trifolium alpestre 9.2 2 --- . --- . --- . --- 1 --- . --- . Trifolium aureum --- 2 --- 1 --- 1 --- . --- . --- 1 --- . Thymus pulegioides subsp. pulegioides --- . --- 1 --- 1 --- . --- . --- . --- 1 Tragopogon ucrainicus --- 1 --- . --- . --- . --- . --- . --- . Alchemilla micans --- . --- 1 --- 1 --- . --- 1 --- . --- 1 Festuca gigantea --- . --- 1 --- . --- . --- . --- 1 --- 1 Centaurium erythraea subsp. erythraea --- 1 --- 2 --- 2 --- . --- 2 --- 4 --- . Carex diluta --- . --- . --- 1 --- . --- . --- 1 --- . Cardamine amara subsp. amara --- . --- . --- 1 --- . --- 1 --- 1 --- 1 Menyanthes trifoliata --- . --- . --- . --- . --- 3 --- 2 --- 2 Glechoma hederacea 11.9 24 --- 17 --- 14 --- 16 --- 2 --- 9 --- 15 Calamagrostis neglecta --- . --- . --- . --- 1 --- 1 --- . --- 1 Koeleria macrantha --- 1 --- . --- . --- . --- . --- . --- . Carex lasiocarpa --- . --- . --- . --- . --- 2 --- . --- 1 Calluna vulgaris --- . --- 1 --- . --- . --- 1 --- . --- . Myosotis ramosissima --- 1 --- 1 --- . --- 1 --- . --- . --- 1 Apera spica-venti --- 1 --- 1 --- . --- . --- . --- . --- . Lathyrus tuberosus --- 1 --- . --- . --- . --- . --- . --- . Silene nutans subsp. nutans --- . --- 1 --- . --- . --- . --- . --- . Allium senescens subsp. montanum --- 1 --- . --- . --- . --- . --- . --- . Galium ruthenicum --- . --- 1 --- . --- . --- . --- . --- . Anacamptis coriophora 15.8 6 --- 2 --- 1 --- . --- 1 --- . --- . Cerastium glomeratum --- . --- . --- . --- . --- . --- 1 --- . Pastinaca sativa subsp. pratensis --- . --- 1 --- 1 --- . --- . --- . --- . Herniaria glabra subsp. glabra 18.5 7 --- 2 --- 1 --- . --- . --- 1 --- . Campanula sibirica subsp. sibirica --- . --- . --- . --- . --- 1 --- . --- . Pedicularis sceptrum-carolinum --- . --- . --- . --- . 13.7 2 --- . --- . Campanula rapunculus --- 1 --- 1 --- . --- . --- . --- . --- . Trollius europaeus --- . --- . --- . --- . 13.7 2 --- . --- . Carex chordorrhiza --- . --- . --- . --- . 13.7 2 --- . --- . Comarum palustre --- . --- . --- 1 --- 2 13.1 8 --- 4 --- 4 Salvia verticillata --- . --- 1 --- . --- . --- . --- . --- . Trifolium medium subsp. medium --- . --- 1 --- . --- . 12.8 4 --- 2 --- 1 Bistorta officinalis --- 1 --- 2 --- 2 --- . 12.7 9 --- 4 --- 6 Salix repens --- . --- . --- . --- . 13.7 2 --- . --- . Campanula glomerata subsp. glomerata --- 1 --- 1 --- . --- 1 19.8 8 --- . --- 1 Eriophorum latifolium --- . --- . --- . --- . 19.5 5 --- . --- 1 Betula humilis --- . --- . --- . --- . 19.5 4 --- . --- . Thyselium palustre --- . --- 1 --- 1 --- 4 19.0 10 --- 1 --- 1 Carex buxbaumii --- 1 --- 1 --- . --- . 18.0 7 --- . --- 1 Carex flacca subsp. flacca --- . --- 2 --- 1 --- . 18.8 9 --- 1 --- 2 Danthonia decumbens subsp. decumbens --- 3 --- 4 --- 1 --- 1 19.0 10 --- . --- . Viola canina --- 2 --- 1 --- 1 --- 6 18.7 11 --- . --- 1 Artemisia marschalliana --- . --- 1 --- . --- . --- . --- . --- . Scorzonera humilis --- . --- 1 --- . --- . --- . --- . --- . Mentha pulegium --- . --- . --- . --- . --- . --- . --- 1 Centaurium pulchellum --- . --- . --- . --- . --- . --- 1 --- . Galeopsis speciosa --- . --- 1 --- 1 --- . --- . --- . --- 1 Hypericum montanum --- . --- 1 --- . --- . --- . --- . --- . Crepis capillaris --- . --- 1 --- . --- . --- . --- . --- . Ajuga reptans --- . --- 1 --- . --- . --- 1 --- . --- . Juncus bufonius --- 2 --- 1 --- 1 --- . --- . --- 2 --- . Euphrasia stricta --- 4 --- 1 --- 4 --- 4 14.8 10 --- 1 --- 1 Nardus stricta --- 1 14.9 9 --- 1 --- . 14.0 9 --- . --- 1 Agrostis capillaris 10.7 24 15.3 28 --- 12 --- 2 14.6 27 --- 6 --- 3 Thelypteris palustris --- . --- . --- 1 --- . 14.3 5 --- 1 --- 2 Iris sibirica --- 3 --- 1 --- 1 --- 1 13.8 5 --- . --- . Swertia perennis --- . --- . --- . --- . 13.7 2 --- . --- . Tofieldia calyculata --- . --- . --- . --- . 13.7 2 --- . --- . Ranunculus lingua --- . --- . --- . --- . 13.7 2 --- . --- . Descurainia sophia --- . --- 1 --- 1 --- . --- . --- . --- . Echium vulgare subsp. vulgare --- 2 --- 1 --- 1 --- . --- . --- . --- . Anthemis ruthenica --- 1 --- 1 --- . --- . --- . --- . --- .

