Phytocoenological and Ecogeographical study of Salvia nutans in Romania

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It is vulnerable to overgrazing and the conversion of grasslands into arable land. Due to these anthropogenic impacts, the species is endangered in some locations in Romania. In this context, the study aims to find the plant associations in which the species occurs and the environmental factors that explain the floristic composition. For the vegetation analysis, 870 relevés were used, and the data were analyzed in JUICE programme, applying the modified TWINSPAN algorithm for clustering. The floristic composition data and environmental variables were used for the multivariate analyses. Salvia nutans was recorded most frequently in the Festuco-Brometea class, the Festucetalia valesiacae order, the Stipion lessingianae alliance, and the Jurineo transylvanicae-Stipetum pulcherrimae and Allio albidi-Stipetum lessingianae associations. Canonical Correlation Analysis indicates that annual precipitation (BIO12) explained most of the variation in floristic composition. S. nutans occurs predominantly on steep slopes with southwest aspects and at an average elevation of 359 m. The species occurs in moderately acidic to slightly alkaline soils, with moderate phosphorus and high potassium concentrations. Salvia nutans is related to topographic and edaphic conditions, showing its role as an indicator species for continental, xerophilic grasslands. Given its vulnerability to land-use changes, S. nutans should be included in long-term monitoring programs to assess population dynamics. anthropogenic impact biogeography dry grasslands habitat requirement Romanian flora Salvia nutans steppe relict Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Highlights is a steppe relict species vulnerable to land-use changes. The species occurs mainly in grasslands on steep, sunny slopes. Its abundance is influenced by annual precipitation and temperature. Biogeographic patterns suggest distinct regional lineages in Transylvania. Long-term monitoring is needed due to habitat degradation and hybridization risks. Introduction Biodiversity decline represents a significant ecological challenge, affecting ecosystem services and human well-being (Dirzo and Raven 2003 ; Ceballos, García, and Ehrlich 2010 ; Isbell et al. 2023 ). Current species extinction rates far exceed historical natural rates, reaching levels that suggest the sixth major extinction event on Earth is underway (Barnosky et al. 2011 ; Ceballos, García, and Ehrlich 2017). This alarming acceleration underscores a conservation crisis where identifying the mechanisms driving biodiversity decline is a key priority for conservation biology (Didham et al. 2007 ). The causes of this decline are multiple, involving direct human activities and indirect environmental changes (Adla et al. 2022 ). Habitat loss, overexploitation, climate change, pollution, and the introduction of invasive species are the primary factors contributing to biodiversity decline, alongside other secondary threats (Maxwell et al. 2016 ; Caro et al. 2022 ). In particular, invasive species are frequently classified as the second most significant source of threat to biodiversity (Dueñas et al. 2018). One of the plant species with restricted habitats is Salvia nutans L., a steppe relict (Gusztáv 2005 ; Sramkó and Laczkó 2020 ) and postglacial in the Carpathian Basin (Németh 1990 ; Simon 1992 ; Đakić, Knežević, and Boža 2012 ). The species S. nutans belongs to the Lamiaceae Martinov family (Sârbu, Ștefan, and Oprea 2013 ) and grows in a restricted range of habitats. Its habitat includes steppe and forest-steppe grasslands, where it co-occurs with various plants, ranging from herbaceous species to dicotyledons. These areas are characterized by chernozem soils on loess and a climate with continental influences. In natural conditions, the habitat exhibits a rich diversity of species, whereas, in degraded conditions, this diversity decreases, with the predominance of monocotyledons and grasses (Soó 1968 ; Csurös 1973, 1974; Gusztáv 2005 ). The species is also found in rocky areas and rock outcrops (Golevich, 2012 ), as well as in grasslands and scrublands (Sârbu, Ștefan, and Oprea 2013 ). From its phytocoenotic perspective, S. nutans occupies subdominant positions or is a central component of rank 1–2 in plant communities (Ziman 1976 ). There are places where this species dominates the vegetation layer (Kondratyuk et al. 1985 ). Salvia nutans is found in Central and Eastern Europe, from the Danube Plain in the west to the Pre-Caucasus in the east (Golevich 2012 ). Thus, S. nutans occurs in Ukraine's southern and central regions, extending from the Crimean Peninsula and the northern Caucasus to the Volga bend. It then continues along the Ural River and the Ural Mountains to the Kama River region (Golevich 2012 ). Its distribution also extends to the outer edges of the Carpathians, from Galicia to the Danube (Csapody 1982 ), with occurrences in the Transylvanian Basin, Vojvodina, and the Great Plain. The western limit of its distribution reaches Hungary (Sramkó and Laczkó 2020 ). The species is also widespread in eastern Romania, more significant in the Transylvanian Basin (Soó 1942 ; Beldie et al. 1961 ) and in Serbia. Historical data indicate its presence in specific locations in Hungary and Romania, suggesting a broader distribution in the past (Gusztáv 2005 ). A wider distribution of the species S. nutans has been recorded throughout the Pannonian Plain, including areas between the Danube and the Tisza, as well as Vojvodina (Soó 1942 ). In Serbia, the species is no longer found (Jovanović and Lakušić 2006 ; Đakić, Knežević, and Boža 2012 ). However, in the Transylvanian Basin, the species has been confirmed in some locations (Soó 1942 ; Ruprecht et al. 2009 ). S. nutans reaches eastwards to the southern limits of the Ural Mountains, although it is no longer part of the current flora of Kazakhstan (Sramkó and Laczkó 2020 ). The main threat to the survival of S. nutans is the ongoing transformation of its natural habitat, especially grasslands. Also, the collection of this species for its ornamental value is a major contributor to the population decline. The decline in the number of individuals resulting from these combined pressures also poses additional risks, including genetic hazards, which may impact the long-term viability of the population (Gusztáv 2005 ). Salvia nutans has not been extensively evaluated for its conservation status at the global and European levels. However, given its presence across various European regions, local assessments might provide insight into its vulnerability or stability within specific habitats. In Romania, for instance, the species may be categorized under specific conservation statuses based on local biodiversity studies, mirroring the approach used for other flora and fauna. For hypothetical context, let us consider that S. nutans is vulnerable in specific habitats due to environmental pressures or anthropogenic factors. In Central and Southeastern Europe, where S. nutans naturally occurs, agricultural expansion and changes in land management practices could influence its conservation status. The study aims to investigate the habitat and phytocoenological preferences of the S. nutans in Romania. The objectives of the study include (i) identification of the habitats and plant associations in which S. nutans is found, (ii) identification of the environmental factors that explain the variation of the floristic composition, and (iii) recommending conservation measures for the protection of the species. The hypothesis of this study is as follows: Due to the transformation of grasslands into agricultural lands and overgrazing, the habitat area of the S. nutans species in Romania has been reduced. In this case, these factors have led to a decline in the population and number of individuals of this species, especially in Transylvania. We also consider that hybridization with Salvia nemorosa L. has an essential role in reducing this species. The development of measures to protect this species is also important for a unique butterfly species, namely Pseudophilotes bavius subsp. hungarica (Diószeghy, 1913) is endemic to Transylvania and dependent on the host plant S. nutans (Crișan 2012). The degradation and loss of the S. nutans species threaten the habitat of this butterfly. The life cycle of the butterfly, which involves laying eggs on host plants and developing into larvae, is directly affected by the conservation status of the S. nutans species (Crișan 2012; Crișan, Vizauer, and Rákosy 2023). In this case, the lack of adequate measures for the populations in Romania affects both the plant and the butterfly. Materials and methods Study area The study areas were represented by the historical regions of Bucovina, Crișana, Dobrogea, Moldova, Muntenia, and Transylvania (Fig. 1 ) from Romania. Bucovina and Moldova are located in the northeastern part of Romania and are characterized by a temperate continental climate with moderate summers, cold winters, and abundant precipitation. Dobrogea is characterized by a moderate continental climate with maritime influences and reduced precipitation compared to the other regions. Transylvania features a continental climate characterized by significant thermal variations and abundant precipitation. In contrast, Crișana and Muntenia are characterized by a temperate continental climate, fertile soils, and favourable annual mean temperatures. The lowest elevation was recorded in Dobrogea, and the highest elevation was recorded in Transylvania (Fick and Hijmans 2017 ). Salvia nutans , a species associated with xeric and mesoxeric grasslands in the steppe and forest-steppe zones, is linked to the typical soils characteristic of these ecosystems. On plateaus and gently inclined slopes in the steppe area, this species colonizes Kastanozems, Calcic, and Haplic Chernozems. In the same situation in the forest-steppe, the species is found on Haplic and Luvic Chernozems, as well as on various types of Phaeozems. It is interesting that in forest-steppe areas and at the borders of vicinal nemoral forest areas, the species can be frequently encountered on steep, sunny slopes with shallow soils, such as Lithic, Skeletic, Calcic, Brunic, and Calcaric Leptosols, as well as on short-profiled Kastanozems. According to our observations, the species typically avoids colonizing barren rocks (IUSS Working Group WRB 2022 ). Study species Salvia nutans (Fig. 2 ) is a perennial species (Ciocârlan 2009 ) with a Pannonian-Pontic distribution (Golevich 2012 ). The plant height ranges from 20 cm to 100 cm (Gusztáv 2005 ) or 120 cm (Beldie et al. 1961 ). The stem is erect, almost leafless in the upper part (Sârbu, Ștefan, and Oprea 2013 ) or sometimes with 1 or 2 pairs of small, simple leaves (Beldie et al. 1961 ; Ciocârlan 2009 ). Usually, 1–2 stems originate from the same rhizome (Beldie et al. 1961 ). The basal leaves are cordate-ovate in shape (Ciocârlan 2009 ; Sârbu, Ștefan, and Oprea 2013 ), with lengths ranging from 3 cm to 16 cm and widths ranging from 2 cm to 12 cm (Beldie et al. 1961 ). The leaf tips are either slightly pointed or rounded, and the margins are crenate. The leaf surface is either glabrous or partially hairy along the central veins. On the underside, the leaves are densely tomentose, with short hairs and numerous sessile glandular hairs of yellow-orange color. The petioles are longer than the leaf blades, with broad bases, and are covered with long, multicellular, dispersed hairs. The stem leaves are small, either sessile or short-petiolate, and lanceolate or occasionally ovate. They have a length from 15 cm to 30 cm and a width from 6 cm to 10 cm. Bracteoles are equal in size or slightly longer than the floral pedicels (Beldie et al. 1961 ). Inflorescence is pendulous at the tip before flowering (Ciocârlan 2009 ). The flowering period extends from late May to early July (Kondratyuk et al. 1985 ). Although most individuals flower in the third year of growth, there are cases in which flowering occurs as early as the second year (Udvardy 2002 ; Gusztáv 2005 ). Vegetation data A total of 870 relevés, comprising 1,421 taxa, were used for the vegetation analysis. Of these, 492 relevés were sourced from the Romanian Grassland Database (RGD; Vassilev et al. 2018 ), while the remaining 378 relevés were collected between May 2017 and June 2024 (personal data). The size of each individual plot was 100 m², while the relevés from the Romanian Grassland Database ranged from 4 m² to 200 m². Environmental variables In the multivariate analysis, abiotic variables (elevation – m a.s.l., aspect – °, slope – °, mean annual temperature – ℃, and annual precipitation – mm) and edaphic variables (P – mg kg − 1 , K – mg kg − 1 , and pH) were included. Soil analyses The values for the chemical elements P, K, and soil pH (Ballabio et al. 2019 ) were extracted from the European Soil Database & soil properties – ESDAC (European Soil Data Centre 2019 ). Climatic data The climatic data (mean annual temperature and annual precipitation) were obtained from the WorldClim database (Fick and Hijmans 2017 ) at a resolution of 30 arc. sec. Elevation, aspect, and slope data were collected in the field. Data preparation for classification analyses Before numerical data analysis, the dataset was standardized according to the following criteria: (i) unification of taxonomy and taxon nomenclature; (ii) species identified only at genus level were eliminated; (iii) unrecognized subspecies were combined into species; (iv) exclusion of bryophyte and lichen species, as they were recorded in only 20 relevés. The classification of plant associations within the alliance and the nomenclature of these associations follow Chifu, Irimia, and Zamfirescu ( 2014 ) and Chifu and Irimia ( 2014a , 2014b ). The nomenclature and taxonomy of plant species follow Euro + Med ( 2025 ), and the nomenclature of higher syntax units follows Mucina et al. ( 2016 ). Additionally EUNIS habitats were determined (Chytrý et al. 2020 ). The final data set included 870 relevés and 904 taxa. Numerical classification and expert system development We used the modified TWINSPAN algorithm (Roleček et al. 2009 ) with three pseudospecies cut levels of 0%, 5%, and 25% cover, minimum group size for division of five relevés, and Whittaker’s beta coefficient as a measure of internal cluster heterogeneity. We assessed the optimal number of clusters using OptimClass 1 (Tichý et al. 2010 ). Diagnostic species for individual clusters were determined based on the phi -coefficient of association applied to virtually standardized cluster sizes (Chytrý et al. 2002 , Tichý and Chytrý 2006 ). We used a fidelity threshold of phi = 0.25 combined with Fisher’s exact test (p < 0.05). We used 25% and 50% constancy as thresholds for determining constant and highly constant species, respectively. The data were represented as mean percentage cover values, according to the scale developed by the Braun-Blanquet ( 1964 ). The mean abundance-dominant interval corresponding to the notes was defined as follows (Cristea, Gafta, and Pedrotti 2004 ): r (0.05%); + (0.5%); 1 (5%); 2 (17.5%); 3 (37.5%); 4 (62.5%); and 5 (87.5%). Vegetation–environment relationship The correlation between floristic composition and environmental variables was analyzed with Detrended Correspondence Analysis (DCA) in the CANOCO v5.1 software (ter Braak & Šmilauer, 2018 ). In this context, DCA analysis was performed to detect floristic gradients, and CCA (Canonical Correspondence Analysis) was applied to quantify the effect of each environmental variable on floristic composition, using the Monte Carlo permutation test (9999 iterations). The average percentage coverage values represented the floristic composition, and the environmental variables were represented by variables with a VIF value of less than 5 (Table 1 ). We used the Variance Inflation Factor (VIF) in SPSS version 23 to assess collinearity between independent variables. Variables with a VIF value greater than 5 were considered multicollinear and were excluded from the model. The relevés distribution map was made in QGIS version 3.34.3 (QGIS Development Team 2025 ). Analyses of differences between syntaxa To determine whether there are statistically significant differences between plant communities regarding environmental variables, the ANOVA test was applied, using R Statistical Software (v4.1.4; R Core Team 2025 ) via the 'tidyverse' v.2.0.0 (Wickham et al. 2019 ), 'car' v.3.1.3 (Fox and Weisberg 2019 ), and 'stats' v.4.5.0 (R Core Team 2025 ). Since significant differences were found (p < 0.05), the post-hoc Tukey test was applied. In this context, communities that did not exhibit significant differences were denoted by the same letter, while communities that did exhibit significant differences were highlighted by different letters. Results Syntaxonomic overview of relevés Salvia nutans populations were found to inhabit eight main phytocoenological classes: Crataego-Prunetea , Digitario sanguinalis-Eragrostietea minoris , Festuco-Brometea , Festuco-Puccinellietea , Molinio-Arrhenatheretea , Quercetea pubescentis , Sisymbrietea , and Trifolio-Geranietea sanguinei . Class : Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 Order : Festucetalia valesiacae Soó 1947 All.: Stipion lessingianae Soó 1947 Ass. : Allio albidi-Stipetum lessingianae (Soó 1947) Coldea et Sârbu in Coldea 2012 Ass. : Jurineo transylvanicae-Stipetum pulcherrimae (Soó 1942) Coldea et Sârbu in Coldea 2012 Ass. : Jurineo arachnoidea-Stipetum lessingianae (Dobrescu 1974) Chifu, Mânzu et Zamfirescu 2006 Ass. : Festucetum rupicolae Burduja et al. 1956 Ass. : Artemisietum ponticae (Soó 1942) Păun 1969 Ass. : Stipo ucrainicae-Festucetum valesiacae (Dihoru 1969) Dihoru et Doniță 1970 Ass. : Convolvulo cantabrici-Stipetum capillatae (Horeanu 1976) Ass. : Agropyro pectinati-Stipetum capillatae (Burduja et al. 1956) Chifu, Manzu et Zamfirescu 2006 Ass. : Taraxaco serotinae-Festucetum valesiacae (Burduja et al. 1956, Răvăruț et al. 1956) Sârbu, Coldea et Chifu 1999 Ass. : Agropyro pectinati-Tanacetinetum millefolii (Șerbănescu 1970) Chifu et Țupu 2009 Ass. : Elytrigietum hispidi (Dihoru 1970) Dihoru et Doniță 1970 Ass. : Astero oleifolius-Ephedretum distachyae Horeanu et Vițalariu 1992 Ass. : Cynodonto-Poëtum angustifoliae (Rapaics 1927) Soó 1957 Ass. : Bombycilaeno erecti-Bothriochloetum ischaemi (Dihoru 1970) Dihoru et Doniță 1970 Ass. : Agropyretum pectiniformae (Dihoru 1970) Dihoru et Doniță 1970 All.: Festucion valesiacae Klika 1931 Ass. : Bothriochloetum ischaemi (Kristiansen 1937) I. Pop 1977 Ass. : Salvio-Festucetum rupicolae Zolyomi 1958 corr. Soó 1964 Sub-ass. : paeonietosum tenuifoliae (Mititelu et al. 1990) Ass. : Festucetum valesiacae-rupicolae Csürös et Kovács 1962 Ass. : Stipetum capillatae (Hueck 1963) Krausch 1961 Ass. : Agrostio-Festucetum valesiacae Borisavljevič et al. 1955 Ass. : Agropyro pectinati-Kochietum prostratae Zolyomi 1958 Ass. : Taraxaco serotinae-Bothriochloetum ischaemi (Burduja et al. 1956) Sârbu, Coldea et Chifu 1999 Sub-ass. : koelerietosum macranthae (Răvăruț et al. 1956) Chifu, Mânzu et Zamfirescu 2006 Ass. : Medicagini minimae-Festucetum valesiacae Wagner 1941 Ass. : Festuceto rupicolae-Caricetum humilis Polgár 1933 Sub-ass. : thymio-salvietosum nutantis Order : Brachypodietalia pinnati Korneck 1974 All .: Cirsio-Brachypodion pinnati Hadač et Klika in Klika et Hadač 1944 Ass. : Cariceto humilis-Brachypodietum pinnati Soó 1947 Sub-ass. : dichathietosum ischaemi Chifu et al.. 2014 Ass. : Helianthemo cani-Seslerietum heuflerianae (Borza 1959) Popescu et Sanda 1992 Ass. : Festuco rupicolae-Brachypodietum pinnati Mahn 1965 Ass. : Jurineo transsilvanicae-Chrysopogonetum grylli (Csürös et Niedermaier 1966) Coldea et Sârbu in Coldea 2012 Ass. : Danthonio alpinae-Stipetum stenophyllae Ghișa 1941 Ass. : Carici humilis-Stipetum joannis Pop et Hodișan 1985 Ass. : Convolvulo cantabrici-Chrysopogonetum grylli (Dihoru 1970) in Chifu et al. 2014 Ass. : Valerianello lasiocarpae-Chrysopogonetum grylli (Dihoru 1970) Coldea et Sârbu in Coldea 2012 Ass. : Ferrulago campestris-Caricetum humilis (Ciocârlan 1969) Coldea et al. 2010 Ass. : Galio octonarii-Stipetum tirsae (Ciocârlan 1969) Popescu et Sanda 1992 Ass. : Rhinantho rumelici-Brometum erecti Sanda et Popescu 1999 Ass. : Thymo comosi-Caricetum humilis Morariu et Danciu 1977 Order : Stipo pulcherrimae-Festucetalia pallentis Pop 1968 All .: Pimpinello-Thymion zygoidis Dihoru et Doniţa 1970 Ass. : Saturejetum caeruleae Cristurean et Ionescu-Țeculescu 1970 Ass. : Koelerio lobatae-Artemisietum lerchianae (Dihoru 1970) Dihoru et Doniță 1970 Ass. : Agropyro pontici-Thymetum zygioidis (Dihoru 1970) Dihoru et Doniță 1970 Ass. : Festucetum callierii (Şerbănescu 1965) Dihoru et Doniță 1970 Class : Crataego-Prunetea Tx. 1962 Order : Prunetalia spinosae Tx. 1952 All. : Prunion fruticosae Tx. 1952 Ass. : Prunetum tenellae Soó 1951 All. : Berberidion vulgaris Br.