235 Phytosociological and ecological peculiarities of Festuca pallens Host in Ukraine Abstract This article deals with phytosociological and ecological requirements of the vulnerable species Festuca pallens in Ukraine. Based on the analysis of 51 relevés, we have recognized the occurrence of Festuca pallens in three associations within the Festuco-Brometea class. Some communities where Festuca pallens occurs have transitional stages of succession to forest edge vegetation according to the expert system classification. In addition, we evaluated the ecological differences among occupied habitats using Didukh’s phytoindication scales and concluded that the most important ecological factors are soil humidity, nitrogen content, soil aeration, salt regime, carbonate content, thermoregime, and climate continentality. Finally, we evaluated the threats to the existence of these communities with the presence of Festuca pallens, taking into account climate change and other impacts of anthropogenic activities (illegal mining, recreation, afforestation). Izvleček V članku obravnavamo fitocenološke in ekološke potrebe ranljive vrste Festuca pallens v Ukrajini. Z analizo 51 vegetacijskih popisov smo potrdili pojavljanje vrste Festuca pallens v treh asociacijah iz razreda Festuco-Brometea. Nekatere združbe, v katerih se pojavlja Festuca pallens, so po klasifikaciji z ekspertnim sistemom sukcesijski stadij prehoda v gozdni rob. Dodatno smo ovrednotili ekološke razlike med habitati s fitoindikacijskimi vrednostmi po Didukhu in ugotovili, da so najpomembnejši dejavniki vlažnost tal, vsebnost dušika, prezračenost tal, slanost, vsebnost karbonatov, toplotni režim in kontinentalnost klime. Za konec smo ovrednotili ogroženost združb z vrsto Festuca pallens v povezavi s klimatskimi spremembami in drugimi antropogenimi aktivnostmi (nedovoljeno rudarjenje, rekreacija, pogozdovanje). Key words: phytosociological assessment, phytoindication, dry grasslands, vegetation, Ukraine. Ključne besede: fitocenološka ocena, fitoindikacija, suha travišča, vegetacija, Ukrajina. Corresponding author: Iuliia Vasheniak E-mail: arrhenatherum@gmail.com Received: 24. 8. 2021 Accepted: 12. 2. 2022 Yakiv Didukh1, Iuliia Vasheniak2 & Iryna Bednarska3 DOI: 10.2478/hacq-2022-000421/2 • 2022, 235–252 1 M.G. Kholodny Institute of Botany, NAS of Ukraine, Kyiv, Ukraine. 2 Vasyl’ Stus Donetsk National University, Vinnytsia, Ukraine. 3 Institute of Ecology of the Carpathians, NAS of Ukraine, Lviv, Ukraine. 21/2 • 2022, 235–252 236 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Introduction Festuca pallens is a characteristic species of Central and Southern Europe (Markgraf-Dannenberg, 1980; Di Pi- etro & Catonica, 1999), but in Ukraine it is located at the eastern limit of its range and is clearly separated from the main part of the species’ range by a disjunction. This species is extremely rare in the Ukrainian flora and is rep- resented by a few isolated populations in a particularly re- stricted area, which naturally led to its endangerment and inclusion in the Red Data Book (Didukh et al., 2009). Festuca pallens is an obligate calcareous, xerophilous spe- cies that occurs on limestone and gypsum rocks, dry cliffs rock shelves and steep walls, and on chalk outcrops with low erosion. According to ecological strategy, this spe- cies is typically stress tolerant, although it also grows in shaded mesophitic communities that have developed on initial rendzic soils. Communities dominated by Festuca pallens are listed in the Green Book of Ukraine (2009) with the presence of Carex humilis, Teucrium chamaedrys, Gypsophila fastigiata, Helianthemum canum, Aurinia saxa- tilis, and steppe grass Stipa capillata (Zaverukha, 1978; Stoiko et al., 1998). According to the National Habitat Catalogue (Kuzemko et al., 2018), habitats with the oc- currence of Festuca pallens are listed as T.1.2.2 Petrophyte steppes on carbonate substrata (Vasheniak, 2018) and as E:2.213 habitats with the dominance of Festuca pallens on carbonate slopes and outcrops (Korotchenko, 2011). Therefore, the species and communities with their par- ticipation in Ukraine must be protected, which requires a meticulous and detailed study. This will allow to pre- dict the behavior and the nature of further development of communities in modern conditions of environmental change. At the same time, such studies imply the condi- tions of existence and phytosociological peculiarities on the eastern border of their distribution in Ukraine, which differ from the characteristics of Central and Southern Eu- rope. Finally, the culmination of these discoveries has led us to critically review the available information on the dis- tribution of Festuca pallens and to study the ecological and phytosociological peculiarities of the respective communi- ties, as well as to predict the possibility of future changes. Systematic position Festuca pallens belongs to a systematically complex group of narrow-leaved fescues, which is interpreted differently by authors. Consequently, the species has received differ- ent nomenclature in the course of scientific development. For example, in different periods the species was consid- ered a subspecies or variety in the composition of Festuca cinerea s.l. (Rauschert, 1961; Toman, 1974; Rothmaler, 1994), Festuca glauca s.l. (Hackel, 1882; Kozłowska, 1926; Soó, 1956; Auquier P. & Kerguélen M., 1978), or as an independent species (Markgraf-Dannenberg, 1980; Šmarda et al., 2007), especially in the series Psam- mophillae (Pawlus, 1985). In Ukraine, Festuca pallens is represented by the typical subspecies of Festuca pallens Host subsp. pallens. Festuca pallens was first mentioned for the territory of Ukraine (at that time Galicia was part of Poland) in the works of Zapałowicz (1906) and A. Kozłowska (1926). There were very few works that mentioned Festuca pallens. Those that contained general systematic treatises on the genus were written by Alekseev (1975) and Tsvelev (1977) on the territory of the former USSR, and 2 works by Ukrainian authors (Tveretinova, 1977; Zaveruha, 1978). In addition, the vegetation with the occurrence of Festuca pallens was considered in the work of Onyshchenko (2001), describing the association Minuartio auctae-Festucetum pallentis within the order Alysso-Sedetalia, class Sedo-Scleranthetea, which occurs in the Nature Reserve “Medobory”. Based on the above facts, we conducted a complex review of the known litera- ture and herbarium data (Supplement 1) of sites with the occurrence of Festuca pallens and the geobotanical relevés of Festuca pallens communities (Figure 1). Materials and methods Study area According to the geobotanical zoning, the study area in- cludes the sub-province of Western Podillia, Central Eu- ropean province and Forest region, and is divided into three geobotanical districts: Opillia, Kremenets Ridge, and Volyn Upland (Figure 1, Appendix). According to Zaverukha (1978), Festuca pallens is most widespread in the vicinity of Podillia (Ivano-Frankivsk region, Galych district). This species grows on extreme- ly steep slopes of gypsum rock up to 10 meters high in various locations, while in other areas it mainly colonizes south-facing slopes. The area most densely colonized by Festuca pallens is on the open southern slopes with the participation of Sedum hispanicum, which is also rare for the continental Ukraine and is located at the eastern limit of its distribution. On the other hand, it was also found in shady, mesophytic communities with abundant mosses and ferns (Cystopteris fragilis, Polypodium vulgare), where Festuca pallens grows much less “vigorously” and forms small lawns, at times only in vegetative state, with- out generative organs, with green leaf blades in contrast to gray-leaved fescues growing in light habitats. In addi- tion, it should be mentioned that Festuca pallens occurs on horizontal or gently sloping soils with floristically 21/2 • 2022, 235–252 237 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine degraded communities. The reason for this could be the accumulation of a considerable amount of soil enriched with organic matter in the swards. Such a substrate is gradually penetrated by other plant species that displace Festuca pallens. However, such a strategy is justified only on gentle slopes, whereas Festuca pallens forms long-term stable communities on steep cliffs (Medwecka-Kornaś & Kornaś, 1959). In addition, this fescue is found near trampled paths, where grass cover is sparse. Near Mezhy- hirtsi in the Ivano-Frankivsk region there is a population of this species, which occurs in the form of small sites on gypsum rocks. Plowing of the area around