Epipactis helleborine subsp. helleborine --- . --- 1 --- . --- . --- . --- . --- . Stachys recta subsp. recta --- . --- . --- 1 --- . --- . --- 1 --- . Eriophorum vaginatum --- . --- . --- . --- . --- 1 --- . --- . Myagrum perfoliatum --- . --- 1 --- . --- . --- . --- . --- . Veronica beccabunga --- . --- . --- 1 --- . --- . --- 1 --- 1 Pastinaca sylvestris --- . --- 1 --- 1 --- . --- . --- . --- . Dianthus trifasciculatus subsp. pseudoba --- 1 --- . --- . --- . --- . --- . --- . Poa trivialis subsp. sylvicola --- . --- . --- 1 --- . --- . --- 1 --- . Elytrigia intermedia subsp. intermedia --- 1 --- 1 --- . --- . --- . --- . --- 1 Nepeta cataria --- . --- . --- . --- . --- . --- 1 --- . Acinos arvensis --- . --- . --- . --- . --- . --- 1 --- . Bromopsis erecta subsp. erecta --- . --- 1 --- . --- . --- . --- . --- . Scleranthus annuus subsp. annuus 13.1 3 --- 1 --- . --- . --- . --- . --- . Butomus umbellatus --- . --- . --- . --- . --- . --- 1 --- 1 Plantago arenaria --- . --- . --- 1 --- . --- . --- . --- . Bromus arvensis subsp. arvensis --- 2 --- 2 --- 1 --- . --- . --- . --- . Sinapis arvensis --- . --- 1 --- . --- . --- . --- 1 --- . Peucedanum oreoselinum --- . --- . --- 1 --- . --- 2 --- 1 --- . Petasites spurius --- 1 --- . --- . --- . --- . --- . --- . Stellaria holostea --- 1 --- 1 --- . --- 1 --- . --- . --- . Artemisia abrotanum --- . --- . --- . --- . --- . --- 1 --- 1 Achillea inundata --- . --- . --- 1 --- . --- . --- 1 --- 1 Carex pilosa --- . --- 1 --- . --- . --- . --- . --- . Bromus scoparius --- . --- 1 --- . --- . --- . --- . --- . Lychnis flos-cuculi --- 27 --- 20 --- 27 --- 30 --- 29 --- 18 --- 33 Solidago canadensis --- 1 --- . --- . --- . --- 1 --- . --- . Myosoton aquaticum --- . --- 1 --- 1 --- . --- . --- 1 --- 2 Erodium cicutarium subsp. cicutarium --- . --- 1 --- . --- . --- . --- . --- . Stellaria crassifolia --- 1 --- . --- . --- . --- . --- . --- . Viola tricolor subsp. tricolor --- . --- 1 --- . --- . --- . --- 1 --- . Trifolium campestre 10.4 3 --- 1 --- . --- . --- 1 --- . --- . Gypsophila muralis 13.4 6 --- 1 --- . --- 2 --- 1 --- 1 --- . Trifolium arvense 15.9 5 --- 2 --- . --- . --- . --- . --- . Scirpus radicans --- . --- . --- . --- 1 --- . --- . --- 1 Pedicularis palustris --- . --- . --- . --- 1 --- . --- . --- . Hylotelephium telephium subsp. telephium 8.9 3 --- 1 --- . --- 1 --- 1 --- . --- . Veronica spicata subsp. spicata 13.3 8 --- 2 --- 1 --- 2 --- 4 --- 1 --- . Echinocystis lobata --- . --- . --- . --- . --- . --- . --- 1 Melampyrum cristatum --- 1 --- 1 --- . --- 1 --- 2 --- . --- . Spergula arvensis --- 1 --- 1 --- 1 --- . --- . --- 1 --- . Xanthium albinum subsp. albinum --- . --- 1 --- 1 --- . --- . --- . --- . Rumex ucranicus --- . --- . --- . --- . --- . --- 1 --- . Bunias orientalis --- . --- 1 --- 1 --- . --- . --- . --- 1 Inula salicina 8.7 4 --- 1 --- . --- . --- 3 --- 2 --- . Carex atherodes --- . --- . --- . --- 1 --- . --- . --- 1 Polygala comosa --- 2 --- 4 --- 2 --- . --- 3 --- 1 --- . Carex humilis --- 1 --- . --- . --- . --- . --- . --- . Verbena officinalis --- . --- 1 --- . --- . --- . --- . --- . Senecio jacobea --- 3 --- 2 --- 1 --- 1 --- . --- 1 --- . Atriplex rosea --- 1 --- . --- . --- . --- . --- . --- . Vicia angustifolia subsp. angustifolia --- 1 --- . --- . --- 1 --- . --- . --- . Arabidopsis thaliana --- . --- 1 --- . --- . --- . --- . --- . Mentha x verticillata --- . --- . --- . --- . --- . --- 1 --- . Galium verticillatum --- 1 --- 1 --- . --- . --- 1 --- . --- 1 Rhinanthus minor --- 8 9.6 11 10.9 12 --- 2 --- 3 --- 1 --- 1 Rorippa sylvestris --- 3 --- 3 13.8 11 --- 6 --- . --- 5 --- 2 Viola hirta --- . --- 1 --- . --- . --- . --- . --- . Rumex confertus 15.3 26 --- 10 --- 15 --- 20 --- . --- 12 --- 10 Fragaria vesca 9.2 2 --- . --- . --- . --- 1 --- . --- . Silene multiflora 14.8 3 --- . --- . --- . --- . --- . --- . Anagallis arvensis subsp. arvensis 9.2 2 --- . --- . --- . --- 1 --- . --- . Sedum sexangulare 10.9 2 --- 1 --- . --- . --- . --- . --- . Ficaria verna --- . --- . --- . --- . --- . --- . --- 1 Arctium tomentosum --- . --- 1 --- . --- . --- . --- . --- 1 Cardamine parviflora --- . --- . --- . --- . --- . --- . --- 1 Nepeta pannonica --- . --- . --- . --- . --- . --- . --- 1 Glyceria notata --- . --- 1 --- . --- . --- . --- 1 --- 1 Serratula lycopifolia --- . --- . --- . --- 1 --- . --- . --- . Silene viscosa --- 1 --- 1 --- . --- . --- . --- . --- . Geum urbanum --- 1 --- . --- 1 --- . --- . --- . --- . Carex secalina --- . --- . --- . --- . --- . --- 1 --- . Galium odoratum --- 2 --- . --- . --- 2 --- . --- . --- . Tussilago farfara --- . --- 1 --- 2 --- . --- 3 --- 1 --- 3 Arabis hirsuta --- . --- 1 --- . --- . --- 1 --- . --- . Jacobaea paludosa subsp. paludosa --- . --- . --- 1 --- . --- . --- 1 --- 1 Anthriscus sylvestris subsp. sylvestris --- . --- 1 --- 1 --- . --- . --- 1 --- 1 Senecio erucifolius --- . --- 1 --- 1 --- . --- . --- . --- . Conyza canadensis --- 4 --- 2 --- 1 --- . --- . --- 2 --- . Aster tripolium --- . --- . --- 1 --- . --- . --- 1 --- . Dactylorhiza maculata subsp. maculata --- 1 --- . --- 1 --- . --- 3 --- . --- 1 Elymus caninus --- . --- . --- 1 --- . --- . --- 1 --- . Eleocharis ovata --- . --- . --- 1 --- . --- . --- . --- . Centaurea phrygia subsp. phrygia --- . --- . --- . --- . --- 1 --- . --- . Crepis paludosa --- . --- . --- . --- . --- 1 --- . --- . Salvia officinalis --- . --- . --- . --- . --- 1 --- . --- . Tephroseris palustris --- . --- 1 --- . --- . --- . --- . --- 1 Limonium alutaceum --- 1 --- 1 --- . --- . --- . --- . --- . Iva xanthiifolia --- . --- . --- 1 --- . --- . --- . --- . Verbascum lychnitis --- . --- 1 --- . --- . --- . --- 1 --- . Veronica officinalis --- 1 --- 1 --- . --- . --- . --- . --- . Inula helenium --- . --- 1 --- 1 --- . --- 1 --- 1 --- 1 Cicuta virosa --- . --- . --- . --- 1 --- . --- 1 --- 1 Carex pseudocyperus --- . --- . --- . --- . --- . --- 1 --- . Fritillaria meleagroides --- 1 --- . --- 1 --- . --- . --- . --- . Odontites vernus --- . --- . --- 1 --- . --- . --- . --- . Althaea officinalis --- 2 --- 1 --- 1 --- 2 --- . --- . --- 1 Hypericum maculatum subsp. maculatum --- . --- . --- . --- . --- 1 --- . --- . Lathyrus sativus --- . --- . --- . --- . --- . --- 1 --- . Matricaria recutita --- . --- . --- 1 --- . --- . --- 1 --- . Holosteum umbellatum subsp. umbellatum --- . --- . --- 1 --- . --- . --- . --- . Vicia parviflora --- 1 --- 1 --- . --- . --- . --- 1 --- . Viola reichenbachiana --- . --- 1 --- 1 --- . --- 1 --- . --- . Potentilla norvegica --- . --- . --- 1 --- . --- . --- 1 --- .