-Bl. ex Tx. 1952 Ass. : Pruno spinosae-Crataegetum Hueck 1931 Class : Trifolio-Geranietea sanguinei T. Müller 1962 Order : Antherico ramosi-Geranietalia sanguinei Julve ex Dengler in Dengler et al. 2003 All. : Geranion sanguinei Tx. in T. Müller 1962 Ass. : Inulo ensifoliae-Peucedanetum cervariae Kozlowska 1925 em. Van Gils et Kovács 1977 Ass. : Geranio-Dictamnetum Wendelberger ex T. Müller 1962 Ass. : Clematido rectae-Laserpitietum latifolii Schneider-Binder 1984 Class : Molinio-Arrhenatheretea Tx. 1937 Order : Arrhenatheretalia elatioris Tx. 1931 All. : Cynosurion cristati Tx. 1947 Ass. : Anthoxantho-Agrostetum capillaris Sillinger 1933 All. : Arrhenatherion elatioris Koch 1926 Ass. : Arrhenatheretum elatioris Br.-Bl. ex Scherrer 1925 Order : Potentillo-Polygonetalia avicularis Tx. 1947 All. : Potentillion anserinae Tx. 1947 Ass. : Rorippo austriacae-Agropyretum repentis (Timar 1947) R. Tx. 1950 Class : Festuco-Puccinellietea Soó ex Vicherek 1973 Order : Puccinellietalia Soó 1947 All. : Festucion pseudovinae Soó 1933 Ass. : Artemisio-Festucetum pseudovinae Soó (1927) 1945 Class : Sisymbrietea Gutte et Hilbig 1975 Order : Sisymbrietalia sophiae J. Tx. ex Görs 1966 All. : Sisymbrion officinalis Tx. et al. ex von Rochow 1951 Ass. : Bromo squarrosi-Xeranthemetum annui Coroi 2001 Ass. : Hordeo murini-Cynodontetum dactyloni Felföldy ex Borhidi 1949 Class : Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959 Order : Quercetalia pubescenti-petraeae Klika 1933 All. : Quercion petraeae Issler 1931 Ass. : Corno-Quercetum pubescentis Mathé et Kovács 1962 Ass. : Paeonio peregrinae-Quercetum pubescentis (Sârbu 1982) Sanda et Popescu 1999 Class : Digitario sanguinalis-Eragrostietea minoris Mucina, Lososová et Šilc in Mucina et al. 2016 Order : Eragrostietalia J. Tx. ex Poli 1966 All. : Salsolion ruthenicae Philippi ex Oberd. 1983 Community : Cynodon dactylon EUNIS habitats The identified plant associations were assigned to eight major EUNIS habitat types, namely (at level 2 of the EUNIS hierarchy): R1 – Dry grasslands ( R1A Semi-dry perennial calcareous grassland - meadow steppe, R1B Continental dry grassland - true steppe, R16 Perennial rocky grassland of Central and South-Eastern Europe); R2 – Mesic grasslands (R21 Mesic permanent pasture of lowlands and mountains, and R22 Low and medium altitude hay meadow); R3 – Seasonally wet and wet grasslands (R36 Moist or wet mesotrophic to eutrophic pasture); R5 – Woodland fringes and clearings and tall forb stands (R51 Thermophilous forest fringe of base-rich soils); R6 – Inland salt steppes (R62 Continental inland salt steppe); S3 – Temperate and Mediterranean montane scrub (S35 Temperate and submediterranean thorn scrub); T1 – Broadleaved deciduous forests (T19 Temperate and submediterranean thermophilous deciduous forest); and V1 – Arable land and market gardens (V15 Bare tilled, fallow or recently abandoned arable land). Cluster analysis The cluster analysis results were presented as a dendrogram (Fig. 3 ), which illustrated the hierarchical relationships between the vegetation units, and a synoptic table (Supplementary Material 1). The analyzed vegetation was classified into six clusters, which were correlated with different syntax groups recognized in the specialized literature. Cluster 1: Cl. Festuco-Brometea , Ord. Festucetalia valesiacae , Al. Stipion lessingianae , Ass. Stipo ucrainicae-Festucetum valesiacae (17 relevés) Structure and composition of plant community Phlomis herba-venti had a fidelity of 58% and was the species most strongly associated with this community. Other important species for this community that had relatively significant contributions were Centaurea orientalis (53.2%) and Cytisus austriacus (52.6%). The number of species per 100 m 2 ranged from 13 to 44, and vegetation cover varied between 48% and 83%. Salvia nutans had a cover ranging from 0.5–5%. Ecology and distribution The communities analysed were recorded at elevations varied between 22 to 122 m a.s.l. (above sea level). The climate is moderate, with annual mean temperatures ranging from 10.9 to 11.9°C. Annual precipitation was also moderate, ranging from 432 to 470 mm. The temperatures in May and June varied between 15.6 and 16.1°C. They ranged from 20.2 to 20.7°C in June, while precipitation was lower in May (37–45 mm) compared to June (48–55 mm), suggesting a gradual increase in humidity during the warm season. Soils ranged from neutral to slightly alkaline, with moderate phosphorus and potassium concentrations. The communities were recorded on gentle slopes with a northeastern aspect in the grasslands of Constanța County. Cluster 2: Cl. Festuco-Brometea , Ord. Festucetalia valesiacae , Al. Stipion lessingianae , Ass. Taraxaco serotinae-Festucetum valesiacae and Jurineo arachnoidea-Stipetum lessingianae ( 36 relevés ) Structure and composition of plant community Galium glaucum had a fidelity of 63.2%, the species most strongly associated with this community. Similarly, Festuca stricta subsp. sulcata (60.6%) and Carex humilis (60.1%) contributed significantly to the structure and functioning of the community. Another species contributing to the community structure was Astragalus monspessulanus , which had a fidelity of 56.6%, similar to Thymus odoratissimus , which had a fidelity of 54%. Similarly, Peucedanum ruthenicum is a representative species for this community, although it had a lower fidelity (51%). The number of plant species per 100 m 2 ranged from 13 to 95, while vegetation cover varied between 51% and 100%. Salvia nutans had a cover between 0.05% and 5%. Ecology and distribution The annual mean temperatures range from 6.9 to 11.6°C, indicating a moderate to cold climate. The annual precipitation ranges from 428 to 590 mm, indicating moderate humidity conditions. The mean monthly temperatures in May and June range from 12.6 to 16.1°C in May and 16 to 20.7°C in June. The monthly precipitation ranges from 41 to 72 mm in May and from 50 to 95 mm in June. The soils had a pH ranging from weakly alkaline to neutral, characterized by moderate phosphorus and rich in potassium concentrations. The communities in this cluster were recorded at elevations ranging from 31 to 630 m a.s.l., on slopes that varied from gentle to very steep, predominantly with a northeastern aspect across the grasslands of Alba, Brașov, Constanța, Iași, Suceava, and Vaslui counties. Cluster 3: Cl. Festuco-Brometea , Ord. Festucetalia valesiacae , All. Stipion lessingianae , Pimpinello-Thymion zygoidis , Ass. Taraxaco serotinae-Festucetum valesiacae and Agropyro pontici-Thymetum zygioidis ( 57 relevés ) Structure and composition of plant community Convolvulus cantabrica was the species most strongly associated with this community, with a fidelity of 61.3%. Erysimum diffusum (59%) and Koeleria lobata (58.6%) had an essential role in these communities, contributing significantly to the community structure. Also, other species with relatively significant fidelity were Euphorbia seguierana (55.9%), Satureja coerulea (54.2%) and Achillea coarctata (54.2%). Agropyron cristatum recorded a fidelity of 50.7%. The number of plant species per 100 m 2 ranged from 15 to 85, and vegetation cover ranged from 45–100%. Salvia nutans had a cover ranging from 0.05–5%. Ecology and distribution The climate was characterized by moderate temperatures and precipitation. Mean monthly temperatures ranged from 14.8 to 16.8°C in May and from 19.1 to 20.7°C in June. Monthly precipitation was moderate, from 38 to 55 mm in May and from 47 to 67 mm in June. The soils associated with these communities had a pH ranging from slightly acidic to neutral and were characterized by moderate phosphorus and potassium concentrations. Communities in this cluster were recorded at elevations ranging from 10 to 286 m a.s.l., on slopes ranging from gentle to very steep, with a predominantly northeastern aspect in the grasslands of Constanța and Tulcea counties. Cluster 4: Cl. Festuco-Brometea , Ord. Festucetalia valesiacae , All. Stipion lessingianae , Ass. Jurineo transylvanicae-Stipetum pulcherrimae , Allio albidi-Stipetum lessingianae , and Prunetum tenellae (562 relevés) Structure and composition of plant community Crataegus monogyna (66.9%) and Potentilla taurica (66.3%) had the highest fidelity values, characteristic species for this community. Valeriana coronata (65.2%) and Rosa canina (62.2%) were other important species, significantly contributing to the community structure. Also, the species Marrubium vulgare (61%) and Salvia revelata (55.9%) recorded a relatively high fidelity. Species such as Thalictrum aquilegiifolium (54.4%), Achillea ochroleuca (51.6%), Potentilla reptans (50.8%), and Euphrasia stricta (50%) recorded a moderate to high fidelity. The number of species per 100 m 2 ranged from 9 to 99, and the vegetation cover ranged from 29–100%. Salvia nutans had a cover ranging from 0.05–37.5%. Ecology and distribution The communities included in this cluster were recorded at elevations from 95 to 610 m a.s.l., on very steep slopes with a southwest aspect. Annual precipitation ranged from 575 to 731 mm, and annual mean temperatures ranged from 6.2 to 10 ℃. The soils had a pH ranging from slightly acidic to weakly alkaline, characterized by moderate phosphorus and rich potassium concentrations in the grasslands of Alba, Bistrita-Năsăud, Brașov, Cluj, Mureș, Sălaj, Sibiu, and Suceava counties. ° Cluster 5: Cl. Festuco-Brometea , Ord. Brachypodietalia pinnati , All. Cirsio-Brachypodion pinnati , Ass. Cariceto humilis-Brachypodietum pinnati ( 157 relevés ) Structure and composition of plant community Brachypodium pinnatum recorded a fidelity of 62.2%, the species most strongly associated with this community. The number of species per 100 m 2 ranged from 45 to 58, and the vegetation cover ranged from 30–100%. Within this community, Salvia nutans recorded a relatively variable cover, ranging from 0.05–37.5%. Ecology and distribution The communities analysed were recorded at elevations ranging from 90 to 1553 m a.s.l., located on slopes ranging from moderate to very steep with a predominantly southern and western aspect. The soils presented a pH ranging from slightly acidic to neutral, characterised by moderate phosphorus and potassium concentrations. Annual precipitation ranged from 449 to 990 mm, and annual mean temperatures ranged from 2.4 to 10.5 ℃ in the grasslands of Alba, Bacău, Bihor, Brașov, Cluj, Covasna, Harghita, Mureș, Sălaj, Sibiu, and Suceava counties. Cluster 6: Cl. Festuco-Brometea , Ord. Festucetalia valesiacae , All. Stipion lessingianae , Ass. Taraxaco serotinae-Bothriochloetum ischaemi and Astero oleifolius-Ephedretum distachyae ( 41 relevés ) Structure and composition of plant community Galium humifusum was the species most strongly associated with this community, with a fidelity of 59.9%. The other species, such as Astragalus onobrychis (39.5%), Ephedra distachya (39.3%), and Diplotaxis muralis (39.3%), had lower fidelity. The number of species per 100 m 2 ranged from 7 to 61, and the vegetation cover ranged from 47–98%. Salvia nutans presented a cover range of 0.5–5%. Ecology and distribution The communities included in this cluster were recorded at elevations ranging from 5 to 387 m a.s.l., on moderate slopes with a northeastern aspect. Climatic conditions were characterized by annual precipitation ranging from 429 to 609 mm and annual mean temperatures ranging from 8.6 to 12 ℃ in the grasslands of Constanța, Iași, Sibiu, Suceava, Tulcea, and Vaslui counties. Table 1 Measured values of bioclimatic, soil chemical, and topographic variables were analysed. Values are means ± standard deviations (SD). VEGC = vegetation cover; BIO12 = annual precipitation; BIO1 = annual mean temperature; Aspect: S – south, SSW – south-southwest, SE – southeast, SSE – south-southeast, SW – southwest, W – west; P = phosphorus, K = potassium. Cluster Cluster 1 Cluster 2 Cluster 3 Cluster 4 Cluster 5 Cluster 6 No. of species/relevés 28 ± 8.5 43 ± 20 48 ± 14.5 40 ± 16.8 38 ± 16.6 35 ± 15.3 Vegetation cover (%) 61 ± 9.1 87 ± 12.1 83 ± 13.4 83 ± 12.9 82 ± 14.7 77 ± 14.9 Aspect W SSE SE SW SW SSW Slope (°) 5.6 ± 3.1 14.5 ± 15.3 8.1 ± 6 30.6 ± 14.5 22.1 ± 14.4 12.4 ± 10 Elevation (m a.s.l.) 76.6 ± 25.8 204 ± 142.7 91.5 ± 85 392.6 ± 67.3 450.6 ± 138.8 178.3 ± 99.4 pH 7.1 ± 0.2 6.3 ± 0.4 7 ± 0.3 6 ± 0.6 5.7 ± 1.2 6.4 ± 1.1 P (mg kg − 1 ) 27.2 ± 7.5 28.2 ± 6 27 ± 7.3 26.0 ± 6.5 24.5 ± 8.8 28.4 ± 7.3 K (mg kg − 1 ) 263.3 ± 71.9 342.1 ± 99.4 278.5 ± 76.7 297.4 ± 56.4 281.2 ± 73 331.4 ± 132 BO1 (℃) 11.5 ± 0.3 9.4 ± 0.9 11.1 ± 0.7 8.5 ± 0.4 8.2 ± 0.8 9.8 ± 0.9 BIO12 (mm) 453.7 ± 16.7 549.7 ± 34.8 443.9 ± 15.6 608.3 ± 19.1 608.5 ± 43.5 524.2 ± 53.1 Mean temperature – May 15.7 ± 0.1 15.4 ± 0.9 15.8 ± 0.4 14.5 ± 0.4 14 ± 1 15.4 ± 0.4 Mean temperature – June 20.3 ± 0.1 18.9 ± 1 20.3 ± 0.5 17.3 ± 0.4 16.9 ± 1 19.2 ± 0.9 Mean precipitation – May 41.5 ± 2.2 62.8 ± 7.2 43.5 ± 3.8 72.7 ± 1.7 73.5 ± 4.6 58.5 ± 10 Mean precipitation – June 52.1 ± 2.1 85.2 ± 10.5 53.8 ± 4.4 93.5 ± 2 93.7 ± 5.9 76.6 ± 14.9 Salvia nutans occurs predominantly on steep slopes (from 21° to 35°; 31%), followed by very steep slopes (> 35°), with 26%. The species also occurs on gentle slopes (3–10°; 23%), moderate slopes (11–20°; 15%), and very gentle slopes (0–3°; 5%). The dominant aspect in which S. nutans occurs is southwest (39%), followed by southern (23%), and western (13%). The species also occurs on secondary aspects, such as southeast (8%) and northeast (7%). In contrast, on the northwest (5%), eastern (3%), and northern (3%) aspects, the species recorded a reduced presence. In terms of elevation, S. nutans prefers low elevations. The average elevation was 359 m, ranging from 5 to 850 m a.s.l. (Fig. 4 ). The soils were predominantly moderately acidic (pH range of 5.81 to 6.80, 59%). The species was also identified on slightly acidic soils (pH 5.01–5.80, 27%), neutral soils (pH 6.81–7.20, 11%), slightly alkaline soils (pH 7.21–8.40, 2%), and moderately strongly acidic soils (pH 4.31–5.00, 1%). Based on the soil phosphorus concentration, the species distribution indicates a preference for soils with low phosphorus concentrations (ranging from 18.1 to 36 mg kg -1 , or 81%). The presence of the species in soils with moderate phosphorus concentrations (ranging from 8.1 to 18 mg kg -1 , 13%) and high phosphorus levels (ranging from 36.1 to 72 mg kg -1 , 6%) was reduced. Regarding potassium concentration, a preference for soils very rich in potassium (from 265.1 to 400 mg kg -1 ; 69%) was observed among the species. The presence of the species in soils rich in potassium (ranging from 200.1 to 265 mg kg -1 , 25%) was moderate. In soils with moderate (ranging from 132.1 to 200 mg kg -1 ; 3.8%), excessive (> 400 mg kg -1 ; 2.1%), and poor potassium levels (ranging from 66.1 to 132 mg kg -1 ; 0.2%), the species exhibited a reduced presence (Fig. 5 ). The annual mean temperature (BIO1) ranged from 2.5 to 11.9°C, with an average of 8.78°C. The average annual precipitation was moderate, ranging from 426 to 748 mm, with an average of 588 mm. The relationship between floristic composition and environmental variables The DCA data showed that Axis 1 was the most important. It explained the most variation and had the most extended gradient length (Table 2 ; Fig. 6 ). The other axes made a minor contribution. Table 2 Summary of the DCA analysis. Axis 1 Axis 2 Axis 3 Axis 4 Eigenvalues 0.6026 0.4672 0.3416 0.2854 Explained variation (cumulative) 2.57 4.57 6.02 7.24 Gradient length 4.43 3.07 4.36 3.99 Pseudo-canonical correlation (suppl.) 