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1 The species composition of fragmented semi-natural grasslands may change over time due to stochastic local extinction and colonization events, successional change and/or as a response to changing management or abiotic conditions. The resistance of vegetation to change may be mediated through the effects of topography (slope and aspect) on soils and microclimate. 2 To assess long-term vegetation change in British chalk grasslands, 92 plots first surveyed by F. H. Perring in 1952-53, and distributed across four climatic regions, were re-surveyed during 2001-03. Changes in vegetation since the original survey were assessed by comparing local colonization and extinction rates at the plot scale, and changes in species frequency at the subplot scale. Vegetation change was quantified using indirect ordination (Detrended Correspondence Analysis; DCA) and Ellenberg indicator values. 3 Across all four regions, there was a significant decrease in species number and a marked decline in stress-tolerant species typical of species-rich calcareous grasslands, both in terms of decreased plot occupancy and decreased frequency within occupied plots. More competitive species typical of mesotrophic grasslands had colonized plots they had not previously occupied, but had not increased significantly in frequency within occupied plots. 4 A significant increase in Ellenberg fertility values, which was highly correlated with the first DCA axis, was found across all regions. The magnitude of change of fertility and moisture values was found to decrease with angle of slope and with a topographic solar radiation index derived from slope and aspect. 5 The observed shift from calcareous grassland towards more mesotrophic grassland communities is consistent with the predicted effects of both habitat fragmentation and nutrient enrichment. It is hypothesized that chalk grassland swards on steeply sloping ground are more resistant to invasion by competitive grass species than those on flatter sites due to phosphorus limitation in shallow minerogenic rendzina soils, and that those with a southerly aspect are more resistant due to increased magnitude and frequency of drought events.

QUESTIONS: What is the time‐scale for natural regeneration of calcareous grassland? Is this time‐scale the same for individual plant species, plant community composition and functional traits? LOCATION: Defence Training Estate Salisbury Plain, Wiltshire, UK. METHODS: We investigated the rate of natural regeneration of species‐rich calcareous grassland across a 20 000‐ha landscape. We combined a large‐scale botanical survey with historic land‐use data (6–150 yrs before present) and examined differences between grassland age classes in the occurrence of individual plant species, plant community composition and plant community functional traits. RESULTS: Many species showed a significant association with grasslands over 100 yrs old. These included the majority of those defined elsewhere as calcareous grassland indicators, although some appeared on grasslands <10 yrs old. Plant community composition showed increasing similarity to the oldest grasslands with increased grassland age, with the exception of very recently ex‐agricultural grasslands. Most functional traits showed clear trends with grassland age, with dispersal ability differing most strongly between recent and older grasslands, whilst soil fertility and pH tolerance were more influential over longer time‐scales. CONCLUSIONS: Even in a well connected landscape, natural regeneration to a community resembling ancient calcareous grassland in terms of functional traits and plant community composition takes over a century, although changes at the level of individual species may occur much earlier. These findings confirm the uniqueness of ancient calcareous grassland. They also suggest that the targets of re‐establishment efforts should be adjusted to account for the likely time‐scale of full community re‐assembly.

Although it is well known that arbuscular mycorrhizal fungi (AMF) play a key role in the functioning of natural ecosystems, the underlying drivers determining the composition of AMF communities remain unclear. In this study, we established 138 sampling plots at 46 grassland sites, consisting of 26 acidic grasslands and 20 calcareous grasslands spread across eight European countries, to assess the relative importance of abiotic and biotic filtering in driving AMF community composition and structure in both the grassland soils and in the roots of 13 grassland plant species. Soil AMF communities differed significantly between acidic and calcareous grasslands. In root AMF communities, most variance was attributable to soil variables while very little variation was explained by host plant identity. Root AMF communities in host plant species occurring in only one grassland type closely resembled the soil AMF communities of that grassland type and the root AMF communities of other host plant species occurring in the same grassland type. The observed AMF–host plants networks were not modular but nested. Our results indicate that abiotic conditions, rather than biotic filtering through host plant specificity, are the most important drivers in shaping AMF communities in European semi-natural grasslands.

