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Aim: Our knowledge of Pleistocene refugia and post‐glacial recolonization routes of forest understorey plants is still very limited. The geographical ranges of these species are often rather narrow and show highly idiosyncratic, often fragmented patterns indicating either narrow and species‐specific ecological tolerances or strong dispersal limitations. However, the relative roles of these factors are inherently difficult to disentangle. Location: Central and south‐eastern Europe. Time period: 17,100 BP – present. Major taxa studied: Five understorey herbs of European beech forests: Aposeris foetida, Cardamine trifolia, Euphorbia carniolica, Hacquetia epipactis and Helleborus niger. Methods: We used spatio‐temporally explicit modelling to reconstruct the post‐glacial range dynamics of the five forest understorey herbs. We varied niche requirements, demographic rates and dispersal abilities across plausible ranges and simulated the spread of species from potential Pleistocene refugia identified by phylogeographical analyses. Then we identified the parameter settings allowing for the most accurate reconstruction of their current geographical ranges. Results: We found a largely homogenous pattern of optimal parameter settings among species. Broad ecological niches had to be combined with very low but non‐zero rates of long‐distance dispersal via chance events and low rates of seed dispersal over moderate distances by standard dispersal vectors. However, long‐distance dispersal events, although rare, led to high variation among replicated simulation runs. Main conclusions: Small and fragmented ranges of many forest understorey species are best explained by a combination of broad ecological niches and rare medium‐ and long‐distance dispersal events. Stochasticity is thus an important determinant of current species ranges, explaining the idiosyncratic distribution patterns of the study species despite strong similarities in refugia, ecological tolerances and dispersal abilities. [ABSTRACT FROM AUTHOR]

The occurrence and origin of dry grasslands and their rich biota in the moderately humid Central‐European climate have fascinated scientists for over a century. Modern palaeoecological and phylogeographical data support earlier hypotheses that these grasslands are late Pleistocene relicts and can therefore be considered part of the Eurasian forest–steppe biome. However, it is still unclear which factors fostered the maintenance of steppe patches in Central Europe throughout the Holocene. Here, we provide an overview of the main hypotheses, which stress, respectively, the effects of climate, edaphic conditions and disturbances. We then develop a general conceptual framework on how these three factors interact to form forest–steppe mosaics. We thereby emphasize the role of topography as a crucial control on forest–steppe patterns at the landscape scale. Topography is related to several mechanistic drivers that influence vegetation processes, such as near‐surface microclimate and soil formation. Consequently, topographic variation allows both forest and steppe patches to occur beyond their macroclimatic niche, favouring the development of forest–steppe mosaics. To illustrate our framework, we demonstrate the interactive effect of macroclimate and topography on the occurrence of steppe patches at 108 selected Central European forest–steppe sites. Although we developed our framework focusing on the current distribution of Central European forest–steppe, we suggest that it contributes to the understanding of similar transitions between temperate forest and steppe biomes in the past as well as elsewhere in the world. [ABSTRACT FROM AUTHOR]

Aim: The steppe grasslands of eastern Central Europe are exceptionally species rich and valuable from a nature conservation point of view. However, their historical biogeography is still poorly understood. Here we use the regional diversity of habitat specialists and chloroplast DNA data to investigate potential long-term refugia of steppe species in this region. Location: Pannonian Basin and adjacent regions; SW Russia. Taxon: Vascular plants. Methods: After identifying habitat specialists of the three main steppe types (meadow steppes, grass steppes and rocky steppes), we compiled their regional presence--absence in grid cells of 75 km x 75 km. We analysed the dependency of habitat specialist diversity to climate, topographic heterogeneity and geographical distance to potential refugia. For genetic analysis, we sampled three or four habitat specialists of each steppe type and used cpDNA markers to investigate intraspecific diversity and geographical distribution of haplotypes. We also tested for correspondence between the number of habitat specialists and haplotype diversity. Results: Climate and topography explained between 40% and 63% of the variance in habitat specialist diversity. Adding geographical distance to potential refugia increased the explained variance in the models for all steppe types. Chloroplast haplotypes featured a complex pattern across the study area. Several species showed a strong geographical differentiation, suggesting migration waves from multiple refugia with only limited subsequent genetic intermixture. Maximum haplotype diversity in a region showed a better correlation with the number of habitat specialists per steppe type than mean haplotype diversity. Main conclusions: We can clearly reject the scenario of a late-Holocene immigration of steppe species from areas outside the Pannonian Basin. Most species must have been present in the region since at least the early Holocene, highlighting the importance of the lower mountain ranges surrounding the Pannonian Basin as long-term refugia for European steppe species. Dispersal limitation and resulting migration lags seem to have a strong influence on the distribution of steppe species in Central Europe. [ABSTRACT FROM AUTHOR]