0.8254 0.2208 0.4329 0.3782 CCA data showed that annual precipitation (BIO12) explained most of the variation in the floristic composition of S. nutans communities, followed by slope (Table 3 ). Table 3 Results of the CCA ordination. Variable Explains % Contribution % pseudo-F P P(adj) Annual Precipitation (BIO12) 1.86 53.09 16.3 0.002 0.014 Slope (°) 0.34 9.57 3 0.002 0.014 Elevation (m a.s.l.) 0.31 8.93 2.8 0.002 0.014 Aspect 0.3 8.66 2.7 0.002 0.014 Potassium (K) 0.24 6.71 2.1 0.002 0.014 pH 0.24 6.88 2.1 0.008 0.056 Phosphorus (P) 0.22 6.15 1.9 0.002 0.014 In locations (50 locations in Tulcea and Constanța counties) with precipitation ranging from 400 to 450 mm/year, S. nutans has a low cover, ranging from 0.5–5%. In these locations, the species occurs at low elevations (54 m a.s.l.) on gentle slopes (9°). The soils are neutral, with an average phosphorus and high potassium. The dominant associations were Agropyro pontici-Thymetum zygioidis and Taraxaco serotinae-Bothriochloetum ischaemi . In contrast, the temperature is higher (11.3 ℃) compared to the other precipitation intervals (Fig. 7 ). In the range of 451 and 500 mm/year, S. nutans had a similar cover, from 0.5–5%, but was identified in fewer locations (38 locations in Tulcea, Constanța and Galați counties). Within these locations, S. nutans was recorded at moderate elevations (127 m a.s.l.), on gentle slopes (5°) and at lower temperatures of 11.03 ℃. Competition with other species may be one reason why the cover with S. nutans was lower. The soils are neutral, with medium phosphorus concentrations and very high potassium. The dominant association was Stipo ucrainicae-Festucetum valesiacae (Fig. 7 ). In the interval from 501 to 550 mm/year, S. nutans had the same cover, from 0.5–5%. In these locations (19 locations in Vaslui, Iași, and Bacău counties), S. nutans was observed at elevations higher than 158 m a.s.l., on moderate slopes (12.7°). The mean annual temperature was lower (9.7℃). The soils are moderately acidic, with a medium phosphorus concentration and very high potassium levels. The dominant associations were Astero oleifolius-Ephedretum distachyae and Taraxaco serotinae-Festucetum valesiacae (Fig. 7 ). In 325 locations across Vaslui, Iași, Suceava, Covasna, and Alba counties, with annual precipitation ranging from 551 to 600 mm/year, S. nutans had a cover ranging from 0.05–62.5%. The elevations were higher (362 m a.s.l.), and steep slopes (24°). The temperatures were lower (8.7°C). The soils were moderately acidic, with medium phosphorus and very high potassium concentrations. The dominant associations were Allio albidi-Stipetum lessingianae , Danthonio alpinae-Stipetum stenophyllae, Festuceto rupicolae-Caricetum humilis , and Festucetum rupicolae (Fig. 7 ). Most observations of the species were recorded in locations with annual precipitation from 601 to 650 mm/year. In these locations (412 locations in Sălaj, Bistrița, Cluj, and Mureș counties), S. nutans had a cover from 0.05–37.5%. Elevations were high (416 m a.s.l.), steep slopes (32°) and much lower temperatures (8.3°C). The soils were moderately acidic, with medium phosphorus and very high potassium concentrations (Fig. 7 ). In locations (20 locations in Cluj and Bistrița-Năsăud counties) with precipitation higher than 650 mm/year, S. nutans had a cover ranging from 0.05–17.5%. In these locations, the highest elevations (604 m) and the lowest average annual temperatures (7.52°C) were recorded on steep slopes (21°). The soils are weakly acidic, with medium to high phosphorus concentrations (Fig. 7 ). Discussion General aspects Salvia nutans is widely distributed in steppe grasslands across Ukraine, Russia, and other Eastern European regions (Budak 1999 ; Gusztáv 2005 ; Đakić, Knežević, and Boža 2012 ). The results indicate a high presence of the species in Transylvania and the plains of Eastern Romania, confirming its presence in the marginal areas of its European distribution (Soó 1968 ; Csapody 1982 ). In Romania (Oprea 2005 ) and Hungary (Gusztáv 2005 ), the species was much more widespread in the past. Due to overgrazing and the transformation of grasslands into agricultural land, the habitat of the species is threatened in Hungary (Gusztáv 2005 ) and Romania. Syntaxonomical scheme and EUNIS habitats This study confirms that S. nutans occurs predominantly in dry grasslands of the Festuco - Brometea class (827 relevés, representing 95% of the total), which is in agreement with the data in the literature (Soó 1968 ; Sârbu, Ștefan, and Oprea 2013 ), where the species is described as a xerophytic species characteristic of open habitats. The high percentage of 95% compared to the presence of the species in other habitats shows a clear preference for this type of grassland, where the soils are well-drained. In the vegetation classes Crataego-Prunetea (19 relevés, representing 2% of the total) and Trifolio-Geranietea (13 relevés, representing 2% of the total), the presence of the species was reduced, indicating that S. nutans also colonizes less typical habitats. In contrast, the presence of the species in the classes Digitario sanguinalis-Eragrostietea minoris (one relevé, representing 0.1% of the total), Festuco-Puccinellietea (one relevé, representing 0.1% of the total), Molinio - Arrhenatheretea (four relevés, representing 0.5% of the total), Quercetea pubescentis (two relevés, representing 0.2% of the total), and Sisymbrietea (three relevés, representing 0.3% of the total) was significantly reduced, indicating a marginal presence, not being characteristic for these communities. Some of these data are also confirmed by specialized literature (Horeanu 1974 ; Kovács 2003 ; Germany 2013 ), which shows that S. nutans has been recorded in other vegetation classes as well. At the order level, the results indicated that S. nutans was associated with Festucetalia valesiacae (659 relevés, representing 77% of the total). The second most frequent order was Brachypodietalia pinnati (127 relevés, representing 15% of the total). In the orders Stipo pulcherrimae-Festucetalia pallentis (24 relevés, representing 3% of the total), Prunetalia spinosae (19 relevés, representing 2% of the total), Antherico ramosi-Geranietalia sanguinei (13 relevés, representing 2% of the total), S. nutans had a reduced presence. In the remaining orders ( Arrhenatheretalia elatioris , Eragrostietalia , Potentillo-Polygonetalia avicularis , Puccinellietalia , Quercetalia pubescenti-petraeae , and Sisymbrietalia sophiae ), S. nutans had a significantly reduced presence. At the alliance level, S. nutans was recorded most frequently in Stipion lessingianae (468 relevés, representing 54% of the total), Festucion valesiacae (191 relevés, representing 22% of the total), and Cirsio-Brachypodion pinnati (144 relevés, representing 17% of the total). In the alliances Pimpinello-Thymion zygoidis (24 relevés, representing 3% of the total) and Geranion sanguinei (13 relevés, representing 2% of the total), S. nutans had a low presence. In the remaining alliances ( Arrhenatherion elatioris , Berberidion vulgaris , Cynosurion cristati , Festucion pseudovinae , Potentillion anserinae , Quercion petraeae , Salsolion ruthenicae , and Sisymbrion officinalis ), S. nutans had a very low presence. According to the literature, S. nutans is characteristic of Stipion lessingianae (Ruprecht et al. 2009 ; Sârbu, Ștefan, and Oprea 2013 ) and Festucion valesiacae alliances (Kovács 2011 ). Regarding plant associations, S. nutans occurs most frequently in the associations Jurineo transylvanicae-Stipetum pulcherrimae (159 relevés, representing 18% of the total) and Allio albidi-Stipetum lessingianae (123 relevés, representing 18% of the total). Also, other associations in which S. nutans occurs were Festucetum rupicolae (81 relevés, representing 9% of the total), Salvio-Festucetum rupicolae (54 relevés, 6%), Festuceto rupicolae-Caricetum humilis (49 relevés, 6%), Taraxaco serotinae-Festucetum valesiacae relevés (49 relevés, 6%), Cariceto humilis-Brachypodietum pinnati (45 relevés, 5%). The presence of S. nutans species in Salvio-Festucetum rupicolae Zolyomi 1958 corr. Soó 1964, Jurineo transylvanicae-Stipetum pulcherrimae and Festuceto rupicolae-Caricetum humilis in Transylvania was reported by Soó ( 1942 , 1968 ), Cśurös (1973, 1974) and Gusztáv ( 2005 ). Also, Irimia and Mânzu ( 2013 ) mention the presence of the species in Taraxaco serotinae-Festucetum valesiacae . At the same time, the presence of the species in Crataego monogynae-Cerasetum mahaleb (Pînzaru, 2021 ) shows the adaptability of the species in transitional areas between grasslands and shrub vegetation. Moreover, S. nutans has also been reported in associations such as Elytrigietum hispidi , Jurineo arachnoideae-Stipetum lessingianae , Stipo ucrainicae-Festucetum valesiacae , and Thymio pannonici-Chrysopogonetum grylli (Popescu et al., 2008 ). In addition, C ariceto humulis-Festucetum rupicolae and Bothriochloetum ischaemi (Kovács 2011 ) indicate a consistent presence in the Festucion valesiacae alliance. The species is also found in Medicagini minimae-Festucetum (Aniței and Mititelu 1997), an association specific to xerophilous mesophilous grasslands. Habitat degradation in the loess grasslands of the Crișana region is described in the literature (Gusztáv 2005 ) as leading to a simplification of diversity and the dominance of grasses, e.g. Festuca valesiaca subsp. parviflora (Hack.) Tracey. The relationship between floristic composition and environmental variables The most influential environmental factor explaining the variation in the floristic composition of S. nutans phytocenoses was annual precipitation (BIO12). This finding is consistent with previous research demonstrating the influence of edaphic and climatic factors on the formation of grassland communities (Klimeš and Doležal 2010 rök et al. 2018 ; Bai et al. 2021 ). The abundance and distribution of S. nutans may be affected by changes in climate patterns, such as increased precipitation and prolonged droughts. Topographic variables had a significant role in explaining the variation in floristic composition. Similar findings have been recorded in other grassland communities, where competition between species, environmental conditions, and soil water retention are influenced by topographic diversity (Moeslund et al. 2013 ). The species' preference for very steep slopes in locations with precipitation from 551 to 650 mm/year may be related to the reduced competition with mesophytic species (Grime 2001 ). Soil chemical variables had a secondary role in explaining the variation in floristic composition. In general, soil pH and phosphorus availability varied slightly, while potassium concentrations were very high. This means that S. nutans can tolerate a broader range of soil conditions, which is a characteristic of species adapted to semi-arid grasslands (Janišová et al. 2014 ). Additionally, in locations with high precipitation, soil pH is typically lower, suggesting an interaction between soil acidity and plant communities (Chytrý et al. 2015 ). Bioclimatic variables, primarily precipitation and topographic factors, played a significant role in the formation of S. nutans communities. Considering the species' vulnerability to water availability, future climate variations may lead to changes in its abundance and distribution. In this context, in locations where the species is threatened by anthropogenic impact, it is important to develop conservation and monitoring measures. The biogeographic, ecological, and evolutionary context of the species Salvia nutans is a member of the section Plethiosphace Benth. in the subgenus Sclarea (Moench). Benth. The section has a Palearctic range and was considered monophyletic in a single molecular analysis addressing it (Will and Classen-Bockhoff 2017 ), which, however, involved only a few species, none of which were Salvia nutans . According to the authors, the evolutionary relationships among the species in this section remain unknown. In Transylvania, Romania, the hybrid S. nutans × nemorosa is very frequently encountered wherever the two species come into contact. At the same time, S. nemorosa seems to actively displace S. nutans in many locations, including well-preserved steppe grasslands (personal observations). This is a serious problem for S. nutans conservation, at least in Romania, but it also implies that the two species are closely related evolutionarily and ecologically. The other hybrid, known as S. nutans , is S. × simonkaiana Borb. ( nutans × pratensis ) it is far rarer. No other hybrids are known between S. nutans and any other species in the section Plethiosphace that occur within its range. Salvia nutans is a West Palearctic steppe and forest-steppe element (Pobedinova 1954 ; Ozenda 1994 ; Sramkó et al. 2020) with a wide range extending continuously from the Lower Danube forest-steppe areas to the steppe and forest-steppe of the Ufa-Samara-Orenburg region immediately west of the Ural Mountains. Isolated populations are found in the insular forest-steppes of Thrace, the Pannonian Basin, and the Transylvanian Basin. While in the Transylvanian Basin the species is frequent in the mesoxeric and xeric grasslands (as it is presented here) from the forest-steppe and the nemoral adjacent area (here only on steep sunny slopes) in the Pannonian Basin it is scarce – two locations only in Hungary (Sramkó et al. 2020) while in Vojvodina, all the previously known five populations are extinct nowadays (Đakić, Knežević, and Boža 2012 ). Interestingly, two distinct sampled populations from the Transylvanian Basin exhibit different genetic affinities; one is more closely related to Hungarian populations west of the region, while the other is more closely related to populations north of the Black Sea (Sramkó et al. 2020). This implies a biogeographically interesting heterogeneity for Salvia nutans from Transylvania, with one possible explanation being that the species migrated into this region from two directions: from the west (Pannonia) and the east (Moldova), presumably in different and distant periods. Conclusions Salvia nutans was found in eight main phytocoenological classes, showing its versatility in various habitats. This species was most frequently recorded in dry grasslands, with the highest abundance reported in the associations Jurineo transylvanicae-Stipetum pulcherrimae and Allio albidi-Stipetum lessingianae . The species was recorded in areas with moderate elevation, on gentle to very steep slopes, being affected by overgrazing and the transformation of grasslands into arable land. The abundance of the species varies significantly in the six clusters, highlighting different environmental conditions that influence its distribution. Thus, the highest average cover of the species S. nutans was recorded in cluster 4 (75%). A moderate abundance of the species was reported in cluster 5 (19%). In contrast, low abundance was recorded in clusters 1 and 2 (2%), 3, and 6 (1%). Canonical Correspondence Analysis (CCA) revealed that the mean annual temperature (BIO1) accounted for the majority of the variation in floristic composition. In locations with precipitation ranging from 400 to 550 mm/year, S. nutans had a low cover, ranging from 0.05–0.5%. Thus, once the precipitation increases to 650 mm/year, the cover increases to 62.5%. At intervals higher than 650 mm/year, cover decreased. This suggests that the abundance of the species may be limited by excess moisture and lower temperatures. From a biogeographical perspective, S. nutans is characteristic of the West-Palaearctic steppe and forest-steppe. The species has a continuous range from the forest-steppe areas of the Lower Danube to the steppe and forest-steppe areas of the Ufa-Samara-Orenburg region. In the Transylvanian region, S. nutans is frequent, while in the Pannonian Basin, it is very rare. Declarations ORCID iDs Simona Dumitrița Chirilă https://orcid.org/0000-0003-3397-1834 Alexandru Sabin Bădărău https://orcid.org/0000-0001-5113-2802 Mihai Doroftei https://orcid.org/0000-0002-8388-087X Kiril Vassilev https://orcid.org/0000-0003-4376-5575 Iuliia Vasheniak https://orcid.org/0000-0003-1020-3007 Author contributions Simona Dumitrița Chirilă : Conceptualization, Methodology, Investigation, Software, Data curation, Writing- Original draft preparation. Alexandru Sabin Bădărău : Data curation, Visualization, Investigation, Writing- Original draft preparation, Supervision. Mihai Doroftei : Data curation, Visualization, Investigation, Writing - review and editing, Supervision. Kiril Vassilev : Data curation, Visualization, Writing - review and editing, Supervision; Iuliia Vasheniak : Visualization, Software, Data curation, Writing- Original draft preparation, Supervision. Conflicts of Interest The authors declare no conflicts of interest. 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Annali di Bot 1:19–28 Maxwell SL, Fuller RA, Brooks TM, Watson JE (2016) Biodiversity: The ravages of guns, nets and bulldozers. Nature 536(7615):143–145. https://doi.org/10.1038/536143a Moeslund JE, Arge L, Bøcher PK, Dalgaard T, Odgaard MV, Nygaard B, Svenning JC (2013) Topographically controlled soil moisture is the primary driver of local vegetation patterns across a lowland region. Ecosphere 4(7):1–26. https://doi.org/10.1890/ES13-00102.1 Mucina L, Bültmann H, Dierßen K, Tichý L et al (2016) Vegetation of Europe: hierarchical floristic classification system of vascular plant, bryophyte, lichen and algal communities. Appl Veg Sci 19:3–264. https://doi.org/10.1111/avsc.12257 Németh F (1990) Száras növények. In: Rakonczay Z (ed.) Vörös Könyv. Akadémiai Kiadó, Budapest: 265–321 Oprea A (2005) Lista critică a plantelor vasculare din România. Editura Universității Alexandru Ioan Cuza, Iași Ozenda P (1994) Végétation du continent européen. Delachaux et Niestlé Pînzaru P (2021) Crataego monogynae-Cerasetum mahaleb Pînzaru ( Berberidion vulgaris Br.-Bl. ex Tx. 1952) – asociație nouă în vegetația de stâncării din Republica Moldova. Revista Botanică 23(2):32–43 Pobedinova EG (1954) Genus Salvia L. In: Shishkin BK (ed) Flora of the USSR, vol 21. Akademii Nauk USSR, Moscow Popescu A, Doniță N, Roșca V, Băjenaru B (2008) Vegetația Parcului Național „Munții Măcinului. Editura Silvică, București, 240 pp. https://editurasilvica.ro/wp-content/uploads/2023/05/Vegetatia-Parcului-National-Muntii-Macinului-integral.pdf . Accessed on 23 March 2025 QGIS Development Team (2025) QGIS Geographic Information System. Version 3.34.3. Open Source Geospatial Foundation Project. https://qgis.org/ Accessed on 4 August 2025 R Core Team (2025) R: A Language and Environment for Statistical Computing. 