Aims The class Crypsietea aculeatae comprises pioneer ephemeral dwarf-graminoid vegetation occurring in periodically flooded saline habitats. Although numerous regional studies have described this vegetation, a unified classification across Europe has been lacking. This study aims to establish a formalized, continent-wide classification at the association level, identify its biogeographical patterns, and assess relationships between species composition and environmental gradients.Location Europe (except Armenia, Azerbaijan, Georgia, and Russia).Methods We compiled a dataset of 2538 vegetation plots from the European Vegetation Archive (EVA), private databases, and literature. A hierarchical expert system was developed using formal definitions of syntaxa based on species cover. Nonmetric multidimensional scaling (NMDS) was applied to evaluate compositional differences and reveal biogeographical patterns. Distance-based redundancy analysis (db-RDA) and unweighted community means of Ellenberg-type indicator values (EIVs) were used to assess environmental drivers.Results We present a revised syntaxonomic framework for the class Crypsietea aculeatae and propose updates to the EuroVegChecklist. One alliance (Cypero-Spergularion marinae) and four associations (Crypsietum aculeatae, Crypsietum schoenoidis, Cyperetum pannonici, and Oxybasio chenopodioidis-Crypsietum aculeatae) were formally defined, with basic descriptions of their species composition, ecological characteristics, and geographical distribution. The main gradients in vegetation variability within the class Crypsietea aculeatae are driven by climatic gradient and edaphic factors, including soil moisture, soil acidity, salinity, and nutrient availability.Conclusions We established the first formal classification of annual vegetation in European saline seasonal wetlands. The syntaxonomic nomenclature was revised, and modifications to the EuroVegChecklist were proposed. The resulting classification system, accompanied by an expert system, enables consistent application in nature conservation and ecological research.

The current knowledge on vegetation of salt-affected habitats has been advanced, calling for a supra-regional assessment. We evaluate the common and distinct features of inland saline/alkaline landscapes of temperate Europe in terms of floristic composition, vegetation types, and abiotic conditions to find out what are the main drivers of their spatial variability and diversity. We delineate 13 subregions with a high occurrence of inland saline/alkaline habitats and by utilizing extensive field surveys in the past 20 years we analyze factors presumably affecting their variability: the size of the area, its proximity to the seacoast, and bioclimatic variables. We subjected them to descriptive statistics and ANOVA; principal components analysis was performed to reduce the number of dimensions for each dataset; correlation analysis was conducted to identify the statistical dependence between the diversity of subregions and observed factors. Despite the general uniformity typical for saline habitats, we observed that the subregions exhibit significant dissimilarity. Among the 107 autochtonous plant specialists, they have in common only one obligate and five facultative halophytes (Puccinellia distans agg.; Carex distans, Juncus gerardi, Lotus tenuis, Schoenoplectus lacustris subsp. glaucus and Trifolium fragiferum). The size of the subregion and its distance from the nearest seas did not affect the overall variability. Higher halophyte richness is driven by the broader range of abiotic and biotic prerequisites, especially the specific climate featuring summer evaporation causing various salinization levels in the soil, which is the most pronounced in the central subregions of the Pannonian Lowland. In its peripheries, the effect of specific conditions is lower, generating a reduced richness of halophytes, and in the subregions of the North German and Polish Plain and the Transylvanian Basin, the edaphic conditions (salt springs from salt deposits) take the main role, resulting also in a decreased halophyte richness and variability.

The first comprehensive phytosociological classification of all vegetation types in Europe (EuroVegChecklist; Applied Vegetation Science, 2016, 19, 3-264) contained brief descriptions of each type. However, these descriptions were not standardized and mentioned only the most distinct features of each vegetation type. The practical application of the vegetation classification system could be enhanced if users had the option to select sets of vegetation types based on various combinations of structural, ecological, and biogeographical attributes. Based on a literature review and expert knowledge, we created a new database that assigns standardized categorical attributes of 12 variables to each of the 1106 alliances dominated by vascular plants defined in EuroVegChecklist. These variables include dominant life form, phenological optimum, substrate moisture, substrate reaction, salinity, nutrient status, soil organic matter, vegetation region, elevational vegetation belt, azonality, successional status and naturalness. The new database has the potential to enhance the usefulness of phytosociological classification for researchers and practitioners and to help understand this classification to non-specialists.