Anthemis arvensis subsp. arvensis --- . --- 1 --- . --- . --- . --- . --- . Origanum vulgare subsp. vulgare --- . --- 1 --- . --- . --- . --- . --- . Scabiosa ochroleuca --- . --- 1 --- . --- . --- 1 --- . --- . Barbarea stricta --- 1 --- . --- . --- . --- . --- 1 --- . Carex hartmanii --- 1 --- 1 --- . --- . --- . --- . --- 1 Jasione montana var. montana --- 1 --- . --- . --- . --- . --- . --- . Leonurus marrubiastrum --- . --- . --- . --- . --- . --- 1 --- . Hieracium caespitosum --- . --- 1 --- . --- . --- . --- . --- . Euphorbia helioscopia --- . --- 1 --- . --- . --- . --- . --- . Rumex crispus --- 9 --- 9 --- 9 --- 10 --- 1 --- 11 --- 14 Elytrigia elongata --- . --- 1 --- . --- . --- . --- . --- . Peucedanum cervaria --- . --- . --- 1 --- . --- . --- . --- . Ranunculus cassubicus --- . --- . --- 1 --- . --- . --- 1 --- . Bromopsis riparia --- . --- . --- . --- . --- . --- . --- 1 Papaver rhoeas --- . --- 1 --- 1 --- . --- . --- . --- . Naumburgia thyrsiflora --- . --- . --- 1 --- . --- 1 --- 1 --- 1 Ranunculus arvensis --- . --- . --- 1 --- . --- . --- . --- . Odontites vulgaris --- 2 --- 2 --- 1 --- 1 --- 4 --- 4 --- 2 Juncus filiformis --- . --- . --- 1 --- 1 --- . --- . --- . Hieracium x floribundum --- . --- 1 --- 1 --- . --- . --- . --- . Trisetum flavescens subsp. flavescens --- . --- 1 --- . --- . --- . --- . --- 1 Carex riparia --- . --- . --- . --- 2 --- . --- 2 --- 2 Erucastrum gallicum --- . --- . --- 1 --- . --- . --- . --- . Cyperus pannonicus --- . --- . --- . --- . --- . --- 1 --- . Schoenoplectus lacustris --- . --- . --- . --- 1 --- . --- 1 --- 1 Bidens radiatus --- . --- . --- 1 --- . --- . --- 1 --- 1 Vincetoxicum hirundinaria subsp. hirundi --- 1 --- 1 --- . --- . --- . --- . --- . Potentilla reptans --- 5 --- 14 --- 17 --- 15 --- 1 --- 14 --- 14 Rumex stenophyllus --- 1 --- . --- . --- . --- . --- 1 --- . Campanula rapunculoides --- . --- 1 --- . --- . --- . --- . --- . Dianthus armeria --- 1 --- . --- . --- . --- . --- . --- . Senecio erraticus --- . --- 1 --- 1 --- . --- . --- . --- . Minuartia viscosa --- . --- . --- 1 --- . --- . --- 1 --- . Galium trifidum --- . --- . --- . --- . --- . --- . --- 1 Veronica dillenii --- 1 --- . --- 1 --- . --- . --- . --- . Juncus bulbosus --- . --- . --- . --- . --- 1 --- . --- 1 Anacamptis morio subsp. morio --- . --- 1 --- . --- . --- . --- . --- . Capsella bursa-pastoris --- 3 --- 5 --- 3 --- . --- . --- 3 --- 1 Solanum dulcamara --- . --- . --- . --- 1 --- . --- . --- 1 Cardamine hirsuta --- . --- 1 --- . --- . --- . --- 1 --- 1 Maianthemum bifolium --- . --- . --- . --- . --- 1 --- . --- . Erysimum hieraciifolium --- 1 --- . --- . --- . --- . --- . --- . Sedum acre 12.7 7 12.1 7 --- 2 --- 1 --- . --- . --- . Lythrum tribracteatum --- . --- . --- . --- . --- . --- 1 --- . Pinguicula vulgaris --- . --- . --- . --- . --- 2 --- 1 --- . Artemisia austriaca --- 2 --- 1 --- 1 --- . --- . --- . --- . Juncus compressus --- 3 --- 2 --- 3 --- 1 --- 2 --- 3 --- 1 Ranunculus auricomus --- 4 --- . --- 1 11.1 5 --- 1 --- . --- 1 Scirpoides holoschoenus --- . --- 1 --- . --- . --- 2 --- . --- . Carex colchica --- 1 --- . --- . --- . --- . --- . --- . Carduus crispus subsp. crispus --- 1 --- 5 --- 6 --- 2 --- 1 --- 1 --- 4 Verbascum nigrum subsp. nigrum --- . --- . --- . --- . --- . --- . --- 1 Frangula alnus subsp. alnus --- . --- . --- . --- . --- 1 --- . --- . Centaurea stoebe subsp. pseudomaculosa --- . --- 1 --- . --- . --- . --- . --- . Achillea nobilis subsp. nobilis --- . --- 1 --- . --- . --- . --- . --- . Juncus tenuis --- 1 12.1 5 --- 1 --- . --- 2 --- 1 --- 1 Primula elatior subsp. elatior --- . --- 1 --- . --- . --- 1 --- . --- . Cirsium alatum --- 1 --- . --- . --- . --- 1 --- 2 --- 1 Epilobium parviflorum --- . --- . --- 1 --- . --- 1 --- 1 --- 1 Fritillaria ruthenica --- 1 --- . --- . --- 2 --- . --- . --- . Gladiolus tenuis --- 2 --- 2 --- 1 --- 2 --- . --- 1 --- 1 Ranunculus sardous --- . --- 2 --- 2 --- . --- . --- 1 --- . Carduus nutans subsp. nutans --- . --- . --- 1 --- . --- . --- 1 --- . Sesleria caerulea subsp. caerulea --- . --- . --- . --- . --- 1 --- . --- . Scorzonera parviflora --- 1 --- . --- . --- . --- . --- 1 --- . Euphorbia stricta --- . --- . --- . --- 1 --- 1 --- . --- . Carex echinata --- . --- 1 --- 1 --- . --- 3 --- . --- . Leonurus cardiaca --- . --- . --- . --- . --- . --- . --- 1 Chamaecytisus ruthenicus --- 1 --- . --- . --- . --- . --- . --- . Digitaria ischaemum --- . --- 1 --- . --- . --- . --- . --- . Setaria viridis --- 1 --- . --- . --- . --- . --- 1 --- . Erysimum cheiranthoides --- . --- . --- 1 --- . --- . --- . --- . Sagina nodosa --- 2 --- 1 --- 5 --- 2 --- 5 --- 4 --- 1 Lolium multiflorum --- . --- . --- 1 --- . --- . --- . --- 1 Rorippa palustris --- . --- 1 --- 1 --- 2 --- . --- 2 --- 1 Veratrum nigrum --- . --- 1 --- 1 --- . --- 2 --- . --- 1 Melampyrum pratense --- . --- . --- 1 --- . --- . --- 1 --- . Aethusa cynapium subsp. cynapium --- 1 --- . --- . --- . --- . --- . --- 1 Tripleurospermum inodorum --- 1 --- 4 --- 3 --- . --- 1 --- 2 --- . Rumex obtusifolius subsp. silvestris --- . --- 1 --- . --- . --- . --- . --- 1 Holcus mollis subsp. mollis --- . --- 1 --- . --- . --- . --- . --- . Poa bulbosa subsp. bulbosa --- 1 --- . --- . --- . --- . --- . --- . Spergularia rubra var. rubra --- 1 --- 1 --- 1 --- . --- . --- 1 --- . Achillea ptarmica --- . --- . --- . --- . --- . --- 1 --- . Typha angustifolia --- . --- . --- . --- . --- . --- 1 --- . Melilotus albus --- 1 --- 1 --- . --- . --- . --- . --- . Barbarea vulgaris subsp. vulgaris --- . --- . --- 1 --- . --- . --- . --- . Eriophorum polystachyon --- . --- . --- 1 --- . --- 2 --- 1 --- 1 Linaria vulgaris --- 4 --- 4 --- 1 --- . --- 2 --- 1 --- 1 Berteroa incana 10.0 4 8.4 3 --- 1 --- . --- . --- 1 --- . Carex elata --- . --- . --- . --- 1 --- . 10.6 3 --- 1 Persicaria lapathifolia subsp. lapathifo --- . --- 1 --- 2 --- . --- . 10.7 4 --- 1 Euphorbia cyparissias --- 2 --- 1 --- . --- . --- . --- 1 --- . Leersia oryzoides --- . --- . --- . --- . --- . 10.6 2 --- 1 Anacamptis palustris subsp. palustris --- . --- . --- 1 --- . --- . 11.8 3 --- 1 Blysmus compressus --- . --- 1 --- 1 --- . --- . 12.3 4 --- 1 Chenopodium album subsp. album --- 1 --- 1 --- . --- . --- . 11.0 3 --- 1 Cirsium esculentum --- . --- 1 --- 1 --- . --- . 11.6 2 --- . Cynosurus cristatus --- 1 15.3 10 --- 4 --- . --- 7 --- 1 --- 1 Alisma lanceolatum --- . --- . --- . 14.7 2 --- . --- . --- . Centaurea diffusa --- 1 --- 1 --- . --- . --- . --- . --- . Leucanthemum vulgare subsp. vulgare 8.2 18 19.6 27 --- 10 --- 2 --- 19 --- 1 --- 3 Hypericum perforatum subsp. perforatum --- 2 18.0 9 --- 1 --- . --- 4 --- 1 --- .

