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Dolines are depressions in karst landscapes that are of high value for conservation, providing habitats and supporting species not found in the surrounding landscape. This is due to their high microhabitat diversity and ability to decouple microclimate from regional climate changes, making them potential refugia for biodiversity. Nevertheless, local anthropogenic disturbances have had considerable impact on the species composition and vegetation structure of many dolines. Here we investigate the conservation value of dolines in three European karst areas, where different levels and types of anthropogenic disturbances have been shaping the vegetation for centuries, using the number of plant species that are cool-adapted, moist-adapted and of high conservation importance (i.e. vulnerable species) as indicators. We found that anthropogenic disturbances generally have a negative impact, reducing the number of vulnerable species supported by dolines. However, more cool-adapted and moist-adapted species were found in some dolines planted with non-native Picea abies than in less disturbed dolines, indicating that anthropogenic disturbances can also have positive consequences for biodiversity. We conclude that anthropogenic disturbances alter the capacity of dolines to support vulnerable species, and that this will impact survival of species in landscapes under global warming. In this context, the effects of various disturbances on species composition and diversity need to carefully considered to determine the best conservation and/or management options.

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. Refereed/Peer-reviewed