AimIn salt-affected environments, salinity shapes ecosystem functions and species composition. Apart from salinity, however, we know little about how soil chemical factors affect plant species. We hypothesized that specific ions, most of which contribute to salinity, co-determine plant niche differentiation. We asked if the importance of ions differs for species with (halophytes) and without (associated species) physiological adaptations to saline soils.LocationCarpatho-Pannonian region (Central and Eastern Europe).Time period2005-2021.Major taxa studiedVascular plants.MethodsWe recorded species occurrences and collected soil samples in 433 plots in saline habitats. We measured pH, salinity (electrical conductivity), and concentrations of Ca2+, K+, Mg2+, Na+, SO42- Cl-, CO32- and mineral nitrogen (mN) and calculated the sodium adsorption ratio (SAR). For 88 species, we fitted response curves with Huisman-Olff-Fresco (HOF) models. To study ions' effects on species composition and ions' variance, we compared unconstrained and constrained ordinations and performed a principal component analysis. We used random forests to analyse the importance of ions for individual species and created two-dimensional species niche plots for key ions.ResultsIon concentration niches varied among species and did not necessarily correspond to soil salinity or alkalinity. We frequently observed monotonic, sigmoidal model responses, while skewed unimodal responses were rare. Ions explained a considerable proportion of species compositional variation. Particularly, Na+, SO42-, Cl-, and CO32- contributed to the ions' variance. Na+, followed by SO42-, Cl-, CO32-, Ca2+, Mg2+, and mN, was most important for the occurrence of individual species. Compared to associated species, Na+, SO42-, and mN were significantly less important for halophytes, whereas Cl- and CO32- played a significant role.Main conclusionsWe show that ion composition co-determines niche differentiation in saline soils, suggesting evolved physiological adaptations in halophytes. Our study calls for incorporating high-resolution data on soil ion composition in ecological research.

Numerous features of the great diversity of saline environments in Romania (Southeastern Europe) have been investigated, although less explored regions still require field research. Tectonic movements and abundant fossil salt deposits have led to various isolated salt fountains, salty rocks and hypersaline ponds in the contact zone between the Eastern Carpathians and the Transylvanian Basin. According to the European Habitat Directive, these natural salt bodies belong to a priority habitat. In the counties of Harghita and Braşov, salt springs have been used as cold-water spas for decades, often without regard for their fragility. Their unique flora and vegetation remain largely unexplored. Based on numerical analysis of relevés collected from twelve salt spring locations in eastern Transylvania, we provide a comprehensive survey of saline vegetation. Compared to the large saline areas in western Transylvania, these are smaller, and only three euhalophytic plant associations make up the following vegetation zonation: Scorzonero parviflorae-Juncetum gerardi , Puccinellietum limosae , and Salicornietum prostratae . Their physiognomy and species composition resemble the well-studied travertine fens of the Western Carpathians (Slovakia). Joint analysis of data from rich fens (association Glauco-Trichophoretum pumili ) in the Spiš basin revealed the insular character of sub-mountain halophilic vegetation in both regions. The construction of new tourist resorts poses a threat to the mineral salt springs. Fragmentation, waste depositing and water withdrawal for swimming pools have destroyed many sites. Our results can help to estimate the recent area and conservatuion status of temperate inland salt marshes in Romania.

Understanding large-scale drivers of biodiversity in palustrine wetlands is challenging due to the combined effects of macroclimate and local edaphic conditions. In boreal and temperate fen ecosystems, the influence of macroclimate on biodiversity is modulated by hydrological settings across habitats, making it difficult to assess their vulnerability to climate change. Here, we investigate the influence of macroclimate and edaphic factors on three Essential Biodiversity Variables across eight ecologically defined habitats that align with ecosystem classifications and red lists. We used 27,555 vegetation plot samples from European fens to assess the influence of macroclimate and groundwater pH predictors on the geographic distribution of each habitat type. Additionally, we modeled the relative influence of macroclimate, water pH, and water table depth on community species richness and composition, focusing on 309 plant specialists. Our models reveal strong effects of mean annual temperature, diurnal thermal range, and summer temperature on biodiversity variables, with contrasting differences among habitats. While macroclimatic factors primarily shape geographic distributions and species richness, edaphic factors emerge as the primary drivers of composition for vascular plants and bryophytes. Annual precipitation exhibits non-linear effects on fen biodiversity, with varying impact across habitats with different hydrological characteristics, suggesting a minimum requirement of 600 mm of annual precipitation for the occurrence of fen ecosystems. Our results anticipate potential impacts of climate warming on European fens, with predictable changes among habitat types and geographic regions. Moreover, we provide evidence that the drivers of biodiversity in boreal and temperate fens are closely tied to the ecological characteristics of each habitat type and the dispersal abilities of bryophytes and vascular plants. Given that the influence of macroclimate and edaphic factors on fen ecosystems is habitat specific, climate change research and conservation actions should consider ecological differentiation within functional IUCN ecosystems at continental and regional scales.