the slope where Festuca pallens grows, as well as grazing, resulted in degradation of dry grasslands. In addition, ruderal ele- ments invade and the rocks are overgrown by shrubs. In this situation, the lack of refuges and the species’ inability to compete with other dominant species led to a signifi- cant reduction in its population (single individuals on the slope) and a decrease in the overall size of the plants. The third local population of Festuca pallens in Opillia was es- tablished near the village of Pidkamin (Ivano-Frankivsk region, Rogatyn district) on the steep, rugged gypsum walls of the northern and southern exposure at a height of 2–5 m, with the presence of Sedum acre, S. hispanicum, Gypsophila thyraica and Thymus pannonicus. The Kremenets Ridge is one of the most important refugia where local populations of Festuca pallens have been preserved at the eastern limit of the species’ range. It has been observed in a few places on Bona Hill (Zam- kova Hill), Divocha Hill (Divochi Skeli) and Strakhova Hill. Festuca pallens is one of the dominant species in the petrophytic communities on these hills, which are mainly south-facing, with medium to very steep, open rocky slopes composed of Sarmatian-age limestones, characterized by a porous or multistepped surface form (Zaverukha, 1978). It is worth noting that the Minuar- Figure 1: Locations of the relevés with the presence of Festuca pallens in Ukraine. Slika 1: Lokacije vegetacijskih popisov s navzočnostjo vrste Festuca pallens v Ukrajini. 52°N 51°N 50°N 49°N 48°N 47°N 46°N 45°N 44°N 43°N 42°N 41°N 40°N 52°N 51°N 50°N 49°N 48°N 47°N 46°N 45°N 44°N 43°N 42°N 41°N 40°N 23°N 24°N 25°N 26°N 27°N 28°N 29°N 30°N 31°N 32°N 33°N 34°N 35°N 36°N 37°N 38°N 39°N 40°N 41°N 42°N 43°N 21/2 • 2022, 235–252 238 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine tio auctae-Festucetum pallentis association (Onyshchenko, 2001; Vinnichenko & Oliiar, 2004) on Divocha Hill is associated with the presence of both Festuca pallens and Alyssum montanum, Artemisia campestris, Carex humilis, Dianthus arenarius subsp. pseudoserotinus, Festuca makut- rensis, Minuartia aucta, Sedum acre and Thymus species oc- curs. On Strakhova Hill, Festuca pallens grows together with Carex humilis and Peucedanum oreoselinum on the eastern slope with a slight inclination, which is partially overgrown by shrubs (Frangula alnus, Euonymus verru- cosa) (Vinnichenko & Oliiar, 2004). The herb layer is formed by the dominant species Carex humilis and Carex montana, Lembotropis nigricans, Peucedanum oreoselinum, Polygonatum odoratum, Pulsatilla patens, Ranunculus brey- ninus, Teucrium chamaedrys and Thalictrum minus. In ad- dition, these communities are atypical of Festuca pallens and rich in species of forest edge vegetation of the class Trifolio-Geranietea and in the successional stage of shrub vegetation (Berberidion). A special habitat for Festuca pallens is a sand quarry in the town of Lysa near the village of Kulykiv (Ternopil re- gion, Kremenets district). It is worth noting that Festuca pallens rarely grows on sandstone, even if it is very cal- careous, except for a few isolated sites in Poland, where Jurassic sandstones are unusually hard and can serve as a stony substrate for fescue (Motyka, 1947). It is also inter- esting to note that Festuca pallens never grows on marls, which are essentially conglomerates (limestones and sandstones), whereas many plants can grow on both sub- strates. The most plausible explanation is that marls retain more moisture than sandstones, which is critical for Festu- ca pallens. Development of the quarry in Lysa is currently halted, but insignificant quarrying operations are occa- sionally carried out in its area. As a result, there is a whole range of transitional states from vegetation-free outcrops and pioneer communities to secondary shrub cover and coenotically unformed communities. The northern pop- ulation of Festuca pallens was discovered on the Volyn Upland near the village of Mala Mylcha (Rivne Region, Dubno District), where the species occurs on chalk hills at elevations up to 350 m on the slopes of southern ex- posures. This fescue actively colonizes chalk outcrops and steppe communities with the dominance of Carex humilis and a sparse herb layer. Festuca pallens thus grows in dif- ferent habitats under different conditions, but the most important characteristic is the carbonate content of the substrates and the sparse herb layer, even in disturbed or converted areas with weak ecological competition. It should be noted, therefore, that excessive recreational use or other anthropogenic impacts can trigger the extinction of a species. Such behavior is characteristic of many rare species, which we interpret as an “effect of repulsion of relics” (Didukh, 1988). In disturbed areas, Festuca pallens usually forms normal, full-sized populations; however, the viability of its individuals remains lower compared to plants of natural habitats. In particular, the vast majority of the species’ morphometric indicators decline, includ- ing the number and height of generative stems and seed productivity under unfavorable conditions (Bednarska, 2011). Thus, seed reproduction is activated by the re- moval of coenotic pressure and competition from other species at the periphery of populations. Incidentally, this is observed both on natural landslides and outcrops and under disturbed conditions, which are sometimes com- pletely uncharacteristic for Festuca pallens. When coe- notic competition decreases, the behavior of the species is focused on recovery and establishment, while in closed coenoses it is about the self-sufficiency of the population. Vegetation data collection We have used 51 phytosociological relevés with a uniform plot size of 10 m² according to the Braun-Blanquet ap- proach (Braun-Blanquet, 1964) with the modification of the EDGG method, personally collected in Ukraine in 2016 and 2020 with specific vascular plants, bryophytes and epigeic lichens. Nomenclature for vascular plant species is according to the Euro+Med Database check- list (http://www.emplantbase.org), while bryophytes are named according to Boiko (2008) and lichens according to Kondratiuk (1998). Cluster analysis and expert system processing Data were analyzed using JUICE software (Tichý, 2002). We used the modified TWINSPAN algorithm (Hill, 1979): three pseudospecies cut levels -0, 5, and 25, Whit- taker method to distinguish clusters. Diagnostic species for each cluster were estimated based on the phi- coef- ficient (the threshold for fidelity greater than 25%) and tested using Fisher’s exact test (p≥0.01) (Chytrý et al., 2002), with sizes of all groups standardized to equal size (Tichý & Chytrý, 2006). For identification of highly con- stant species, we set the threshold for constancy at more than 50% and for constant species at more than 25%. Class- and order-level identification was performed us- ing the EuroVegChecklist Expert System (Mucina et al., 2016) and additionally using the Slovak Expert System (Janišová et al., 2007) to identify clusters with question- able syntaxonomy. All unidentified relevés or the relevés with low diversity of taxa (less than 10 taxa) were exclud- ed from the dataset. As a result, we obtained 44 relevés, which we analyzed thoroughly. 21/2 • 2022, 235–252 239 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Detrended Corresponded Analysis using Didukh’s scales A Detrended Correspondence Analysis (DCA) was conducted to evaluate the effects of environmental fac- tors on plant communities. The phytoindication scales of Y. Didukh (Didukh, 2011) are comparable to other environmental indicator values (Ellenberg) and reflect amplitude indices of species characterized by the follow- ing dimensions: soil humidity (Hd – 23 grades ), vari- ability of damping (fH – 11 grades ), soil acidity (Rc – 15 grades), total salt regime (Sl – 19 grades ), carbonate con- tent (Ca – 13 grades ), nitrogen content (Nt – 11 grades), aeration of the soil (Ae – 15 grades ), thermoregime of the climate (Tm – 17 grades), humidity of the climate (Om – 23 grades), continentality of the climate (Kn – 17 grades), cryoregime of the climate (Cr – 15 grades) and lightness in the community (Lc – 9 grades). Phytoindica- tion values were passively projected onto a DCA graph. We used the program R (R Core Team, 2013) to visual- ize the DCA analysis. Assessing of climate change impact on Festuca pallens communities To assess the impact of climate change, point calcula- tions of relevant indicators of 12 leading environmen- tal factors were performed according to the scales and methods of Didukh (2021). Based on statistical