Aims Phytosociological classification of fen vegetation (Scheuchzerio palustris‐Caricetea fuscae class) differs among European countries. Here we propose a unified vegetation classification of European fens at the alliance level, provide unequivocal assignment rules for individual vegetation plots, identify diagnostic species of fen alliances, and map their distribution. Location Europe, western Siberia and SE Greenland. Methods 29 049 vegetation‐plot records of fens were selected from databases using a list of specialist fen species. Formal definitions of alliances were created using the presence, absence and abundance of Cocktail‐based species groups and indicator species. DCA visualized the similarities among the alliances in an ordination space. The ISOPAM classification algorithm was applied to regional subsets with homogeneous plot size to check whether the classification based on formal definitions matches the results of unsupervised classifications. Results The following alliances were defined: Caricion viridulo‐trinervis (sub‐halophytic Atlantic dune‐slack fens), Caricion davallianae (temperate calcareous fens), Caricion atrofusco‐saxatilis (arcto‐alpine calcareous fens), Stygio‐Caricion limosae (boreal topogenic brown‐moss fens), Sphagno warnstorfii‐Tomentypnion nitentis (Sphagnum‐brown‐moss rich fens), Saxifrago‐Tomentypnion (continental to boreo‐continental nitrogen‐limited brown‐moss rich fens), Narthecion scardici (alpine fens with Balkan endemics), Caricion stantis (arctic brown‐moss rich fens), Anagallido tenellae‐Juncion bulbosi (Ibero‐Atlantic moderately rich fens), Drepanocladion exannulati (arcto‐boreal‐alpine non‐calcareous fens), Caricion fuscae (temperate moderately rich fens), Sphagno‐Caricion canescentis (poor fens) and Scheuchzerion palustris (dystrophic hollows). The main variation in the species composition of European fens reflected site chemistry (pH, mineral richness) and sorted the plots from calcareous and extremely rich fens, through rich and moderately rich fens, to poor fens and dystrophic hollows. ISOPAM classified regional subsets according to this gradient, supporting the ecological meaningfulness of this classification concept on both the regional and continental scale. Geographic/macroclimatic variation was reflected in the second most important gradient. Conclusions The pan‐European classification of fen vegetation was proposed and supported by the data for the first time. Formal definitions developed here allow consistent and unequivocal assignment of individual vegetation plots to fen alliances at the continental scale.

The pollen and seeds of 3 paleoendemic resurrection species, Ramonda myconi (L.) Rchb. (Iberian Peninsula), R. nathaliae Pančić & Petrović, and R. serbica Pančić (Balkan Peninsula), as well as of natural hybrids between the 2 last species, have been analyzed using light and scanning electron microscopy. Their general structural differences, taxonomic and phylogenetic significance, pollen viability, and seed germination capacity, as well as the correlation of pollen and seed characteristics and plant ploidy levels, have been studied. The pollen grains are small (R. myconi, R. nathaliae) to medium (R. serbica) in size, ranging from 10 to 28 µm, and 3-colporate, isopolar monads with microreticulate-perforate exine. Seeds are small, 309 to 1000 µm long and 80 to 425 µm wide, elongated, with a reticulate surface and auriculate ornamentation. Micromorphologies of the pollen exine ornamentation and seed surface revealed significant differences among the 3 species. Pollen from hybrid individuals was heterogeneous in size and morphology, and the germination of their tiny seeds was very low. A strong correlation was found between pollen size, DNA content, and chromosome number. The environmental influence on pollen and seeds of 3 species and especially of the R. nathaliae population growing on serpentine is also discussed.