There is a long tradition in Europe of assigning ecological indicator values to plants and using these values in ecological research. A special case is that of the salt-tolerant species. Saline soils are extremely heterogeneous and their physical and chemical properties vary significantly with microrelief and between alternating dry/wet seasons. The complexity of such soils suggests using salt indicator values. This study resulted in the first multi-country database of vascular plants occurring in inland salt marshes and on salt steppes in temperate Europe based on expert revision of the literature and field experience. The inventory of the 190 salt-tolerant species was carried out according to their quantitative representation in saline and non-saline habitats. These species were each classified into one of three categories of salt tolerance (obligate halophytes, facultative halophytes, accessory/associated species) assigned salt numbers on a nine-point scale reflecting their individual preference for soil salinity based on their calculated halophytic value. Saline soils are reliably indicated by the presence of obligate halophytes; these specialists grow exclusively in natural saline habitats. Only 45 species are assigned to this group, while 61 species make up the group of facultative halophytes and 84 are accessory species with a wide ecological niche, occurring more or less accidentally in saline habitats. Their number is likely to increase since every plant (non-halophyte) recorded in a saline habitat can be considered to be an accessory species. The obtained salt numbers showed a close consistency with the recently used salinity indicator values estimated by Ellenberg, Borhidi and Breckle; in terms of categories of salt tolerance, only slight differences were detected.

The class Montio-Cardaminetea includes vegetation of springs with constant water flow. These habitats, which function as islands for highly specialized and sensitive biota, are endangered by ongoing landscape and climatic changes. Although a harmonized classification into vegetation units is necessary for effective habitat conservation, there is currently no synthetic classification of the class Montio-Cardaminetea. Here a large set of vegetation-plot records was obtained from national and private databases. The aim was to validate the EuroVegChecklist classification scheme, search for additional ecologically meaningful vegetation types and develop an automatic system for classifying new plots from Europe. We formally defined the cores of eight of the ten EuroVegChecklist alliances: Caricion remotae, Cratoneurion commutati, Lycopodo europaei-Cratoneurion commutati, Epilobio nutantis-Montion, KoenigioMicrojuncion, Mniobryo-Epilobion hornemanii, Philonotidion seriatae (Cardamino-Montion) and Swertio perennis-Anisothecion squarrosi, which were also reproduced by unsupervised classifications. Both unsupervised and semi-supervised classifications further suggested two alliances not previously recognized in the EuroVegChecklist: Anthelion julaceae (liverwort dominated subalpine to alpine springs in highly oceanic regions in Britain) and Cratoneuro filicini-Calthion laetae (mesotrophic herb-rich subalpine and alpine springs). The unsupervised classifications mainly reflected the base saturation gradient, distinguishing between calcareous and non-calcareous springs. Therefore, it is suggested the order Montio-Cardaminetalia, which is broadly delimited in the EuroVegChecklist, be divided into two separate orders and the following three orders should be distinguished within the class Montio-Cardaminetea: CardaminoChrysosplenietalia (non-calcareous forest springs; Caricion remotae), Cardamino-Cratoneuretalia (calcareous springs; Cratoneurion commutati, Lycopodo europaei-Cratoneurion commutati) and Montio-Cardaminetalia (non-calcareous non-forest springs; all other alliances).