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Akadémiai Kiadó, Budapest, p 506 Sramkó G, Laczkó L (2020) A hazai bókoló zsálya ( Salvia nutans L.) populációk konzervációgenetikai összehasonlítása erdélyi és oroszországi populációkkal = Conservation genetic analysis of Transylvanian and Russian Salvia nutans L. populations in comparison with the relict Hungarian populations. Crisicum: Körös-Maros Nemzeti Park Igazgatóság Időszaki Kiadványa 11:111–125 ter Braak CJF, Šmilauer P (2018) Canoco reference manual and user's guide: Software for ordination (version 5.10). Biometris. Wageningen University & Research Tichý L, Chytrý M (2006) Statistical determination of diagnostic species for site groups of unequal size. J Veg Sci 17:809–818. https://doi.org/10.1111/j.1654-1103.2006.tb02504.x Tichý L, Chytrý M, Hájek M et al (2010) OptimClass: Using species-to-cluster fidelity to determine the optimal partition in classification of ecological communities. J Veg Sci 21(2):287–299 Török P, Janišová M, Kuzemko A, Rūsiņa S, Stevanović ZD (2018) Grasslands, their threats and management in Eastern Europe. In: Squires VR, Dengler J, Feng H, Hua L (eds) Grasslands of the World. CRC, pp 78–102. https://doi.org/10.1201/9781315156125-11 Udvardy L (2002) Szaporodásbiológiai megfigyelések a Salvia nutans veszélyeztetettségének megítéléséhez. In: Az I. Magyar Természetvédelmi Biológiai Konferencia Program és Absztrakt kötete: 216 Vassilev K, Ruprecht E, Alexiu V, Dengler J et al (2018) The Romanian Grassland Database (RGD): historical background, current status and future perspectives. Phytocoenologia 48:91–100. https://doi.org/10.1127/phyto/2017/0229 Wickham H, Averick M, Bryan J, Chang W, McGowan LD, François R, Grolemund G, Hayes A, Henry L, Hester J, Kuhn M, Pedersen TL, Miller E, Bache SM, Müller K, Ooms J, Robinson D, Seidel DP, Spinu V, Takahashi K, Vaughan D, Wilke C, Woo K, Yutani H (2019) Welcome to the tidyverse. J Open Source Softw 4(43):1686. https://doi.org/10.21105/joss.01686 Will M, Classen-Bockhoff R (2017) Time to split Salvia s.l. (Lamiaceae) – New insights from Old World Salvia phylogeny. Mol Phylogenet Evol 109:33–58. https://doi.org/10.1016/j.ympev.2016.12.041 Ziman SN (1976) Life forms and biology of steppe plants of Donbass. Naukova Dumka, Kyiv, p 191 Supplementary Files TableSupplementary1.xlsx Cite Share Download PDF Status: Published Journal Publication published 24 Apr, 2026 Read the published version in Biologia → Version 1 posted Editorial decision: Major revisions 16 Sep, 2025 Reviewers agreed at journal 26 May, 2025 Reviewers invited by journal 07 May, 2025 Editor assigned by journal 30 Apr, 2025 First submitted to journal 29 Apr, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6556566","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":453189164,"identity":"f3304823-746f-4e4f-bbe8-a3babdd75581","order_by":0,"name":"Simona Dumitrița Chirilă","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA/ElEQVRIiWNgGAWjYDACZgiVwMbA2HgAyJBjYOAhXksDSIsxYS1QkAAiQFoSGwhpkW/nTnxcUGOXxyd9uOHAjz926RuOnz344AODnZxuA3YtBod5NxvPOJZczMaX2HCwty05d8OZvGTDGQzJxmYHcGhh5t0mzcN2ILGNB+gX3oYDuRsO5JhJ8zAcSNyGQ4t8M0jLP4iWg3/+HEg3OP8GvxaGw0AtvG0QLYeB1iUY3CBgC9gvvH3JEC2ybcmGM2+8MTacYYDbL/L9Zzc+5vlmlzi/h/3hwzd/7OT5zucYPvhQYSeHSwsmUACrNCBWOdjeBlJUj4JRMApGwUgAAIcxX0ccs6qbAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0003-3397-1834","institution":"Danube Delta National Institute for Research and Development: Institutul National de Cercetare-Dezvoltare Delta Dunarii","correspondingAuthor":true,"prefix":"","firstName":"Simona","middleName":"Dumitrița","lastName":"Chirilă","suffix":""},{"id":453189165,"identity":"2145214d-08ed-472f-bba3-0d5fd6b37c69","order_by":1,"name":"Alexandru Sabin Bădărău","email":"","orcid":"","institution":"UBB: Universitatea Babes-Bolyai","correspondingAuthor":false,"prefix":"","firstName":"Alexandru","middleName":"Sabin","lastName":"Bădărău","suffix":""},{"id":453189166,"identity":"6fbebf0f-b1dc-4b42-81fa-0cdf35ac660d","order_by":2,"name":"Mihai Doroftei","email":"","orcid":"","institution":"Danube Delta National Institute for Research and Development: Institutul National de Cercetare-Dezvoltare Delta Dunarii","correspondingAuthor":false,"prefix":"","firstName":"Mihai","middleName":"","lastName":"Doroftei","suffix":""},{"id":453189167,"identity":"fa0c1bdc-003a-4587-b82e-22af2eb8befe","order_by":3,"name":"Kiril Vassilev","email":"","orcid":"","institution":"Bulgarian Academy of Sciences: B'lgarska akademia na naukite","correspondingAuthor":false,"prefix":"","firstName":"Kiril","middleName":"","lastName":"Vassilev","suffix":""},{"id":453189168,"identity":"f39ef651-fddb-4ba7-a91d-c6f3d4e3ef85","order_by":4,"name":"Iuliia Vasheniak","email":"","orcid":"","institution":"National Academy of Sciences of Ukraine: Nacional'na akademia nauk Ukraini","correspondingAuthor":false,"prefix":"","firstName":"Iuliia","middleName":"","lastName":"Vasheniak","suffix":""}],"badges":[],"createdAt":"2025-04-29 12:44:01","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6556566/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6556566/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11756-026-02192-7","type":"published","date":"2026-04-24T15:58:28+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":82512581,"identity":"c4c4e592-a742-4fd7-9a83-f54859c12d6b","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":997555,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the study area in Romania, showing the location of the grid cells (10 × 10 km) with markers from the literature and personal observations (Map: S. D. Chirilă).\u003c/p\u003e","description":"","filename":"Figure1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/e2f14dc2b8e4465d33942b2d.jpg"},{"id":82512586,"identity":"6348e0a3-6733-4fcb-8285-a4d6abf9486b","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":5052062,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eSalvia nutans\u003c/em\u003e L.: A, B, C – the flowering stage. (Photographs: S. Chirilă, May 2024).\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/d28fbbf4bd73c4955dc26376.jpg"},{"id":82512578,"identity":"a40dfcec-2729-4ee9-afd1-0ef35ae06928","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":90628,"visible":true,"origin":"","legend":"\u003cp\u003eDendrogram of relevés with \u003cem\u003eSalvia nutans\u003c/em\u003e in Romania.\u003c/p\u003e","description":"","filename":"Figure3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/c77c6cfad6ec52747f07cb1a.jpg"},{"id":82512582,"identity":"bccb38e8-3341-4a15-84e3-9ceb939a9f8d","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":210325,"visible":true,"origin":"","legend":"\u003cp\u003eTopographic preferences of \u003cem\u003eSalvia nutans.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Figure4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/1c448df7d0473dcede9dafc1.jpg"},{"id":82513676,"identity":"35404a0e-5d76-48dc-9f39-dc2435431818","added_by":"auto","created_at":"2025-05-12 11:12:42","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":346841,"visible":true,"origin":"","legend":"\u003cp\u003eSoil chemical preferences of \u003cem\u003eSalvia nutans.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Figure5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/7f599cedcafb2bd8959dd8a7.jpg"},{"id":82512587,"identity":"563c1a06-4630-4b89-b772-87098b22b453","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":278678,"visible":true,"origin":"","legend":"\u003cp\u003eDCA ordination diagrams of the 870 vegetation plots\u003c/p\u003e","description":"","filename":"Figure6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/2b9dcf00032981c11e596ea7.jpg"},{"id":82513679,"identity":"7f7aa33f-e37e-4b57-ad7b-08690468329c","added_by":"auto","created_at":"2025-05-12 11:12:42","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":458769,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of \u003cem\u003eSalvia nutans\u003c/em\u003e according to annual precipitation.\u003c/p\u003e","description":"","filename":"Figure7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/9a456a05f1ac64ab50078cc8.jpg"},{"id":107928170,"identity":"9f669f1e-2eb5-48d6-9b65-5a37ef5c8b0b","added_by":"auto","created_at":"2026-04-27 16:08:46","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":8173010,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/97240367-6ba5-4ee8-9c0e-b6de8a60f4e8.pdf"},{"id":82512592,"identity":"abc6043a-0bee-42cb-9dc4-f978dd0ad96f","added_by":"auto","created_at":"2025-05-12 11:04:42","extension":"xlsx","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":20861,"visible":true,"origin":"","legend":"","description":"","filename":"TableSupplementary1.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-6556566/v1/b1799fb4fd8048227810de62.xlsx"}],"financialInterests":"","formattedTitle":"Phytocoenological and Ecogeographical study of Salvia nutans in Romania","fulltext":[{"header":"Highlights","content":"\u003cp\u003e is a steppe relict species vulnerable to land-use changes.\u003c/p\u003e\u003cp\u003eThe species occurs mainly in grasslands on steep, sunny slopes.\u003c/p\u003e\u003cp\u003eIts abundance is influenced by annual precipitation and temperature.\u003c/p\u003e\u003cp\u003eBiogeographic patterns suggest distinct regional lineages in Transylvania.\u003c/p\u003e\u003cp\u003eLong-term monitoring is needed due to habitat degradation and hybridization risks.\u003c/p\u003e"},{"header":"Introduction","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eBiodiversity decline represents a significant ecological challenge, affecting ecosystem services and human well-being (Dirzo and Raven \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Ceballos, Garc\u0026iacute;a, and Ehrlich \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Isbell et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Current species extinction rates far exceed historical natural rates, reaching levels that suggest the sixth major extinction event on Earth is underway (Barnosky et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Ceballos, Garc\u0026iacute;a, and Ehrlich 2017). This alarming acceleration underscores a conservation crisis where identifying the mechanisms driving biodiversity decline is a key priority for conservation biology (Didham et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The causes of this decline are multiple, involving direct human activities and indirect environmental changes (Adla et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Habitat loss, overexploitation, climate change, pollution, and the introduction of invasive species are the primary factors contributing to biodiversity decline, alongside other secondary threats (Maxwell et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Caro et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In particular, invasive species are frequently classified as the second most significant source of threat to biodiversity (Due\u0026ntilde;as et al. 2018).\u003c/p\u003e \u003cp\u003eOne of the plant species with restricted habitats is \u003cem\u003eSalvia nutans\u003c/em\u003e L., a steppe relict (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Sramk\u0026oacute; and Laczk\u0026oacute; \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and postglacial in the Carpathian Basin (N\u0026eacute;meth \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Simon \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Đakić, Knežević, and Boža \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The species \u003cem\u003eS. nutans\u003c/em\u003e belongs to the Lamiaceae Martinov family (S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and grows in a restricted range of habitats. Its habitat includes steppe and forest-steppe grasslands, where it co-occurs with various plants, ranging from herbaceous species to dicotyledons. These areas are characterized by chernozem soils on loess and a climate with continental influences. In natural conditions, the habitat exhibits a rich diversity of species, whereas, in degraded conditions, this diversity decreases, with the predominance of monocotyledons and grasses (So\u0026oacute; \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1968\u003c/span\u003e; Csur\u0026ouml;s 1973, 1974; Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The species is also found in rocky areas and rock outcrops (Golevich, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), as well as in grasslands and scrublands (S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). From its phytocoenotic perspective, \u003cem\u003eS. nutans\u003c/em\u003e occupies subdominant positions or is a central component of rank 1\u0026ndash;2 in plant communities (Ziman \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e1976\u003c/span\u003e). There are places where this species dominates the vegetation layer (Kondratyuk et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1985\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is found in Central and Eastern Europe, from the Danube Plain in the west to the Pre-Caucasus in the east (Golevich \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Thus, \u003cem\u003eS. nutans\u003c/em\u003e occurs in Ukraine's southern and central regions, extending from the Crimean Peninsula and the northern Caucasus to the Volga bend. It then continues along the Ural River and the Ural Mountains to the Kama River region (Golevich \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Its distribution also extends to the outer edges of the Carpathians, from Galicia to the Danube (Csapody \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1982\u003c/span\u003e), with occurrences in the Transylvanian Basin, Vojvodina, and the Great Plain. The western limit of its distribution reaches Hungary (Sramk\u0026oacute; and Laczk\u0026oacute; \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The species is also widespread in eastern Romania, more significant in the Transylvanian Basin (So\u0026oacute; \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1942\u003c/span\u003e; Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e) and in Serbia. Historical data indicate its presence in specific locations in Hungary and Romania, suggesting a broader distribution in the past (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). A wider distribution of the species \u003cem\u003eS. nutans\u003c/em\u003e has been recorded throughout the Pannonian Plain, including areas between the Danube and the Tisza, as well as Vojvodina (So\u0026oacute; \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1942\u003c/span\u003e). In Serbia, the species is no longer found (Jovanović and Lakušić \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Đakić, Knežević, and Boža \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). However, in the Transylvanian Basin, the species has been confirmed in some locations (So\u0026oacute; \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1942\u003c/span\u003e; Ruprecht et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). \u003cem\u003eS. nutans\u003c/em\u003e reaches eastwards to the southern limits of the Ural Mountains, although it is no longer part of the current flora of Kazakhstan (Sramk\u0026oacute; and Laczk\u0026oacute; \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe main threat to the survival of \u003cem\u003eS. nutans\u003c/em\u003e is the ongoing transformation of its natural habitat, especially grasslands. Also, the collection of this species for its ornamental value is a major contributor to the population decline. The decline in the number of individuals resulting from these combined pressures also poses additional risks, including genetic hazards, which may impact the long-term viability of the population (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e has not been extensively evaluated for its conservation status at the global and European levels. However, given its presence across various European regions, local assessments might provide insight into its vulnerability or stability within specific habitats. In Romania, for instance, the species may be categorized under specific conservation statuses based on local biodiversity studies, mirroring the approach used for other flora and fauna. For hypothetical context, let us consider that \u003cem\u003eS. nutans\u003c/em\u003e is vulnerable in specific habitats due to environmental pressures or anthropogenic factors. In Central and Southeastern Europe, where \u003cem\u003eS. nutans\u003c/em\u003e naturally occurs, agricultural expansion and changes in land management practices could influence its conservation status.\u003c/p\u003e \u003cp\u003eThe study aims to investigate the habitat and phytocoenological preferences of the \u003cem\u003eS. nutans\u003c/em\u003e in Romania. The objectives of the study include (i) identification of the habitats and plant associations in which \u003cem\u003eS. nutans\u003c/em\u003e is found, (ii) identification of the environmental factors that explain the variation of the floristic composition, and (iii) recommending conservation measures for the protection of the species. The hypothesis of this study is as follows: Due to the transformation of grasslands into agricultural lands and overgrazing, the habitat area of the \u003cem\u003eS. nutans\u003c/em\u003e species in Romania has been reduced. In this case, these factors have led to a decline in the population and number of individuals of this species, especially in Transylvania. We also consider that hybridization with \u003cem\u003eSalvia nemorosa\u003c/em\u003e L. has an essential role in reducing this species.\u003c/p\u003e \u003cp\u003eThe development of measures to protect this species is also important for a unique butterfly species, namely \u003cem\u003ePseudophilotes bavius\u003c/em\u003e subsp. \u003cem\u003ehungarica\u003c/em\u003e (Di\u0026oacute;szeghy, 1913) is endemic to Transylvania and dependent on the host plant \u003cem\u003eS. nutans\u003c/em\u003e (Crișan 2012). The degradation and loss of the \u003cem\u003eS. nutans\u003c/em\u003e species threaten the habitat of this butterfly. The life cycle of the butterfly, which involves laying eggs on host plants and developing into larvae, is directly affected by the conservation status of the \u003cem\u003eS. nutans\u003c/em\u003e species (Crișan 2012; Crișan, Vizauer, and R\u0026aacute;kosy 2023). In this case, the lack of adequate measures for the populations in Romania affects both the plant and the butterfly.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe study areas were represented by the historical regions of Bucovina, Crișana, Dobrogea, Moldova, Muntenia, and Transylvania (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) from Romania. Bucovina and Moldova are located in the northeastern part of Romania and are characterized by a temperate continental climate with moderate summers, cold winters, and abundant precipitation. Dobrogea is characterized by a moderate continental climate with maritime influences and reduced precipitation compared to the other regions. Transylvania features a continental climate characterized by significant thermal variations and abundant precipitation. In contrast, Crișana and Muntenia are characterized by a temperate continental climate, fertile soils, and favourable annual mean temperatures. The lowest elevation was recorded in Dobrogea, and the highest elevation was recorded in Transylvania (Fick and Hijmans \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e, a species associated with xeric and mesoxeric grasslands in the steppe and forest-steppe zones, is linked to the typical soils characteristic of these ecosystems. On plateaus and gently inclined slopes in the steppe area, this species colonizes Kastanozems, Calcic, and Haplic Chernozems. In the same situation in the forest-steppe, the species is found on Haplic and Luvic Chernozems, as well as on various types of Phaeozems. It is interesting that in forest-steppe areas and at the borders of vicinal nemoral forest areas, the species can be frequently encountered on steep, sunny slopes with shallow soils, such as Lithic, Skeletic, Calcic, Brunic, and Calcaric Leptosols, as well as on short-profiled Kastanozems. According to our observations, the species typically avoids colonizing barren rocks (IUSS Working Group WRB \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eStudy species\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) is a perennial species (Cioc\u0026acirc;rlan \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) with a Pannonian-Pontic distribution (Golevich \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The plant height ranges from 20 cm to 100 cm (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) or 120 cm (Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e). The stem is erect, almost leafless in the upper part (S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) or sometimes with 1 or 2 pairs of small, simple leaves (Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e; Cioc\u0026acirc;rlan \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Usually, 1\u0026ndash;2 stems originate from the same rhizome (Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e). The basal leaves are cordate-ovate in shape (Cioc\u0026acirc;rlan \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), with lengths ranging from 3 cm to 16 cm and widths ranging from 2 cm to 12 cm (Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e). The leaf tips are either slightly pointed or rounded, and the margins are crenate. The leaf surface is either glabrous or partially hairy along the central veins. On the underside, the leaves are densely tomentose, with short hairs and numerous sessile glandular hairs of yellow-orange color. The petioles are longer than the leaf blades, with broad bases, and are covered with long, multicellular, dispersed hairs. The stem leaves are small, either sessile or short-petiolate, and lanceolate or occasionally ovate. They have a length from 15 cm to 30 cm and a width from 6 cm to 10 cm. Bracteoles are equal in size or slightly longer than the floral pedicels (Beldie et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1961\u003c/span\u003e). Inflorescence is pendulous at the tip before flowering (Cioc\u0026acirc;rlan \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The flowering period extends from late May to early July (Kondratyuk et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). Although most individuals flower in the third year of growth, there are cases in which flowering occurs as early as the second year (Udvardy \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eVegetation data\u003c/h3\u003e\n\u003cp\u003eA total of 870 relev\u0026eacute;s, comprising 1,421 taxa, were used for the vegetation analysis. Of these, 492 relev\u0026eacute;s were sourced from the Romanian Grassland Database (RGD; Vassilev et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), while the remaining 378 relev\u0026eacute;s were collected between May 2017 and June 2024 (personal data). The size of each individual plot was 100 m\u0026sup2;, while the relev\u0026eacute;s from the Romanian Grassland Database ranged from 4 m\u0026sup2; to 200 m\u0026sup2;.