Introduction: The recovery of habitats following restoration requires that suitable environmental conditions, propagule sources, and adequate active management all co-occur. Identifying the extent to which these different factors co-exist can support post-restoration actions and pinpoint possible locations for colonization by plant species at a landscape scale. We studied a 400 ha landscape undergoing restoration to identify the suitable habitats for the establishment of grassland communities. Methods: We combined environmental data for variables which are key for plant species occurrence (vegetation structure, soil moisture, slope range, cover of bedrock) with observational layers of connectivity (proximity to propagule source and livestock activity by using global positioning system collars in the landscape) to classify restored sites according to their potential to support establishment of grassland patches. Using 320 plant inventory plots (1 × 1 m) distributed over the area, we tested the relationship between suitability values from models and aspects of plant community composition. Results: We found that plant community composition was best explained by the interactions between environmental conditions and the location of propagule source and livestock activity. Greater proximity to propagule sources and the activity of grazing animals increased the likelihood for grassland species to colonize. Conclusions: Applying suitability maps by using spatial layers in restoration efforts can be a useful tool for locating candidate sites and planning post-restoration management actions. It would allow prioritizing certain locations where restoration could be most effective, for instance by steering the grazing regime to enhance colonization of target species.

Habitat fragmentation increases the proportion of forest borders in the landscape and many forest borders lose their structural complexity due to modern forestry practices. However, remnants of structurally complex deciduous forests can remain as ecotones between plantations and agricultural fields. In this study we used terrestrial laser scanning to measure structural complexity of different forest borders, measured microclimate, and surveyed bats and macro-moths to understand how these taxa are affected. Our aim is to disentangle the main drivers, direct or indirect, that influence bat and moth assemblages. We studied 79 forest borders, and surrounding landscapes and compared them with adjacent agricultural fields and coniferous plantations. Overall, we found less bat activity and lower macro-moth diversity in simple compared to complex borders. Using structural equation modelling, we show the contrasting responses of forest-specialist bats and moths to structural complexity; with bats responding positively and moths negatively. We found similar divergent results in relation to understorey openness; with increasing forest-specialist bat activity but a lower diversity of forest-specialist moths in more open borders. Understorey vegetation also appears to regulate microclimate with more open borders being warmer and less humid. This has a potential knock-on effect for bats as they favoured borders that were warmer and more humid. Surrounding land-cover was more important than structural complexity for generalist species; with increasing generalist bat activity due to a higher proportion of local deciduous forest cover and increasing generalist moth diversity in landscapes with more forest borders. Overall, these complex relationships between forest structure, microclimate and landscape factors, coupled with divergent responses of both taxa highlight their diverse ecological needs. Therefore, we highlight the importance of managing forest borders to retain complexity and connectivity within multifunctional landscapes.

Sympatric bat species can co-exist and avoid interspecific competition via niche differentiation e.g. diet. Detecting dietary differences can be achieved by comparing dietary niches of sympatric and allopatric populations. If dietary overlap is higher in sympatry versus allopatry, co-occurrence may be altering the dietary niche of the species. Our study region offers an excellent opportunity to investigate this because two species, Eptesicus nilssonii and Pipistrellus pygmaeus, occur sympatrically across southern Sweden and E. nilssonii occurs allopatrically in the north. We analysed droppings from 19 roosts during two months: six P. pygmaeus, seven E. nilssonii located in the known distribution of P. pygmaeus (sympatric) and six roosts located outside the known range of P. pygmaeus (allopatric). We used DNA metabarcoding to assess dietary overlap and dietary niche breadth in both species. We found a high dietary overlap between E. nilssonii and P. pygmaeus at all taxonomic levels of prey. The dominant prey orders in the three bat populations were Ephemeroptera (mayflies) and one Diptera species (cranefly); five species from these orders accounted for > 70% of all reads. When using quantitative relative read abundance data at the species level interspecific overlap was higher than intraspecific overlap. We find a consistent but not significant pattern that E. nilssonii populations co-occurring with P. pygmaeus have the narrowest dietary niche breadth. This could indicate that the co-occurrence with P. pygmaeus is having an impact on prey selection for sympatric E. nilssonii populations. However, confirming competition between these species would require sampling of the insect population near to the colonies. Both bat species' diets are also heavily dominated by only five species. These are all currently common insect species and most likely indicates opportunistic foraging on abundant insect populations.