Knautia arvensis subsp. arvensis --- 1 16.3 5 --- 1 --- . --- 1 --- . --- . Scutellaria hastifolia --- 2 --- 1 --- . 12.8 5 --- 1 --- 1 --- . Veronica scutellata --- 1 --- . --- 1 16.6 8 --- 1 --- 4 --- 1 Persicaria hydropiper --- 1 --- . --- 4 --- 7 --- . 14.7 12 --- 7 Echinochloa crus-galli subsp. crus-galli --- . --- . --- 1 --- . --- . 14.9 3 --- . Sonchus asper subsp. asper --- . --- . --- . --- . --- . 14.4 2 --- . Taraxacum bessarabicum --- . --- . --- . --- . --- . 14.4 2 --- . Triglochin maritima --- . --- . --- 1 --- . --- . 19.8 5 --- . Juncus articulatus --- . --- 5 --- 7 --- 3 --- 13 19.9 22 --- 10 Sium latifolium --- 1 --- 1 --- 1 --- 8 --- 1 16.6 12 --- 6 Xanthium strumarium subsp. strumarium --- 1 --- . --- . --- . --- . 17.4 4 --- . Carduus acanthoides --- . --- 2 --- 1 --- . --- . 12.6 4 --- 1 Ranunculus sceleratus --- 1 --- 1 --- . --- . --- . 13.0 3 --- . Epilobium hirsutum --- . --- . --- . --- . --- 1 12.4 5 --- 3 Alopecurus arundinaceus --- 1 --- 1 --- 1 --- 1 --- . 12.5 7 --- 5 Oenanthe aquatica --- . --- 1 --- 1 --- 1 --- . 14.4 5 --- 2 Glaux maritima --- . --- . --- . --- . --- . 14.4 2 --- . Polygonum aviculare --- 3 --- 1 --- 1 --- . --- . 14.2 5 --- . Juncus gerardii --- 1 --- 2 8.0 8 --- 2 --- 1 14.3 11 --- 3 Festuca rubra aggr. 11.6 35 17.2 41 --- 26 --- 2 --- 32 --- 15 --- 9 Veronica verna subsp. verna --- 4 14.5 5 --- . --- . --- . --- . --- . Silene vulgaris subsp. vulgaris --- 1 11.0 2 --- . --- . --- . --- . --- 1 Erigeron acris subsp. acris --- . 11.0 4 --- 1 --- . --- 3 --- . --- . Thymus serpyllum subsp. serpyllum --- . 14.5 4 --- . --- . --- 2 --- . --- . Consolida regalis subsp. regalis --- . 15.8 3 --- . --- . --- . --- . --- . Silene latifolia subsp. alba --- 1 15.1 5 --- 1 --- . --- . --- . --- 1 Arenaria leptoclados subsp. leptoclados --- . 10.7 2 --- . --- . --- . --- 1 --- . Cerastium arvense subsp. arvense --- 1 9.9 4 --- 2 --- . --- 1 --- 1 --- 1 Allium scorodoprasum --- . --- . --- . 16.9 3 --- . --- . --- . Vicia cracca subsp. cracca 12.7 33 --- 17 --- 17 19.9 40 --- 22 --- 1 --- 12 Thalictrum aquilegiifolium --- . --- 1 --- 2 12.0 3 --- . --- . --- . Thalictrum flavum subsp. flavum --- 2 --- 1 --- 2 17.0 13 --- 7 --- 1 --- 6 Eleocharis uniglumis --- 1 --- . --- 1 12.1 6 --- 1 --- 3 --- 1 Rhinanthus angustifolius 12.9 18 8.9 15 --- 7 --- 4 --- 12 --- 3 --- 2 Hierochloe odorata --- 1 --- 1 --- 1 10.9 4 --- . --- . --- 1 Thalictrum lucidum 9.2 9 --- 4 --- 2 9.8 9 --- 1 --- . --- 5 Equisetum arvense subsp. arvense 11.3 23 17.0 28 --- 14 --- 8 --- 12 --- 6 --- 4 Polygala vulgaris subsp. vulgaris --- 6 17.2 11 --- 2 --- 1 --- 4 --- . --- . Pilosella officinarum subsp. officinarum --- 4 19.9 12 --- 1 --- 1 --- 4 --- . --- . Ajuga genevensis --- . 11.2 1 --- . --- . --- . --- . --- . Artemisia vulgaris --- 3 16.0 6 --- 1 --- . --- . --- . --- 1 Calamagrostis epigejos 19.2 20 --- 3 --- 1 8.1 12 --- 12 --- 1 --- 2 Serratula coronata --- 1 --- . --- . --- . --- . --- . --- . Plantago media 16.1 18 13.9 17 --- 9 --- 3 --- 3 --- 3 --- 1 Agrimonia eupatoria --- 4 15.7 7 --- 1 --- . --- 1 --- . --- . Carex ericetorum --- . --- . --- 1 --- . --- . --- . --- . Tragopogon pratensis --- 1 --- . --- . --- . --- . --- . --- . Carex muricata --- 6 8.9 8 --- 2 --- 2 --- 1 --- 2 --- 4 Picris hieracoides subsp. hieracioides --- 1 12.8 3 --- 1 --- . --- . --- . --- . Luzula campestris --- 1 19.7 14 --- 5 --- . --- 9 --- . --- 2 Hypochaeris radicata subsp. radicata --- . 