process- ing of geobotanical data, the amplitude of their values for the corresponding syntaxon, average (X0), minimum (X0min) and maximum (X0max), as well as the corre- lation between their change for each of the factors, es- pecially in relation to the temperature regime, were de- termined. Further measures were based on the fact that the increase in the average annual temperature by +1, +2, +2.5 and + 3.0 °C were determined on the basis of the correlation dependence equations and the corresponding values of the indicators for each of the factors were calcu- lated. These values were taken as average values (X + 1, X + 2, X + 2.5 and X + 3), in relation to which the am- plitude limits (±2σ) were calculated. The next analysis consisted in evaluating the overlap of the amplitudes of the obtained values in relation to the original, reflecting the current growth conditions of Festuca pallens. In the case of a direct ratio between the factors, this overlap was estimated in relation to the maximum (X0max), and in the case of an inverse ratio in relation to the minimum (X0min). If the mean values (X + 1, X + 2, X + 2.5 and X + 3) do not exceed the amplitude (X0min -X0max) and only the maximum (or minimum) values are outside this amplitude, then there is no threat to the existence of the species. If the mean values (X + 1, X + 2, X + 2.5 and X + 3) are outside the amplitude (X0min–X0max), but the minimum or maximum values (X + 1, X + 2, X + 2.5, and X + 3) are still within (X0min–X0max), there is a significant risk of habitat loss. If all values are outside the amplitude (X0min–X0max), then conditions have changed drastically, making the existence of Festuca pallens impossible and the original habitat has become extinct. The next step was to evaluate the prediction of resis- tance, the nature of habitat development based on the behavior of species reflected in the environmental strat- egy of Grime (CRS). The calculation of the number of species of each type of strategy for specific relevés was carried out according to the formula: SS = S + (CS/2) + (SR/2) + (CRS/3); CC = C + (CS/2) + (CR/2) + (CRS/3); RR = R + (CR/2) + (RS/2) + CTS/3). The means and squared deviations (±2σ) for distinguished syntaxa, as well as the relationship between indicators were calcu- lated: SS:CC; RR:CC; SS:RR and ratio C:S:R = 1.0:X:Y. The obtained indicators (as a percentage) of the ratios were plotted on the sides of Grime›s triangle, where the vertices correspond to 100%. On the line C-S the point is SS = 100-CC and the position is CC = 100CC/(CC + SS). On the line C-R the point CC = 100-RR, then the position is RR = 100RR/(RR + CC). On the line S-R, the point RR = 100-SS and the position is SS = 100SS/ (SS + RR). The points were placed on the sides of the triangle and connected. On the basis of these numbers, the «centers of their masses» of the triangle were defined, corresponding to an intersection of its medians confined to the corresponding fields. On the basis of the average values of the ratios of the different types of strategies (%) for a certain type of syntaxon, a three-dimensional matrix was created, illustrating the possible way of development of the communities. This system of applying different methods allowed us to aim at a comprehensive evalua- tion of Festuca pallens communities and the ecological conditions of their existence, as well as to make certain predictions about possible changes, which is the core of developing measures for their conservation. Results The vegetation data of 44 relevés, consisting of 201 vascu- lar plant and cryptogamic species, are divided into three main groups (Table 1), all belonging to dry grasslands of the class Festuco-Brometea, two associations of the order Stipo pulcherrimae-Festucetalia pallentis, Galio campanu- lati-Poion versicoloris alliance, and one syntaxon without clear syntaxonomic affiliation. 21/2 • 2022, 235–252 240 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Number of relevés 18 15 11 Cluster 1 2 3 Festuca pallens 100 100 100 Ass. Schivereckio podolicae-Seselietum libanotidis Gypsophila thyraica 61 7 45 Thymus pannonicus 50 . 18 Campanula rotundifolia 50 13 . Astragalus onobrychis 33 20 9 Arabidopsis arenosa 33 0 18 Ass. Minuartio auctae-Festucetum pallentis Clinopodium acinos 6 80 9 Thymus serpyllum . 73 18 Alyssum gmelinii . 67 9 Minuartia setacea subsp. setacea 6 67 18 Medicago falcata 22 53 9 Centaurea stoebe 11 53 18 Anthyllis vulneraria 6 33 . Saxifraga tridactylites . 33 . Silene eugeniae 11 27 . transitional communities from Festuco-Brometea to Trifolio-Geranietea Anthericum ramosum . 27 82 Teucrium chamaedrys 6 7 64 Stipa capillata . 7 55 Salvia verticillata 11 . 45 Geranium sanguineum 11 . 45 Cytisus paczoskii . . 45 Galium boreale subsp. exoletum 17 13 45 Teucrium montanum 6 20 45 Thalictrum minus . 13 45 Galium mollugo 11 . 36 Pimpinella saxifraga . 7 36 Stachys recta . 7 36 Cytisus albus 6 . 27 Lembotropis nigricans . . 27 All. Galio campanulati-Poion versicoloris, Ord. Stipo pulcherrimae-Festucetalia pallentis Potentilla incana 39 67 73 Allium lusitanicum 67 53 27 Sedum acre 50 80 9 Helianthemum canum 11 33 73 Carex humilis 11 33 64 Melampyrum arvense 11 33 9 Table 1: Synoptic table of communities in which Festuca pallens (Ukraine) occurs, indicating the constancy of species expressed by their percentage frequency to the respective three clusters. Shaded species are ranked by decreasing constancy: dark shading ≥50%, light shading ≥25%. Phi coefficient values are not shown, but species with phi- value greater than 0.25 are accepted as differential for the alliances. Species of each syntaxon with a constancy of 15% or less as well as other taxa with a constancy of 15% or less are not shown in the table. Tabela 1: Sinoptična tabela združb z vrsto Festuca pallens v Ukrajini. Prikazana je stalnost vrst s frekvenco v odstotkih v treh klastrih. Zasenčene vrste so urejene po padajoči stalnosti: temno osenčeno ≥ 50%, svetlo osenčeno ≥ 25%. Vrednosti fi koeficienta so prikazane samo za vrste z vrednostjo večjo od 0,25, kar smo upoštevali kot razlikovalnice za zvezo. Značilne vrste za posameze sintaksone s stalnostjo, manjšo od 15%, in ostale vrste s stalnostjo, manjšo od 15%, v tabeli niso prikazane. 21/2 • 2022, 235–252 241 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Number of relevés 18 15 11 Inula ensifolia 17 7 27 Galium glaucum 17 . 27 Iris aphylla subsp. hungarica 11 . 18 Sempervivum ruthenicum 22 7 . Cl. Festuco-Brometea Asperula cynanchica 61 40 55 Artemisia campestris 56 67 9 Festuca stricta subsp. sulcata 39 47 9 Campanula sibirica 28 27 36 Elytrigia intermedia 28 40 36 Euphorbia cyparissias 56 20 73 Arenaria serpyllifolia 33 33 . Scabiosa ochroleuca . 67 55 Koeleria macrantha 39 33 27 Hypericum perforatum 11 7 36 Euphorbia seguieriana 11 7 36 Other species Veronica spicata 33 27 18 Asplenium ruta-muraria 28 27 . Galium verum 22 13 27 Echium vulgare 6 20 27 Vincetoxicum hirundinaria . 20 27 Centaurea phrygia . . 27 Hieracium umbellatum 22 . . Geranium robertianum 17 7 . Peucedanum cervaria 17 7 9 Seseli libanotis subsp. intermedium 17 . 9 Cryptogam species (in alphabetical order) Abietinella abietina 22 67 27 Barbula unguiculata . 40 9 Brachythecium albicans 22 . . Bryum argenteum . 20 18 Bryum kunzei . 67 18 Ceratodon purpureus 11 53 9 Cladonia pocillum . 20 9 Cladonia pyxidata 17 13 9 Collema tenax 17 27 9 Ditrichum flexicaule . 60 27 Encalypta vulgaris 6 47 9 Homalothecium lutescens . 20 9 Homalothecium sericeum . 47 9 Hypnum vaucheri 11 27 . Pseudocrossidium hornschuchianum . 20 . Rhytidium rugosum . 27 . Tortula calcicolens 0 27 . Tortella inclinata . 33 27 Tortella tortuosa . 13 27 Tortula ruralis 6 47 18 Weissia sp. . . 27 21/2 • 2022, 235–252 242 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Figure 2: Pictures of the communities with the presence of Festuca pallens. a) Minuartio auctae-Festucetum pallentis, Krements Ridge, Divochi Skeli Hill, Ukraine; b) Schivereckio podolicae-Seselietum libanotidis, Velyki Goldy Hills, c) transitional communities from Festuco-Brometea to Trifolio-Geranietea, Krements Ridge, Strakhova Hill, photos made by Iuliia Vasheniak. Slika 2: Slike združb z vrsto Festuca pallens. a) Minuartio auctae- Festucetum pallentis, greben Krements, hrib Divochi Skeli, Ukrajina, foto Vasheniak Iullia; b) Schivereckio podolicae-Seselietum libanotidis, hribovje Velyki Goldy, c) prehodna združba med razredoma Festuco- Brometea in Trifolio-Geranietea, greben Krements, hrib Strakhova, fotografije Iuliia Vasheniak. On Figure 2 some examples of communities of distin- guished vegetation types are presented. a b c Figure 3: Detrended Correspondence Analysis of the distinguished syn- taxa. 1-Schivereckio podolicae-Seselietum libanotidis, 2-Minuartio auctae- Festucetum pallentis, 3-transitional communities from Festuco-Brometea to Trifolio-Geranietea. Acronyms: Hd (soil humidity), fH (variability of damping), Rc (soil acidity), Sl – total salt content in soil, Ca – calcium/ magnesium content in soil, Nt – nitrogen content in soil, Ae – soil aeration, Tm – thermal climate, Om – ombroregime, Kn – continental climate, Cr – cryoclimate, Lc – light. Figure 3: Korespondenčna analiza z odstranjenim trendom obravnavanih sintaksonov. 