Aims: Eurasian forest‐steppes are among the most complex non‐tropical terrestrial ecosystems. Despite their considerable scientific, ecological and economic importance, knowledge of forest‐steppes is limited, particularly at the continental scale. Here we provide an overview of Eurasian forest‐steppes across the entire zone: (a) we propose an up‐to‐date definition of forest‐steppes, (b) give a short physiogeographic outline, (c) delineate and briefly characterize the main forest‐steppe regions, (d) explore forest‐steppe biodiversity and conservation status, and (e) outline forest‐steppe prospects under predicted climate change.Location: Eurasia (29°–56°N, 16°–139°E).Results and Conclusions: Forest‐steppes are natural or near‐natural vegetation complexes of arboreal and herbaceous components (typically distributed in a mosaic pattern) in the temperate zone, where the co‐existence of forest and grassland is enabled primarily by the semi‐humid to semi‐arid climate, complemented by complex interactions of biotic and abiotic factors operating at multiple scales. This new definition includes lowland forest–grassland macromosaics (e.g. in Eastern Europe), exposure‐related mountain forest‐steppes (e.g. in Inner Asia), fine‐scale forest–grassland mosaics (e.g. in the Carpathian Basin) and open woodlands (e.g. in the Middle East). Using criteria of flora, physiognomy, relief and climate, nine main forest‐steppe regions are identified and characterized. Forest‐steppes are not simple two‐phase systems, as they show a high level of habitat diversity, with forest and grassland patches of varying types and sizes, connected by a network of differently oriented edges. Species diversity and functional diversity may also be exceptionally high in forest‐steppes. Regarding conservation, we conclude that major knowledge gaps exist in determining priorities at the continental, regional, national and local levels, and in identifying clear target states and optimal management strategies. When combined with other threats, climate change may be particularly dangerous to forest‐steppe survival, possibly resulting in compositional changes, rearrangement of the landscape mosaic or even the latitudinal or altitudinal shift of forest‐steppes.

Recent advances in ecology and biogeography demonstrate the importance of fire and large herbivores – and challenge the primacy of climate – to our understanding of the distribution, stability, and antiquity of forests and grasslands. Among grassland ecologists, particularly those working in savannas of the seasonally dry tropics, an emerging fire–herbivore paradigm is generally accepted to explain grass dominance in climates and on soils that would otherwise permit development of closed‐canopy forests. By contrast, adherents of the climate–soil paradigm, particularly foresters working in the humid tropics or temperate latitudes, tend to view fire and herbivores as disturbances, often human‐caused, which damage forests and reset succession. Towards integration of these two paradigms, we developed a series of conceptual models to explain the existence of an extensive temperate forest–grassland mosaic that occurs within a 4.7 million km² belt spanning from central Europe through eastern Asia. The Eurasian forest‐steppe is reminiscent of many regions globally where forests and grasslands occur side‐by‐side with stark boundaries. Our conceptual models illustrate that if mean climate was the only factor, forests should dominate in humid continental regions and grasslands should prevail in semi‐arid regions, but that extensive mosaics would not occur. By contrast, conceptual models that also integrate climate variability, soils, topography, herbivores, and fire depict how these factors collectively expand suitable conditions for forests and grasslands, such that grasslands may occur in more humid regions and forests in more arid regions than predicted by mean climate alone. Furthermore, boundaries between forests and grasslands are reinforced by vegetation–fire, vegetation–herbivore, and vegetation–microclimate feedbacks, which limit tree establishment in grasslands and promote tree survival in forests. Such feedbacks suggest that forests and grasslands of the Eurasian forest‐steppe are governed by ecological dynamics that are similar to those hypothesised to maintain boundaries between tropical forests and savannas. Unfortunately, the grasslands of the Eurasian forest‐steppe are sometimes misinterpreted as deforested or otherwise degraded vegetation. In fact, the grasslands of this region provide valuable ecosystem services, support a high diversity of plants and animals, and offer critical habitat for endangered large herbivores. We suggest that a better understanding of the fundamental ecological controls that permit forest–grassland coexistence could help us prioritise conservation and restoration of the Eurasian forest‐steppe for biodiversity, climate adaptation, and pastoral livelihoods. Currently, these goals are being undermined by tree‐planting campaigns that view the open grasslands as opportunities for afforestation. Improved understanding of the interactive roles of climate variability, soils, topography, fire, and herbivores will help scientists and policymakers recognise the antiquity of the grasslands of the Eurasian forest‐steppe.

Woody plants in water‐limited ecosystems affect their environment on multiple scales: locally, natural stands can create islands of fertility for herb layer communities compared to open habitats, but afforestation has been shown to negatively affect regional water balance and productivity. Despite these contrasting observations, no coherent multiscale framework has been developed for the environmental effects of woody plants in water‐limited ecosystems. To link local and regional effects of woody species in a spatially explicit model, we simultaneously measured site conditions (microclimate, nutrient availability and topsoil moisture) and conditions of regional relevance (deeper soil moisture), in forests with different canopy types (long, intermediate and short annual lifetime) and adjacent grasslands in sandy drylands. All types of forests ameliorated site conditions compared to adjacent grasslands, although natural stands did so more effectively than managed ones. At the same time, all forests desiccated deeper soil layers during the vegetation period, and the longer the canopy lifetime, the more severe the desiccation in summer and more delayed the recharge after the active period of the canopy. We conclude that the site‐scale environmental amelioration brought about by woody species is bound to co‐occur with the desiccation of deeper soil layers, leading to deficient ground water recharge. This means that the cost of creating islands of fertility for sensitive herb layer organisms is an inevitable negative impact on regional water balance. The canopy type or management intensity of the forests affects the magnitude but not the direction of these effects. The outlined framework of the effects of woody species should be considered for the conservation, restoration or profit‐oriented use of forests as well as in forest‐based carbon sequestration and soil erosion control projects in water‐limited ecosystems.