14.0 5 --- . --- . --- 3 --- . --- . Artemisia absinthium --- 1 12.9 3 --- 1 --- . --- . --- 1 --- . Oenothera biennis --- 3 --- 2 --- . --- . --- . --- . --- . Anchusa officinalis --- . --- 1 --- . --- . --- . --- . --- . Bidens cernuus --- . --- . --- 1 --- . --- . --- 1 --- 1 Pulicaria vulgaris --- . --- 1 --- . --- . --- . --- . --- . Helictotrichon pubescens subsp. pubescen --- . --- 1 --- . --- . --- 2 --- . --- 1 Gnaphalium uliginosum --- . --- . --- . --- . --- . --- 1 --- . Lycopus exaltatus --- 3 --- 1 --- 1 --- 2 --- 1 --- 2 --- 3 Melilotus officinalis --- . --- 1 --- . --- . --- . --- 1 --- . Geranium collinum --- 1 --- . --- . --- . --- . --- 1 --- . Poa trivialis subsp. trivialis --- 8 --- 5 --- 9 --- 12 --- 2 --- 13 --- 8 Alchemilla species --- . --- 1 --- . --- . --- 1 --- . --- . Thymus x tschernjajevii --- 1 --- . --- . --- . --- . --- . --- . Carlina vulgaris --- . --- 1 --- . --- . --- . --- . --- . Atriplex prostrata --- . --- . --- . --- . --- . --- 1 --- . Trisetum sibiricum --- 1 --- . --- . --- . --- . --- . --- . Plantago maritima subsp. maritima --- 1 --- . --- . --- . --- . --- . --- . Chaerophyllum aromaticum --- . --- 1 --- . --- . --- . --- . --- . Tragopogon dubius subsp. dubius --- . --- 1 --- . --- . --- . --- . --- . Arctium lappa --- . --- . --- 1 --- . --- . --- 1 --- 1 Saponaria officinalis --- 1 --- . --- . --- . --- . --- . --- . Dryopteris cristata --- . --- . --- . --- . --- . --- . --- 1 Sonchus oleraceus --- . --- 1 --- 1 --- . --- 1 --- 2 --- 2 Festuca arundinacea subsp. orientalis --- 3 9.7 9 --- 6 --- . --- 2 --- 6 --- 4 Veronica prostrata --- 1 --- . --- . --- . --- . --- . --- . Arenaria serpyllifolia --- 1 --- . --- . --- . --- . --- . --- . Acorus calamus --- . --- . --- 1 --- . --- . --- 3 --- 1 Hydrocotyle vulgaris --- . --- . --- . --- . --- 1 --- . --- . Carex michelii --- 1 --- . --- . --- . --- . --- . --- . Milium effusum subsp. effusum --- 1 --- 1 --- . --- . --- . --- . --- . Vincetoxicum rossicum --- 1 --- . --- . --- . --- . --- . --- . Luzula pilosa --- . --- 1 --- 1 --- . --- . --- . --- . Humulus lupulus --- 1 --- . --- . --- . --- . --- 1 --- 1 Chartolepis glastifolia --- 1 --- . --- . --- . --- . --- . --- . Thalictrum simplex subsp. simplex --- 4 --- 1 --- 1 --- 2 --- . --- . --- 1 Persicaria minor --- . --- 1 --- . --- . --- . --- . --- . Carex melanostachya 12.1 3 --- . --- . --- 1 --- . --- . --- . Verbascum phlomoides --- . --- 1 --- . --- . --- . --- . --- . Fallopia convolvulus --- 1 --- 1 --- 1 --- . --- . --- 1 --- . Carex buekii --- . --- . --- . --- . --- . --- . --- 1 Atriplex calotheca --- . --- . --- . --- . --- . --- 1 --- . Rorippa brachycarpa --- 1 --- . --- . --- 2 --- . --- . --- . Helictotrichon pratense subsp. pratense --- . --- 1 --- . --- . --- . --- . --- . Luzula pallidula --- 1 --- . --- . --- . --- . --- . --- . Cruciata glabra --- 1 --- 1 --- . --- . --- 3 --- . --- . Glaucium corniculatum --- . --- 1 --- . --- . --- . --- . --- . Scutellaria altissima --- . --- 1 --- . --- . --- . --- . --- . Onopordum acanthium subsp. acanthium --- . --- 1 --- . --- . --- . --- . --- . Erysimum leucanthemum --- . --- . --- . --- . --- . --- 1 --- . Salix viminalis --- . --- . --- . --- . --- 1 --- . --- 1 Listera ovata --- . --- . --- . --- . --- . --- . --- 1 Ambrosia artemisiifolia --- . --- 1 --- . --- . --- . --- . --- .