Question: What are the long-term implications of former fertilisation for the ecological restoration of calcareous grasslands? Location: Gerendal, Limburg, The Netherlands. Methods: In 1970, ten permanent plots were established in just abandoned agricultural calcareous grassland under a regime of annual mowing in August. From 1971 to 1979, two different fertiliser treatments were applied twice a year to a subset of the plots (artificial fertiliser with different proportions of nitrogen and phosphorus). The vegetation of the plots was recorded yearly and vegetation biomass samples were taken for peak standing crop and total amounts of nitrogen, phosphorus and potassium. Species composition and floristic diversity were analysed over the research period (1970–2006) and between the treatments, including the use of multivariate techniques (Detrended Correspondence Analysis). Results: In terms of species number, there is a clear optimum 10 to 20 years after fertilisation has been terminated. Afterwards, there is a slow decrease; no new species appear and species of more nutrient-rich conditions gradually disappear. For the fertilised plots that received a relatively high proportion of N, effects are found only in the first years, whereas, for the plots that received a relatively high proportion of P, long-term after-effects are found in species composition, peak standing crop, total amounts of phosphorus in biomass, and in soil phosphorus data. Conclusions: The effect of artificial fertiliser with a large amount of nitrogen disappears in less than ten years when mown in August, including removal of the hay. This is a promising result for restoration of N-enriched calcareous grasslands, as the applied dose of nitrogen in this experiment largely exceeds the extra input of nitrogen via atmospheric deposition. Application of fertiliser with a large amount of phosphorus, however, has effects even more than 25 years after the last addition. There are no prospects that this effect will become reduced in the near future under the current mowing management. Nomenclature: van der Meijden (2005) for vascular plant species and Schamineé et al. (1996) for plant communities.

The use of hyperspectral (HS) and LiDAR acquisitions has a great potential to enhance mapping and monitoring practices of endangered grasslands habitats, beyond conventional botanical field surveys. In this study we assess the potentiality of recursive feature elimination (RFE) in combination with random forest (RF) classification in extracting the main HS and LiDAR features needed to map selected Natura 2000 grasslands along Polish lowland river valleys, in particular alluvial meadows 6440, lowland hay meadows 6510, and xeric and calcareous grasslands 6120. We developed an automated RFE-RF system capable to combine the potentials of both techniques and applied it to multiple acquisitions. Several LiDAR-based products and different spectral indices (SI) were computed and used as input in the system, with the aim of shedding light on the best-to-use features. Results showed a remarkable increase in classification accuracy when LiDAR and SI products are added to the HS dataset, strengthening in particular the importance of employing LiDAR in combination with HS. Using only the 24 optimal features selection generalized over the three study areas, strongly linked to the highly heterogeneous characteristics of the habitats and landscapes investigated, it was possible to achieve rather high classification results (K around 0.7–0.77 and habitats F1 accuracy around 0.8–0.85), indicating that the selected Natura 2000 meadows and dry grasslands habitats can be automatically mapped by airborne HS and LiDAR data. Similar approaches might be considered for future monitoring activities in the context of habitats protection and conservation.

The risk of ecosystem function degradation with biodiversity loss has emerged as a major scientific concern in recent years. Possible relationships between taxonomic diversity and magnitude and stability of ecosystem processes build upon species’ functional characteristics, which determine both susceptibility to environmental change and contribution to ecosystem properties. The functional diversity within communities thus provides a potential buffer against environmental disturbance, especially for properties emerging from interactions among species. In complex plant–pollinator networks, distantly related taxa spanning a great trait diversity shape network architecture. Here, we address the question of whether network properties are maintained after habitat loss by complementary contributions of phylogenetically distant pollinator taxa. We quantified contributions of wild bees and hoverflies to network structure (connectance, network specialization, specialization asymmetry) in 32 calcareous grassland fragments varying in size. Although hoverflies are often regarded less susceptible to environmental change than wild bees, species richness of both taxa was similarly affected by habitat loss. The associated loss of 80% of interactions resulted in small and tightly connected networks, which was more strongly attributed to wild bee loss than hoverfly loss. Networks in small fragments were less specialized due to equivalent losses of species and interactions in both pollinators and plants. Because wild bee and hoverfly loss contributed similarly to declining network specialization, we conclude that trait diversity among distantly related pollinators does not necessarily provide insurance against functional homogenization during community disassembly following habitat loss.