As key components of landscapes, edges have received considerable scientific attention in anthropogenic ecosystems. However, edges in natural and semi-natural forest–grassland mosaics have received less attention, despite the fact that they cover a considerable proportion of these mosaic ecosystems. We studied forest edges in a semi-natural forest–grassland mosaic ecosystem of the Samobor Mountains (Croatia). Our aim was to compare the species composition, diversity and ecological indicator values of forest edges to those of the interior parts of the adjacent forest and grassland habitats. The vegetation was studied in 80 plots established in forest patch interiors, north-facing forest edges, south-facing forest edges and grassland interiors. We found that edges had a unique species composition, containing species from both the forest and the grassland interiors plus their own edge-related species (i.e. species that significantly preferred the edge habitat). These local edge-related species did not correspond to regionally-identified edge-related species. Compared to the forest and the grassland interiors, we revealed increased species richness in north-facing edges but not in south-facing edges. The mean light availability and nutrient supply indicator values of the edges were intermediate between those of the forest interiors and the grasslands. The mean soil moisture indicator values of the edges were similar to those of the grasslands. Our results show that edges form a unique component of forest–grassland mosaics, and they contribute considerably to landscape complexity, which should be taken into account during conservation decisions and habitat management.

Ecosystems with forest and grassland patches as alternative stable states usually contain various closed, semi‐open and open habitats, which may be aligned along a vegetation cover gradient. Taxonomic diversity usually peaks near the middle of the gradient, but our knowledge on functional and phylogenetic diversity trends along gradients is more limited. We investigated the eight main habitats of Hungarian forest‐grassland mosaics, representing various vegetation cover values, and compared their species composition as well as their taxonomic, functional and phylogenetic diversity. We found a compositional gradient ranging from large forest patches through smaller‐sized forest patches and edges to closed and open grasslands. Species richness peaked at the middle of the gradient (at edges). Shannon diversity was high near the middle and at the open end of the gradient. Functional diversity was high throughout woody habitats (in forests and at edges) and was significantly lower in grasslands. When considering all species, phylogenetic diversity tended to peak at north‐facing forest edges. When excluding non‐angiosperms, this peak disappeared. The high taxonomic diversity at the middle of the gradient is in line with the edge‐effect theory. Our results suggest that community assembly in grasslands may be dominated by environmental filtering, while competition may be decisive in woody habitats. The low phylogenetic diversity of grassland habitats can be explained by their young evolutionary age compared to forests. Synthesis. Functional and phylogenetic diversity do not necessarily coincide with taxonomic diversity along vegetation cover gradients. In ecosystems where forest and grassland patches represent alternative stable states, the trends of taxonomic diversity may be similar to those revealed here, but functional diversity patterns may be more system‐specific for some traits. Trends in phylogenetic diversity may vary according to the evolutionary age of the habitats.

Questions: Woody encroachment affects many open habitats from semi‐deserts to wetlands and grasslands. We aimed to study the effect of native woody encroachment on grassland plant biodiversity in loess steppe fragments by analysing the vegetation composition of grasslands subjected to increasing levels of encroachment. We studied both ancient and restored grasslands with the following research hypotheses: (a) increasing woody encroachment decreases total diversity and the species richness of dry‐grassland species; and (b) the effect of woody cover on grassland biodiversity differs between ancient and restored grasslands.Location: South‐Eastern part of Transdanubia, Hungary, Central Europe.Methods: Altogether 63 loess grassland fragments were selected for the study. The percentage cover of trees, shrubs and herbaceous vegetation were recorded in 400‐m²‐sized plots (n = 110). The effects of woody encroachment and grassland age on diversity, total species richness of the herb layer, and richness of dry‐grassland species were analysed.Results: We found that woody encroachment affected the total richness of the herb layer and the species richness of dry‐grassland species. For most of the listed variables, lowest values were found for the highest woody encroachment groups. Grassland age affected the species richness of the herb layer and that of dry‐grassland species; lower values were detected in restored grasslands. In restored grasslands, Shannon diversity and species evenness were lower, while Berger–Parker dominance was higher than in ancient grasslands. Species composition and richness displayed a relatively high resistance to moderate woody encroachment; the highest decrease in diversity was detected at a high level of woody encroachment (>52% of woody cover).Conclusions: We can conclude that low to moderate woody encroachment cannot be simplistically regarded as degradation. Results suggest that moderately encroached loess grasslands can be easily restored by the suppression of woody species, as their species pool still contains many dry‐grassland species targeted for restoration.