\u003c/p\u003e\n\u003ch3\u003eEnvironmental variables\u003c/h3\u003e\n\u003cp\u003eIn the multivariate analysis, abiotic variables (elevation \u0026ndash; m a.s.l., aspect \u0026ndash; \u0026deg;, slope \u0026ndash; \u0026deg;, mean annual temperature \u0026ndash; ℃, and annual precipitation \u0026ndash; mm) and edaphic variables (P \u0026ndash; mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, K \u0026ndash; mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and pH) were included.\u003c/p\u003e\n\u003ch3\u003eSoil analyses\u003c/h3\u003e\n\u003cp\u003eThe values for the chemical elements P, K, and soil pH (Ballabio et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) were extracted from the European Soil Database \u0026amp; soil properties \u0026ndash; ESDAC (European Soil Data Centre \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eClimatic data\u003c/h2\u003e \u003cp\u003eThe climatic data (mean annual temperature and annual precipitation) were obtained from the WorldClim database (Fick and Hijmans \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) at a resolution of 30 arc. sec. Elevation, aspect, and slope data were collected in the field.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eData preparation for classification analyses\u003c/h3\u003e\n\u003cp\u003eBefore numerical data analysis, the dataset was standardized according to the following criteria: (i) unification of taxonomy and taxon nomenclature; (ii) species identified only at genus level were eliminated; (iii) unrecognized subspecies were combined into species; (iv) exclusion of bryophyte and lichen species, as they were recorded in only 20 relev\u0026eacute;s.\u003c/p\u003e \u003cp\u003eThe classification of plant associations within the alliance and the nomenclature of these associations follow Chifu, Irimia, and Zamfirescu (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and Chifu and Irimia (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2014a\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014b\u003c/span\u003e). The nomenclature and taxonomy of plant species follow Euro\u0026thinsp;+\u0026thinsp;Med (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), and the nomenclature of higher syntax units follows Mucina et al. (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Additionally EUNIS habitats were determined (Chytr\u0026yacute; et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The final data set included 870 relev\u0026eacute;s and 904 taxa.\u003c/p\u003e\n\u003ch3\u003eNumerical classification and expert system development\u003c/h3\u003e\n\u003cp\u003eWe used the modified TWINSPAN algorithm (Roleček et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) with three pseudospecies cut levels of 0%, 5%, and 25% cover, minimum group size for division of five relev\u0026eacute;s, and Whittaker\u0026rsquo;s beta coefficient as a measure of internal cluster heterogeneity. We assessed the optimal number of clusters using OptimClass 1 (Tich\u0026yacute; et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Diagnostic species for individual clusters were determined based on the \u003cem\u003ephi\u003c/em\u003e-coefficient of association applied to virtually standardized cluster sizes (Chytr\u0026yacute; et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2002\u003c/span\u003e, Tich\u0026yacute; and Chytr\u0026yacute; \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). We used a fidelity threshold of \u003cem\u003ephi\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.25 combined with Fisher\u0026rsquo;s exact test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). We used 25% and 50% constancy as thresholds for determining constant and highly constant species, respectively.\u003c/p\u003e \u003cp\u003eThe data were represented as mean percentage cover values, according to the scale developed by the Braun-Blanquet (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1964\u003c/span\u003e). The mean abundance-dominant interval corresponding to the notes was defined as follows (Cristea, Gafta, and Pedrotti \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2004\u003c/span\u003e): r (0.05%); + (0.5%); 1 (5%); 2 (17.5%); 3 (37.5%); 4 (62.5%); and 5 (87.5%).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eVegetation\u0026ndash;environment relationship\u003c/h2\u003e \u003cp\u003eThe correlation between floristic composition and environmental variables was analyzed with Detrended Correspondence Analysis (DCA) in the CANOCO v5.1 software (ter Braak \u0026amp; Šmilauer, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In this context, DCA analysis was performed to detect floristic gradients, and CCA (Canonical Correspondence Analysis) was applied to quantify the effect of each environmental variable on floristic composition, using the Monte Carlo permutation test (9999 iterations). The average percentage coverage values represented the floristic composition, and the environmental variables were represented by variables with a VIF value of less than 5 (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We used the Variance Inflation Factor (VIF) in SPSS version 23 to assess collinearity between independent variables. Variables with a VIF value greater than 5 were considered multicollinear and were excluded from the model. The relev\u0026eacute;s distribution map was made in QGIS version 3.34.3 (QGIS Development Team \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eAnalyses of differences between syntaxa\u003c/h2\u003e \u003cp\u003eTo determine whether there are statistically significant differences between plant communities regarding environmental variables, the ANOVA test was applied, using R Statistical Software (v4.1.4; R Core Team \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) via the 'tidyverse' v.2.0.0 (Wickham et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), 'car' v.3.1.3 (Fox and Weisberg \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), and 'stats' v.4.5.0 (R Core Team \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Since significant differences were found (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), the post-hoc Tukey test was applied. In this context, communities that did not exhibit significant differences were denoted by the same letter, while communities that did exhibit significant differences were highlighted by different letters.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\n \u003ch2\u003eSyntaxonomic overview of relev\u0026eacute;s\u003c/h2\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003e\u003cem\u003eSalvia nutans\u003c/em\u003e populations were found to inhabit eight main phytocoenological classes:\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eCrataego-Prunetea\u003c/em\u003e, \u003cem\u003eDigitario sanguinalis-Eragrostietea minoris\u003c/em\u003e, \u003cem\u003eFestuco-Brometea\u003c/em\u003e, \u003cem\u003eFestuco-Puccinellietea\u003c/em\u003e, \u003cem\u003eMolinio-Arrhenatheretea\u003c/em\u003e, \u003cem\u003eQuercetea pubescentis\u003c/em\u003e, \u003cem\u003eSisymbrietea\u003c/em\u003e, and \u003cem\u003eTrifolio-Geranietea sanguinei\u003c/em\u003e.\u003c/p\u003e\n \u003c/div\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eFestuco-Brometea\u003c/em\u003e Br.-Bl. et Tx. ex So\u0026oacute; 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003eFestucetalia valesiacae\u003c/em\u003e So\u0026oacute; 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.:\u0026nbsp;\u003c/strong\u003e\u003cem\u003eStipion lessingianae\u003c/em\u003e So\u0026oacute; 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAllio albidi-Stipetum lessingianae\u003c/em\u003e (So\u0026oacute; 1947) Coldea et S\u0026acirc;rbu in Coldea 2012\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/em\u003e (So\u0026oacute; 1942) Coldea et S\u0026acirc;rbu in Coldea 2012\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eJurineo\u003c/em\u003e \u003cem\u003earachnoidea-Stipetum lessingianae\u003c/em\u003e (Dobrescu 1974) Chifu, M\u0026acirc;nzu et Zamfirescu 2006\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eFestucetum rupicolae\u003c/em\u003e Burduja et al. 1956\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eArtemisietum ponticae\u003c/em\u003e (So\u0026oacute; 1942) Păun 1969\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eStipo ucrainicae-Festucetum valesiacae\u003c/em\u003e (Dihoru 1969) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eConvolvulo cantabrici-Stipetum capillatae\u003c/em\u003e (Horeanu 1976)\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgropyro pectinati-Stipetum capillatae\u003c/em\u003e (Burduja et al. 1956) Chifu, Manzu et Zamfirescu 2006\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/em\u003e (Burduja et al. 1956, Răvăruț et al. 1956) S\u0026acirc;rbu, Coldea et Chifu 1999\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgropyro pectinati-Tanacetinetum millefolii\u003c/em\u003e (Șerbănescu 1970) Chifu et Țupu 2009\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eElytrigietum hispidi\u003c/em\u003e (Dihoru 1970) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAstero oleifolius-Ephedretum distachyae\u003c/em\u003e Horeanu et Vițalariu 1992\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eCynodonto-Po\u0026euml;tum angustifoliae\u003c/em\u003e (Rapaics 1927) So\u0026oacute; 1957\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eBombycilaeno erecti-Bothriochloetum ischaemi\u003c/em\u003e (Dihoru 1970) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgropyretum pectiniformae\u003c/em\u003e (Dihoru 1970) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.:\u0026nbsp;\u003c/strong\u003e\u003cem\u003eFestucion valesiacae\u003c/em\u003e Klika 1931\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eBothriochloetum ischaemi\u003c/em\u003e (Kristiansen 1937) I. Pop 1977\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eSalvio-Festucetum rupicolae\u003c/em\u003e Zolyomi 1958 corr. So\u0026oacute; 1964\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Sub-ass.\u003c/strong\u003e: \u003cem\u003epaeonietosum tenuifoliae\u003c/em\u003e (Mititelu et al. 1990)\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eFestucetum valesiacae-rupicolae\u003c/em\u003e Cs\u0026uuml;r\u0026ouml;s et Kov\u0026aacute;cs 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eStipetum capillatae\u003c/em\u003e (Hueck 1963) Krausch 1961\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgrostio-Festucetum valesiacae\u003c/em\u003e Borisavljevič et al. 1955\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgropyro pectinati-Kochietum prostratae\u003c/em\u003e Zolyomi 1958\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eTaraxaco serotinae-Bothriochloetum ischaemi\u003c/em\u003e (Burduja et al. 1956) S\u0026acirc;rbu, Coldea et Chifu 1999\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Sub-ass.\u003c/strong\u003e: \u003cem\u003ekoelerietosum macranthae\u003c/em\u003e (Răvăruț et al. 1956) Chifu, M\u0026acirc;nzu et Zamfirescu 2006\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eMedicagini minimae-Festucetum valesiacae\u003c/em\u003e Wagner 1941\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eFestuceto rupicolae-Caricetum humilis\u003c/em\u003e Polg\u0026aacute;r 1933\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Sub-ass.\u003c/strong\u003e: \u003cem\u003ethymio-salvietosum nutantis\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eOrder\u003c/strong\u003e: \u003cem\u003eBrachypodietalia pinnati\u003c/em\u003e Korneck 1974\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e.: \u003cem\u003eCirsio-Brachypodion pinnati\u003c/em\u003e Hadač et Klika in Klika et Hadač 1944\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eCariceto humilis-Brachypodietum pinnati\u003c/em\u003e So\u0026oacute; 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eSub-ass.\u003c/strong\u003e: \u003cem\u003edichathietosum ischaemi\u0026nbsp;\u003c/em\u003eChifu et al.. 2014\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eHelianthemo cani-Seslerietum heuflerianae\u003c/em\u003e (Borza 1959) Popescu et Sanda 1992\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eFestuco rupicolae-Brachypodietum pinnati\u003c/em\u003e Mahn 1965\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eJurineo transsilvanicae-Chrysopogonetum grylli\u003c/em\u003e (Cs\u0026uuml;r\u0026ouml;s et Niedermaier 1966) Coldea et S\u0026acirc;rbu in Coldea 2012\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eDanthonio alpinae-Stipetum stenophyllae\u003c/em\u003e Ghișa 1941\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eCarici humilis-Stipetum joannis\u003c/em\u003e Pop et Hodișan 1985\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eConvolvulo cantabrici-Chrysopogonetum grylli\u003c/em\u003e (Dihoru 1970) in Chifu et al. 2014\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eValerianello lasiocarpae-Chrysopogonetum grylli\u003c/em\u003e (Dihoru 1970) Coldea et S\u0026acirc;rbu in Coldea 2012\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eFerrulago campestris-Caricetum humilis\u003c/em\u003e (Cioc\u0026acirc;rlan 1969) Coldea et al. 2010\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eGalio octonarii-Stipetum tirsae\u003c/em\u003e (Cioc\u0026acirc;rlan 1969) Popescu et Sanda 1992\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eRhinantho rumelici-Brometum erecti\u003c/em\u003e Sanda et Popescu 1999\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eThymo comosi-Caricetum humilis\u003c/em\u003e Morariu et Danciu 1977\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eOrder\u003c/strong\u003e: \u003cem\u003eStipo pulcherrimae-Festucetalia pallentis\u003c/em\u003e Pop 1968\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e.: \u003cem\u003ePimpinello-Thymion zygoidis\u003c/em\u003e Dihoru et Doniţa 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eSaturejetum caeruleae\u003c/em\u003e Cristurean et Ionescu-Țeculescu 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eKoelerio lobatae-Artemisietum lerchianae\u003c/em\u003e (Dihoru 1970) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eAgropyro pontici-Thymetum zygioidis\u003c/em\u003e (Dihoru 1970) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eFestucetum callierii\u003c/em\u003e (Şerbănescu 1965) Dihoru et Doniță 1970\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eCrataego-Prunetea\u003c/em\u003e Tx. 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003ePrunetalia spinosae\u003c/em\u003e Tx. 1952\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003ePrunion fruticosae\u003c/em\u003e Tx. 1952\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003ePrunetum tenellae\u003c/em\u003e So\u0026oacute; 1951\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eBerberidion vulgaris\u003c/em\u003e Br.-Bl. ex Tx. 1952\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003ePruno spinosae-Crataegetum\u003c/em\u003e Hueck 1931\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eTrifolio-Geranietea sanguinei\u003c/em\u003e T. M\u0026uuml;ller 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eOrder\u003c/strong\u003e: \u003cem\u003eAntherico ramosi-Geranietalia sanguinei\u003c/em\u003e Julve ex Dengler in Dengler et al. 2003\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAll.\u003c/strong\u003e: \u003cem\u003eGeranion sanguinei\u003c/em\u003e Tx. in T. M\u0026uuml;ller 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eInulo ensifoliae-Peucedanetum cervariae Kozlowska\u0026nbsp;\u003c/em\u003e1925 em. Van Gils et Kov\u0026aacute;cs 1977\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eGeranio-Dictamnetum\u003c/em\u003e Wendelberger ex T. M\u0026uuml;ller 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eAss.\u003c/strong\u003e: \u003cem\u003eClematido rectae-Laserpitietum latifolii\u003c/em\u003e Schneider-Binder 1984\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eMolinio-Arrhenatheretea\u003c/em\u003e Tx. 1937\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003eArrhenatheretalia elatioris\u003c/em\u003e Tx. 1931\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eCynosurion cristati\u003c/em\u003e Tx. 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eAnthoxantho-Agrostetum capillaris\u003c/em\u003e Sillinger 1933\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eArrhenatherion elatioris\u003c/em\u003e Koch 1926\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eArrhenatheretum elatioris\u003c/em\u003e Br.-Bl. ex Scherrer 1925\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003ePotentillo-Polygonetalia avicularis\u003c/em\u003e Tx. 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003ePotentillion anserinae\u003c/em\u003e Tx. 