Forest understories play a vital role in ecosystem functioning and the provision of ecosystem services. However, the extent to which environmental conditions drive dominant ecological strategies in forest understories at the continental scale remains understudied. Here, we used ~29 500 forest vegetation plots sampled across Europe and classified into 25 forest types to explore the relative role of macroclimate, soil pH and tree canopy cover in driving abundance‐weighted patterns in the leaf economic spectrum (LES) and plant size spectrum (PSS) of forest understories (shrub and herb layers). We calculated LES using specific leaf area (SLA) and leaf dry matter content (LDMC) and PSS using plant height and seed mass of vascular plant species found in the understories. We found that forest understories had more conservative leaf economics in areas with more extreme mean annual temperatures (mainly Fennoscandia and the Mediterranean Basin), more extreme soil pH and under more open canopies. Warm and summer‐dry regions around the Mediterranean Basin and areas of Atlantic Europe also had taller understories with heavier seeds than continental temperate or boreal areas. Understories of broadleaved deciduous forests, such as Fagus forests on non‐acid soils, or ravine forests, more commonly hosted species with acquisitive leaf economics. In contrast, some coniferous forests, such as Pinus, Larix and Picea mire forests, or Pinus sylvestris light taiga and sclerophyllous forests, more commonly hosted species with conservative leaf economics. Our findings highlight the importance of macroclimate and soil factors in driving trait variation of understory communities at the continental scale and the mediator effect of canopy cover on these relationships. We also provide the first maps and analyses of LES and PSS of forest understories across Europe and give evidence that the understories of European forest types are differently positioned along major axes of trait variation. [ABSTRACT FROM AUTHOR]

Aims: The effect of biogeographical processes on the spatial turnover component of beta‐diversity over large spatial extents remains scarcely understood. Here, we aim at disentangling the roles of environmental and historical factors on plant taxonomic and phylogenetic turnover, while controlling for the effects of species richness and rarity. Location: European beech (Fagus sylvatica) forests in Europe. Methods: We aggregated plant species occurrences from vegetation plots in spatial grid cells of 0.25º × 0.25º to calculate the spatial turnover component of taxonomic (TBDturn) and phylogenetic (PBDturn) beta‐diversity for each cell. We also calculated the deviation of PBDturn given TBDturn (PBDdev‐turn), which measures the importance of phylogenetic turnover after factoring out taxonomic turnover. Beta‐diversity was calculated for each grid cell as the mean pairwise dissimilarity between the focal cell and all other cells. We used structural equation modeling (SEM) to examine the relationships between environmental (climate, soil pH, and distance from the geographical distribution limit of beech) and historical (distance from beech glacial refugia) predictors and beta‐diversity metrics. Results: We found a geographically consistent variation in taxonomic and phylogenetic turnover. Overall, TBDturn and PBDturn increased significantly toward more extreme climatic conditions, on more acidic soils, and toward the margins of beech distribution. The effects of environmental variables and the distance from glacial refugia on beta‐diversity metrics were mediated by species richness and rarity. Phylogenetic turnover was low in relation to taxonomic turnover (i.e., high PBDdev‐turn) in areas closer to glacial refugia. Conclusions: Continental‐scale patterns of beta‐diversity in European beech forests are the result of complementary ecological and evolutionary processes. In general, beech forests are taxonomically and phylogenetically more distinct in climatically marginal areas of their European range. However, the spatial variation of beta‐diversity in European beech forest flora is still strongly characterized by the distribution of groups of closely related species that evolved or survived in glacial refugia. [ABSTRACT FROM AUTHOR]

Aims: To link the Braun-Blanquet units of the EuroVegChecklist (EVC) with the upper levels of the International Vegetation Classification (IVC), and to propose a division level classification for Europe. Study area: Europe. Methods: We established a tabular linkage between EVC classes and IVC formations and identified mismatches between these two levels. We then proposed IVC division level units to organize EVC classes. Results: We organized the EVC classes into 21 formations and 30 divisions. We flagged classes that did not fit comfortably within an existing formation, either because its content corresponded to more than one formation or because it did not fit any formation description. In a few cases, we split EVC classes because they seemed too heterogenous to be assigned to a single formation. Conclusions: The IVC approach adds a set of physiognomic and ecological criteria that effectively organizes the EVC classes, which are already being increasingly informed by physiognomy. Therefore, the formation concepts are relatively natural extensions of concepts already embedded in the classes. However, physiognomic placement of Braun-Blanquet classes can be difficult when the sampling of the vegetation is at finer grain than usual in the respective formation (tall-scrub, annual pioneer communities). Some EVC classes seem too heterogenous to fit into the IVC formation system. Delimitation of these classes has often been a matter of debate for many decades, and the IVC perspective might help to solve these intricate issues. In other cases, mismatches between phytosociological classes and IVC formations might better be solved by emending the current formation concepts. [ABSTRACT FROM AUTHOR]

