How do management and restoration needs of mountain grasslands depend on moisture regime? Experimental study from north‐western Slovakia (Western Carpathians)

Question: How does species composition change in traditionally managed meadows after mowing has ceased, and in abandoned meadows after re‐introduction of mowing? Are there differences in the dynamics of dry and moderately wet meadows? Location: Zázrivá‐Ple?ivá (19°11′N, 49°16′E), north‐western Slovakia, western Carpathians. Methods: Pairs of experimental plots (mown and unmown) were established to replicate each combination of dry/wet and traditionally managed/abandoned meadows. Changes in species composition were studied over 5 years. The data on changes in species composition was analysed by constrained and unconstrained ordinations, and visualized using Principal Response Curves. Results: Species composition of newly abandoned wet grasslands was changing towards the corresponding long‐abandoned plots even in the first year of abandonment. Similarly, newly established restoration mowing in abandoned dry grasslands rapidly shifted the stand species composition towards that of traditionally managed ones. Nevertheless, 4 year after reintroduction of mowing, the species composition of the restored plots was still far from the target composition. The effect of mowing in abandoned wet grasslands and abandonment in dry grasslands was much less pronounced and slower. Conclusions: Moisture regime is a very important factor determining the management needs of various grassland types. Wet grasslands are much more sensitive to abandonment, with a rapid degradation rate and limited possibilities for restoration, which can be extremely slow. Even in the dry grasslands, that quickly responded to restoration mowing, restoration is a long‐term process.     

Evaluating alternative mowing regimes for conservation management of Central European mesic hay meadows: A field experiment

Conservation status of hay meadows highly depends on their management. The main goal of this study was to assess the efficiency of different mowing regimes in maintenance of plant species richness and diversity of mesic hay meadows. The field experiment was carried out on a species rich, mesic hay meadow in Western Hungary. We evaluated the effects of four alternative types of management on the plant community after 7 years of continuous treatment: (1) mowing twice a year, typical traditional management, (2) mowing once a year in May, most practised currently by local farmers, (3) mowing once a year in September, often proposed for conservation management and (4) abandonment of mowing. Traditional mowing resulted in significantly higher number and higher diversity of vascular plant species than other mowing regimes. Mowing twice a year was the only efficient way to control the spread of the invasive Solidago gigantea, and mowing in September was more successful in it than mowing in May. We conclude that the traditional mowing regime is the most suitable to maintain botanical diversity of mesic hay meadows; however, other regimes should also be considered if certain priority species are targeted by conservation.     

The influence of environmental factors and management methods on the vegetation of mesic grasslands in a central European mountain range

We examined how environmental factors combined with the diverse ways of the extensive management of mesic mountain grasslands affect species composition and diversity in the Polish part of Central Sudetes Mts. Based on the data from 100 research plots altitude, organic matter content and exchangeable magnesium in the soil, as well as the maximum soil water capacity and amount of sand fraction significantly affected species composition. Among the management methods, a significant effect on the differentiation of species composition was observed following the cessation of usage and mowing. Mown meadows had the highest share of forbs in the biomass, whereas those abandoned ones were dominated by tall and expansive grasses, mainly Calamagrostis epigejos. The species richest grasslands occurred in areas with soils of high water capacity, containing high concentrations of calcium and low amounts of total nitrogen. No significant effect of management methods on the total number of species as well as on the number of forb species was observed. Mown pastures had the highest mean value of the Shannon–Wiener diversity index.     

Influence of abandonment on syrphid assemblages in mountainous meadows

Grasslands are often managed with different intensities in the European Alps. Studies have shown that a medium management can benefit plant and animal biodiversity in these ecosystems. However, in recent decades abandonment of extensively managed meadows is an ongoing global challenge. Syrphids (hoverflies) have been recognized as a threatened group due to environmental drivers, and these managed grasslands provide preferred habitats for them. Yet, at what extent syrphids respond to grassland management strategies is little known. We investigated whether abandonment of managed mountain meadows (mown once a year, no fertilizer use) affects syrphid abundance, richness and species assemblages. Moreover, the influence of plant richness, flower frequency and surrounding landscape on syrphids were assessed. Four managed meadows were compared with four abandoned meadows in a mountainous region in Austria. Adult syrphids were sampled over two consecutive years (June and August 2015–2016) by sweep netting using line transects and observation plots. Syrphid abundance was significantly higher in managed than abandoned meadows and increased with increasing plant richness and flower frequency across management types. Management and sampling time also significantly affected syrphid species assemblages. Syrphid richness and abundance were not influenced by surrounding landscape. We conclude that both abandoned and managed meadows provide important habitat types for syrphids. Therefore, efforts should be made to maintain both managed and abandoned meadows in order to protect syrphid species within mountainous landscapes.     