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eRorippo austriacae-Agropyretum repentis\u003c/em\u003e (Timar 1947) R. Tx. 1950\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eFestuco-Puccinellietea\u003c/em\u003e So\u0026oacute; ex Vicherek 1973\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003ePuccinellietalia\u003c/em\u003e So\u0026oacute; 1947\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eFestucion pseudovinae\u003c/em\u003e So\u0026oacute; 1933\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eArtemisio-Festucetum pseudovinae\u003c/em\u003e So\u0026oacute; (1927) 1945\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eSisymbrietea\u003c/em\u003e Gutte et Hilbig 1975\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003eSisymbrietalia sophiae\u003c/em\u003e J. Tx. ex G\u0026ouml;rs 1966\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eSisymbrion officinalis\u003c/em\u003e Tx. et al. ex von Rochow 1951\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eBromo squarrosi-Xeranthemetum annui\u003c/em\u003e Coroi 2001\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eHordeo murini-Cynodontetum dactyloni\u003c/em\u003e Felf\u0026ouml;ldy ex Borhidi 1949\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eQuercetea pubescentis\u003c/em\u003e Doing-Kraft ex Scamoni et Passarge 1959\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003eQuercetalia pubescenti-petraeae\u003c/em\u003e Klika 1933\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eQuercion petraeae\u003c/em\u003e Issler 1931\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003eCorno-Quercetum pubescentis\u003c/em\u003e Math\u0026eacute; et Kov\u0026aacute;cs 1962\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Ass.\u003c/strong\u003e: \u003cem\u003ePaeonio peregrinae-Quercetum pubescentis\u003c/em\u003e (S\u0026acirc;rbu 1982) Sanda et Popescu 1999\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eClass\u003c/strong\u003e: \u003cem\u003eDigitario sanguinalis-Eragrostietea minoris\u003c/em\u003e Mucina, Lososov\u0026aacute; et \u0026Scaron;ilc in Mucina et al. 2016\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;Order\u003c/strong\u003e: \u003cem\u003eEragrostietalia\u003c/em\u003e J. Tx. ex Poli 1966\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;All.\u003c/strong\u003e: \u003cem\u003eSalsolion ruthenicae\u003c/em\u003e Philippi ex Oberd. 1983\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Community\u003c/strong\u003e: \u003cem\u003eCynodon dactylon\u003c/em\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e\n \u003ch2\u003eEUNIS habitats\u003c/h2\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003eThe identified plant associations were assigned to eight major EUNIS habitat types, namely (at level 2 of the EUNIS hierarchy): R1 \u0026ndash; \u003cstrong\u003eDry grasslands (\u003c/strong\u003eR1A Semi-dry perennial calcareous grassland - meadow steppe, R1B Continental dry grassland - true steppe, R16 Perennial rocky grassland of Central and South-Eastern Europe); R2 \u0026ndash; \u003cstrong\u003eMesic grasslands\u003c/strong\u003e (R21 Mesic permanent pasture of lowlands and mountains, and R22 Low and medium altitude hay meadow); R3 \u0026ndash; \u003cstrong\u003eSeasonally wet and wet grasslands\u003c/strong\u003e (R36 Moist or wet mesotrophic to eutrophic pasture); R5 \u0026ndash;\u003cstrong\u003eWoodland fringes and clearings and tall forb stands\u003c/strong\u003e (R51 Thermophilous forest fringe of base-rich soils); R6 \u0026ndash; \u003cstrong\u003eInland salt steppes\u003c/strong\u003e (R62 Continental inland salt steppe); S3 \u0026ndash; \u003cstrong\u003eTemperate and Mediterranean montane scrub\u003c/strong\u003e (S35 Temperate and submediterranean thorn scrub); T1 \u0026ndash;\u003cstrong\u003eBroadleaved deciduous forests\u003c/strong\u003e (T19 Temperate and submediterranean thermophilous deciduous forest); and V1 \u0026ndash; \u003cstrong\u003eArable land and market gardens\u003c/strong\u003e (V15 Bare tilled, fallow or recently abandoned arable land).\u003c/p\u003e\n \u003c/div\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\n \u003ch2\u003eCluster analysis\u003c/h2\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003eThe cluster analysis results were presented as a dendrogram (Fig. \u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e), which illustrated the hierarchical relationships between the vegetation units, and a synoptic table (Supplementary Material 1). The analyzed vegetation was classified into six clusters, which were correlated with different syntax groups recognized in the specialized literature.\u003c/p\u003e\n \u003c/div\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 1: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eFestucetalia valesiacae\u003c/strong\u003e, \u003cstrong\u003eAl.\u003c/strong\u003e \u003cstrong\u003eStipion lessingianae\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eStipo ucrainicae-Festucetum valesiacae\u003c/strong\u003e \u003cstrong\u003e(17 relev\u0026eacute;s)\u003c/strong\u003e\u003c/p\u003e\n \u003c/div\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003ePhlomis herba-venti\u003c/em\u003e had a fidelity of 58% and was the species most strongly associated with this community. Other important species for this community that had relatively significant contributions were \u003cem\u003eCentaurea orientalis\u003c/em\u003e (53.2%) and \u003cem\u003eCytisus austriacus\u003c/em\u003e (52.6%). The number of species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 13 to 44, and vegetation cover varied between 48% and 83%. \u003cem\u003eSalvia nutans\u003c/em\u003e had a cover ranging from 0.5\u0026ndash;5%.\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe communities analysed were recorded at elevations varied between 22 to 122 m a.s.l. (above sea level). The climate is moderate, with annual mean temperatures ranging from 10.9 to 11.9\u0026deg;C. Annual precipitation was also moderate, ranging from 432 to 470 mm. The temperatures in May and June varied between 15.6 and 16.1\u0026deg;C. They ranged from 20.2 to 20.7\u0026deg;C in June, while precipitation was lower in May (37\u0026ndash;45 mm) compared to June (48\u0026ndash;55 mm), suggesting a gradual increase in humidity during the warm season. Soils ranged from neutral to slightly alkaline, with moderate phosphorus and potassium concentrations. The communities were recorded on gentle slopes with a northeastern aspect in the grasslands of Constanța County.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 2: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eFestucetalia valesiacae\u003c/strong\u003e, \u003cstrong\u003eAl.\u003c/strong\u003e \u003cstrong\u003eStipion lessingianae\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/strong\u003e \u003cstrong\u003eand\u003c/strong\u003e \u003cstrong\u003eJurineo arachnoidea-Stipetum lessingianae\u003c/strong\u003e \u003cstrong\u003e(\u003c/strong\u003e36 relev\u0026eacute;s\u003cstrong\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eGalium glaucum\u003c/em\u003e had a fidelity of 63.2%, the species most strongly associated with this community. Similarly, \u003cem\u003eFestuca stricta\u003c/em\u003e subsp. \u003cem\u003esulcata\u003c/em\u003e (60.6%) and \u003cem\u003eCarex humilis\u003c/em\u003e (60.1%) contributed significantly to the structure and functioning of the community. Another species contributing to the community structure was \u003cem\u003eAstragalus monspessulanus\u003c/em\u003e, which had a fidelity of 56.6%, similar to \u003cem\u003eThymus odoratissimus\u003c/em\u003e, which had a fidelity of 54%. Similarly, \u003cem\u003ePeucedanum ruthenicum\u003c/em\u003e is a representative species for this community, although it had a lower fidelity (51%). The number of plant species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 13 to 95, while vegetation cover varied between 51% and 100%. \u003cem\u003eSalvia nutans\u003c/em\u003e had a cover between 0.05% and 5%.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe annual mean temperatures range from 6.9 to 11.6\u0026deg;C, indicating a moderate to cold climate. The annual precipitation ranges from 428 to 590 mm, indicating moderate humidity conditions. The mean monthly temperatures in May and June range from 12.6 to 16.1\u0026deg;C in May and 16 to 20.7\u0026deg;C in June. The monthly precipitation ranges from 41 to 72 mm in May and from 50 to 95 mm in June. The soils had a pH ranging from weakly alkaline to neutral, characterized by moderate phosphorus and rich in potassium concentrations. The communities in this cluster were recorded at elevations ranging from 31 to 630 m a.s.l., on slopes that varied from gentle to very steep, predominantly with a northeastern aspect across the grasslands of Alba, Brașov, Constanța, Iași, Suceava, and Vaslui counties.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 3: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eFestucetalia valesiacae\u003c/strong\u003e, \u003cstrong\u003eAll.\u003c/strong\u003e \u003cstrong\u003eStipion lessingianae\u003c/strong\u003e, \u003cstrong\u003ePimpinello-Thymion zygoidis\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/strong\u003e \u003cstrong\u003eand\u003c/strong\u003e \u003cstrong\u003eAgropyro pontici-Thymetum zygioidis\u003c/strong\u003e \u003cstrong\u003e(\u003c/strong\u003e57 relev\u0026eacute;s\u003cstrong\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eConvolvulus cantabrica\u003c/em\u003e was the species most strongly associated with this community, with a fidelity of 61.3%. \u003cem\u003eErysimum diffusum\u003c/em\u003e (59%) and \u003cem\u003eKoeleria lobata\u003c/em\u003e (58.6%) had an essential role in these communities, contributing significantly to the community structure. Also, other species with relatively significant fidelity were \u003cem\u003eEuphorbia seguierana\u003c/em\u003e (55.9%), \u003cem\u003eSatureja coerulea\u003c/em\u003e (54.2%) and \u003cem\u003eAchillea coarctata\u003c/em\u003e (54.2%). \u003cem\u003eAgropyron cristatum\u003c/em\u003e recorded a fidelity of 50.7%. The number of plant species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 15 to 85, and vegetation cover ranged from 45\u0026ndash;100%. \u003cem\u003eSalvia nutans\u003c/em\u003e had a cover ranging from 0.05\u0026ndash;5%.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe climate was characterized by moderate temperatures and precipitation. Mean monthly temperatures ranged from 14.8 to 16.8\u0026deg;C in May and from 19.1 to 20.7\u0026deg;C in June. Monthly precipitation was moderate, from 38 to 55 mm in May and from 47 to 67 mm in June. The soils associated with these communities had a pH ranging from slightly acidic to neutral and were characterized by moderate phosphorus and potassium concentrations. Communities in this cluster were recorded at elevations ranging from 10 to 286 m a.s.l., on slopes ranging from gentle to very steep, with a predominantly northeastern aspect in the grasslands of Constanța and Tulcea counties.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 4: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eFestucetalia valesiacae\u003c/strong\u003e, \u003cstrong\u003eAll.\u003c/strong\u003e \u003cstrong\u003eStipion lessingianae\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/strong\u003e, \u003cstrong\u003eAllio albidi-Stipetum lessingianae\u003c/strong\u003e, \u003cstrong\u003eand\u003c/strong\u003e \u003cstrong\u003ePrunetum tenellae\u003c/strong\u003e (562 relev\u0026eacute;s)\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eCrataegus monogyna\u003c/em\u003e (66.9%) and \u003cem\u003ePotentilla taurica\u003c/em\u003e (66.3%) had the highest fidelity values, characteristic species for this community. \u003cem\u003eValeriana coronata\u003c/em\u003e (65.2%) and \u003cem\u003eRosa canina\u003c/em\u003e (62.2%) were other important species, significantly contributing to the community structure. Also, the species \u003cem\u003eMarrubium vulgare\u003c/em\u003e (61%) and \u003cem\u003eSalvia revelata\u003c/em\u003e (55.9%) recorded a relatively high fidelity. Species such as \u003cem\u003eThalictrum aquilegiifolium\u003c/em\u003e (54.4%), \u003cem\u003eAchillea ochroleuca\u003c/em\u003e (51.6%), \u003cem\u003ePotentilla reptans\u003c/em\u003e (50.8%), and \u003cem\u003eEuphrasia stricta\u003c/em\u003e (50%) recorded a moderate to high fidelity. The number of species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 9 to 99, and the vegetation cover ranged from 29\u0026ndash;100%. \u003cem\u003eSalvia nutans\u003c/em\u003e had a cover ranging from 0.05\u0026ndash;37.5%.\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe communities included in this cluster were recorded at elevations from 95 to 610 m a.s.l., on very steep slopes with a southwest aspect. Annual precipitation ranged from 575 to 731 mm, and annual mean temperatures ranged from 6.2 to 10 ℃. The soils had a pH ranging from slightly acidic to weakly alkaline, characterized by moderate phosphorus and rich potassium concentrations in the grasslands of Alba, Bistrita-Năsăud, Brașov, Cluj, Mureș, Sălaj, Sibiu, and Suceava counties.\u003c/p\u003e\n \u003cp\u003e\u0026deg;\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 5: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eBrachypodietalia pinnati\u003c/strong\u003e, \u003cstrong\u003eAll.\u003c/strong\u003e \u003cstrong\u003eCirsio-Brachypodion pinnati\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eCariceto humilis-Brachypodietum pinnati\u003c/strong\u003e \u003cstrong\u003e(\u003c/strong\u003e157 relev\u0026eacute;s\u003cstrong\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eBrachypodium pinnatum\u003c/em\u003e recorded a fidelity of 62.2%, the species most strongly associated with this community. The number of species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 45 to 58, and the vegetation cover ranged from 30\u0026ndash;100%. Within this community, \u003cem\u003eSalvia nutans\u003c/em\u003e recorded a relatively variable cover, ranging from 0.05\u0026ndash;37.5%.\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe communities analysed were recorded at elevations ranging from 90 to 1553 m a.s.l., located on slopes ranging from moderate to very steep with a predominantly southern and western aspect. The soils presented a pH ranging from slightly acidic to neutral, characterised by moderate phosphorus and potassium concentrations. Annual precipitation ranged from 449 to 990 mm, and annual mean temperatures ranged from 2.4 to 10.5 ℃ in the grasslands of Alba, Bacău, Bihor, Brașov, Cluj, Covasna, Harghita, Mureș, Sălaj, Sibiu, and Suceava counties.\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eCluster 6: Cl.\u003c/strong\u003e \u003cstrong\u003eFestuco-Brometea\u003c/strong\u003e, \u003cstrong\u003eOrd.\u003c/strong\u003e \u003cstrong\u003eFestucetalia valesiacae\u003c/strong\u003e, \u003cstrong\u003eAll.\u003c/strong\u003e \u003cstrong\u003eStipion lessingianae\u003c/strong\u003e, \u003cstrong\u003eAss.\u003c/strong\u003e \u003cstrong\u003eTaraxaco serotinae-Bothriochloetum ischaemi\u003c/strong\u003e \u003cstrong\u003eand\u003c/strong\u003e \u003cstrong\u003eAstero oleifolius-Ephedretum distachyae\u003c/strong\u003e \u003cstrong\u003e(\u003c/strong\u003e41 relev\u0026eacute;s\u003cstrong\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eStructure and composition of plant community\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eGalium humifusum\u003c/em\u003e was the species most strongly associated with this community, with a fidelity of 59.9%. The other species, such as \u003cem\u003eAstragalus onobrychis\u003c/em\u003e (39.5%), \u003cem\u003eEphedra distachya\u003c/em\u003e (39.3%), and \u003cem\u003eDiplotaxis muralis\u003c/em\u003e (39.3%), had lower fidelity. The number of species per 100 m\u003csup\u003e2\u003c/sup\u003e ranged from 7 to 61, and the vegetation cover ranged from 47\u0026ndash;98%. \u003cem\u003eSalvia nutans\u003c/em\u003e presented a cover range of 0.5\u0026ndash;5%.\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eEcology and distribution\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003eThe communities included in this cluster were recorded at elevations ranging from 5 to 387 m a.s.l., on moderate slopes with a northeastern aspect. Climatic conditions were characterized by annual precipitation ranging from 429 to 609 mm and annual mean temperatures ranging from 8.6 to 12 ℃ in the grasslands of Constanța, Iași, Sibiu, Suceava, Tulcea, and Vaslui counties.\u003c/p\u003e\n \u003ctable id=\"Tab1\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eMeasured values of bioclimatic, soil chemical, and topographic variables were analysed. Values are means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviations (SD). VEGC\u0026thinsp;=\u0026thinsp;vegetation cover; BIO12\u0026thinsp;=\u0026thinsp;annual precipitation; BIO1\u0026thinsp;=\u0026thinsp;annual mean temperature; Aspect: S \u0026ndash; south, SSW \u0026ndash; south-southwest, SE \u0026ndash; southeast, SSE \u0026ndash; south-southeast, SW \u0026ndash; southwest, W \u0026ndash; west; P\u0026thinsp;=\u0026thinsp;phosphorus, K\u0026thinsp;=\u0026thinsp;potassium.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 1\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 2\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 3\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 4\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 5\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCluster 6\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eNo. of species/relev\u0026eacute;s\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28\u0026thinsp;\u0026plusmn;\u0026thinsp;8.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e43\u0026thinsp;\u0026plusmn;\u0026thinsp;20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e48\u0026thinsp;\u0026plusmn;\u0026thinsp;14.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e40\u0026thinsp;\u0026plusmn;\u0026thinsp;16.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e38\u0026thinsp;\u0026plusmn;\u0026thinsp;16.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e35\u0026thinsp;\u0026plusmn;\u0026thinsp;15.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eVegetation cover (%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e61\u0026thinsp;\u0026plusmn;\u0026thinsp;9.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e87\u0026thinsp;\u0026plusmn;\u0026thinsp;12.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e83\u0026thinsp;\u0026plusmn;\u0026thinsp;13.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e83\u0026thinsp;\u0026plusmn;\u0026thinsp;12.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e82\u0026thinsp;\u0026plusmn;\u0026thinsp;14.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e77\u0026thinsp;\u0026plusmn;\u0026thinsp;14.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eAspect\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSSE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSSW\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSlope (\u0026deg;)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.6\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e14.5\u0026thinsp;\u0026plusmn;\u0026thinsp;15.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e8.1\u0026thinsp;\u0026plusmn;\u0026thinsp;6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e30.6\u0026thinsp;\u0026plusmn;\u0026thinsp;14.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e22.1\u0026thinsp;\u0026plusmn;\u0026thinsp;14.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eElevation (m a.s.l.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e76.6\u0026thinsp;\u0026plusmn;\u0026thinsp;25.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e204\u0026thinsp;\u0026plusmn;\u0026thinsp;142.