1-Schivereckio podolicae-Seselietum libanotidis, 2-Minuartio auctae-Festucetum pallentis, 3- prehodna združba med razredoma Festuco- Brometea in Trifolio-Geranietea. Okrajšave: Hd (vlažnost tal), fH (variabil- nost vlažnosti), Rc (kislost tal), Sl – vsebnost soli v tleh, Ca – vsebnost kalcija/magnezija v tleh, Nt – vsebnost dušika v tleh, Ae – prezračenost tal, Tm – toplota klime, Om – ombrorežim, Kn – kontinentalnost klime, Cr – krioklima, Lc – svetloba. Ecological peculiarities of the communities with the presence of Festuca pallens As we can see from Figure 3, three syntaxa were clearly distinguished on the DCA visualization. The main axis (DCA 1) distinguishes the differences of Minuartio auctae- Festucetum pallentis association compared to other syntaxa 21/2 • 2022, 235–252 243 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine according to most environmental factors (soil humid- ity, nitrogen content, aeration of the soil, salt regime, carbonate content, thermoregime, and continuity of climate) and transitional communities, which include different stages of digression according to variability of damping, humidity of the climate and light in the com- munity. Although the second axis (DCA 2) does not show significant differences between the associations, it reflects the difference between the Schivereckio podolici- Seselietum libanotidis association and the previous ones in terms of soil acidity and cryoregime of the climate. The transitional communities are located along the vec- tors of variability of damping, humidity and light in the community. averaged min max T-0 T+1 % T+2 % T+2,5 % T+3 % Tm-A 9.21 8.67 9.75 9.21 9.9 102 10.6 109 10.9 112 11.3 116 Tm-B 9.21 8.67 9.75 9.21 9.86 101 10.5 108 10.8 111 11.2 114 Cr-A 8.71 8.08 9.33 8.8 9.12 97.7 9.43 101 9.59 103 9.75 105 Cr-B 8.71 8.08 9.33 8.8 9.11 97.6 9.49 102 9.65 103 9.86 106 Kn-A 8.69 7.97 9.42 8.74 8.98 95.4 9.22 97.9 9.34 99.2 9.46 100 Kn-B 8.69 7.97 9.42 8.74 8.97 95.2 9.2 97.6 9.29 98.6 9.43 100 Om-A 12 11.2 12.8 12 11.3 99.6 10.5 106 10.2 110 9.8 115 Om-B 12 11.2 12.8 12 11.2 99.8 10.7 105 10.5 107 10.1 111 Hd-A 8.77 7.8 9.73 8.83 8.16 95.7 7.5 104 7.17 109 6.83 114 Hd-B 8.77 7.8 9.73 8.83 8.19 95.2 7.57 103 7.3 107 6.94 112 Fh-A 6.13 5.27 6.99 6.19 6.33 90.7 6.48 92.7 6.55 93.7 6.62 94.8 Fh-B 6.13 5.27 6.99 6.19 6.33 90.5 6.46 92.5 6.52 93.3 6.6 94.4 Nt-A 4.19 3.37 5.02 4.25 3.95 85.2 3.66 92 3.51 95.9 3.36 100 Nt-B 4.19 3.37 5.02 4.25 3.97 84.8 3.69 91.3 3.57 94.3 3.41 98.8 Ae-A 5.2 4.6 5.8 5.23 4.99 92.3 4.75 96.9 4.64 99.3 4.51 102 Ae-B 5.2 4.6 5.8 5.23 5 92 4.78 96.4 4.68 98.3 4.55 101 Rc-A 8.7 7.96 9.44 8.7 9.32 98.7 9.93 105 10.2 108 10.5 112 Rc-B 8.7 7.96 9.44 8.7 9.28 98.3 9.86 104 10.1 107 10.4 111 Sl-A 8 7.03 8.96 8.01 8.56 95.5 9.1 102 9.37 105 9.65 108 Sl-B 8 7.03 8.96 8.01 8.57 95.7 8.98 100 9.15 102 9.38 105 Ca-A 8.91 7.45 10.4 8.88 9.41 90.7 9.93 95.8 10.2 98.3 10.5 101 Ca-B 8.91 7.45 10.4 8.88 9.43 90.9 9.82 94.7 9.99 96.3 10.2 98.5 Lc-A 7.64 7.24 8.04 7.61 7.79 96.9 7.96 99 8.05 100 8.14 101 Lc-B 7.64 7.24 8.04 7.61 7.78 96.7 7.94 98.8 8.01 99.6 8.11 101 Table 3: Mean indicator values, amplitude of environmental factors and their possible changes at increase of average annual temperatures by +1, +2, +2.5 and +3.0 °C. The minimum values, in relation to which the limits are calculated, are marked by a green color, and the maximum values by a blue color. The zone of increasing risk is marked by a yellow color; the risk of loss is marked by a red color. Variants were calculated depending on the degree of correlation between indicators of different (A) and petrophytic types (B) of Western Podillia groups. Table 3: Povprečne indikatorske vrednosti ekoloških dejavnikov in njihove možne spremembe ob povprečnem porastu temperature za +1, +2, +2,5 and +3,0 °C. Minimalne vrednosti, s katerimi smo izračunali mejne vrednosti, so označene z zeleno, maksimalne vrednosti z modro. Razpon povečanja ogoženosti je obarvan z rumeno, območje z možnostjo izgube pa z rdečo. Različice smo izračunali v odvisnosti od razmerja med indikatorji različnih (A) in petrofitskih (B) tipov za skupine iz zahodnega Podillia. Acronyms: Hd (soil humidity), fH (variability of damping), Rc (soil acidity), Sl – total salt content in soil, Ca – calcium/magnesium content in soil, Nt – nitrogen content in soil, Ae – soil aeration, Tm – thermal climate, Om – ombroregime, Kn – continental climate, Cr – cryoclimate, Lc – light in the communities. 21/2 • 2022, 235–252 244 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Based on the obtained indicators of species participa- tion in specific communities (Table 4), we calculate the average values for associations and the proportions of their ratio, where the category of competitors (C) = 1. uartio auctae-Festucetum pallentis associations of the Galio campanulati-Poion versicoloris alliance, Stipo pulcherrimae- Festucetalia pallentis order. The third group reflects deriva- tive stages of developing dry grassland communities, but we could not classify them and will refer to them as tran- sitional communities from Festuco-Brometea to Trifolio- Geranietea. According to the results we propose syntaxo- nomic scheme of the communities with the occurrence of Festuca pallens Cl. Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 Ord. Stipo pulcherrimae-Festucetalia pallentis Pop 1968 All. Galio campanulati-Poion versicoloris Kukovitsa et al. 1994 ex Didukh & Vasheniak 2018 Ass. Schivereckio podolicae-Seselietum libanoti- dis Didukh & Vasheniak 2018 Ass. Minuartio auctae-Festucetum pallentis Onyshchenko 2001 Com. transitional communities from Festuco- -Brometea to Trifolio-Geranietea The Schivereckio podolicae-Seselietum libanotidis associa- tion has already been known in the Western and Central Podillia (Didukh & Vasheniak, 2018), but without men- tion of Festuca pallens. According to our present results, these communities with this fescue species develop ex- plicitly on Tortonian gypsum rocks exclusively in Opillia (geomorphological part of Western Podillia). The Minuartio auctae-Festucetum pallentis associa- tion (Table S2) was described by Onyshchenko (2001) from Krements Ridge as part of the Medobory Nature Reserve within the Alysso-Sedion alliance, order Alysso- Sedetalia, class Sedo-Scleranthetea. We (Vasheniak et al., 2021), however, transferred it to the class Festuco-Brom- etea and it is now considered within the alliance Galio campanulati-Poion versicoloris, order Stipo pulcherrimae- Festucetalia pallentis, because perennial plants dominate (Allium lusitanicum, Asperula cynanchica, Carex humilis, Elytrigia intermedia, Euphorbia cyparissias, Festuca pal- lens, Festuca stricta subsp. sulcata, Koeleria macrantha, Medicago falcata, Potentilla incana, Sedum acre, and Sca- biosa ochroleuca), although therophytes are present in with high frequency (Arenaria serpyllifolia, Clinopodium Syntaxon Proportion C S R C S R Schivereckio podolicae-Seselietum libanotidis 9 5 1 1 : 0.6 : 0.2 Minuartio auctae-Festucetum pallentis 10 6 2 1 : 0.6 : 0.2 Transitional communities from Festuco-Brometea to Trifolio-Geranietea 15 6 1 1 : 0.4 : 0.1 Average values 11.3 6 1 1 : 0.5 : 0.06 Table 4: Numbers of species and proportions in distinguished syntaxa by the Grime’s CSR strategy. Tabela 4: Število vrst in deleži v posameznih sintaksonih glede na CSR strategije po Grimu. Figure 4: Distribution of species by proportions according to the indicators of Grime’s CSR strategy (A) and the position of the “center of their masses” (B), which determines the patterns of development of the communities. 1-Schivereckio podolicae-Seselietum libanotidis, 2-Minuartio auctae-Festucetum pallentis, 3-transitional communities from Festuco-Brometea to Trifolio-Geranietea. Slika 4: Deleži vrst glede na CSR strategije po Grimu (A) in položaj njihovega »težišča«, ki opredeljuje vzorec razvoja teh združb. 