Agri-environment incentive payments and plant species richness under different management intensities in mountain meadows of Switzerland

We investigated whether agri-environmental incentive payments help to maintain biodiversity. We studied the effect of agricultural management intensity on vascular plant species richness and plant assemblages of mountain meadows in Switzerland. Other factors such as slope, altitude or accessibility (distance from farmyard) were also taken into account. Vegetation sampling was conducted at 69 sites representing five different management types, differing with respect to nutrient input and soil moisture: (i) dry extensive meadows; (ii) extensive meadows; (iii) dry low-intensive meadows; (iv) low-intensive meadows; (v) intensive meadows. There was a significant negative relationship between plant species richness and management intensity: The mean number of plant species per management type declined markedly when management intensity increased, although dry sites harboured slightly more species regardless of management intensity (dry extensive > dry low intensive > extensive > low intensive >> intensive meadows). Species richness was clearly affected by management intensity, but not so by slope, altitude or accessibility. There was a gradual shift in plant assemblages among management types with only intensive meadows differing from the other four types of differently managed meadows. We therefore found, in contrast to many studies done in the European lowlands, positive effects of incentive payments on plant species richness.     

Modelling biomass of mountainous grasslands by including a species composition map

High mountain grasslands offer multiple goods and services to society but are severely threatened by improper land use practices such as abandonment or rapid intensification. In order to reduce abandonment and strengthen the common extensive agricultural practice a sustainable land use management of high mountain grasslands is needed. A spatially detailed yield assessment helps to identify possible meadows or, on the contrary, areas with a low carrying capacity in a region, making it easier to manage these sites. Such assessments are rarely available for remote and inaccessible areas. Remotely sensed vegetation indices are able to provide valuable information on grassland properties. These indices tend, however, to saturate for high biomass. This affects their applicability to assessments of high-yield grasslands.The main aim of this study was to model a spatially explicit grassland yield map and to test whether saturation issues can be tackled by consideration of plant species composition in the modelling process. The high mountain grassland of the subalpine belt (1800 – 2500 m a.s.l.) in the Kazbegi region, Greater Caucasus, Georgia, was chosen as test site for its strong species composition and yield gradients.We first modelled the species composition of the grassland described as metrically scaled gradients in the form of ordination axes by random forest regression. We then derived vegetation indices from Rapid Eye imagery, and topographic variables from a digital elevation model, which we used together with the multispectral bands as predictive variables. For comparison, we performed two yield models, one excluding the species composition maps and one including the species composition map as predictors. Moreover, we performed a third individual model, with species composition as predictors and a split dataset, to produce the final yield map.Three main grassland types were found in the vegetation analysis: Hordeum violaceum-meadows, Gentianella caucasea-grassland and Astragalus captiosus-grassland. The three random forest regression models for the ordination axes explained 64%, 33% and 46% of the variance in species composition. Independent validation of modelled ordination scores against a validation data set resulted in an R² of 0.64, 0.32 and 0.46 for the first, second and third axes, respectively. The model based on species composition resulted in a R² = 0.55, whereas the benchmark model showed weaker relationships between yield and the multispectral reflectance, vegetation indices, and topographical parameters (R² = 0.42). The final random forest yield model used to derive the yield map resulted in 62% variance explained and an R² = 0.64 between predicted and observed biomass. The results further indicate that high yields are generally difficult to predict with both models.The benefit of including a species composition map as a predictor variable for grassland yield lies in the preservation of ecologically meaningful features, especially the occurrence of high yielding vegetation type of Hordeum violaceum meadows is depicted accurately in the map. Even though we used a gradient based design, sharp boundaries or immediate changes in productivity were visible, especially in small structures such as arable fields or roads (Fig. 6b), making it a valuable tool for sustainable land use management. The saturation effect however, was mitigated by using species composition as predictor variables but is still present at high yields.     

The effects of mowing and fertilization on carbohydrate reserves and regrowth of grasses: do they promote plant coexistence in species-rich meadows?

After the cessation of regular management and after fertilization a single clonal species tends to dominate in many types of grasslands, whereas in regularly managed meadows these potential dominants usually attain a low cover. It has been hypothesized that plants reaching a high dominance in abandoned and fertilized meadows are selectively suppressed by mowing so that a balanced competition is maintained and competitive exclusion is postponed. We compared regeneration capacity and carbohydrate reserves accumulated by three species of clonal grasses, which markedly increase their dominance in irregularly mown, un-mown or fertilized meadows. Above-ground biomass and the amount of storage carbohydrates of the two largest species, (Molinia arundinacea, Calamagrostis epigejos) were reduced in a mown meadow. This effect was weaker in Bromus erectus, which produces smaller shoots. Shoots of Molinia were most impacted by mowing but their regeneration was efficient due to the large carbohydrate reserves in the shoot bases. Fertilization did not affect Bromus and Calamagrostis. In contrast, fertilized plants of Molinia produced larger storage organs and accumulated more carbohydrates. We conclude that plant size and growth form are important features promoting the ability of potential dominants to tolerate mowing and regenerate after it. Our results confirm that taller plants are selectively suppressed by this type of disturbance, thereby potentially promoting plant coexistence.     