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e91.5\u0026thinsp;\u0026plusmn;\u0026thinsp;85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e392.6\u0026thinsp;\u0026plusmn;\u0026thinsp;67.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e450.6\u0026thinsp;\u0026plusmn;\u0026thinsp;138.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e178.3\u0026thinsp;\u0026plusmn;\u0026thinsp;99.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003epH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e6.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eP (mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e27.2\u0026thinsp;\u0026plusmn;\u0026thinsp;7.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28.2\u0026thinsp;\u0026plusmn;\u0026thinsp;6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e27\u0026thinsp;\u0026plusmn;\u0026thinsp;7.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e26.0\u0026thinsp;\u0026plusmn;\u0026thinsp;6.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e24.5\u0026thinsp;\u0026plusmn;\u0026thinsp;8.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28.4\u0026thinsp;\u0026plusmn;\u0026thinsp;7.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eK (mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e263.3\u0026thinsp;\u0026plusmn;\u0026thinsp;71.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e342.1\u0026thinsp;\u0026plusmn;\u0026thinsp;99.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e278.5\u0026thinsp;\u0026plusmn;\u0026thinsp;76.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e297.4\u0026thinsp;\u0026plusmn;\u0026thinsp;56.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e281.2\u0026thinsp;\u0026plusmn;\u0026thinsp;73\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e331.4\u0026thinsp;\u0026plusmn;\u0026thinsp;132\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBO1 (℃)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e8.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBIO12 (mm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e453.7\u0026thinsp;\u0026plusmn;\u0026thinsp;16.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e549.7\u0026thinsp;\u0026plusmn;\u0026thinsp;34.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e443.9\u0026thinsp;\u0026plusmn;\u0026thinsp;15.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e608.3\u0026thinsp;\u0026plusmn;\u0026thinsp;19.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e608.5\u0026thinsp;\u0026plusmn;\u0026thinsp;43.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e524.2\u0026thinsp;\u0026plusmn;\u0026thinsp;53.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eMean temperature \u0026ndash; May\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e14.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e14\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eMean temperature \u0026ndash; June\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e18.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e17.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e19.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eMean precipitation \u0026ndash; May\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e41.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e62.8\u0026thinsp;\u0026plusmn;\u0026thinsp;7.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e43.5\u0026thinsp;\u0026plusmn;\u0026thinsp;3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e72.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e73.5\u0026thinsp;\u0026plusmn;\u0026thinsp;4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e58.5\u0026thinsp;\u0026plusmn;\u0026thinsp;10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eMean precipitation \u0026ndash; June\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e52.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.2\u0026thinsp;\u0026plusmn;\u0026thinsp;10.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e53.8\u0026thinsp;\u0026plusmn;\u0026thinsp;4.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e93.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e93.7\u0026thinsp;\u0026plusmn;\u0026thinsp;5.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e76.6\u0026thinsp;\u0026plusmn;\u0026thinsp;14.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003e\u003cem\u003eSalvia nutans\u003c/em\u003e occurs predominantly on steep slopes (from 21\u0026deg; to 35\u0026deg;; 31%), followed by very steep slopes (\u0026gt;\u0026thinsp;35\u0026deg;), with 26%. The species also occurs on gentle slopes (3\u0026ndash;10\u0026deg;; 23%), moderate slopes (11\u0026ndash;20\u0026deg;; 15%), and very gentle slopes (0\u0026ndash;3\u0026deg;; 5%). The dominant aspect in which \u003cem\u003eS. nutans\u003c/em\u003e occurs is southwest (39%), followed by southern (23%), and western (13%). The species also occurs on secondary aspects, such as southeast (8%) and northeast (7%). In contrast, on the northwest (5%), eastern (3%), and northern (3%) aspects, the species recorded a reduced presence. In terms of elevation, \u003cem\u003eS. nutans\u003c/em\u003e prefers low elevations. The average elevation was 359 m, ranging from 5 to 850 m a.s.l. (Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\n \u003c/div\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003eThe soils were predominantly moderately acidic (pH range of 5.81 to 6.80, 59%). The species was also identified on slightly acidic soils (pH 5.01\u0026ndash;5.80, 27%), neutral soils (pH 6.81\u0026ndash;7.20, 11%), slightly alkaline soils (pH 7.21\u0026ndash;8.40, 2%), and moderately strongly acidic soils (pH 4.31\u0026ndash;5.00, 1%). Based on the soil phosphorus concentration, the species distribution indicates a preference for soils with low phosphorus concentrations (ranging from 18.1 to 36 mg kg\u003csup\u003e-1\u003c/sup\u003e, or 81%). The presence of the species in soils with moderate phosphorus concentrations (ranging from 8.1 to 18 mg kg\u003csup\u003e-1\u003c/sup\u003e, 13%) and high phosphorus levels (ranging from 36.1 to 72 mg kg\u003csup\u003e-1\u003c/sup\u003e, 6%) was reduced. Regarding potassium concentration, a preference for soils very rich in potassium (from 265.1 to 400 mg kg\u003csup\u003e-1\u003c/sup\u003e; 69%) was observed among the species. The presence of the species in soils rich in potassium (ranging from 200.1 to 265 mg kg\u003csup\u003e-1\u003c/sup\u003e, 25%) was moderate. In soils with moderate (ranging from 132.1 to 200 mg kg\u003csup\u003e-1\u003c/sup\u003e; 3.8%), excessive (\u0026gt;\u0026thinsp;400 mg kg\u003csup\u003e-1\u003c/sup\u003e; 2.1%), and poor potassium levels (ranging from 66.1 to 132 mg kg\u003csup\u003e-1\u003c/sup\u003e; 0.2%), the species exhibited a reduced presence (Fig. \u003cspan class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e\n \u003c/div\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003eThe annual mean temperature (BIO1) ranged from 2.5 to 11.9\u0026deg;C, with an average of 8.78\u0026deg;C. The average annual precipitation was moderate, ranging from 426 to 748 mm, with an average of 588 mm.\u003c/p\u003e\n \u003c/div\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e\n \u003ch2\u003eThe relationship between floristic composition and environmental variables\u003c/h2\u003e\n \u003cp\u003eThe DCA data showed that Axis 1 was the most important. It explained the most variation and had the most extended gradient length (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e; Fig. \u003cspan class=\"InternalRef\"\u003e6\u003c/span\u003e). The other axes made a minor contribution.\u0026nbsp;\u003c/p\u003e\n \u003ctable id=\"Tab2\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eSummary of the DCA analysis.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAxis 1\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAxis 2\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAxis 3\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAxis 4\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eEigenvalues\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.6026\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.4672\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.3416\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.2854\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eExplained variation (cumulative)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.57\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.57\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e7.24\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eGradient length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.43\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3.99\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePseudo-canonical correlation (suppl.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.8254\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.2208\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.4329\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.3782\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003cp\u003e\u003c/p\u003e\n \u003cp\u003eCCA data showed that annual precipitation (BIO12) explained most of the variation in the floristic composition of \u003cem\u003eS. nutans\u003c/em\u003e communities, followed by slope (Table \u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e). \u0026nbsp;\u003c/p\u003e\n \u003ctable id=\"Tab3\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eResults of the CCA ordination.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eVariable\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eExplains %\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eContribution %\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003epseudo-F\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eP\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eP(adj)\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eAnnual Precipitation (BIO12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.86\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e53.09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSlope (\u0026deg;)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e9.57\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eElevation (m a.s.l.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eAspect\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePotassium (K)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.71\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003epH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.008\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.056\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePhosphorus (P)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.014\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cdiv class=\"BlockQuote\"\u003e\n \u003cp\u003eIn locations (50 locations in Tulcea and Constanța counties) with precipitation ranging from 400 to 450 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e has a low cover, ranging from 0.5\u0026ndash;5%. In these locations, the species occurs at low elevations (54 m a.s.l.) on gentle slopes (9\u0026deg;). The soils are neutral, with an average phosphorus and high potassium. The dominant associations were \u003cem\u003eAgropyro pontici-Thymetum zygioidis\u003c/em\u003e and \u003cem\u003eTaraxaco serotinae-Bothriochloetum ischaemi\u003c/em\u003e. In contrast, the temperature is higher (11.3 ℃) compared to the other precipitation intervals (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eIn the range of 451 and 500 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e had a similar cover, from 0.5\u0026ndash;5%, but was identified in fewer locations (38 locations in Tulcea, Constanța and Galați counties). Within these locations, \u003cem\u003eS. nutans\u003c/em\u003e was recorded at moderate elevations (127 m a.s.l.), on gentle slopes (5\u0026deg;) and at lower temperatures of 11.03 ℃. Competition with other species may be one reason why the cover with \u003cem\u003eS. nutans\u003c/em\u003e was lower. The soils are neutral, with medium phosphorus concentrations and very high potassium. The dominant association was \u003cem\u003eStipo ucrainicae-Festucetum valesiacae\u003c/em\u003e (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eIn the interval from 501 to 550 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e had the same cover, from 0.5\u0026ndash;5%. In these locations (19 locations in Vaslui, Iași, and Bacău counties), \u003cem\u003eS. nutans\u003c/em\u003e was observed at elevations higher than 158 m a.s.l., on moderate slopes (12.7\u0026deg;). The mean annual temperature was lower (9.7℃). The soils are moderately acidic, with a medium phosphorus concentration and very high potassium levels. The dominant associations were \u003cem\u003eAstero oleifolius-Ephedretum distachyae\u003c/em\u003e and \u003cem\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/em\u003e (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eIn 325 locations across Vaslui, Iași, Suceava, Covasna, and Alba counties, with annual precipitation ranging from 551 to 600 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e had a cover ranging from 0.05\u0026ndash;62.5%. The elevations were higher (362 m a.s.l.), and steep slopes (24\u0026deg;). The temperatures were lower (8.7\u0026deg;C). The soils were moderately acidic, with medium phosphorus and very high potassium concentrations. The dominant associations were \u003cem\u003eAllio albidi-Stipetum lessingianae\u003c/em\u003e, \u003cem\u003eDanthonio alpinae-Stipetum stenophyllae, Festuceto rupicolae-Caricetum humilis\u003c/em\u003e, and \u003cem\u003eFestucetum rupicolae\u003c/em\u003e (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eMost observations of the species were recorded in locations with annual precipitation from 601 to 650 mm/year. In these locations (412 locations in Sălaj, Bistrița, Cluj, and Mureș counties), \u003cem\u003eS. nutans\u003c/em\u003e had a cover from 0.05\u0026ndash;37.5%. Elevations were high (416 m a.s.l.), steep slopes (32\u0026deg;) and much lower temperatures (8.3\u0026deg;C). The soils were moderately acidic, with medium phosphorus and very high potassium concentrations (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eIn locations (20 locations in Cluj and Bistrița-Năsăud counties) with precipitation higher than 650 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e had a cover ranging from 0.05\u0026ndash;17.5%. In these locations, the highest elevations (604 m) and the lowest average annual temperatures (7.52\u0026deg;C) were recorded on steep slopes (21\u0026deg;). The soils are weakly acidic, with medium to high phosphorus concentrations (Fig. \u003cspan class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\n \u003c/div\u003e\n\u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eGeneral aspects\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is widely distributed in steppe grasslands across Ukraine, Russia, and other Eastern European regions (Budak \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Đakić, Knežević, and Boža \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The results indicate a high presence of the species in Transylvania and the plains of Eastern Romania, confirming its presence in the marginal areas of its European distribution (So\u0026oacute; \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1968\u003c/span\u003e; Csapody \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1982\u003c/span\u003e). In Romania (Oprea \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) and Hungary (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), the species was much more widespread in the past. Due to overgrazing and the transformation of grasslands into agricultural land, the habitat of the species is threatened in Hungary (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) and Romania.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003eSyntaxonomical scheme and EUNIS habitats\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThis study confirms that \u003cem\u003eS. nutans\u003c/em\u003e occurs predominantly in dry grasslands of the \u003cem\u003eFestuco\u003c/em\u003e-\u003cem\u003eBrometea\u003c/em\u003e class (827 relev\u0026eacute;s, representing 95% of the total), which is in agreement with the data in the literature (So\u0026oacute; \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1968\u003c/span\u003e; S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), where the species is described as a xerophytic species characteristic of open habitats. The high percentage of 95% compared to the presence of the species in other habitats shows a clear preference for this type of grassland, where the soils are well-drained.\u003c/p\u003e \u003cp\u003eIn the vegetation classes \u003cem\u003eCrataego-Prunetea\u003c/em\u003e (19 relev\u0026eacute;s, representing 2% of the total) and \u003cem\u003eTrifolio-Geranietea\u003c/em\u003e (13 relev\u0026eacute;s, representing 2% of the total), the presence of the species was reduced, indicating that \u003cem\u003eS. nutans\u003c/em\u003e also colonizes less typical habitats. In contrast, the presence of the species in the classes \u003cem\u003eDigitario sanguinalis-Eragrostietea minoris\u003c/em\u003e (one relev\u0026eacute;, representing 0.1% of the total), \u003cem\u003eFestuco-Puccinellietea\u003c/em\u003e (one relev\u0026eacute;, representing 0.1% of the total), \u003cem\u003eMolinio\u003c/em\u003e-\u003cem\u003eArrhenatheretea\u003c/em\u003e (four relev\u0026eacute;s, representing 0.5% of the total), \u003cem\u003eQuercetea pubescentis\u003c/em\u003e (two relev\u0026eacute;s, representing 0.2% of the total), and \u003cem\u003eSisymbrietea\u003c/em\u003e (three relev\u0026eacute;s, representing 0.3% of the total) was significantly reduced, indicating a marginal presence, not being characteristic for these communities. Some of these data are also confirmed by specialized literature (Horeanu \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Kov\u0026aacute;cs \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Germany \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), which shows that \u003cem\u003eS. nutans\u003c/em\u003e has been recorded in other vegetation classes as well.\u003c/p\u003e \u003cp\u003eAt the order level, the results indicated that \u003cem\u003eS. nutans\u003c/em\u003e was associated with \u003cem\u003eFestucetalia valesiacae\u003c/em\u003e (659 relev\u0026eacute;s, representing 77% of the total). The second most frequent order was \u003cem\u003eBrachypodietalia pinnati\u003c/em\u003e (127 relev\u0026eacute;s, representing 15% of the total). In the orders \u003cem\u003eStipo pulcherrimae-Festucetalia pallentis\u003c/em\u003e (24 relev\u0026eacute;s, representing 3% of the total), \u003cem\u003ePrunetalia spinosae\u003c/em\u003e (19 relev\u0026eacute;s, representing 2% of the total), \u003cem\u003eAntherico ramosi-Geranietalia sanguinei\u003c/em\u003e (13 relev\u0026eacute;s, representing 2% of the total), \u003cem\u003eS. nutans\u003c/em\u003e had a reduced presence. In the remaining orders (\u003cem\u003eArrhenatheretalia elatioris\u003c/em\u003e, \u003cem\u003eEragrostietalia\u003c/em\u003e, \u003cem\u003ePotentillo-Polygonetalia avicularis\u003c/em\u003e, \u003cem\u003ePuccinellietalia\u003c/em\u003e, \u003cem\u003eQuercetalia pubescenti-petraeae\u003c/em\u003e, and \u003cem\u003eSisymbrietalia sophiae\u003c/em\u003e), \u003cem\u003eS. nutans\u003c/em\u003e had a significantly reduced presence.