1-Schivereckio podolicae-Seselietum libanotidis, 2-Minuartio auctae- Festucetum pallentis, 3- prehodna združba med razredoma Festuco- Brometea in Trifolio-Geranietea. Discussion Phytosociological peculiarities of communities with the presence of Festuca pallens As we can see from Table 1, the communities are divided into three groups. The first and second groups reflect the communities of the Festuco-Brometea class, presented by the Schivereckio podolicae-Seselietum libanotidis and Min- 21/2 • 2022, 235–252 245 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine acinos, and Saxifraga tridactylites) and herbaceous layers with sparse cover. These communities occur mainly on Sarmatian limestones of the Volyn-Podillia Upland. The floristic composition of the third group is very diverse. Although most species belong to the Festuco-Brometea class, they are acutely disturbed and characterize the stages of colonization of carbonate deposits where erosio- philes are present (Anthericum ramosum, Echium vulgare, Inula ensifolia, Salvia verticillata, Teucrium chamaedrys, T. montanum, and Vincetoxicum hirundinaria), along with forest edge species (Geranium sanguineum, Polygo- natum odoratum, Vicia cracca) and even shrubs (Cytisus albus, C. paczoskii, Geranium sanguineum, Lembotropis nigricans). It is obvious that there are two possibilities: closed steppe communities and derivative stages of their formation, in which Festuca pallens thrives in the absence of competition or with less competition. To confirm the violation of living conditions, the initial stages of com- munity formation indicate the results of processing by the expert system HES (Janišová et al., 2007), where three relevés from the group were recognized as Trifolio- Geranietea class. On the other hand, the EuroVegCheck- list Expert System (Mucina et al., 2016) shows that all relevés belong to the Festuco-Brometea class (Table S2). In our view, these communities lose distinct syntaxo- nomical affiliation and move to transitional communi- ties, but they retain the floristic core of steppe species and can be classified within Festucion valesiacae alliance, order Festucetalia valesiacae, at this current stage. Moreover, the core of the communities with the occur- rence of Festuca pallens is considered as Galio campanula- ti-Poion versicoloris alliance, which according to Didukh et al. (2021) is vicarious to Bromo pannonici-Fectucion csikhegyensis alliance occurring in the Pannonian province (Svydovets Mountains in Ukraine). Moreover, the highest concentration of relevés from the second cluster was rec- ognized as Bromo pannonici-Fectucion csikhegyensis alliance by the Slovak expert system HES (Janišová et al., 2007), confirming our previous point. Thus, these petrophytic communities are clearly different from the communities that occur in Central Europe (Didukh et al., 2021). It is noteworthy that communities with the presence of Fes- tuca pallens are more widespread in Central Europe than in Eastern Europe within the Bromo pannonici-Fectucion csikhegyensis and Diantho lumnitzeri-Seslerion albicantis alliance. Although these alliances have been mentioned in Ukraine (Janišová et al., 2014; Didukh et al., 2021), this is probably due to the fact that the communities with Festuca pallens are located at the eastern limit of the range and belong to other alliances that differ in their environ- mental conditions from the communities within the opti- mum environment in Central or Southern Europe. Ecological assessment of Festuca pallens habitats, threats to the existence of the species and forecast of possible changes Most habitats of Festuca pallens are vulnerable and could change under the influence of various ecological factors. The communities of the Minuartio auctae-Festucetum pal- lentis and Schivereckio podolicae-Seselietum libanotidis asso- ciations grow on initial soils that developed on gypsum and limestone and are highly dependent on the chemical com- position of the parent rock (Didukh & Vasheniak, 2018). Obviously, initial soils differ by their acidity, because Schiv- ereckio podolicae-Seselietum libanotidis communities grow on gypsols with higher acidity than on epilithic soils where Minuartio auctae-Festucetum pallentis communities occur. It should be added that calcium carbonate on limestone rocks is easily converted to soluble forms of calcium bicar- bonate that reduce acidity, which is not typical of gypsum formed from poorly soluble calcium sulfate. Moreover, ac- cording to our analysis, the communities of the association Schivereckio podolicae-Seselietum libanotidis occur in the southern part of the Volyn-Podillia Upland in the Velyki Goldy Ridge and Opillia and Minuartio auctae-Festucetum pallentis in the northern part of the Volyn-Podillia Upland in the Krements Ridge and Volyn Upland. This fact causes them to differ in cryoregime vectors (Figure 3). On the other hand, the shaded transitional communities grow on more or less developed rendzic leptosols with a thickness of more than 5 cm, rich in organic matter and in which there are soluble forms of calcium bicarbonate and other anions. Not only herbaceous plants, but also shrubs (Cytisus albus, C. paczoskii) and young trees (Pinus nigra) can settle in such groups, indicating a successive change in the direction of afforestation of these particular grasslands. The next step of our analysis was to evaluate the threats to the existence of these communities. Since they belong to the E1.2 petrophytic type habitats (Perennial calcareous grasslands and basic steppes), they are not threatened by any anthropogenic impacts, except for possible mechani- cal destruction or overgrowth of shrubs (Berberidion) (Korotchenko, 2011). Nevertheless, the question of pos- sible changes related to climatic factors is important. It is known that differentiation, function and development of communities are determined by hydrothermal conditions (temperature and precipitation). Based on the research of climatologists and the forecasts developed by them, an in- crease in average annual temperatures is observed, while changes in annual precipitation, both in the past and in the future, have a fluctuating character (Krakovskaia et al., 2009). Considering this fact, as well as the peculiarity of 21/2 • 2022, 235–252 246 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine carbonate substrates, where precipitation is not delayed, the peculiarities of the groups adapted to the conditions of moisture deficiency, as well as the high resistance to en- vironmental changes, we focused our calculations on the assessment of temperature changes. At the same time, we assumed that such climate changes act as a trigger mech- anism that affects a change in edaphic properties of the soil, and such an indirect effect is much more significant than direct effects (Didukh et al., 2016). Based on the correlations between the changes in average annual tem- peratures, thermoregime of humidity of the climate, and continentality cryoregime, as well as indicators of edaphic factors (soil humidity and salt regime), corresponding cal- culations were carried out on the change of indicators at an increase in average annual temperatures by +1, +2, +2, 5, and +3 °C (Table 3) (Didukh, 2021). At an increase of +1 °C, the average thermoregime is already beyond the amplitude of the modern existence of these communities, and with changes in temperature conditions, these groups fall into the risk zone of reduction. They are also in such a zone at a temperature increase of +2 °C, and at an increase to +2.5 °C and +3 °C they fall into the risk zone of exist- ence and are threatened with extinction. Accordingly, they fall into the risk zone of reduction in terms of cryoclimate indicators, when the temperature rises +2 and +3 °C, and the humidity of the climate and the acidity of the soil at +2, and losses - /+ 3 °C. The risk zone of reduction in rela- tion to the change of salt regime is much broader and is established when the temperature rises above 2 and 3 °C. On the other hand, indicators of factors such factors as continentality, variability of damping, carbonate and ni- trogen content, soil aeration with increasing temperature do not limit these communities. In contrast to more closed communities formed on de- veloped soils with stronger competition (forest, steppe, meadow), where the impact of climate change is indirect and manifested by changes in edaphic factors, the direct climate in this case shows a stronger impact of change, especially in terms of increasing temperatures and PAR (phytosynthetically active radiation) with the change in edaphic factors. To develop predictions of potential changes in these communities, we use the Grime spe- cies ecosystem indicators (CSR), which reflect ecosystem functioning, development, and sustainability based on as- similation, accumulation, and