Topographic depressions can provide climate and resource microrefugia for biodiversity.

Microrefugia are often located within topographically complex regions where stable environmental conditions prevail. Most of the studies concerning the distributions of climate change-sensitive species have emphasized the dominance of cold air pooling over other environmental factors, such as resource availability. There is a shortage of information on the relationships among topography-related microsite diversity, microclimate, resource availability, and species composition in microrefugia. To fill this knowledge gap, we studied the effects of microclimatic conditions and soil resources on plant species occurrence within and adjacent to 30 large topographic depressions (i.e., dolines) in two distant karst regions. Our results showed that both microclimate and soil resource availability may play a key role in maintaining climate change-sensitive species and biodiversity in dolines; therefore, they may simultaneously act as climate and resource microrefugia. Establishing climate-smart conservation priorities and strategies is required to maintain or increase the refugial capacity of such safe havens.

PADAPT 1.0 - the Pannonian Dataset of Plant Traits.

The existing plant trait databases' applicability is limited for studies dealing with the flora and vegetation of the eastern and central part of Europe and for large-scale comparisons across regions, mostly because their geographical data coverage is limited and they incorporate records from several different sources, often from regions with markedly different climatic conditions. These problems motivated the compilation of a regional dataset for the flora of the Pannonian region (Eastern Central Europe). PADAPT, the Pannonian Dataset of Plant Traits relies on regional data sources and collates data on 54 traits and attributes of the plant species of the Pannonian region. The current version covers approximately 90% of the species of the region and consists of 126,337 records on 2745 taxa. By including species of the eastern part of Europe not covered by other databases, PADAPT can facilitate studying the flora and vegetation of the eastern part of the continent. Although data coverage is far from complete, PADAPT meets the longstanding need for a regional database of the Pannonian flora.

Plant invasion and fragmentation indirectly and contrastingly affect native plants and grassland arthropods.

Plant invasion and habitat fragmentation have a detrimental effect on biodiversity in nearly all types of ecosystems. We compared the direct and indirect effects of the invasion of the common milkweed (Asclepias syriaca) on biodiversity patterns in different-sized Hungarian forest-steppe fragments. We assessed vegetation structure, measured temperature and soil moisture, and studied organisms with different ecological roles in invaded and non-invaded sites of fragments: plants, bees, butterflies, flower-visiting wasps, flies, true bugs, and spiders. Temperature and soil moisture were lower in invaded than in non-invaded area. Milkweed had a positive effect on plant species richness and flower abundance. In contrast, we mainly found indirect effects of invasion on arthropods through alteration of physical habitat characteristics and food resources. Pollinators were positively affected by native flowers, thus, milkweed indirectly supported pollinators. Similarly, we found higher species richness of herbivores in invaded sites than control sites, as species richness of true bugs also increased with increasing plant species richness. Predators were positively affected by complex vegetation structure, higher soil moisture and lower temperature. Furthermore, increasing fragment size had a strong negative effect on spider species richness of non-invaded sites, but no effect in invaded sites. Especially, grassland specialist spiders were more sensitive to fragment size than generalists, whereas generalist spider species rather profited from invasion. Although milkweed invades natural areas, we did not identify strong negative effects of its presence on the diversity of the grassland biota. However, the supportive effect of milkweed on a few generalist species homogenises the communities. The rate of invasion might increase with increasing fragmentation, therefore we recommend eliminating invasive plants from small habitat fragments to preserve the native biota. Focusing also on generalist species and revealing the indirect effects of invasions are essential for understanding the invasion mechanisms and would support restoration efforts.

Environmental heterogeneity increases the conservation value of small natural features in karst landscapes.

Local biodiversity hotspots are often located within regions where extreme and variable environmental - e.g., climatic and soil - conditions occur. These areas are conservation priorities. Although environmental heterogeneity is recognised as an important determinant of biodiversity, studies focusing on the effects of multiple environmental heterogeneity components in the same ecosystem are scarce. Here we investigate how topography and related microclimatic variables and soil properties may influence the biodiversity and conservation value of karst landscapes. Karst landscapes of the world contain millions of dolines (i.e. bowl- or funnel-shaped depressions) that may function as 'small natural features' with a disproportionately large role in maintaining biodiversity relative to their size. We assessed the diversity of microclimates, soils and vegetation and their relationships in six microhabitats (south-facing slopes, east-facing slopes, west-facing slopes, north-facing slopes and bottoms of dolines, and the adjacent plateau) for nine large dolines in a grassland ecosystem. Although there were remarkable differences among the conservation value of these microhabitats (e.g., representation of different species groups, presence of 'climate relicts'), each microhabitat had an important role in maintaining species that are rare or absent in other microhabitats in the landscape. We found that the studied dolines exhibited highly variable environmental conditions and promoted a high diversity of vegetation types with unique species composition, contributing to the topographic, climatic, soil, vegetation and land cover heterogeneity of karst landscapes. Therefore, our findings highlight that dolines may function as local biodiversity hotspots and have a crucial conservation importance. As dolines are widespread topographic features in many karst landscapes throughout the world, our results could be directly applied to other regions as well. An integrated approach is urgently needed to provide guidelines for landscape management, promoting the retention of the microhabitat diversity of small natural features for species vulnerable to climate change and/or various disturbances.