Abstract. Water resources and associated ecosystems are becoming highly endangered due to ongoing global environmental changes. Spatial ecological modelling is a promising toolbox for understanding the past, present and future distribution and diversity patterns in groundwater-dependent ecosystems, such as fens, springs, streams, reed beds or wet grasslands. Still, the lack of detailed water chemistry maps prevents the use of reasonable models to be applied on continental and global scales. Being major determinants of biological composition and diversity of groundwater-dependent ecosystems, groundwater pH and calcium are of utmost importance. Here we developed an up-to-date European map of groundwater pH and Ca, based on 7577 measurements of near-surface groundwater pH and calcium distributed across Europe. In comparison to the existing European groundwater maps, we included several times more sites, especially in the regions rich in spring and fen habitats, and filled the apparent gaps in eastern and southeastern Europe. We used random forest models and regression kriging to create continuous maps of water pH and calcium at the continental scale, which is freely available also as a raster map (Hájek et al., 2020b; https://doi.org/10.5281/zenodo.4139912). Lithology had a higher importance than climate for both pH and calcium. The previously recognised latitudinal and altitudinal gradients were rediscovered with much refined regional patterns, as associated with bedrock variation. For ecological models of distribution and diversity of many terrestrial ecosystems, our new map based on field groundwater measurements is more suitable than maps of soil pH, which mirror not only bedrock chemistry but also vegetation-dependent soil processes.

The northern temperate genus Dracocephalum consists of approximately 70 species mainly distributed in the steppe-desert biomes of Central and West Asia and the alpine region of the Qinghai-Tibetan Plateau (QTP). Previous work has shown that Dracocephalum is not monophyletic and might include Hyssopus and Lallemantia. This study attempts to clarify the phylogenetic relationships, diversification patterns, and the biogeographical history of the three genera (defined as Dracocephalum s.l.). Based on a sampling of 66 taxa comprising more than 80% from extant species of Dracocephalum s.l., morphological, phylogenetic (maximum parsimony, likelihood, and Bayesian inference based on nuclear ITS and ETS, plastid rpl32-trnL, trnL-trnF, ycf1, and ycf1-rps15, and two low-copy nuclear markers AT3G09060 and AT1G09680), molecular dating, diversification, and ancestral range estimation analyses were carried out. Our results demonstrate that both Hyssopus and Lallemantia are embedded within Dracocephalum and nine well-supported clades can be recognized within Dracocephalum s.l. Analyses of divergence times suggest that the genus experienced an early rapid radiation during the middle to late Miocene with major lineages diversifying within a relatively narrow timescale. Ancestral area reconstruction analyses indicate that Dracocephalum s.l. originated in Central and West Asia and southern Siberia, and dispersed from Central and West Asia into the QTP and adjacent areas twice independently during the Pliocene. The aridification of the Asian interior possibly promoted the rapid radiation of Dracocephalum within this region, and the uplift of the QTP appears to have triggered the dispersal and recent rapid diversification of the genus in the QTP and adjacent regions. Combining molecular phylogenetic and morphological evidence, a revised infrageneric classification of Dracocephalum s.l. is proposed, which recognizes nine sections within the genus.

Abstract. Water resources and associated ecosystems are becoming highly endangered due to ongoing global environmental changes. Spatial ecological modelling is a widely used tool for understanding the past, present and future distribution and diversity patterns in groundwater-dependent ecosystems, such as fens, springs, streams, reed beds or wet grasslands. Still, the lack of detailed water chemistry maps prevents their reasonable use on continental and global scales. Being major determinants of biological composition and diversity of groundwater-dependent ecosystems, groundwater pH and calcium are of utmost importance. Here we developed the up-to-date European map of groundwater pH and Ca, based on 7,577 measurements of near-surface groundwater pH and calcium distributed across Europe. In comparison to the existing European groundwater maps, we included a several times larger number of sites, especially in the regions rich in spring and fen habitats, and filled the apparent gaps in Eastern and Southeastern Europe. We used Random Forest models and regression kriging to create continuous maps of water pH and calcium at the continental scale, which is freely available also as a raster map (Hájek et al. 2020; https://doi.org/10.5281/zenodo.4139912). Lithology had higher importance than climate for both pH and calcium. The previously recognised latitudinal and altitudinal gradients were rediscovered with much refined regional patterns, as associated with bedrock variation. For ecological models of distribution and diversity of groundwater-dependent, but also other terrestrial, ecosystems, the new map is more suitable than previously used maps of soil pH, unlike which it mirrors bedrock chemistry more than vegetation-dependent soil processes.