transfer of environmental resources, energy, conservation, and transmission of ge- netic information (Didukh et al., 2016). As shown in Figure 4, all indicators in the CS zone are somewhat closer to C (competitors) than to S (stress toler- ants) and very far from R (ruderal species). This could be due to the fact that their development is possible due to successional mechanisms, but limited by extremes. This is due to the carbonate substrate where resource use is limited and calcareous species are narrowly specialized (Didukh, 2011). It should be mentioned that the location of the Schivereckio podolici-Seselietum libanotidis and Festuco- Brometea to Trifolio-Geranietea transitional communities is closer to the apex C of the triangle than the location of the typical petrophytic communities of the Minuartio auctae- Festucetum pallentis association. The following specializa- tion of calcareous stress-tolerant adaptive mechanisms, including the introduction of new species and successional changes, could determine the structure and development of these communities existing in extreme conditions. Conclusions Festuca pallens is distributed only in the Volyn-Podillia Upland in Ukraine and thrives on the eastern limit of its range. Populations of small sizes in isolated locations have been found on dry gypsum, limestone and chalk sub- strates. The communities in which Festuca pallens occurs used to belong to the associations Schivereckio podolicae- Seselietum libanotidis and Minuartio auctae-Festucetum pallentis, to the alliance Galio campanulati-Poion versicolo- ris, to the order Stipo pulcherrimae-Festucetalia pallentis and to communities in derivative stages of development of steppe vegetation. These communities are ecologically distinct from those found in Central and Southern Eu- rope. According to the method of synphytoindication, the environmental conditions of the communities were presented and it was also calculated that in the case of a + 2 °C increase in the average annual temperature, a change in the living conditions is possible, which will lead to a decrease in the population of the area, and in the case of a +3 °C increase, to its loss. Successive changes in these communities were limited by extreme conditions caused by carbonate substrates that limit resource use. Festuca pallens is a rare species in Ukraine and is listed in the Red Data Book of Ukraine, and the communities where Fes- tuca pallens occurs are listed in the Green Data Book of Ukraine. In addition, the habitats are listed in Resolution 4 of the Bern Convention as E1.2 Perennial calcareous grasslands and basic steppes and in Annex I of the Habitat Directive 92/43 as 6190 Rupicolous pannonic grasslands (Stipo-Festucetalia pallentis) and need to be protected. In our opinion, management measures should include further steps: the introduction of sustainable grazing by cows, goats and horses on Festuca pallens habitats; the im- plementation of an information campaign against spring burning; the removal and control of invasive plants in the habitats; the prevention of inappropriate afforestation of steppe slopes; the fight against illegal mining of limestone and gypsum on Festuca pallens habitats. 21/2 • 2022, 235–252 247 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Yakiv P. Didukh  https://orcid.org/0000-0002-5661-3944 Iuliia Vasheniak  https://orcid.org/0000-0003-1020-3007 Iryna Bednarska  https://orcid.org/0000-0002-2520-0511 Acknowledgements We are sincerely thankful to Dr. Svitlana Nyporko for de- termination of the bryophytes and Dr. Valerii Darmostuk for the determination of the lichens from our expeditions. 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Appendix List of localities of Festuca pallens Host according to her- barium funds (asterisk indicates the localities where the current growth of the Festuca pallens is not confirmed). Ternopil region, Krements Ridge: Bozha hill* (s. coll. 1858 LE; Rogovych 1898 LE; Zapiatova 1961 KW, s. coll. 1961 KW; Bednarska 2005 LWKS IB-896), Divochi Skeli tract (Divocha hill) (Mądalski 1936 LWS 32733; Gryn 1940 KW; Kotov, Temko 1954 LE; Klokov, Zaverukha 1958 KW; Prokudin, Shustova 1958 CWU; Lazebna 1976 LWS 32734, 32735, 32736; Kagalo 1983 LW 011664; Kagalo 1984 LW; Kagalo, Gynda 1988 LWKS 01630; Kagalo, Bednarska 1998 LWKS 8492; Bednarska 2005 LWKS IB-893), Chercha hill (Mryts 1936 LWS 32948; Zapiatova 1961 KW), vicinity of Zholoby and Pidlistsi vil- lages, Strakhova hill (Gryn 1940 KW; Iermachenko 1958 CWU; Klokov, Zaverukha 1958 KW; Zaverukha 1959 KW 009190, 005497, 005498; Zaverukha 1959 LWKS 201; Prokudin, Tveretynova 1964 LE; Bednarska 2005 LWKS IB-901), vicinity of Krements town (Rogovicz, 1952 LE; Barbarych, Kucheriava 1958 KW; s. coll. 1893 LE), vicin- ity of Kulykiv village, Lysa hill (Bednarska 2005 LWKS IB- 1593; Kagalo, Skibitska, Resler 2006 LWKS 25205; Bed- narska, Kagalo 2007 LWKS IB-1167, IB-1168), vicinity of Mykolayiv village, Lysa hill (Klymyshyn 1987 LWS 18439, 18440), Pidvolochysk district, Ostapie village* (Wołoszczak 1889 LW 012060:“Ostapie, in montibus Miodobory. Ex- empl. cultur Leopoli”; Błocki s. dat. LWS 10.507), Shumsk district, Tsetsenivka village* (s. coll. 1958 KW). Ivano-Frankivsk region, Galych district: Mezhygirtsi vil- lage [single plants in semi-degraded habitat] (Kuziarin 2000 LWS), Podillia village (Zaverukha 1976 KW 1 sheet without number + sheet № 005835; Kagalo, Bednarska 1998 LWKS 8896; Bednarska, Nakonechyi 2000 LWKS IB-181, 11598; Kagalo, Bednarska 2014 LWKS IB-1677); Rogatyn district: Luchyntsi village, Velyki Goldy tract (Shevchuk 1961 LWS 10.566; Lazebna 1978 LWS 32740, 32741; Kagalo 1988 LWKS 020546; Kagalo, Bednarska 1998 LWKS 8489; Bed- narska, Nakonechyi 2000 LWKS IB-191). Rivne region, Dubno district: Mylcha village (Didukh 2000 KW 008689; Bednarska, Kagalo 2007 LWKS IB- 1173c). 21/2 • 2022, 235–252 249 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Ta bl e 2: P hy to so ci ol og ic al ta bl e of th e co m m un iti es w ith th e pr es en ce o f F est uc a pa lle ns . S pe ci es a re a rr an ge d ac co rd in g to d ec re as in g ph i-v al ue s a nd a cc or di ng to d ec re as in g ov er al l c on sta nc y fo r n on -d ia gn os tic sp ec ie s. Ta be la 2 : F ito ce no lo šk a ta be la zd ru žb z vr sto F est uc a pa lle ns . V rs te so ra zv rš čn en e pa da jo če g le de n a fi vr ed no st, n ez na či ln e vr ste p a pa da jo če g le de n a sk up no st al no st. Ta bl e nu m be r 33 34 32 45 46 35 28 43 44 42 29 30 31 23 26 27 24 25 18 21 19 22 17 16 6 10 5 8 4 7 1 2 3 41 15 11 12 14 9 36 38 40 39 37 Re le ve n um be r ( T U R BO V EG ) 70 71 69 204 208 72 65 202 203 201 66 67 68 60 63 64 61 62 55 58 56 59 54 53 43 47 42 45 41 44 38 39 40 200 52 48 49 51 46 97 99 101 100 98 C ou nt ry c od e U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A U A N r. ta bl e in p ub l. 16158 16159 16157 16161 16149 16153 16154 16155 7021 7040 7048 7035 7037 N r. re le vé in ta bl e 1 1 1 1 1 1 1 1 1 1 1 1 1 56 61 59 62 58 57 46 50 45 48 44 47 41 42 43 55 51 52 54 49 10 0 10 3 10 5 10 4 10 1 C ov er a bu nd an ce sc al e 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 D at e (y ea r/ m on th /d ay ) 20160806 20160806 20160806 20160806 20160806 20160806 20160805 20160806 20160806 20160806 20160805 20160805 201608 20070502 20070620 20070620 20070620 20070620 20180628 20180628 20180628 20180628 20180628 20180628 20180606 20180627 20180606 20180606 20180606 20180606 20180606 20180606 20180606 20160806 20180627 20180627 20180627 20180627 20180606 20200813 20200813 20200813 20200813 20200813 Re le vé a re a (m 2) 16 16 16 10 10 16 16 10 10 10 16 16 16 16 16 16 16 16 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 Al tit ud e (m ) 200 200 205 299 342 340 265 237 277 273 270 278 314 290 205 205 200 200 371 391 387 376 374 368 371 381 367 379 375 363 373 372 375 230 370 391 377 378 373 214 235 212 234 231 As pe ct (d eg re es ) 350 350 180 40 205 270 340 310 330 180 270 270 180 270 230 180 180 260 298 40 140 130 160 270 280 230 210 240 240 240 70 200 220 200 220 170 200 280 275 170 210 250 229 Sl op e (d eg re es ) 90 90 80 40 90 60 60 45 30 30 5 15 10 75 75 10 15 20 35 5 70 5 5 3 27 5 5 3 5 30 15 25 30 15 21 15 10 11 20 C ov er to ta l ( % ) 70 50 40 60 60 40 40 60 60 60 30 50 50 30 30 30 40 30 80 50 80 80 80 80 70 80 80 80 90 80 90 90 90 80 80 80 80 80 80 30 60 60 60 60 C ov er sh ru b la ye r ( % ) 5 5 5 5 5 C ov er h er b la ye r ( % ) 70 70 40 40 50 70 40 45 40 40 30 15 40 30 30 30 40 30 60 20 60 30 60 60 50 60 60 50 85 60 70 70 60 60 60 50 75 60 70 20 40 50 55 55 C ov er m os s l ay er (% ) 20 20 20 20 10 20 5 15 20 20 40 20 20 30 20 50 20 20 20 10 20 30 5 20 20 