1. Agricultural intensification, afforestation and land abandonment are major drivers of biodiversity loss in semi-natural grasslands across Europe. Reversing these losses requires the reinstatement of plant–animal interactions such as pollination. Here we assessed the differences in species composition and patterns of plant-pollinator interactions in ancient and restored grasslands and how these patterns are influenced by landscape connectivity, across three European regions (Belgium, Germany and Sweden).
2. We evaluated the differences in pollinator community assemblage, abundance and interaction network structure between 24 ancient and restored grasslands. We then assessed the effect of surrounding landscape functional connectivity (i.e. green infrastructure, GI) on these variables and tested possible consequences on the reproduction of two model plants, Lotus corniculatus and Salvia pratensis.
3. Neither pollinator richness nor species composition differed between ancient and restored grasslands. A high turnover of interactions across grasslands was detected but was mainly due to replacement of pollinator and plant species. The impact of grassland restoration was consistent across various pollinator functional groups, whereas the surrounding GI had differential effects. Notably, bees, butterflies, beetles, and dipterans (excluding hoverflies) exhibited the most significant responses to GI variations. Interestingly, networks in restored grasslands were more specialised (i.e. less functionally redundant) than in ancient ones and also showed a higher number of insect visits to habitat-generalist plant species. Landscape connectivity had a similar effect, with habitat-specialist plant species receiving fewer visits at higher GI values.
4. Fruit set in S. pratensis and L. corniculatus was unaffected by grassland type or GI. However, the fruit set in the specialist S. pratensis increased with the number of pollinator visits, indicating a positive correlation between pollinator activity and reproductive success in this particular species.
5. Synthesis and applications. Our findings provide evidence of the necessity to enhance ecosystem functions while avoiding biotic homogenisation. Restoration programs should aim at increasing landscape connectivity which influences plant communities, pollinator assemblages and their interaction patterns. To avoid generalist species taking over from specialists in restored grasslands, we suggest reinforcing the presence of specialist species in the latter, for instance by means of introductions, as well as increasing the connectivity to source populations.

In recent years, restoration on former grassland sites has been widely encouraged globally, aiming to address the historical loss of 90% of ancient species-rich grasslands, and to mitigate the associated threat to grassland biodiversity. The objective of our study was to investigate on a small-scale how plant species spontaneously colonize restored grasslands. We inventoried 275 permanent plots twice (in 2019 and 2021) in a restored grassland, following the removal of a conifer plantation. Species richness and vegetation cover in surveyed plots were dependent on grazing activity and distance to adjacent grassland. Plant species associated with forest habitats declined, while the occurrence of generalist species together with a few grassland specialists generally increased. However, not all grassland specialists gained occurrence and the colonization pattern was not consistent over time, possibly due to the lack of continuous seed arrival and low livestock activity and hence lack of disturbance. These results suggest that successful colonization of plant species benefits from links to species-rich sites adjacent to the restoration target, with spatial dispersal and improved conditions for species establishment being key to species occurrence. Both dispersal and establishment potential are likely facilitated through the presence of grazing livestock with access to both species-rich grasslands and restoration targets. However, the shift towards a more typical grassland community takes place gradually, with vulnerable populations of early colonizing grassland species prone to local extinction in short-term. As a result, continued functional connectivity provided by grazing animals is necessary to improve the diversity of the restored site and ensure the establishment of grassland specialists and to maintain the plant community composition.

Questions: Baltic coastal meadows are ecologically unique habitats that have been severely impacted by habitat loss and environmental change. To determine the effects of habitat loss and isolation on their plant communities, we analysed the relationships between species richness and habitat size and amount. Because coastal meadows host species with a vast array of traits, we expected responses to vary between species groups. Location: Swedish Baltic coast. Methods: We inventoried the presence of vascular plant species in twenty-eight 1-m2 plots placed along edaphically defined transects in fifteen coastal meadows. We determined the richness of three species groups: all species, halophytes and inland grassland specialists. We then mapped the habitat for coastal grassland plants using GIS overlay analysis. Using this habitat map, we calculated two variables: “habitat size” and “habitat amount”. We tested correlations between species richness measures and habitat variables, as well as determining the distribution of species traits within meadows. Results: We recorded 174 plant species, of which 6 were halophytes and 35 were inland grassland specialists. Species traits coincided with edaphic sea-to-land gradients. Halophyte and inland grassland specialist richness were significantly correlated with both habitat variables (r = 0.52–0.71). No correlations were found with total species richness. Our habitat map showed that there are 8,900 ha of managed Baltic coastal meadow left in Sweden, mostly in the south. Conclusions: Species traits and distribution play a major role in determining persistence in the face of habitat loss and environmental change. This is especially true for some halophyte populations, which are more susceptible to habitat size and isolation because of their specialisation. Furthermore, they risk being squeezed between the dual threats of encroaching succession and sea-level rise. Preventing habitat loss, restoring meadows and increasing connectivity is crucial for the persistence of specialist plant species.