Oxalis stricta --- . --- 1 --- . --- . --- . --- 1 --- 1 Calamagrostis canescens --- . --- . --- 1 --- 2 --- 3 --- 1 --- 2 Rumex acetosa subsp. acetosa --- 24 9.4 29 6.8 27 --- 18 --- 26 --- 5 --- 11 Herniaria polygama --- . --- 1 --- . --- . --- . --- . --- . Hypericum tetrapterum --- . --- . --- 1 --- . --- 1 --- 1 --- 1 Epilobium roseum --- . --- . --- . --- . --- . --- 1 --- . Persicaria mitis --- . --- . --- . --- . --- . --- 1 --- . Geum aleppicum --- . --- . --- 1 --- . --- . --- . --- . Gladiolus imbricatus --- 1 --- 1 --- . --- 1 --- 1 --- . --- 1 Aconitum variegatum subsp. variegatum --- . --- . --- . --- . --- . --- . --- 1 Echinops sphaerocephalus --- 1 --- 1 --- . --- . --- . --- 1 --- 1 Securigera varia --- 1 --- 1 --- . --- . --- . --- . --- . Angelica archangelica --- . --- . --- 1 --- 1 --- . --- . --- 1 Arabis planisiliqua subsp. planisiliqua --- . --- . --- . --- . --- 1 --- . --- . Geranium pratense --- 11 --- 2 --- 9 --- 2 --- 3 --- 7 --- 5 Myosotis arvensis --- 1 10.6 2 --- . --- . --- . --- . --- . Cirsium arvense --- 5 --- 11 --- 14 --- 2 --- 2 --- 15 --- 11 Phleum nodosum --- 1 --- 1 --- . --- . --- . --- . --- . Cardaminopsis arenosa subsp. arenosa --- . --- 1 --- 1 --- . --- 1 --- . --- . Scrophularia nodosa --- 1 --- 1 --- 1 --- 2 --- . --- 1 --- . Silene viscaria --- 1 --- . --- 1 --- . --- . --- . --- . Angelica palustris --- . --- 1 --- 1 --- . --- 2 --- 2 --- 1 Aegopodium podagraria --- . --- 1 --- 1 --- . --- 1 --- 1 --- . Hieracium echioides --- . --- . --- . --- . --- 1 --- 1 --- . Achillea setacea --- 1 --- . --- . --- . --- . --- 1 --- . Galeopsis tetrahit --- . --- . --- . --- . --- . --- . --- 1 Rhinanthus alectorolophus subsp. alector --- 1 --- 1 --- 1 --- 1 --- . --- . --- . Sparganium emersum --- . --- . --- . --- . --- . --- 1 --- . Salix triandra --- . --- . --- 1 --- 1 --- 2 --- 2 --- 1 Pedicularis kaufmannii --- 1 --- . --- . --- . --- . --- . --- . Tragopogon podolicus --- 1 --- . --- . --- . --- . --- . --- . Thalictrum minus subsp. minus --- . --- . --- . --- . --- 1 --- . --- . Epilobium angustifolium --- . --- . --- . --- . --- . --- 1 --- . Iris humilis subsp. arenaria --- . --- 1 --- 1 --- 1 --- . --- . --- . Equisetum hyemale --- . --- . --- . --- . --- . --- . --- 1 Bidens frondosus --- . --- 1 --- 1 --- . --- . --- . --- . Petasites hybridus --- . --- . --- 1 --- . --- . --- . --- . Alchemilla monticola --- . --- . --- 1 --- . --- . --- . --- . Carex brizoides --- . --- . --- 1 --- . --- . --- . --- 1 Chenopodium glaucum --- . --- . --- 1 --- . --- . --- 1 --- . Potentilla arenaria --- . --- . --- . --- . --- . --- . --- 1 Geranium sanguineum --- . --- . --- . --- . --- 1 --- . --- . Crepis praemorsa --- . --- 1 --- . --- . --- . --- . --- . Primula veris subsp. veris --- . --- 1 --- . --- . --- . --- . --- . Equisetum pratense --- 3 --- 4 --- 6 --- 5 --- 9 --- 2 --- 5 Carex ligerica --- . --- . --- . --- . --- 1 --- . --- . Filago vulgaris --- 1 --- . --- . --- . --- . --- . --- . Silaum silaus --- 1 --- . --- . --- 1 --- . --- . --- . Rorippa anceps --- . --- 1 --- . --- . --- . --- . --- . Dianthus deltoides subsp. deltoides 19.6 13 9.0 8 --- 3 --- . --- 3 --- . --- . Fragaria viridis --- 2 9.4 2 --- . --- . --- . --- . --- . Festuca ovina subsp. ovina --- 3 11.3 3 --- . --- . --- . --- . --- . Arrhenatherum elatius subsp. elatius --- 1 11.5 4 --- . --- . --- 2 --- . --- 1 Thlaspi arvense --- 1 --- 1 --- . --- . --- . --- . --- . Alchemilla subcrenata --- . --- . --- . --- . --- . --- 1 --- 1 Veronica arvensis --- 1 9.5 3 --- 1 --- . --- 1 --- . --- . Carex disticha --- . --- . --- 1 --- 3 --- 1 --- 2 --- 3 Trifolium rubens --- . --- 1 --- . --- . --- . --- . --- . Carex elongata --- 1 --- . --- 1 --- . --- . --- 1 --- 1 Ranunculus polyanthemos subsp. polyanthe 9.3 10 9.3 10 --- 4 --- 2 --- 5 --- 3 --- 1 Centaurea cyanus --- 1 --- 1 --- . --- . --- . --- . --- . Viola arvensis subsp. arvensis --- 1 12.7 3 --- . --- . --- 1 --- . --- . Vicia sepium subsp. sepium --- . 13.6 4 --- 1 --- . --- 1 --- . --- 1 Bromus squarrosus --- 1 --- . --- . --- . --- . --- . --- . Salvia pratensis subsp. pratensis --- 1 10.6 2 --- . --- . --- . --- . --- . Carex lepidocarpa subsp. lepidocarpa --- . --- 1 --- . --- . --- . --- . --- 1 Aristolochia clematitis --- 1 --- . --- . --- 1 --- . --- . --- . Callitriche palustris --- . --- 1 --- . --- . --- . --- . --- . Linum perenne --- 1 --- . --- . --- . --- . --- . --- . Vicia villosa subsp. villosa --- 3 --- 1 --- 1 --- 3 --- 1 --- . --- 1 Senecio doria subsp. doria --- 1 --- . --- . --- . --- . --- . --- . Veronica anagallis-aquatica --- . --- . --- . --- 2 --- . --- 2 --- 1 Poa compressa --- 2 --- 1 --- 1 --- . --- . --- 1 --- . Puccinellia distans subsp. distans --- . --- 1 --- . --- . --- . --- 1 --- . Ranunculus bulbosus subsp. bulbosus --- . --- . --- . --- . --- . --- 1 --- . Equisetum sylvaticum --- . --- 1 --- . --- . --- . --- . --- . Omalotheca sylvatica subsp. sylvatica --- . --- 1 --- . --- . --- . --- . --- . Artemisia campestris subsp. campestris --- . --- 1 --- . --- . --- . --- . --- . Sonchus arvensis subsp. arvensis --- 3 --- 4 --- 4 --- 1 --- 4 --- 7 --- 1 Asperula cynanchica subsp. cynanchica --- 1 --- 1 --- 1 --- . --- . --- . --- . Pulmonaria mollis subsp. mollis --- . --- 1 --- . --- . --- . --- . --- .