20 30 20 20 20 5 20 10 10 20 10 5 5 C ov er li tte r l ay er (% ) 2 2 10 30 10 10 10 10 10 10 10 10 10 C ov er b ar e ro ck (% ) 50 50 50 20 60 70 70 50 70 80 80 70 80 Av er . h ei gh t ( hi gh ) h er bs (c m ) 10 01 00 80 80 10 0 50 10 0 80 15 15 15 15 15 Av er . h ei gh t l ow es t h er bs (c m ) 10 10 20 1 3 30 20 2 2 1 30 20 20 15 10 10 15 10 1 M ax im um h ei gh t h er bs (c m ) 54 80 50 99 30 58 51 67 25 53 53 50 50 20 50 25 50 81 85 50 50 80 80 80 50 80 35 39 10 8 81 61 M os se s i de nt ifi ed (y /n ) Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y N N N N N Li ch en s i de nt ifi ed (y /n ) Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y Y N N N N N lo ng itu de 24.736 24.736 24.736 24.742 24.484 24.770 24.652 24.740 24.740 24.740 24.669 24.669 24.670 24.442 25.724 25.724 25.634 25.725 50.094 50.095 50.094 50.094 50.094 50.093 50.119 50.074 50.118 50.117 50.118 50.118 50.117 50.118 50.118 24.670 50.085 50.080 50.079 50.085 50.080 25.566 25.567 25.567 25.566 25.566 la tit ud e 49.276 49.276 49.276 49.276 49.118 49.120 49.338 49.277 49.277 49.277 49.284 49.284 49.284 49.405 50.118 50.118 50.056 50.117 25.740 25.732 25.733 25.733 25.734 25.737 25.727 25.644 25.727 25.729 25.727 25.728 25.730 25.729 25.729 49.284 25.674 25.645 25.645 25.674 25.644 50.354 50.355 50.356 50.355 50.354 H ES FBB01? FBB01? FBB01? FBB01? TGA01? TGA01? FBB01? FBB01? FBB05? FBB05? FBB05? FBB05? FBC02? FBA02? FBA02? FBA02? FBA02? FBB05? FBB04? FBB01? FBB01? FBB01? FBB01? FBB04? FBB01? FBB05? FBC02? FBB05? FBB02? FBB04? FBC01? FBC01? FBC02? TGA01? TGA01? TGA01? TGA01? FBB05? FBC02? TGA04? FBB01? FBB05? FBB02? FBB05? Eu ro Ve gC he ck lis t FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES SED SED FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES FES Re le ve ID 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 21/2 • 2022, 235–252 250 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Ta bl e nu m be r 33 34 32 45 46 35 28 43 44 42 29 30 31 23 26 27 24 25 18 21 19 22 17 16 6 10 5 8 4 7 1 2 3 41 15 11 12 14 9 36 38 40 39 37 Re le ve n um be r ( T U R BO V EG ) 70 71 69 204 208 72 65 202 203 201 66 67 68 60 63 64 61 62 55 58 56 59 54 53 43 47 42 45 41 44 38 39 40 200 52 48 49 51 46 97 99 101 100 98 Fe stu ca p al len s r + r r r r 2 2 2 1 2 2 2 2 2 1 2 1 2 2 2 2 2 2 2 2 2 r r 2 2 r 2 r 2 2 2 2 r 2 2 3 r r A ss . S ch iv er ec ki o po do lic ae -S es el ie tu m li ba no ti di s G yp so ph ila th yr ai ca . . 2 . + 1 2 2 2 r 2 1 2 1 . . . . . . . . . . . r . . . . . . . r r . . 2 r . . . . 2 Th ym us p an no ni cu s . . . . . . . r + r r r . r r + r . . . . . . . . . . . . . . . . r r . . . . . . . . . C am pa nu la ro tu nd ifo lia + 1 . r . . . r r r . . . r . . r r . 2 r . . . . . . . . . . . . . . . . . . . . . . . As tra ga lu s o no br yc hi s . . r . r + . . r . r r . . . r . . . . . r r r . . . . . . . . . 2 . . . . . . . . . . Ar ab id op sis a re no sa r . . r r r + r . . . . r . . . r . . . . . . . . . . . . . . . . . . . . . . . . . r r A ss . M in ua rt io a uc ta e- Fe st uc et um p al le nt is C lin op od iu m a cin os . . . . . . . . . . . . . . r . . . r r r r r . . r r 2 r . r r r . . . . . . . . . . r Th ym us se rp yll um . . . . . . . . . . . . . . . . . . + 2 r r . . 3 . r + + r r . r . . . . r . . . r . . Al ys su m g m eli ni i . . . . . . . . . . . . . . . . . . . r r r r r . r r r . . . r + . . . . . r . . . . . M in ua rt ia se ta ce a su bs p. se ta ce a . . . . . . . . . . . . . . . + . . . r + r 2 . 2 . . r r 2 r . 2 . r . . r . . . . . . M ed ica go fa lca ta . . . . + + . . r . . . . . . . + . r . . r . r r r . r r . r . . r . . . . . . . . . . C en ta ur ea st oe be . . . . r r . . . . . . . . . . . . . . r r r r . . r r . . r . r . r . . r . . . . . . An th yll is vu ln er ar ia . . . . . . . . . . . . . . . . r . . . . . . . r r r r . r . . . . . . . . . . . . . . Sa xi fra ga tr id ac ty lit es . . . . . . . . . . . . . . . . . . r r + r . . . . . . . . . r . . . . . . . . . . . . Si len e e ug en ia e . . 1 . . . . . . r . . . . . . . . . . . . . . r . r r r . . . . . . . . . . . . . . . tr an si ti on al c om m un it ie s f ro m F es tu co -B ro m et ea to T ri fo lio -G er an ie te a An th er icu m ra m os um . . . . . . . . . . . . . . . . . . . . . . . . . . 2 + + . + . . + 2 2 2 2 2 . . 2 2 2 Te uc riu m ch am ae dr ys . . . . . . . . + . . . . . r . . . . . . . . . . . . r . . . . . . . r r r r r r r . . St ip a ca pi lla ta . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . r r r r r r . . . . . Sa lv ia v er tic ill at a . . . . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r + r + r G er an iu m sa ng ui ne um . . . . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . r r r r r . . . . . . Cy tis us p ac zo sk ii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r r r r 2 . . . . . G al iu m b or ea le su bs p. ex ol et um . . . . . . . r r r . . . . . . . . r . . . r . . . . . . . . . . r . . . . . r . r + r Te uc riu m m on ta nu m . . . . . . . . . . . . 1 . . . . . . . . . . . . r r . . . . . r . . 2 + . r . 2 . 2 . Th al ict ru m m in us . . . . . . . . . . . . . . . . . . r . . . . . . r . . . . . . . . r r . r r . . . . r G al iu m m ol lu go . r . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r r r r . Pi m pi ne lla sa xi fra ga . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . + r . r r St ac hy s r ec ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . r r . r r Cy tis us a lb us . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . . . . . r r . . 2 . . . . . Le m bo tro pi s n ig ric an s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . 2 r . . . . . . A ll. G al io c am pa nu la ti -P oi on v er si co lo ri s, O rd . S ti po p ul ch er ri m ae -F es tu ce ta lia p al le nt is , C l. Fe st uc o- B ro m et ea Po te nt ill a in ca na . . . . 1 1 . . r . . + . r r . . r . . r . r 2 . . r 2 2 2 2 2 r . 2 2 r r . r . + + + Al liu m lu sit an icu m . + + . 1 1 1 r r r 2 2 2 . . . . + . . . . . . 2 2 r 2 . 2 2 2 2 r . r r . . . . . . . Se du m a cr e 1 + . . . . . . . . r r 3 + + + + . 2 + + 2 + . + r . r r 2 + r . . . . . . . . 2 . . . H eli an th em um ca nu m . . . . . . . . . . . . . . + 1 . . . . . . . . r . r . r r . . r . r r r 2 2 . . 2 r r 21/2 • 2022, 235–252 251 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Ta bl e nu m be r 33 34 32 45 46 35 28 43 44 42 29 30 31 23 26 27 24 25 18 21 19 22 17 16 6 10 5 8 4 7 1 2 3 41 15 11 12 14 9 36 38 40 39 37 Re le ve n um be r ( T U R BO V EG ) 70 71 69 204 208 72 65 202 203 201 66 67 68 60 63 64 61 62 55 58 56 59 54 53 43 47 42 45 41 44 38 39 40 200 52 48 49 51 46 97 99 101 100 98 C ar ex h um ili s . . . . . . . . 2 . . . r . . . . . . . . . . . . r 2 2 2 . . . r 2 2 + . 2 2 . . . 2 r M ela m py ru m a rv en se . . . . r r . . . . . . . . . . . . . . . . . . . . r r . . r r r r . . . . . . . . . . In ul a en sif ol ia . . . . . . . . . . + r r . . . . . . . . . . . . r . . . . . . . r . r r . . . . . . . G al iu m g la uc um . . . . . . . . . . + 1 + . . . . . . . . . . . . . . . . . . . . r . r r . . . . . . . Ir is ap hy lla su bs p. h un ga ric a . . . . 2 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . r . . . . . . . Se m pe rv iv um ru th en icu m 2 3 3 2 . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . C l. Fe st uc o- B ro m et ea As pe ru la cy na nc hi ca r . . . r r . r r . r r r . 1 . + r . . . . r r . r r . + . . r . r r r . r r . . . . r Ar te m isi a ca m pe str is . + 2 . 2 2 1 . . . + . r . r r + . r r r r . r . . r r r . r r . + . . . . . . . . . . Fe stu ca st ric ta su bs p. su lca ta 2 2 . r r r . . r r . . . . . . . . . . 2 r . r 2 . . r . . r r . . . . . . . . r . . . C am pa nu la si bi ric a . . . . . . . . . r r r r . . . . r 2 . . . . . . r . . . r . r . . . . . r r . . r . r El yt rig ia in te rm ed ia r . . . + 1 + r . . . . . . . . . . r . . r r r r r . . . . . . . . r r . . . . r . r . Eu ph or bi a cy pa ris sia s . . . . r r 1 r r r 1 + + . . . r . . . . . . . . r . . . . r r . r r . r . . r r r r r Ar en ar ia se rp yll ifo lia 2 1 . r r r + . . . . . . . . . . . . . . r r r . . r . . . . r . . . . . . . . . . . . Sc ab io sa o ch ro leu ca . . . . . . . . . . . . . . . . . . r . r . r . . r . r r r r r r . r . . r r . r . r r Ko ele ria m ac ra nt ha . . . . + + 1 r r r . . . . . . r . . . r r . . r r . . . . r . . . . . . . . . + r . r H yp er icu m p er fo ra tu m . . . . . . . . r . . . . . . . r . . . . . . . . . . r . . . . . . . r . . . r r . r . Eu ph or bi a seg ui er ia na . . . . . . . . . . . . . . . . r + . . . . . . . . . . . r . . . . . r . 