Aim: Current species‐richness patterns are sometimes interpreted as a legacy of landscape history, but historical processes shaping the distribution of species during the Holocene are frequently omitted in biodiversity models. Here, we test their importance in modelling current species richness of vascular plants in forest and grassland vegetation. Location: Western Carpathians and adjacent regions. Taxon: Vascular plants. Methods: Numbers of all species and of habitat specialists were extracted from plot records of forest and grassland vegetation. For each plot, environmental and historical data were derived from thematic maps. Historical data related to the persistence of (a) temperate taxa during the Late Glacial and Early Holocene, (b) open‐landscape taxa during the Middle Holocene and (c) taiga taxa during the Late Holocene were based on 112 fossil pollen profiles. Boosted regression trees were used to model spatial patterns in species richness. Results: Historical variables always appeared among the best predictors of current species richness. In light forests, species richness highly mirrored both the Late Glacial (12.5% contribution) and Middle‐Holocene (8.6%) landscape history. The latter factor became an important predictor also for species richness of steppe grasslands (8.3%) along with temperature seasonality (11.9%). Species richness of dark coniferous forests was best predicted by the Late‐Holocene occurrence of taiga forests (14.8%), which had an even stronger effect on the richness of habitat specialists (20.5%). Main conclusions: Landscape changes since the Last Glacial Maximum are important predictors of current plant species richness. The historical effects were found to be habitat specific and, because they may interact with recent environmental conditions and anthropogenic pressures, they often show a nonlinear relationship with species richness. We provide one possible direction of incorporating past landscape changes to the models of species richness. [ABSTRACT FROM AUTHOR]

A historical overview of the phytosociological method in Europe is presented. Some methodological and procedural differences in the application of the Braun-Blanquet approach, from the selection of the sampling plots to the assignment of relevés to existing or newly described units, are briefly compared. The main advantages and limitations of the phytosociological vegetation classification are reviewed and discussed, also in light of their applications for vegetation mapping and monitoring. [ABSTRACT FROM AUTHOR]

Questions What are the main floristic patterns in European beech forests? Which classification at the alliance and suballiance level is the most convincing? Location Europe and Asia Minor. Methods We applied a TWINSPAN classification to a data set of 24 605 relevés covering the whole range of Fagus sylvatica forests and the western part of Fagus orientalis forests. We identified 24 'operational phytosociological units' (OPUs), which were used for further analysis. The position of each OPU along the soil pH and temperature gradient was evaluated using Ellenberg Indicator Values. Fidelity of species to OPUs was calculated using the phi coefficient and constancy ratio. We compared alternative alliance concepts, corresponding to groups of OPUs, in terms of number and frequency of diagnostic species. We also established formal definitions for the various alliance concepts based on comparison of the total cover of the diagnostic species groups, and evaluated alternative geographical subdivisions of beech forests. Results The first and second division levels of TWINSPAN followed the temperature and soil pH gradients, while lower divisions were mainly geographical. We grouped the 22 OPUs of Fagus sylvatica forests into acidophytic, meso-basiphytic and thermo-basiphytic beech forests, and separated two OPUs of F. orientalis forests. However, a solution with only two ecologically defined alliances of F. sylvatica forests (acidophytic vs basiphytic) was clearly superior with regard to number and frequency of diagnostic species. In contrast, when comparing groupings with three to six geographical alliances of basiphytic beech forests, respectively, we did not find a strongly superior solution. Conclusions We propose to classify F. sylvatica forests into 15 suballiances - three acidophytic and 12 basiphytic ones. Separating these two groups at alliance or order level was clearly supported by our results. Concerning the grouping of the 12 basiphytic suballiances into ecological or geographical alliances, as advocated by many authors, we failed to find an optimal solution. Therefore, we propose a multi-dimensional classification of basiphytic beech forests, including both ecological and geographical groups as equally valid concepts which may be used alternatively depending on the purpose and context of the classification. [ABSTRACT FROM AUTHOR]

