Questions: (1) Does soil acidification in calcareous sandy grasslands lead to loss of plant diversity? (2) What is the relationship between the soil content of lime and the plant availability of mineral nitrogen (N) and phosphorus (P) in sandy grasslands?Location: Sandy glaciofluvial deposits in south-eastern Sweden covered by xeric sand calcareous grasslands (EU habitat directive 6120).Methods: Soil and vegetation were investigated in most of the xeric sand calcareous grasslands in the Scania region (136 sample plots distributed over four or five major areas and about 25 different sites). Environmental variables were recorded at each plot, and soil samples were analysed for exchangeable P and N, as well as limestone content and pH. Data were analysed with regression analysis and canonical correspondence analysis.Results: Plant species richness was highest on weakly acid to slightly alkaline soil; a number of nationally red-listed species showed a similar pattern. Plant species diversity and number of red-listed species increased with slope. Where the topsoil had been acidified, limestone was rarely present above a depth of 30 cm. The presence of limestone restricts the availability of soil P, placing a major constraint on primary productivity in sandy soils.Conclusions: Acidification of sandy grasslands leads to reduced abundance of desirable species, although the overall effect is rather weak between pH 5 and pH 9. Slopes are important for high diversity in sandy grasslands. Calcareous soils cannot be restored through shallow ploughing, but deep perturbation could increase the limestone content of the topsoil and favour of target species.

Drought is a great challenge to agricultural production, and cultivation of drought-tolerant or water use-efficient cultivars is important to ensure high biomass yields for bio-refining and bioenergy. Here, we evaluated drought tolerance of four C 3 species, Dactylis glomerata cvs. Sevenop and Amba, Festuca arundinacea cvs. Jordane and Kora, Phalaris arundinacea cvs. Bamse and Chieftain and Festulolium pabulare cv. Hykor, and two C 4 species Miscanthus 9 giganteus and M. lutarioriparius. Control (irrigated) and drought-treated plants were grown on coarse and loamy sand in 1 m 2 lysimeter plots where rain was excluded. Drought periods started after harvest and lasted until 80% of available soil water had been used. Drought caused a decrease in dry matter yield (DM; P < 0.001) for all species and cultivars during the drought period. Cultivars Sevenop, Kora and Jordane produced DM at equal levels and higher than the other C 3 cultivars in control and drought-treated plots both during and after the drought period. Negative correlations were observed between stomatal conductance (g s ) and leaf water potential (P < 0.01) and positive correlations between g s and DM (P < 0.05) indicating that g s might be suitable for assessment of drought stress. There were indications of positive associations between plants carbon isotope composition and water use efficiency (WUE) as well as DM under well-watered conditions. Compared to control, drought-treated plots showed increased growth in the period after drought stress. Thus, the drought events did not affect total biomass production (DM total ) of the whole growing season. During drought stress and the whole growing season, WUE was higher in drought-treated compared to control plots, so it seems possible to save water without loss of biomass. Across soil types, M. lutarioriparius had the highest DM total (15.0 t ha À1 ), WUE total (3.6 g L À1 ) and radiation use efficiency (2.3 g MJ À1) of the evaluated grasses.

The species-rich calcareous grassland communities in Europe are gradually disappearing due to lack of management such as grazing or cultivation, resulting in decalcification and reduction of gaps in the vegetation. In this study, experimental soil perturbation (deep and shallow) was performed in degenerated sandy grassland in plots with a size of 8 9 8 m, using a randomised block design. The hypothesis that soil perturbation that inverts the soil layers decreases nutrient availability, creates vegetation gaps and thereby selects for desirable species was tested through comparisons with untreated controls as well as with nearby target habitats. The deep perturbation was designed to bring CaCO 3 up to the surface, whereas the shallow perturbation tested the effect of disturbance alone. The effects of soil perturbation on soil chemistry, vegetation and beetle communities were analysed for the 2 years following the treatments. Increased pH and calcium concentration, and decreased nitrogen and phosphorus availability, showed that deep perturbation was successful in restoring the soil chemistry to levels similar to those of the target habitat. Perturbated plots were rapidly colonised by the acid tolerant grass Corynephorus canescens, but the slow colonisation of the threatened calcicole species Koeleria glauca was an indication that the vegetation could be evolving towards the target vegetation. Six red-listed beetle species associated with open, dry grasslands were found, out of which four were found only in perturbated plots, although this could not be statistically tested. In conclusion, it may take many years or even decades for the establishment of desirable flora, and seeding could therefore be a suitable method of increasing the rate of succession.