How to not trade water for carbon with tree planting in water-limited temperate biomes?

The most widespread nature-based solution for mitigating climate change is tree planting. When realized as forest restoration in historically forested biomes, it can efficiently contribute to the sequestration of atmospheric carbon and can also entail significant biodiversity and ecosystem service benefits. Conversely, tree planting in naturally open biomes can have adverse effects, of which water shortage due to increased evapotranspiration is among the most alarming ones. Here we assessed how soil texture affects the strength of the trade-off between tree cover and water balance in the forest-steppe biome, where the global pressure for afforestation is threatening with increasing tree cover above historical levels. Here we monitored vertical soil moisture dynamics in four stands in each of the most common forest types of lowland Hungary on well-drained, sandy (natural poplar groves, and Robinia and pine plantations) and on poorly drained, silty-clayey soils (natural oak stands and Robinia plantations), and neighboring grasslands. We found that forests on sand retain moisture in the topsoil (approx. 20 cm) throughout the year, but a thick dry layer develops below that during the vegetation period, significantly impeding groundwater recharge. Neighboring sandy grasslands showed an opposite pattern, with often dry topsoil but intact moisture reserves below, allowing deep percolation. In contrast, forests on silty-clayey soils did not desiccate lower soil layers compared neighboring grasslands, which in turn showed moisture patterns similar to sandy grasslands. We conclude that, in water-limited temperate biomes where landscape-wide water regime depends on deep percolation, soil texture should drive the spatial allocation of tree-based climate mitigation efforts. On sand, the establishment of new forests should be kept to a minimum and grassland restoration should be preferred. The trade-off between water and carbon is less pronounced on silty-clayey soils, making forest patches and wooded rangelands viable targets for both climate mitigation and ecosystem restoration.

How climate, topography, soils, herbivores, and fire control forest-grassland coexistence in the Eurasian forest-steppe.

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.

Indirect effects of COVID-19 on the environment: How plastic contamination from disposable surgical masks affect early development of plants.

Personal protective equipment, used extensively during the COVID-19 pandemic, heavily burdened the environment due to improper waste management. Owing to their fibrous structure, layered non-woven polypropylene (PP) disposable masks release secondary fragments at a much higher rate than other plastic waste types, thus, posing a barely understood new form of ecological hazard. Here we show that PP mask fragments of different sizes induce morphogenic responses in plants during their early development. Using in vitro systems and soil-filled rhizotrons, we found that several PP mask treatments modified the root growth of Brassica napus (L.) regardless of the experimental system. The environment around the root and mask fragments seemed to influence the effect of PP fabric fragment contamination on early root growth. In soil, primary root length was clearly inhibited by larger PP mask fragments at 1 % concentration, while the two smallest sizes of applied mask fragments caused distinct, concentration-dependent changes in the lateral root numbers. Our results indicate that PP can act as a stressor: contamination by PP surgical masks affects plant growth and hence, warrants attention. Further investigations regarding the effects of plastic pollution on plant-soil interactions involving various soil types are urgently needed.

Post-restoration grassland management overrides the effects of restoration methods in propagule-rich landscapes.

Grassland restoration is gaining momentum worldwide to tackle the loss of biodiversity and associated ecosystem services. Restoration methods and their effects on ecological community reassembly have been extensively studied across various grassland types, while the importance of post-restoration management has so far received less attention. Grassland management is an important surrogate for natural disturbances, with which most ancient grasslands have coevolved. Thus, without the reintroduction of management-related disturbance, restoration targets are unlikely to be achieved in restored grasslands. In this study, we aimed to explore how 20 yr of management by mowing once a year or light cattle grazing affects restoration success in Palearctic meadow-steppe grasslands restored by either sowing native grasses (sown sites), applying Medicago sativa as a nurse plant (Medicago sites), or allowing spontaneous succession (spontaneous sites). We found that, following mowing, sown sites maintained long-lasting establishment limitation, while Medicago sites experienced a delay in succession. These limitations resulted in low total and target species richness, low functional redundancy, and distinct species and functional composition compared to reference data from ancient grasslands. Spontaneous sites that were mowed reached a more advanced successional stage, although they did not reach reference levels regarding most vegetation descriptors. Sown and Medicago sites that were grazed had higher total and target species richness than those that were mowed, and showed restoration success similar to that of spontaneous sites, on which grazing had only moderate further positive effects. Grazed sites, irrespective of the restoration method, were uniformly species rich, functionally diverse, and functionally redundant, and thus became important biodiverse habitats with considerable resilience. We conclude that an optimally chosen post-restoration management may have an impact on long-term community reassembly comparable to the choice of restoration method. Restoration planners may, therefore, need to put more emphasis on future management than on the initial restoration method. However, our findings also imply that if local constraints, such as potentially high invasive propagule pressure, necessitate the application of restoration methods that could also hinder the establishment of target species, the long-term recovery of the grassland can still be ensured by wisely chosen post-restoration management.