2 2 . r . . . O th er sp ec ie s Ve ro ni ca sp ica ta . . . . r r + r r r . . . . . . . . . . . . . . . . . r r . r r . r r . . . . . . . . . As pl en iu m ru ta -m ur ar ia . r . r . . . . r . . . . . . r . + r r r . . . . r . . . . . . . . . . . . . . . . . . G al iu m v er um . . . . r r r . r . . . . . . . . . . . . . r . . . . . . . r . . . . . . . . . r r . 2 Ec hi um v ul ga re . . . . . . . . . . . . . . . . . r . . . r . r . r . . . . . . . . . . r . . r . . r . Vi nc et ox icu m h iru nd in ar ia . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . r r . . . r r . r . . . . . C en ta ur ea p hr yg ia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r r r H ier ac iu m u m be lla tu m . . . . r r . r r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G er an iu m ro be rt ia nu m r + . r . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . . . . . . . Pe uc ed an um ce rv ar ia . . . . . . . r r r . . . . . . . . . . . . . . . . r . . . . . . . . . r . . . . . . . Se sel i l ib an ot is su bs p. in te rm ed iu m r r . r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . . . . . . C ry pt og am sp ec ie s ( in a lp ha be ti ca l o rd er ) Ab iet in ell a ab iet in a . . . 2 2 . . r r . . . . . . . . . . r 2 . . . r 2 2 r . 2 2 2 2 . . . . . . . 2 2 + . Ba rb ul a un gu icu la ta . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 + 1 2 . r . . . . . . 2 . . . . . Br ac hy th ec iu m a lb ica ns . . . 2 . . . r + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Br yu m a rg en te um . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . 2 . r . r . . . r . . . . . . Br yu m k un ze i . . . . . . . . . . . . . . . . . . 2 . 2 2 2 2 . r . 2 2 2 . . 2 . . . 2 + . . . . . . C er at od on p ur pu re us . . . . . . . . r r . . . . . . . . . . 2 2 . r r . + 2 . 2 + . . + . . . . . . . . . . C la do ni a po cil lu m . . . . . . . . . . . . . . . . . . . . . r . . . . . . . 2 r . . . . . r . . . . . . . 21/2 • 2022, 235–252 252 Didukh et al. Phytosociological and ecological peculiarities of Festuca pallens in Ukraine Ta bl e nu m be r 33 34 32 45 46 35 28 43 44 42 29 30 31 23 26 27 24 25 18 21 19 22 17 16 6 10 5 8 4 7 1 2 3 41 15 11 12 14 9 36 38 40 39 37 Re le ve n um be r ( T U R BO V EG ) 70 71 69 204 208 72 65 202 203 201 66 67 68 60 63 64 61 62 55 58 56 59 54 53 43 47 42 45 41 44 38 39 40 200 52 48 49 51 46 97 99 101 100 98 C la do ni a py xi da ta . . . . . . . 2 r r . . . . . . . . . . . . . . . . . . r . . r . . . . . . r . . . . . C ol lem a te na x . . . . . . . 1 2 2 . . . . . . . . 2 . . . . . r . . . . 2 . r . r . . . . . . . . . . D itr ich um fl ex ica ul e . . . . . . . . . . . . . . . . . . . 2 2 2 . . . 2 2 2 r 2 . r . . . 2 2 . + . . . . . En ca lyp ta v ul ga ris . . . . . . . . + . . . . . . . . . 2 . 2 2 2 2 . 2 . . . . . r . . . . 2 . . . . . . . H om al ot he ciu m lu te sce ns . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . 2 2 . . . . . . . . r . . . . H om al ot he ciu m se ric eu m . . . . . . . . . . . . . . . . . . 2 2 2 2 . . . . . . . . 2 r 2 . . 2 . . . . . . . . H yp nu m v au ch er i . . . . . . . . r r . . . . . . . . . 2 . . . . . . + . . 2 2 . . . . . . . . . . . . . Ps eu do cr os sid iu m h or ns ch uc hi an um . . . . . . . . . . . . . . . . . . . . + 2 . . . . . . . . . . 2 . . . . . . . . . . . Rh yt id iu m ru go su m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 + 2 2 . . . . . . . . . . . To rt ell a to rt uo sa . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . 2 . . . 2 . . 2 . . r . . To rt ul a ca lci co len s . . . . . . . . . . . . . . . . . . . . . 2 . . . . . 2 2 . . . 2 . . . . . . . . . . . To rt ell a in cli na ta . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 . 2 2 . . 2 . . . 2 2 2 . . . . . To rt ul a ru ra lis . . . + . . . . . . . . . . . . . . . . . r 2 . . . 2 2 . . 2 2 3 . . . . 2 . . 2 . . . W eis sia sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . 2 . . . r . Sp ec ie s o f v as cu la r pl an ts , r ar el y oc cu rr ed in th e co m m un it ie s: Ac er n eg un do (4 0: r) , A ch ill ea m ill efo liu m (3 4: r) , A do ni s v er na lis (3 4: r) , A lli um p od ol icu m (5 -6 :r; 3 2: r) , A . s tri ctu m (1 8: r) , A lys su m h irs ut um (1 7: r) , A . r os tra tu m (2 9- 31 :r) , A sp len iu m tri ch om an es (1 5: r; 31 :r) , A str ag al us a us tri ac us (2 5: r) , A ur in ia sp . ( 15 :+ ; 1 6: r; 18 :r) , B er be ris v ul ga ris (3 8: r; 44 :r) , B er te ro a in ca na (1 9: +; 2 8: r) , B et ul a kl ok ov ii (3 9: r) , B ot rio ch lo a isc h- ae m um (1 7: r) , B ra ch yp od iu m p in na tu m (4 :r) , B ro m us sq ua rr os us (2 2: r) , B up leu ru m fa lca tu m (3 4: r) , C ar ex h irt a (1 7: r) , C ar lin a ac an th ifo lia su bs p. u tz ka (3 9: r) , C en ta ur ea ja ce a (4 0: r) , C . s ca bi os a (3 5: r) , C er as tiu m a rv en se (2 :r; 1 4: r; 24 :r; 4 1: r) , C on vo lv ul us a rv en sis (4 3: r) , C or nu s s an gu in ea (4 2: r) , C yt isu s b lo ck ia nu s ( 8- 9: r) , C . r ut he ni cu s ( 27 :r; 3 1: r; 32 :r) , D au cu s c ar ot a (4 0: r) , D ia nt hu s a re na riu s s ub sp . p seu do ser ot in us (2 8: r; 32 :r) , D ict am nu s a lb us (3 5: r) , D ra ba v er na (2 0: r) , E pi pa cti s a tro ru be ns (3 8: r) , E ry sim um d iff us um (2 :r; 3 :r; 1 3: r) , E up ho rb ia ni ca ee ns is su bs p. st ep po sa (2 6: r; 27 :r; 2 9: r) , E . s tri cta (8 :r) , F ra ga ria v iri di s ( 40 :r) , F ra ng ul a al nu s ( 35 :r; 3 9: r) , G al iu m h um ifu su m (1 9: r; 24 :r) , H eli an th em um n um m ul ar iu m (3 1: r) , H eli cto tri ch on d ese rt or um (1 8: +) , H ol os te um u m be lla tu m (1 9: r) , H ylo te lep hi um m ax im um (1 :2 ; 2 :r; 7 :r) , H yp er icu m el eg an s ( 34 :r) , J ac ob ae a vu lg ar is (5 -6 :r; 3 4: r) , J ur in ea m ol lis (3 6: r; 39 :r) , L ap pu la sq ua rr os a (2 2: r) , L in ar ia v ul ga ris (4 1: r) , L in um c at ha rt icu m (3 7: r) , L ot us c or ni cu la tu s ( 40 :r; 4 2: r) , M ed ica go m in im a (1 9: r; 44 :r) , M ela m py ru m c ris ta tu m (1 1- 13 :r) , M eli lo tu s a lb us (1 :r) , M . o ffi cin al is (3 2: r) , N oc ca ea p er fo lia ta (1 9: r) , O no br yc hi s a re na ria (4 1: r; 44 :r) , O tit es sp . ( 17 :+ ), Pe uc ed an um or eo sel in um (3 7: r) , P ilo sel la offi cin ar um (4 1: r; 44 :r) , Pi nu s n ig ra (j uv .) (2 7: r; 39 :r) , P oa co m pr ess a (8 :r; 2 2: r; 40 :r) , P ol yg al a co m os a (4 2: r) , P ol yg on at um m ul tifl or um (2 6: r; 35 :r; 3 7: r; 38 :r) , P ol yg on at um o do ra tu m (5 :+ ; 6 :+ ), Pr un ell a gr an - di flo ra (3 7: r) , R hi na nt hu s m in or (5 -6 :+ ), Ro sa to m en to sa (5 :r) , R ub us ca esi us (3 4: r) , S al vi a nu ta ns (3 9: r) , S . p ra te ns is (3 4: r; 44 :r) , S ed um h isp an icu m (2 :r; 3 :r; 1 4: +) , S ol id ag o ca na de ns is (4 2: r) , S tip a pu lch er rim a (3 5: r; 37 :r) , Th esi um li no ph yll on (3 2: r; 39 :r) , T ra go po go n du bi us (2 6: r) , T rin ia m ul tic au lis (3 9: r) , U rt ica d io ica (4 :r) , V al er ia na tu be ro sa (2 6: r) , V er ba scu m ph lo m oi de s ( 23 :r; 2 4: r) , V er on ica p ro str at a (2 4: r) , V ici a cr ac ca (5 :r; 6 :+ ), Vi ol a ca ni na (4 3: r) C ry pt og am sp ec ie s, r ar el y oc cu rr ed in th e co m m un it ie s: Br yu m ca esp iti ciu m (3 2: r) , B ry um sp . ( 10 :r) , C am pt ot he ciu m lu tes ce ns (5 :1 ), C am py liu m ch ry so ph yll um (2 2: r; 37 :2 ), C an de la rie lla v ite lli na (9 :2 ), C la do ni a re i ( 28 :r) , C la do ni a sp . ( 19 - 20 :r) , C . s ub ul at a (8 :r) , C ol lem a cr isp um (2 0: r) , C ten id iu m m ol lu scu m (4 0: +; 4 4: +) , D icr an ell a he ter om al la (2 2: 2) , D icr an um b on jea ni i ( 10 :2 ), D id ym od on fa lla x (2 2: r; 27 :r) , D . v in ea lis (2 2: r; 27 :r) , D ip lo sch ist es m us co ru m ( 20 :r) , E nc al yp ta st re pt oc ar pa ( 26 :2 ), Ev er ni a m eso m or ph a (4 :r; 8 :r) , F iss id en s t ax ifo liu s ( 37 :2 ), G rim m ia p ul vi na ta ( 19 :2 ), G rim m ia s p. ( 28 :r) , G ym no sto m um ca lca re um (2 6: 2) , H yp nu m cu pr ess ifo rm e ( 21 :r; 2 6: r) , L ec an or a m ur al is (2 5: r; 27 :2 ), Le pr ar ia m em br an ac ea (2 2: r) , L ep to gi um te nu iss im um (3 2: +) , L esk ea p ol yc ar pa (3 1: 2) , M yu re lla ju la ce a (2 6: 2) , M . s ib iri ca (2 6: r) , O rth ot ric hu m a ffi ne (5 :+ ), O . c up ul at um (2 7: 2; 3 7: 2) , O rth ot ric hu m sp . ( 31 :2 ), Pe lti ge ra m al ac ea (4 :+ ), P. ru fes ce ns (3 3: r) , P hy sci a ten ell a (4 :+ ), Ra m al in a po lli na ria (4 :2 ; 8 :+ ; 9 :r; 1 0: r) , R . p ol ym or ph a (4 :+ ; 8 :+ ; 9 :2 : 1 0: +) , S ch ist id iu m a po ca rp um (3 3: r) , S ch ist id iu m sp . ( 20 :+ ; 3 1: r) , T on in ia se di fo lia (2 1: r) , T or tu la m ur al is (1 9- 20 :r)