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.

Earth is facing simultaneous biodiversity and climate crises. Climate-change refugia - areas that are relatively buffered from climate change - can help address both of these problems by maintaining biodiversity components when the surrounding landscape no longer can. However, this capacity to support biodiversity is often vulnerable to severe climate change and other stressors. Thus, management actions need to consider the complex and multidimensional nature of refugia. We outline an approach to understand refugia-promoting processes and to evaluate refugial capacity to determine suitable management actions. Our framework applies climate-change refugia as tools to facilitate resistance in modern conservation planning. Such refugia-focused management can reduce extinctions and maintain biodiversity under climate change.

Background and aims Dolines are small- to large-sized bowl-shaped depressions of karst surfaces. They may constitute important microrefugia, as thermal inversion often maintains cooler conditions within them. This study aimed to identify the effects of large- (macroclimate) and small-scale (slope aspect and vegetation type) environmental factors on cool-adapted plants in karst dolines of East-Central Europe. We also evaluated the potential of these dolines to be microrefugia that mitigate the effects of climate change on cool-adapted plants in both forest and grassland ecosystems. Methods We compared surveys of plant species composition that were made between 2007 and 2015 in 21 dolines distributed across four mountain ranges (sites) in Hungary and Romania. We examined the effects of environmental factors on the distribution and number of cool-adapted plants on three scales: (1) regional (all sites); (2) within sites and; (3) within dolines. Generalized linear models and non-parametric tests were used for the analyses. Key Results Macroclimate, vegetation type and aspect were all significant predictors of the diversity of cool-adapted plants. More cool-adapted plants were recorded in the coolest site, with only few found in the warmest site. At the warmest site, the distribution of cool-adapted plants was restricted to the deepest parts of dolines. Within sites of intermediate temperature and humidity, the effect of vegetation type and aspect on the diversity of cool-adapted plants was often significant, with more taxa being found in grasslands (versus forests) and on north-facing slopes (versus south-facing slopes). Conclusions There is large variation in the number and spatial distribution of cool-adapted plants in karst dolines, which is related to large- and small-scale environmental factors. Both macro- and microrefugia are therefore likely to play important roles in facilitating the persistence of cool-adapted plants under global warming.

Fine-scale topographic complexity creates important microclimates that can facilitate species to grow outside their main distributional range and increase biodiversity locally. Enclosed depressions in karst landscapes (‘dolines’) are topographically complex environments which produce microclimates that are drier and warmer (equator-facing slopes) and cooler and moister (pole-facing slopes and depression bottoms) than the surrounding climate. We show that the distribution patterns of functional groups for organisms in two different phyla, Arthropoda (ants) and Tracheophyta (vascular plants), mirror this variation of microclimate. We found that north-facing slopes and bottoms of solution dolines in northern Hungary provided key habitats for ant and plant species associated with cooler and/or moister conditions. Contrarily, south-facing slopes of dolines provided key habitats for species associated with warmer and/or drier conditions. Species occurring on the surrounding plateau were associated with intermediate conditions. We conclude that karst dolines provide a diversity of microclimatic habitats that may facilitate the persistence of taxa with diverse environmental preferences, indicating these dolines to be potential safe havens for multiple phyla under local and global climate oscillations.