Aims: Ellenberg-type indicator values are expert-based rankings of plant species according to their ecological optima on main environmental gradients. Here we extend the indicator-value system proposed by Heinz Ellenberg and co-authors for Central Europe by incorporating other systems of Ellenberg-type indicator values (i.e., those using scales compatible with Ellenberg values) developed for other European regions. Our aim is to create a harmonized data set of Ellenberg-type indicator values applicable at the European scale.Methods: We collected European data sets of indicator values for vascular plants and selected 13 data sets that used the nine-, ten- or twelve-degree scales defined by Ellenberg for light, temperature, moisture, reaction, nutrients and salinity. We compared these values with the original Ellenberg values and used those that showed consistent trends in regression slope and coefficient of determination. We calculated the average value for each combination of species and indicator values from these data sets. Based on species' co-occurrences in European vegetation plots, we also calculated new values for species that were not assigned an indicator value.Results: We provide a new data set of Ellenberg-type indicator values for 8908 European vascular plant species (8168 for light, 7400 for temperature, 8030 for moisture, 7282 for reaction, 7193 for nutrients, and 7507 for salinity), of which 398 species have been newly assigned to at least one indicator value.Conclusions: The newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of the European flora and vegetation or for gap-filling in regional data sets. The European indicator values and the original and taxonomically harmonized regional data sets of Ellenberg-type indicator values are available in the Supporting Information and the Zenodo repository.

Monitoring the occurrence of new alien species among native flora is the first prerequisite for preventing their later invasive spread.During the last decade, we recorded data regarding 25 alien taxa new to Slovakia including 21 casual and 4 locally naturalised species, with ornamental plants predominating significantly.The locally naturalised species Cenchrus flaccidus, Cotoneaster zabelii, Doronicum pardalianches and Panicum virgatum are able to spread to native habitats and thus represent the greatest risk to native plant species in Slovakia.

While the importance of conservation mowing for mesic grasslands is generally accepted, its use for fens and fen grasslands interspersed within agricultural land is still controversial. Although fens may persist naturally, ongoing environmental changes increase productivity and accelerate succession. These processes can be mitigated through conservation management with appropriate settings. However, long-term management experiments are challenging and provide only locally valid results. Here, we analysed vegetation data (bryophytes and vascular plants) from seven management experiments (spanning 3-20 years) conducted in Central European poor, moderately-rich, and calcareous spring fens (Czech Republic, Slovakia). Two of these experiments examined the effects of restoration of abandoned fens, while five experiments examined changes in mowing regimes in managed fens (cessation, intensification, delay to autumn, and litter removal). Data were analysed using unidimensional and multidimensional methods separately for the initial, extended, and entire period. Mowing had a statistically significant effect on species composition except for the shortest (3-year) experiment. Litter removal did not compensate for mowing. Mowing twice or delayed mowing significantly affected the species composition of calcareous fens. In all cases, cessation of mowing significantly reduced the richness of species, especially those of conservation importance. In contrast, any mowing of abandoned fens increased species richness. The effects of mowing intensification or cessation on species richness and composition of a restored calcareous fen were evident in the first 2-3 years. Other effects were initially weak or nonsignificant but later became stronger, such as mowing delay and restoration removal of litter, which became significant only after nearly 20 years. We found that cessation or restoration of mowing usually triggers a rapid vegetation change, whereas it can take decades to detect the response caused by changes in mowing timing. Importantly, mowing can stabilise or even restore vegetation of fen ecosystems that have been weakened by their fragmentation in the temperate agricultural landscapes.

This is a database of the occurrence of 1105 phytosociological alliances in 82 European territorial units and distribution maps generated from this database. The data correspond to those published by Preislerová et al. (2022). This article also contains a description of the methods.

Aims Classification of European bog vegetation (Oxycocco-Sphagnetea class); identification of diagnostic species for the class and vegetation subgroups (orders and alliances); development of an expert system for automatic classification of vegetation plots; and production of distribution maps of the Oxycocco-Sphagnetea class and its alliances. Location Europe. Methods A data set of vegetation-plot records was compiled to include various bog types over most of the European continent. An unsupervised classification (beta-flexible linkage method, Sorensen distance measure) and detrended correspondence analysis (DCA) ordination were applied. Formal definitions of syntaxa based on species presence and covers, and respecting the results of the unsupervised classification, were developed and included in a classification expert system. Results The Oxycocco-Sphagnetea class, its two orders (Sphagno-Ericetalia tetralicis and Sphagnetalia medii) and seven compositionally distinct alliances were formally defined. In addition to the syntaxa included in EuroVegChecklist, three new alliances were distinguished: Rubo chamaemori-Dicranion elongati (subarctic polygon and palsa mires); Erico mackaianae-Sphagnion papillosi (blanket bogs of the northwestern Iberian Peninsula); and Sphagno baltici-Trichophorion cespitosi (boreal bog lawns). The latter alliance is newly described in this article. Conclusions This first pan-European formalized classification of European bog vegetation partially followed the system presented in EuroVegChecklist, but suggested three additional alliances. One covers palsa and polygon mires, one covers Iberian bogs with endemics and one fills the syntaxonomical gap for lawn microhabitats in boreal bogs. A classification expert system has been developed, which allows assignment of vegetation plots to the types described.