\u003c/p\u003e \u003cp\u003eAt the alliance level, \u003cem\u003eS. nutans\u003c/em\u003e was recorded most frequently in \u003cem\u003eStipion lessingianae\u003c/em\u003e (468 relev\u0026eacute;s, representing 54% of the total), \u003cem\u003eFestucion valesiacae\u003c/em\u003e (191 relev\u0026eacute;s, representing 22% of the total), and \u003cem\u003eCirsio-Brachypodion pinnati\u003c/em\u003e (144 relev\u0026eacute;s, representing 17% of the total). In the alliances \u003cem\u003ePimpinello-Thymion zygoidis\u003c/em\u003e (24 relev\u0026eacute;s, representing 3% of the total) and \u003cem\u003eGeranion sanguinei\u003c/em\u003e (13 relev\u0026eacute;s, representing 2% of the total), \u003cem\u003eS. nutans\u003c/em\u003e had a low presence. In the remaining alliances (\u003cem\u003eArrhenatherion elatioris\u003c/em\u003e, \u003cem\u003eBerberidion vulgaris\u003c/em\u003e, \u003cem\u003eCynosurion cristati\u003c/em\u003e, \u003cem\u003eFestucion pseudovinae\u003c/em\u003e, \u003cem\u003ePotentillion anserinae\u003c/em\u003e, \u003cem\u003eQuercion petraeae\u003c/em\u003e, \u003cem\u003eSalsolion ruthenicae\u003c/em\u003e, and \u003cem\u003eSisymbrion officinalis\u003c/em\u003e), \u003cem\u003eS. nutans\u003c/em\u003e had a very low presence. According to the literature, \u003cem\u003eS. nutans\u003c/em\u003e is characteristic of \u003cem\u003eStipion lessingianae\u003c/em\u003e (Ruprecht et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; S\u0026acirc;rbu, Ștefan, and Oprea \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and \u003cem\u003eFestucion valesiacae\u003c/em\u003e alliances (Kov\u0026aacute;cs \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding plant associations, \u003cem\u003eS. nutans\u003c/em\u003e occurs most frequently in the associations \u003cem\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/em\u003e (159 relev\u0026eacute;s, representing 18% of the total) and \u003cem\u003eAllio albidi-Stipetum lessingianae\u003c/em\u003e (123 relev\u0026eacute;s, representing 18% of the total). Also, other associations in which \u003cem\u003eS. nutans\u003c/em\u003e occurs were \u003cem\u003eFestucetum rupicolae\u003c/em\u003e (81 relev\u0026eacute;s, representing 9% of the total), \u003cem\u003eSalvio-Festucetum rupicolae\u003c/em\u003e (54 relev\u0026eacute;s, 6%), \u003cem\u003eFestuceto rupicolae-Caricetum humilis\u003c/em\u003e (49 relev\u0026eacute;s, 6%), \u003cem\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/em\u003e relev\u0026eacute;s (49 relev\u0026eacute;s, 6%), \u003cem\u003eCariceto humilis-Brachypodietum pinnati\u003c/em\u003e (45 relev\u0026eacute;s, 5%). The presence of \u003cem\u003eS. nutans\u003c/em\u003e species in \u003cem\u003eSalvio-Festucetum rupicolae\u003c/em\u003e Zolyomi 1958 corr. So\u0026oacute; 1964, \u003cem\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/em\u003e and \u003cem\u003eFestuceto rupicolae-Caricetum humilis\u003c/em\u003e in Transylvania was reported by So\u0026oacute; (\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1942\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1968\u003c/span\u003e), Cśur\u0026ouml;s (1973, 1974) and Guszt\u0026aacute;v (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Also, Irimia and M\u0026acirc;nzu (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) mention the presence of the species in \u003cem\u003eTaraxaco serotinae-Festucetum valesiacae\u003c/em\u003e. At the same time, the presence of the species in \u003cem\u003eCrataego monogynae-Cerasetum mahaleb\u003c/em\u003e (P\u0026icirc;nzaru, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) shows the adaptability of the species in transitional areas between grasslands and shrub vegetation.\u003c/p\u003e \u003cp\u003eMoreover, \u003cem\u003eS. nutans\u003c/em\u003e has also been reported in associations such as \u003cem\u003eElytrigietum hispidi\u003c/em\u003e, \u003cem\u003eJurineo arachnoideae-Stipetum lessingianae\u003c/em\u003e, \u003cem\u003eStipo ucrainicae-Festucetum valesiacae\u003c/em\u003e, and \u003cem\u003eThymio pannonici-Chrysopogonetum grylli\u003c/em\u003e (Popescu et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). In addition, C\u003cem\u003eariceto humulis-Festucetum rupicolae\u003c/em\u003e and \u003cem\u003eBothriochloetum ischaemi\u003c/em\u003e (Kov\u0026aacute;cs \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) indicate a consistent presence in the \u003cem\u003eFestucion valesiacae\u003c/em\u003e alliance. The species is also found in \u003cem\u003eMedicagini minimae-Festucetum\u003c/em\u003e (Aniței and Mititelu 1997), an association specific to xerophilous mesophilous grasslands. Habitat degradation in the loess grasslands of the Crișana region is described in the literature (Guszt\u0026aacute;v \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) as leading to a simplification of diversity and the dominance of grasses, e.g. \u003cem\u003eFestuca valesiaca\u003c/em\u003e subsp. \u003cem\u003eparviflora\u003c/em\u003e (Hack.) Tracey.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003eThe relationship between floristic composition and environmental variables\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe most influential environmental factor explaining the variation in the floristic composition of \u003cem\u003eS. nutans\u003c/em\u003e phytocenoses was annual precipitation (BIO12). This finding is consistent with previous research demonstrating the influence of edaphic and climatic factors on the formation of grassland communities (Klimeš and Doležal \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2010\u003c/span\u003er\u0026ouml;k et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Bai et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The abundance and distribution of \u003cem\u003eS. nutans\u003c/em\u003e may be affected by changes in climate patterns, such as increased precipitation and prolonged droughts.\u003c/p\u003e \u003cp\u003eTopographic variables had a significant role in explaining the variation in floristic composition. Similar findings have been recorded in other grassland communities, where competition between species, environmental conditions, and soil water retention are influenced by topographic diversity (Moeslund et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The species' preference for very steep slopes in locations with precipitation from 551 to 650 mm/year may be related to the reduced competition with mesophytic species (Grime \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSoil chemical variables had a secondary role in explaining the variation in floristic composition. In general, soil pH and phosphorus availability varied slightly, while potassium concentrations were very high. This means that \u003cem\u003eS. nutans\u003c/em\u003e can tolerate a broader range of soil conditions, which is a characteristic of species adapted to semi-arid grasslands (Janišov\u0026aacute; et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Additionally, in locations with high precipitation, soil pH is typically lower, suggesting an interaction between soil acidity and plant communities (Chytr\u0026yacute; et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBioclimatic variables, primarily precipitation and topographic factors, played a significant role in the formation of \u003cem\u003eS. nutans\u003c/em\u003e communities. Considering the species' vulnerability to water availability, future climate variations may lead to changes in its abundance and distribution. In this context, in locations where the species is threatened by anthropogenic impact, it is important to develop conservation and monitoring measures.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec23\" class=\"Section3\"\u003e \u003ch2\u003eThe biogeographic, ecological, and evolutionary context of the species\u003c/h2\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is a member of the section \u003cem\u003ePlethiosphace\u003c/em\u003e Benth. in the subgenus \u003cem\u003eSclarea\u003c/em\u003e (Moench). Benth. The section has a Palearctic range and was considered monophyletic in a single molecular analysis addressing it (Will and Classen-Bockhoff \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which, however, involved only a few species, none of which were \u003cem\u003eSalvia nutans\u003c/em\u003e. According to the authors, the evolutionary relationships among the species in this section remain unknown. In Transylvania, Romania, the hybrid \u003cem\u003eS. nutans \u0026times; nemorosa\u003c/em\u003e is very frequently encountered wherever the two species come into contact. At the same time, \u003cem\u003eS. nemorosa\u003c/em\u003e seems to actively displace \u003cem\u003eS. nutans\u003c/em\u003e in many locations, including well-preserved steppe grasslands (personal observations). This is a serious problem for \u003cem\u003eS. nutans\u003c/em\u003e conservation, at least in Romania, but it also implies that the two species are closely related evolutionarily and ecologically. The other hybrid, known as \u003cem\u003eS. nutans\u003c/em\u003e, is \u003cem\u003eS.\u003c/em\u003e \u0026times; \u003cem\u003esimonkaiana\u003c/em\u003e Borb. (\u003cem\u003enutans\u003c/em\u003e \u0026times; \u003cem\u003epratensis\u003c/em\u003e) it is far rarer. No other hybrids are known between \u003cem\u003eS. nutans\u003c/em\u003e and any other species in the section \u003cem\u003ePlethiosphace\u003c/em\u003e that occur within its range.\u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is a West Palearctic steppe and forest-steppe element (Pobedinova \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1954\u003c/span\u003e; Ozenda \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Sramk\u0026oacute; et al. 2020) with a wide range extending continuously from the Lower Danube forest-steppe areas to the steppe and forest-steppe of the Ufa-Samara-Orenburg region immediately west of the Ural Mountains. Isolated populations are found in the insular forest-steppes of Thrace, the Pannonian Basin, and the Transylvanian Basin. While in the Transylvanian Basin the species is frequent in the mesoxeric and xeric grasslands (as it is presented here) from the forest-steppe and the nemoral adjacent area (here only on steep sunny slopes) in the Pannonian Basin it is scarce \u0026ndash; two locations only in Hungary (Sramk\u0026oacute; et al. 2020) while in Vojvodina, all the previously known five populations are extinct nowadays (Đakić, Knežević, and Boža \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Interestingly, two distinct sampled populations from the Transylvanian Basin exhibit different genetic affinities; one is more closely related to Hungarian populations west of the region, while the other is more closely related to populations north of the Black Sea (Sramk\u0026oacute; et al. 2020). This implies a biogeographically interesting heterogeneity for \u003cem\u003eSalvia nutans\u003c/em\u003e from Transylvania, with one possible explanation being that the species migrated into this region from two directions: from the west (Pannonia) and the east (Moldova), presumably in different and distant periods.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e was found in eight main phytocoenological classes, showing its versatility in various habitats. This species was most frequently recorded in dry grasslands, with the highest abundance reported in the associations \u003cem\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/em\u003e and \u003cem\u003eAllio albidi-Stipetum lessingianae\u003c/em\u003e. The species was recorded in areas with moderate elevation, on gentle to very steep slopes, being affected by overgrazing and the transformation of grasslands into arable land.\u003c/p\u003e \u003cp\u003eThe abundance of the species varies significantly in the six clusters, highlighting different environmental conditions that influence its distribution. Thus, the highest average cover of the species \u003cem\u003eS. nutans\u003c/em\u003e was recorded in cluster 4 (75%). A moderate abundance of the species was reported in cluster 5 (19%). In contrast, low abundance was recorded in clusters 1 and 2 (2%), 3, and 6 (1%).\u003c/p\u003e \u003cp\u003eCanonical Correspondence Analysis (CCA) revealed that the mean annual temperature (BIO1) accounted for the majority of the variation in floristic composition.\u003c/p\u003e \u003cp\u003eIn locations with precipitation ranging from 400 to 550 mm/year, \u003cem\u003eS. nutans\u003c/em\u003e had a low cover, ranging from 0.05\u0026ndash;0.5%. Thus, once the precipitation increases to 650 mm/year, the cover increases to 62.5%. At intervals higher than 650 mm/year, cover decreased. This suggests that the abundance of the species may be limited by excess moisture and lower temperatures.\u003c/p\u003e \u003cp\u003eFrom a biogeographical perspective, \u003cem\u003eS. nutans\u003c/em\u003e is characteristic of the West-Palaearctic steppe and forest-steppe. The species has a continuous range from the forest-steppe areas of the Lower Danube to the steppe and forest-steppe areas of the Ufa-Samara-Orenburg region. In the Transylvanian region, \u003cem\u003eS. nutans\u003c/em\u003e is frequent, while in the Pannonian Basin, it is very rare.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e "},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eORCID iDs\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSimona Dumitrița Chirilă https://orcid.org/0000-0003-3397-1834\u003c/p\u003e\n\u003cp\u003eAlexandru Sabin Bădărău https://orcid.org/0000-0001-5113-2802\u003c/p\u003e\n\u003cp\u003eMihai Doroftei \u0026nbsp;https://orcid.org/0000-0002-8388-087X\u003c/p\u003e\n\u003cp\u003eKiril Vassilev \u0026nbsp;https://orcid.org/0000-0003-4376-5575\u003c/p\u003e\n\u003cp\u003eIuliia Vasheniak\u003cimg src=\"data:image/png;base64,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\" alt=\"image\"\u003e\u0026nbsp;https://orcid.org/0000-0003-1020-3007\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSimona Dumitrița Chirilă\u003c/strong\u003e: Conceptualization, Methodology, Investigation, Software, Data curation, Writing- Original draft preparation. \u003cstrong\u003eAlexandru Sabin Bădărău\u003c/strong\u003e: Data curation, Visualization, Investigation, Writing- Original draft preparation, Supervision. \u003cstrong\u003eMihai Doroftei\u003c/strong\u003e: Data curation, Visualization, Investigation, Writing - review and editing, Supervision. \u003cstrong\u003eKiril Vassilev\u003c/strong\u003e: Data curation, Visualization, Writing - review and editing, Supervision; \u003cstrong\u003eIuliia Vasheniak\u003c/strong\u003e: Visualization, Software, Data curation, Writing- Original draft preparation, Supervision.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflicts of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNo funding was received for conducting this study.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data that support the findings of this study are included within this paper and its supplementary information. 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Naukova Dumka, Kyiv, p 191\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"anthropogenic impact, biogeography, dry grasslands, habitat requirement, Romanian flora, Salvia nutans, steppe relict","lastPublishedDoi":"10.21203/rs.3.rs-6556566/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6556566/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is a steppe relict species, characteristic of the West Palearctic steppe and forest-steppe. It is vulnerable to overgrazing and the conversion of grasslands into arable land. Due to these anthropogenic impacts, the species is endangered in some locations in Romania. In this context, the study aims to find the plant associations in which the species occurs and the environmental factors that explain the floristic composition.\u003c/p\u003e \u003cp\u003eFor the vegetation analysis, 870 relev\u0026eacute;s were used, and the data were analyzed in JUICE programme, applying the modified TWINSPAN algorithm for clustering. The floristic composition data and environmental variables were used for the multivariate analyses.\u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e was recorded most frequently in the \u003cem\u003eFestuco-Brometea\u003c/em\u003e class, the \u003cem\u003eFestucetalia valesiacae\u003c/em\u003e order, the \u003cem\u003eStipion lessingianae\u003c/em\u003e alliance, and the \u003cem\u003eJurineo transylvanicae-Stipetum pulcherrimae\u003c/em\u003e and \u003cem\u003eAllio albidi-Stipetum lessingianae\u003c/em\u003e associations. Canonical Correlation Analysis indicates that annual precipitation (BIO12) explained most of the variation in floristic composition. \u003cem\u003eS. nutans\u003c/em\u003e occurs predominantly on steep slopes with southwest aspects and at an average elevation of 359 m. The species occurs in moderately acidic to slightly alkaline soils, with moderate phosphorus and high potassium concentrations.\u003c/p\u003e \u003cp\u003e \u003cem\u003eSalvia nutans\u003c/em\u003e is related to topographic and edaphic conditions, showing its role as an indicator species for continental, xerophilic grasslands. Given its vulnerability to land-use changes, \u003cem\u003eS. nutans\u003c/em\u003e should be included in long-term monitoring programs to assess population dynamics.\u003c/p\u003e","manuscriptTitle":"Phytocoenological and Ecogeographical study of Salvia nutans in Romania","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-05-12 11:04:37","doi":"10.21203/rs.3.rs-6556566/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2025-09-16T10:10:16+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-05-26T07:26:24+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-05-07T10:27:01+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-01T01:47:01+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biologia","date":"2025-04-29T08:43:19+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f3b062c4-c8a4-4bbc-8550-caf4390684b2","owner":[],"postedDate":"May 12th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-04-27T16:06:17+00:00","versionOfRecord":{"articleIdentity":"rs-6556566","link":"https://doi.org/10.1007/s11756-026-02192-7","journal":{"identity":"biologia","isVorOnly":false,"title":"Biologia"},"publishedOn":"2026-04-24 15:58:28","publishedOnDateReadable":"April 24th, 2026"},"versionCreatedAt":"2025-05-12 11:04:37","video":"","vorDoi":"10.1007/s11756-026-02192-7","vorDoiUrl":"https://doi.org/10.1007/s11756-026-02192-7","workflowStages":[]},"version":"v1","identity":"rs-6556566","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6556566","identity":"rs-6556566","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}