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European dry thin-soil calcareous grasslands (alvars) are species-rich semi-natural habitats. Cessation of traditional management, such as mowing and grazing, leads to shrub and tree encroachment and the local extinction of characteristic alvar species. While soil microbes are known to play a critical role in driving vegetation and ecosystem dynamics, more information is needed about their composition and function in grasslands of different dynamic stages. Here we assess the composition of soil fungal, prokaryotic, and plant communities using soil environmental DNA from restored alvar grasslands in Estonia. The study areas included grasslands that had experienced different degrees of woody encroachment prior to restoration (woody plant removal and grazing), as well as unmanaged open grasslands. We found that, in general, different taxonomic groups exhibited correlated patterns of between-community variation. Previous forest sites, which had prior to restoration experienced a high degree of woody encroachment by ectomycorrhizal Scots pine, were compositionally most distinct from managed open grasslands, which had little woody vegetation even prior to restoration. The functional structure of plant and fungal communities varied in ways that were consistent with the representation of mycorrhizal types in the ecosystems prior to restoration. Compositional differences between managed and unmanaged open grasslands reflecting the implementation of grazing without further management interventions were clearer among fungal, and to an extent prokaryotic, communities than among plant communities. While previous studies have shown that during woody encroachment of alvar grassland, plant communities change first and fungal communities follow, our DNA-based results suggest that microbial communities reacted faster than plant communities during the restoration of grazing management in alvar grassland. We conclude that while the plant community responds faster to cessation of management, the fungal community responds faster to restoration of management. This may indicate hysteresis, where the eventual pathway back to the original state (grazed ecosystem) differs from the pathway taken towards the alternative state (abandoned semi-natural grassland ecosystem). [ABSTRACT FROM AUTHOR]

The mechanisms driving nonrandom assembly patterns in plant communities have long been of interest in ecological research. Competing ecological theories predict that coexisting species may either be more functionally dissimilar than expected by chance (with functional 'divergence' mainly reflecting niche differentiation) or be functionally more similar than expected (with functional 'convergence' reflecting either the outcome of environmental filtering or weaker-competitor exclusion effects). Assembly patterns are usually assessed at a single scale and disturbance regime, whereas considering different spatial scales and disturbance regimes may clarify the underlying assembly mechanisms., We tested the prediction that convergence and divergence are scale- and disturbance- dependent in grazed and abandoned species-rich dry grasslands within a 22 km2 landscape in south-eastern Sweden. Convergence and divergence were tested for plant species' traits and phylogenetic relationships at three nested spatial scales: within 412 plots (50 × 50 cm, divided into 10 × 10 cm subplots), within 117 grassland patches (from 0.02 to 11.63 ha) and within the whole landscape (across patches)., At the finest scale (10 × 10 cm subplots within plots), coexisting species were more different than expected by chance (divergence), both functionally and phylogenetically, suggesting niche differentiation. At the intermediate scale (50 × 50 cm plots within patches), coexisting species showed convergence, suggesting environmental filtering. No significant deviations from random expectations were detected at the broadest scale (patches within the 22 km2 landscape) - suggesting the prevalence of dispersal limitation at this scale. The fact that nonrandom patterns were particularly evident under grazed conditions is consistent with the prediction that assembly patterns are disturbance dependent., Synthesis. This study shows that multiple trait-based assembly processes operate simultaneously in species-rich communities, across spatial scales and disturbance regimes. The results support earlier theoretical predictions that divergence between coexisting species may be an important driver of community assembly, particularly at finer spatial scales, where species compete for the same local resources. In contrast, environmental filtering is expected at broader spatial scales, where species growing in particular environmental conditions share traits that are adaptive under those conditions. Within given habitat types, dispersal limitation may, however, override environmental filtering at increasing spatial scales of observation. [ABSTRACT FROM AUTHOR]