While the importance of conservation mowing for mesic grasslands is generally accepted, its use for fens and fen grasslands interspersed within agricultural land is still controversial. Although fens may persist naturally, ongoing environmental changes increase productivity and accelerate succession. These processes can be mitigated through conservation management with appropriate settings. However, long-term management experiments are challenging and provide only locally valid results. Here, we analysed vegetation data (bryophytes and vascular plants) from seven management experiments (spanning 3-20 years) conducted in Central European poor, moderately-rich, and calcareous spring fens. Two of these experiments examined the effects of restoration of abandoned fens, while five experiments examined changes in mowing regimes in managed fens (cessation, intensification, delay to autumn, and litter removal). Data were analysed using unidimensional and multidimensional methods separately for the initial, extended, and entire period. Mowing had a statistically significant effect on species composition except for the shortest (3-year) experiment. Litter removal did not compensate for mowing. Mowing twice or delayed mowing significantly affected the species composition of calcareous fens. In all cases, cessation of mowing significantly reduced the richness of species, especially those of conservation importance. In contrast, any mowing of abandoned fens increased species richness. The effects of mowing intensification or cessation on species richness and composition of restored calcareous fen were evident in the first 2-3 years. Other effects were initially weak or nonsignificant but later became stronger, such as mowing delay and restoration removal of litter, which became significant only after nearly 20 years. We found that cessation or restoration of mowing usually triggers a rapid vegetation change, whereas it can take decades to detect the response caused by changes in mowing timing. Importantly, mowing can stabilise or even restore vegetation of fen ecosystems that have been weakened by their fragmentation in the temperate agricultural landscapes.

Classification of European bog vegetation ( class); identification of diagnostic species for the class and vegetation subgroups (orders and alliances); development of an expert system for automatic classification of vegetation plots; and production of distribution maps of the class and its alliances. Europe. A data set of vegetation-plot records was compiled to include various bog types over most of the European continent. An unsupervised classification (beta-flexible linkage method, Sørensen distance measure) and detrended correspondence analysis (DCA) ordination were applied. Formal definitions of syntaxa based on species presence and covers, and respecting the results of the unsupervised classification, were developed and included in a classification expert system. The class, its two orders ( - and ) and seven compositionally distinct alliances were formally defined. In addition to the syntaxa included in EuroVegChecklist, three new alliances were distinguished: - (subarctic polygon and palsa mires); - (blanket bogs of the northwestern Iberian Peninsula); and - (boreal bog lawns). The latter alliance is newly described in this article. This first pan-European formalized classification of European bog vegetation partially followed the system presented in EuroVegChecklist, but suggested three additional alliances. One covers palsa and polygon mires, one covers Iberian bogs with endemics and one fills the syntaxonomical gap for lawn microhabitats in boreal bogs. A classification expert system has been developed, which allows assignment of vegetation plots to the types described.