Natura 2000 Areas, Road, Railway, Water, and Ecological Networks May Provide Pathways for Biological Invasion: A Country Scale Analysis.

Invasive species are a major threat to biodiversity worldwide. Controlling their rapid spread can only be effective if we consider the geographical factors that influence their occurrence. For instance, roads, railway networks, green and blue infrastructure, and elements of ecological networks (e.g., ecological corridors) can facilitate the spread of invasive species. In our study, we mapped the occurrence of five invasive plant taxa (tree of heaven, common milkweed, Russian olive, black locust, and goldenrods) in Hungary, using field photos from the EUROSTAT Land Use and Coverage Area Frame Survey (LUCAS) database from the year 2015. Species point occurrence data were compared with the spatial characteristics of linear transport infrastructure and with the green and blue infrastructure. We found that the occurrence of tree of heaven and Russian olive was strongly related to the road and railway network. The average Euclidean distance of LUCAS points infected with these species from railway embankments and roads was much smaller than that of uninfected points. However, black locust and goldenrods were more common only along the road network. According to our results, the occurrence of some investigated invasive plants was over-represented in the HEN and within Natura 2000 areas of Hungary compared to non-infected points. Our results may provide important information for predicting the rate of invasion and for applying targeted management within the HEN, and Natura 2000 protected areas.

Beyond the Forest-Grassland Dichotomy: The Gradient-Like Organization of Habitats in Forest-Steppes.

Featuring a transitional zone between closed forests and treeless steppes, forest-steppes cover vast areas, and have outstanding conservation importance. The components of this mosaic ecosystem can conveniently be classified into two basic types, forests and grasslands. However, this dichotomic classification may not fit reality as habitat organization can be much more complex. In this study, our aim was to find out if the main habitat types can be grouped into two distinct habitat categories (which would support the dichotomic description), or a different paradigm better fits this complex ecosystem. We selected six main habitats of sandy forest-steppes, and, using 176 relevés, we compared their vegetation based on species composition (NMDS ordination, number of common species of the studied habitats), relative ecological indicator values (mean indicators for temperature, soil moisture, and light availability), and functional species groups (life-form categories, geoelement types, and phytosociological preference groups). According to the species composition, we found a well-defined gradient, with the following habitat order: large forest patches, medium forest patches, small forest patches, north-facing edges, south-facing edges, and grasslands. A considerable number of species were shared among all habitats, while the number of species restricted to certain habitat types was also numerous, especially for north-facing edges. The total (i.e., pooled) number of species peaked near the middle of the gradient, in north-facing edges. The relative ecological indicator values and functional species groups showed mostly gradual changes from the large forest patches to the grasslands. Our results indicate that the widely used dichotomic categorization of forest-steppe habitats into forest and grassland patches is too simplistic, potentially resulting in a considerable loss of information. We suggest that forest-steppe vegetation better fits the gradient-based paradigm of landscape structure, which is able to reflect continuous variations.

Density-Dependent Plant-Plant Interactions Triggered by Grazing.

Plant species performance in rangelands highly depends on the effect of grazing and also on the occurrence of unpalatable benefactor species that can act as biotic refuges protecting neighboring plants from herbivores. The balance between facilitation and competition may changes with the benefactor density. Despite the high number of studies on the role of biotic refuges, the density dependent effects of unpalatable herbaceous plants on the performance of other species, and on the habitat heterogeneity of rangelands are still unclear. Therefore, we performed a study to test the following hypotheses: (i) Performances of understory species follow a humped-back relationship along the density gradient of the unpalatable benefactor species. (ii) Small-scale heterogeneity of the vegetation decreases with increasing benefactor density. We studied meadow steppes with medium intensity cattle grazing in Hungary. We surveyed understory species' performance (number of flowering shoots and cover scores) along the density gradient of a common, native unpalatable species (Althaea officinalis). Our findings supported both hypotheses. We found unimodal relationship between the benefactor cover and both the flowering success and richness of understory species. Moreover, small-scale heterogeneity declined with increasing benefactor cover. In this study we detected a humped-back pattern of facilitation along the density gradient of an herbaceous benefactor in pastures. Indeed, this pattern was predictable based on such conceptual models like "consumer pressure-abiotic stress model," "humped-back model," "intermediate disturbance hypothesis," and "disturbance heterogeneity model"; but until now the validity of these relationships has not been demonstrated for herbaceous species. By the demonstration of this effect between herbaceous species we can better forecast the responses of grasslands to changes in management.

Karst dolines provide diverse microhabitats for different functional groups in multiple phyla.

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.

Large- and small-scale environmental factors drive distributions of cool-adapted plants in karstic microrefugia.

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.