1

Supporting Information to the paper Kuzemko, A.A. Classification of the class Molinio-

Arrhenatheretea in the forest and forest-steppe zones of Ukraine

Supplement S1. A list of species aggregates.

Juncus bufonius aggr. — mainly Juncus bufonius L., very rare Juncus ambiguus Guss.

and Juncus nastanthus V.Krecz. & Gontsch.;

Festuca rubra aggr. — mainly Festuca rubra L., rare Festuca multiflora Hoffm.;

Festuca valesiaca aggr. — mainly Festuca valesiaca aggr., rare Festuca pseudodalmatica

Krajina ex Domin and Festuca pseudovina Hack. ex Wiesb.;

Erigeron acris aggr. — mainly Erigeron acris L., very rare Erigeron podolicus Besser

and Erigeron politus Fr.;

Erigeron annuus aggr. — mainly Erigeron annuus (L.) Pers., rarer Erigeron annuus (L.)

Pers. subsp. septentrionalis (Fernald & Wiegand) Wagenitz and Erigeron annuus (L.) Pers.

subsp. strigosus (Muhl. ex Willd.) Wagenitz;

Arabis hirsuta aggr. — more often Arabis sagittata (Bertol.) DC., rarer Arabis nemorensis

(Hoffm.) W.D.J.Koch;

Cardamine pratensis aggr. — mainly Cardamine pratensis L., rarer Cardamine dentata

Schult.;

Cerastium fontanum aggr. — mainly Cerastium holosteoides Fr., very rare Cerastium

fontanum Baumg.;

Stellaria graminea aggr. — mainly Stellaria graminea L., rarer Stellaria hippoctona

(Czern.) Klokov, very rare Stellaria fragilis Klokov;

Hylotelephium telephium aggr. — mainly Hylotelephium maximum (L.) Holub, rare

Hylotelephium polonicum (Błocki) Holub and Hylotelephium stepposum (Boriss.) Tzvelev,

Anthyllis vulneraria aggr. — mainly Anthyllis macrocephala Wender., very rare Anthyllis

arenaria (Rupr.) Juz., Anthyllis schiwereckii (DC.) Błocki;

Lotus corniculatus aggr. — mainly Lotus corniculatus L., rarer Lotus ucrainicus Klokov,

Lotus ambiguus Besser ex Spreng, Lotus arvensis Pers., Lotus uliginosus Schkuhr;

Medicago falcata aggr. — more often Medicago procumbens Besser, rarer Medicago

romanica Prodán;

Salvia nemorosa aggr. — Salvia illuminata Klokov, Salvia tesquicola Klokov & Pobed.,

Salvia nemorosa L.;

Thymus pulegioides aggr. — mainly Thymus pulegioides L., rare Thymus ucrainicus

(Klokov & Des.-Shost.) Klokov;

2

Persicaria lapathifolia aggr. — more often Persicaria scabra (Moench) Moldenke, rarer

Persicaria lapathifolia (L.) Delarbre;

Euphrasia stricta s.l. — more often Euphrasia stricta D.Wolff ex J.F.Lehm., rarer

Euphrasia brevipila Burn. & Gremli, very rare Euphrasia pectinata Ten., Euphrasia tatarica

Fisch. ex Spreng.;

Melampyrum nemorosum aggr. — mainly Melampyrum nemorosum L., very rare

Melampyrum moravicum Heinr.Braun, Melampyrum polonicum (Beauverd) Soó;

Veronica spicata aggr. — mainly Veronica spicata L., rarer Veronica barrelieri Schott.

Urtica dioica aggr. — more often Urtica dioica L., rarer Urtica galeopsifolia Wierzb. ex

Opiz and Urtica pubescens Ledeb., very rare Urtica kioviensis Rogow.

Supporting Information to the paper Kuzemko, A.A. Classification of the class Molinio-Arrhenatheretea in the forest and forest-steppe zones of Ukraine

Supplement S1. A results of the “crispness of classification” analysis.

Method Number of clustersof cluster analysis 2 3 4 5 6 7 8 9 10 11 12

Sorensen/Flexible beta -25/non-transf 264.67 266.975 271.504 283.512 287.617 283.101 276.833 271.944 274.342 270.718 263.233Sorensen/Flexible beta -25/log 227.131 259.141 244.291 257.673 273.194 272.914 260.99 257.303 254.704 251.562 262.135Rel Sorensen/Flexible beta -25/non-transf 276.073 275.655 270.062 278.386 261.225 256.588 257.987 254.114 245.58 244.161 243.825Rel Sorensen/Flexible beta -25/log 308.436 279.241 277.536 288.334 290.652 287.303 288.361 281.977 274.816 273.908 273.205Euclidean/Ward/non-transf 274.376 228.055 223.01 217.988 205.14 217.621 214.918 207.406 199.848 198.966 206.386Euclidean/Ward/log 253.295 238.364 241.178 233.986 229.418 232.219 229.37 223.849 229.509 223.993 218.675TWINSPAN mod Sorensen 378.75 331.923 297.462 275.966 251.331 227.94 218.057 230.223 217.378 224.278 241.825TWINSPAN mod Simpson 378.75 331.923 297.462 275.966 251.331 227.94 218.057 204.314 217.378 209.286 201.555TWINSPAN mod Jaccard 378.75 331.923 297.462 275.966 251.331 227.94 218.057 230.223 217.378 224.278 241.825TWINSPAN mod Whittaker 378.75 320.316 315.124 318.235 315.684 309.092 303.478 299.765 301.653 297.782 290.782

classification of the class molinio-arrhenatheretea in the

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