Temperate inland salt marshes, formerly used as meadows or pastures, are priority habitats in Europe and are threatened by intensifying anthropogenic activities. They are particularly important because of their biogeographical location on the North German Plain, which is the westernmost hotspot of continental halophytic vegetation in Eurasia. In spite of their remarkably long history of floristic research, there are disproportionately few studies dealing with plant communities. They are traditionally included in the Puccinellion maritimae and Armerion maritimae alliances with a typical distribution along the North and Baltic Sea coasts. The similarity with other inland salt marshes has been overlooked. We surveyed natural and secondary sites with euhalophytic vegetation in central Germany (Saxony-Anhalt and Thuringia) in 2018 and 2020 and analysed 105 phytosociological relevés. This resulted in the identification of three main groups (alliances) containing seven associations: annual hypersaline succulent communities of the alliance Therosalicornion (Salicornietum europaeae, Halimioni pedunculatae-Puccinellietum distantis, Suaedetum maritimae), wet saline meadows of the alliance Juncion gerardi (Triglochino maritimae-Glaucetum maritimae, Scorzonero parviflorae-Juncetum gerardi variant with Glaux maritima) and saline short-grass swards of the alliance Puccinellion limosae (Puccinellietum limosae, Atriplex prostrata community). For each association we defined the typical species composition and calculated Ellenberg indicator values depending on the ecological gradients of soil salinity, nutrients and moisture. To determine possible floristic similarities between the inland and coastal salt marshes, we compared our data with data from other areas of saline vegetation: the southern Baltic Sea coast, Kujawy (Poland), Pannonian Basin (Hungary) and Transylvanian Basin (Romania). We concluded that the salt marshes in inland Germany showed a stronger relation with salt marshes of the remote Pannonian and Transylvanian Basins, and they were well separated from the Baltic salt marshes. Thus, we suggest including inland salt marshes on the North German Plain into the syntaxonomical class of continental halophytic vegetation of Festuco-Puccinellietea.

Rising temperatures may endanger fragile ecosystems because their character and key species show different habitat affinities under different climates. This assumption has only been tested in limited geographical scales. In fens, one of the most endangered ecosystems in Europe, broader pH niches have been reported from cold areas and are expected for colder past periods. We used the largest European‐scale vegetation database from fens to test the hypothesis that pH interacts with macroclimate temperature in forming realized niches of fen moss and vascular plant species. We calibrated the data set (29,885 plots after heterogeneity‐constrained resampling) with temperature, using two macroclimate variables, and with the adjusted pH, a variable combining pH and calcium richness. We modelled temperature, pH and water level niches for one hundred species best characterizing European fens using generalized additive models and tested the interaction between pH and temperature. Fifty‐five fen species showed a statistically significant interaction between pH and temperature (adj p ˂ .01). Forty‐six of them (84%) showed a positive interaction manifested by a shift or restriction of their niche to higher pH in warmer locations. Nine vascular plants and no moss showed the opposite interaction. Mosses showed significantly greater interaction. We conclude that climate significantly modulates edaphic niches of fen plants, especially bryophytes. This result explains previously reported regional changes in realized pH niches, a current habitat‐dependent decline of endangered taxa, and distribution changes in the past. A warmer climate makes growing seasons longer and warmer, increases productivity, and may lower the water level. These effects prolong the duration and intensity of interspecific competition, support highly competitive Sphagnum mosses, and, as such, force niches of specialized fen species towards narrower high‐pH ranges. Recent anthropogenic landscape changes pose a severe threat to many fen species and call for mitigation measures to lower competition pressure in their refugia.

Halophytic vegetation on fossil salt deposits and salt springs is an extreme habitat occurring in isolated continental areas of temperate Europe. It has a relatively rich representation in the Transylvanian Basin of Romania; however, comprehensive research into this highly specialized vegetation is lacking. We provide the first phytosociological survey of recent data of inland salt habitats in Cluj County, where they are widely distributed. Cluster analysis distinguished eight euhalophytic plant communities occupying several zones depending on the micro-topography of the salt-affected area: Scorzonero parviflorae-Juncetum gerardii (including Astero tripolio-Triglochinetum maritime ), Plantagineto cornuti-Agrostetum stoloniferae , Artemisio-Festucetum pseudovinae , Artemisio-Petrosimonietum triandrae , Limonio gmelinii-Artemisietum monogynae , Puccinellietum limosae (each allocated to the class Festuco-Puccinellietea ), then Suaedetum maritimae and Salicornietum prostratae , belonging to the class Therosalicornietea . We revised the nomenclature of these associations, characterized their species composition and ecological preferences based on indicator values for salinity, pH, moisture and nutrients in the soil. We broadly discussed our findings in light of the historical data and documented the current habitat conditions of the sites. Most of them were found to be degraded due to increasing tourism exploiting the curative effects of salt springs. Beyond recognizing the high conservation value of the Transylvanian salt habitats, it is also important to highlight their biogeographical significance, as they fill the gap between the inland salt marshes of Central Europe and the continental salt habitats of Eurasia.