Acidification and N-deposition are continuous processes that alter the composition of plant communities. We investigated vegetation transitions in sandy grasslands and tested two hypotheses: (1) a shift from vegetation dominated by Koeleria glauca to one dominated by Corynephorus canescens is due to acidification and (2) a shift from vegetation dominated by K. glauca to one dominated by Arrhenatherum elatius is due to increased mineral-N. The occurrence of K. glauca and C. canescens followed pH shifts strikingly well. However, the pH varied considerably between sites, and we found that differences in preference for bare sand may be important for the coexistence of these two species at both high and low pH. In 75% of the gradients, the dominance of A. elatius was related to high content of mineral-N. Most species preferred lime-rich soils and we concluded that both N-enrichment and acidification may lead to loss of species in calcareous sandy grasslands.

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We have studied how disturbance by ploughing and rotavation affects the carbon (C) flow to arbuscular mycorrhizal (AM) fungi in a dry, semi-natural grassland. AM fungal biomass was estimated using the indicator neutral lipid fatty acid (NLFA) 16:1ω5, and saprotrophic fungal biomass using NLFA 18:2ω6,9. We labeled vegetation plots with (13)CO(2) and studied the C flow to the signature fatty acids as well as uptake and allocation in plants. We found that AM fungal biomass in roots and soil decreased with disturbance, while saprotrophic fungal biomass in soil was not influenced by disturbance. Rotavation decreased the (13)C enrichment in NLFA 16:1ω5 in soil, but (13)C enrichment in the AM fungal indicator NLFA 16:1ω5 in roots or soil was not influenced by any other disturbance. In roots, (13)C enrichment was consistently higher in NLFA 16:1ω5 than in crude root material. Grasses (mainly Festuca brevipila) decreased as a result of disturbance, while non-mycorrhizal annual forbs increased. This decreases the potential for mycorrhizal C sequestration and may have been the main reason for the reduced mycorrhizal C allocation found in disturbed plots. Disturbance decreased the soil ammonium content but did not change the pH, nitrate or phosphate availability. The overall effect of disturbance on C allocation was that more of the C in AM fungal mycelium was directed to the external phase. Furthermore, the functional identity of the plants seemed to play a minor role in the C cycle as no differences were seen between different groups, although annuals contained less AM fungi than the other groups.

The aim of this study was to ascertain whether there is a relationship between plant species richness and plant-available N, P and water in an environment subject to little anthropogenic disturbance. To accomplish this we studied the vegetation in matorral shrub-lands in northern Patagonia, Argentina. Due to the variation in slope, precipitation and aspect between the sites water status was determined using the 12 C/ 13 C fraction, δ 13 C, to investigate whether this was a confounding factor. The numbers of herb, shrub, liana and tree species were determined at 20 sites along an estimated precipitation gradient. Leaf P and N content and the δ 13 C of Berberis buxifolia were determined, as well as the soil P and N content at the different sites. A negative correlation was found between species richness and Berberis buxifolia foliar P concentration (52% of the species richness variation was accounted for), and a positive correlation was found between plant species richness and Berberis buxifolia foliar N: P ratios (54% of the species richness variation was accounted for). The relationship between species richness and foliar P was seen when all layers of vegetation were included (trees, lianas, shrubs and herbs). Foliar N showed no correlation with species richness, while soil extractable NH 4 showed a weak positive correlation with the number of shrub layer species (lianas, shrubs and trees). The species richness of the shrub layer increased with decreasing values of δ 13 C. Low soil P availability thus affects local species richness in the matorral shrub-lands of Patagonia in Argentina although the growth of vegetation in the area has been shown to be limited by N. We suggest that low P levels increase plant species richness because low soil P concentration is associated with a high P partitioning and high potential for niche separation.