Understanding the establishment of plant species is important to inform management of restored grasslands and to preserve biodiversity in ancient grasslands. In grassland communities, plant species can establish from seeds arriving via spatial dispersal, from seeds in the soil seed bank or through vegetative spread from nearby source individuals. However, this colonization potential and the likelihood of species establishment can vary in grasslands with different land-use history. We investigated the relative importance of local species recruitment sources, such as dispersal in space and time and species presence in adjacent grasslands, in determining establishment of plant species in eight grasslands with different land-use history (paired ancient grasslands continuously managed as pasture vs. restored grasslands on former forest). At each grassland, we established plots (0.25 m2) to monitor seedling emergence from seed dispersal, seed bank, and recorded clonal growth over two growing periods. We found that the likelihood of species establishment was highest from local seed rain, and that species present in the local species pool were more able to germinate and establish in both type of grasslands. Species from the seed bank and clonal growth contributed to a lesser extent to species establishment, but represented a greater proportion of the recolonization and regeneration of species in ancient grasslands. These results demonstrate that surrounding grasslands serve as a source for colonizing species and that dispersal from the adjacent grasslands is the key process in regeneration and colonization of plants. These results imply that the recovery of grasslands depends heavily upon to links to species source in grasslands, especially in restored grasslands. Therefore, management plans should incorporate rotational livestock grazing and larger networks of grassland in restoration efforts, which will enable to desirable species to establish and persist in grasslands.

Ancient semi-natural grasslands in Europe are important for ecosystem service (ES) provision. Often, the surrounding matrix contains ‘Grassland Green Infrastructure’ (GGI) that contain grassland species which have the potential to supplement grassland ES provision across the landscape. Here we investigate the potential for GGI to deliver a set of complementary ES, driven by plant composition.We surveyed 36 landscapes across three European countries comprising core grasslands and their surrounding GGI. We calculated community-level values of plant species characteristics to provide indicators for four ES: nature conservation value, pollination, carbon storage and aesthetic appeal.Inferred ES delivery for GGI was substantially lower than in core grasslands for conservation, pollination and aesthetic appeal indicators, but not for carbon storage. These differences were driven by the GGI having 17% fewer plant species, and compositional differences, with 61% of species unique to the core grasslands. In addition, connectivity to the core, the amount of GGI and inferred seed dispersal distances by livestock, were strongly positively correlated with conservation value, pollination and aesthetic indicators. All ES indicators showed similar responses to the GGI spatial structure and distance to the core, suggesting robust effects of these drivers on ES. We projected that improved landscape-wide delivery of nature conservation value and pollination could be achieved through targeted GGI management. Reductions in the distances seeds would need to disperse, more GGI, along with a diversification of the GGI elements, were predicted to enhance service credits.We conclude that for vegetation-related ES, species surveys can be employed to assess potential ES delivery. Creating and enhancing GGI is a useful landscape management strategy to supplement the ES delivered by ancient grasslands.

Background: Winters are expected to warm more than summers in central and northern Europe, with largely unknown effects on grassland plant communities.

Aims: By studying the interactions between winter warming and summer grazing, we aimed to disentangle their effects and give recommendations for future grassland management.

Methods: Our study area Upper Teesdale, England has winter temperatures close to 0°C and a well-studied vegetation, known for its arctic-alpine species growing at their climatic warm range limits. We set up a winter warming experiment using open top chambers (ca. +0.5°C) from mid-September until mid-May 2019 to 2022 and excluded sheep grazing during summer in a fully factorial design.

Results: Graminoid biomass increased, and bryophyte biomass decreased with winter warming. There was little to no evidence that winter warming affected any of the other plant response variables we measured, neither did grazing nor the interaction between winter warming and grazing.

Conclusions: Our experiment was relatively short in duration and treatments were realistic in magnitude, therefore the plant communities responded only slightly. Nevertheless, our data suggest a change towards more dominant vascular species and less bryophytes with winter warming, which might lead to lasting changes in the plant communities in the longer-term if not buffered by suitable grazing management.

1. One of the main reasons why insect pollinators are declining is a lack of floral resources. In agricultural landscapes, remaining seminatural grasslands play a key role for providing such resources. However, droughts pose an increasing threat to the abundance and continuity of flowers. Soil amendments are a novel management tool for Swedish grasslands aiming to increase carbon sequestration and soil water holding capacity. In this study, we examined how drought is affecting floral resources (i.e. floral units, nectar quantity and nectar continuity) in grasslands with different mowing regimes, and if soil amendments could mitigate potential negative drought effects.
2. In summer 2019, we set up an experiment combining rain-out shelters (‘drought’), soil amendments (‘compost’) and different mowing regimes (‘mown’ vs. ‘abandoned’) in four extensively managed Swedish grasslands (48 plots, size 2 m²). Between May and August 2021, we counted the floral units nine times in each plot. We derived values for the nectar sugar production per floral unit from an existing database.
3. We observed a decrease in floral units under drought in the mown, but not in the abandoned plots. Nectar quantity and continuity over the season were not significantly affected by drought across both mowing regimes—in the abandoned plots the nectar provision even extended slightly in duration (towards late summer). The compost treatment had positive effects on the floral units, nectar quantity and continuity (extending it towards early summer) in the mown, but not in the abandoned plots. The plant species in our study reacted differently to the treatments. Most of the nectar was provided by only few species (mainly Lathyrus pratensis, Vicia cracca and Anthriscus sylvestris).
4. The results are species specific, thus other plant communities might respond differently. However, our experiment shows that nectar provision (based on database values) in grasslands with a native plant community and natural soil conditions remains relatively stable under drought. We also found that soil amendments increase floral resources in managed grasslands.