Rapid environmental changes across Europe include warmer and increasingly variable temperatures, changes in soil nutrient availability, and pollinator decline. These abiotic and biotic changes can affect natural plant populations and force them to optimize resource use against competitors. To date, the evolution of competitive ability in the context of changes in nutrient availability remains understudied. In this study, we investigated whether the common calcareous grassland herb Leontodon hispidus recently evolved its competitive ability and response to nutrient availability. We compared ancestors sampled in 1995 and descendants sampled in 2018 and applied a competition treatment in combination with weekly nutrient treatments (no fertilizer, nitrogen, phosphorus, and both). We found evidence for evolution of increased competitive ability, with descendants producing more vegetative biomass than ancestors when grown under competition. Furthermore, supplementing nutrients (especially N) reduced differences in competitive ability between ancestors and descendants, suggesting that nutrients are a limiting factor in interspecific competition, which could be linked to the decreasing nitrogen emissions into the atmosphere since the 1990s. Our study demonstrates rapid contemporary evolution of competitive ability, but also the complexity of the underlying processes of contemporary evolution, and sheds light on the importance of understudied potential selection agents such as nutrient availability.

Strong declines of grassland species diversity in small and isolated grassland patches have been observed at local and landscape scales. Here, we study how plant–herbivore interaction webs and habitat specialisation of leafhopper communities change with the size of calcareous grassland fragments and landscape connectivity. We surveyed leafhoppers and plants on 14 small (0.1–0.6 ha) and 14 large (1.2–8.8 ha) semi-natural calcareous grassland fragments in Central Germany, differing in isolation from other calcareous grasslands and in the percentage of arable land in the surrounding landscape (from simple to complex landscapes). We quantified weighted trophic links between plants and their phytophagous leafhoppers for each grassland fragment. We found that large and well-connected grassland fragments harboured a high portion of specialist leafhopper species, which in turn yielded low interaction diversity and simple plant-leafhopper food webs. In contrast, small and well-connected fragments exhibited high levels of generalism, leading to higher interaction diversity. In conclusion, food web complexity appeared to be a poor indicator for the management of insect diversity, as it is driven by specialist species, which require high connectivity of large fragments in complex landscapes. We conclude that habitat specialists should be prioritized since generalist species associated with small fragments are also widespread in the surrounding landscape matrix.

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species–area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data‐intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under‐reported.

Elevated CO 2 (eCO 2 ) can stimulate plant productivity and increase carbon (C) input to soils, but nutrient limitation restricts productivity. Despite phosphorus (P)-limited ecosystems increasing globally, it is unknown how nutrient cycling, particularly soil microbial extra cellular enzyme activity (EEA), will respond to eCO 2 in such ecosystems. Long-term nutrient manipulation plots from adjacent P-limited acidic and limestone grasslands were exposed to eCO 2 (600 ppm) provided by a mini-Free Air CO 2 Enrichment system. P-limitation was alleviated (35 kg-P ha −1 y −1 (P35)), exacerbated (35 kg-N ha −1 y −1 (N35), 140 kg-N ha −1 y −1 (N140)), or maintained (control (P0N0)) for > 20 years. We measured EEAs of C-, N- and P-cycling enzymes (1,4-β-glucosidase, cellobiohydrolase, N-acetyl β-D-glucosaminidase, leucine aminopeptidase, and acid phosphatase) and compared C:N:P cycling enzyme ratios using a vector analysis. Potential acid phosphatase activity doubled under N additions relative to P0N0 and P35 treatments. Vector analysis revealed reduced C-cycling investment and increased P-cycling investment under eCO 2 . Vector angle significantly increased with P-limitation (P35 < P0N0 < N35 < N140) indicating relatively greater investment in P-cycling enzymes. The limestone grassland was more C limited than the acidic grassland, characterised by increased vector length, C:N and C:P enzyme ratios. The absence of interactions between grassland type and eCO 2 or nutrient treatment for all enzyme indicators signaled consistent responses to changing P-limitation and eCO 2 in both grasslands. Our findings suggest that eCO 2 reduces C limitation, allowing increased investment in P- and N-cycle enzymes with implications for rates of nutrient cycling, potentially alleviating nutrient limitation of ecosystem productivity under eCO 2 . Graphic abstract