In this paper we present nomenclatural adjustments and validations of syntaxa of the forest vegetation of Europe. We introduce a new, valid name of the class of nemoral coniferous or mixed forests ( Asaro europaei-Abietetea sibiricae) replacing the deciduous Carpino-Fagetea in the easternmost Europe and Siberia. We describe two new orders for birch and birch-poplar woodlands, formerly included in the Betulo pendulae-Populetalia tremulae. We validate the names of two alliances for the deciduous forests of the Southern Urals and the name of an alliance for hemiboreal forest swamps. The suballiance Ostryo-Tilienion, coined to accommodate the xerothermophilous ravine forests of SE Europe, is here elevated to the rank of alliance. Finally, we validate the name Quercion alnifoliae (evergreen oak forests of Cyprus). [ABSTRACT FROM AUTHOR]

Aim Phylogenetic diversity patterns are increasingly being used to better understand the role of ecological and evolutionary processes in community assembly. Here, we quantify how these patterns are influenced by scale choices in terms of spatial and environmental extent and organismic scales. Location European Alps. Methods We applied 42 sampling strategies differing in their combination of focal scales. For each resulting sub-dataset, we estimated the phylogenetic diversity of the species pools, phylogenetic α-diversities of local communities, and statistics commonly used together with null models in order to infer non-random diversity patterns (i.e. phylogenetic clustering versus over-dispersion). Finally, we studied the effects of scale choices on these measures using regression analyses. Results Scale choices were decisive for revealing signals in diversity patterns. Notably, changes in focal scales sometimes reversed a pattern of over-dispersion into clustering. Organismic scale had a stronger effect than spatial and environmental extent. However, we did not find general rules for the direction of change from over-dispersion to clustering with changing scales. Importantly, these scale issues had only a weak influence when focusing on regional diversity patterns that change along abiotic gradients. Main conclusions Our results call for caution when combining phylogenetic data with distributional data to study how and why communities differ from random expectations of phylogenetic relatedness. These analyses seem to be robust when the focus is on relating community diversity patterns to variation in habitat conditions, such as abiotic gradients. However, if the focus is on identifying relevant assembly rules for local communities, the uncertainty arising from a certain scale choice can be immense. In the latter case, it becomes necessary to test whether emerging patterns are robust to alternative scale choices. [ABSTRACT FROM AUTHOR]

Deciduous forests form the dominant natural vegetation of Europe today, but were restricted to small refugia during Pleistocene cold stages, implying an evolutionary past shaped by recurrent range contractions and expansions. Cold-stage forest refugia were probably widespread in southern and central Europe, with the northwestern Balkan Peninsula being of particular importance. However, the actual number and location of deciduous forest refugia, as well as the connections between them, remain disputed. Here, we address the evolutionary dynamics of the deciduous forest understorey species Euphorbia carniolica as a proxy for past forest dynamics. To do so, we obtained genomic and morphometric data from populations representing the species' entire range, investigated phylogenetic position and intraspecific genetic variation, tested explicit demographic scenarios and applied species distribution models. Our data support two disjoint groups linked to separate refugia on the northwestern and central Balkan Peninsula. We find that genetic differentiation between groups started in the early Pleistocene via vicariance, suggesting a larger distribution in the past. Both refugia acted as sources for founder events to the southeastern Alps and the Carpathians; the latter were likely colonised before the last cold stage. In line with traditional views on the pre-Pleistocene origin of many southeastern European deciduous forest species, the origin of E. carniolica was dated to the late Pliocene. The fact that E. carniolica evolved at a time when a period of continuous forestation was ending in much of Eurasia provides an interesting biogeographical perspective on the past links between Eurasian deciduous forests and their biota., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

A central hypothesis of ecology states that regional diversity influences local diversity through species-pool effects. Species pools are supposedly shaped by large-scale factors and then filtered into ecological communities, but understanding these processes requires the analysis of large datasets across several regions. Here, we use a framework of community assembly at a continental scale to test the relative influence of historical and environmental drivers, in combination with regional or local species pools, on community species richness and community completeness. Using 42,173 vegetation plots sampled across European beech forests, we found that large-scale factors largely accounted for species pool sizes. At the regional scale, main predictors reflected historical contingencies related to post-glacial dispersal routes, whereas at the local scale, the influence of environmental filters was predominant. Proximity to Quaternary refugia and high precipitation were the main factors supporting community species richness, especially among beech forest specialist plants. Models for community completeness indicate the influence of large-scale factors, further suggesting community saturation as a result of dispersal limitation or biotic interactions. Our results empirically demonstrate how historical factors complement environmental gradients to provide a better understanding of biodiversity patterns across multiple regions.