Changes of species richness pattern in mountain grasslands: abandonment versus restoration

Changes in plant species richness at various spatial scales were investigated by manipulative experiment in mountain grasslands. The aim of the research was to compare changes in species richness in newly abandoned sites and sites where restoration measures were applied after 20 years of abandonment. The plots were located in two vegetation types with different moisture regime. Species richness decreased significantly after abandonment, mainly at the finest spatial scale of 10 × 10 cm. There was significant increase of species richness on restored sites, but it was apparent mainly at a larger scale. However, even 4 years of regular mowing were not sufficient to restore species richness to the level typical for traditionally managed grasslands in the region. No significant difference was found in the performance of the 2 contrasting vegetation types (wet and dry) in relation to management measures. A significant difference in scale-dependent species richness was only observed. The dry type had a steeper species-area curve, with a lower number of species at the finest spatial scale. According to the results of the experiment, mountain grasslands are very vulnerable habitats, losing their conservation value quickly after abandonment. Restoration is possible due to an extensive species pool in the region, but return to the original species richness at all spatial scales is quite a long process.     

Decline of rare and specialist species across multiple taxonomic groups after grassland intensification and abandonment

Traditionally managed mountain grasslands are declining as a result of abandonment or intensification of management. Based on a common chronosequence approach we investigated species compositions of 16 taxonomic groups on traditionally managed dry pastures, fertilized and irrigated hay meadows, and abandoned grasslands (larch forests). We included faunal above- and below-ground biodiversity as well as species traits (mainly rarity and habitat specificity) in our analyses. The larch forests showed the highest species number (345 species), with slightly less species in pastures (290 species) and much less in hay meadows (163 species). The proportion of rare species was highest in the pastures and lowest in hay meadows. Similar patterns were found for specialist species, i.e. species with a high habitat specificity. After abandonment, larch forests harbor a higher number of pasture species than hay meadows. These overall trends were mainly supported by spiders and vascular plants. Lichens, bryophytes and carabid beetles showed partly contrasting trends. These findings stress the importance to include a wide range of taxonomic groups in conservation studies. All in all, both abandonment and intensification had similar negative impacts on biodiversity in our study, underlining the high conservation value of Inner-Alpine dry pastures.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

Species composition of semi-natural mesic grasslands as a factor influencing the methane yield of plant biomass (Central Europe)

Semi-natural lowland and mountain mesic meadows are grasslands rich in species, and their conservation status depends on treatments such as mowing or grazing livestock. In many countries, the condition of grasslands is deteriorating because of their inappropriate use or abandonment. This study aimed to determine the effects of the species composition of plant communities and functional plant groups on the methane yield from biomass harvested from mesic grasslands in the Sudetes Mountains. Biogas potential analysis was performed based on biomass samples collected from Poland and the Czech Republic. The biogas potential was determined in 40 day-long batch anaerobic digestion tests. The average methane yield obtained from the biomass was 246 ± 16 NL CH₄ kg^?1 VS, whereas the methane yield per hectare was 870 ± 203 m³ CH₄ ha^?1. Plant communities comprising different dominant species had no effect on the methane yield but affected the methane yield per hectare. Additionally, the species composition of grasslands with a higher percentage of forbs had lower biomass yield, resulting in lower methane yields per hectare. The continuity of the low-intensity management of mountain grassland, which can be provided by the utilization of their biomass for bioenergy production, sustains high biodiversity and ensures appropriate meadow conservation.     

Airborne hyperspectral data predict Ellenberg indicator values for nutrient and moisture availability in dry grazed grasslands within a local agricultural landscape

Species-based ecological indices, such as Ellenberg indicators, reflect plant habitat preferences and can be used to describe local environment conditions. One disadvantage of using vegetation data as a substitute for environmental data is the fact that extensive floristic sampling can usually only be carried out at a plot scale within limited geographical areas. Remotely sensed data have the potential to provide information on fine-scale vegetation properties over large areas. In the present study, we examine whether airborne hyperspectral remote sensing can be used to predict Ellenberg nutrient (N) and moisture (M) values in plots in dry grazed grasslands within a local agricultural landscape in southern Sweden. We compare the prediction accuracy of three categories of model: (I) models based on predefined vegetation indices (VIs), (II) models based on waveband-selected VIs, and (III) models based on the full set of hyperspectral wavebands. We also identify the optimal combination of wavebands for the prediction of Ellenberg values. The floristic composition of 104 (4 m × 4 m grassland) plots on the Baltic island of Öland was surveyed in the field, and the vascular plant species recorded in the plots were assigned Ellenberg indicator values for N and M. A community-weighted mean value was calculated for N (mN) and M (mM) within each plot. Hyperspectral data were extracted from an 8 m × 8 m pixel window centred on each plot. The relationship between field-observed and predicted mean Ellenberg values was significant for all three categories of prediction models. The performance of the category II and III models was comparable, and they gave lower prediction errors and higher R² values than the category I models for both mN and mM. Visible and near-infrared wavebands were important for the prediction of both mN and mM, and shortwave infrared wavebands were also important for the prediction of mM. We conclude that airborne hyperspectral remote sensing can detect spectral differences in vegetation between grassland plots characterised by different mean Ellenberg N and M values, and that remote sensing technology can potentially be used to survey fine-scale variation in environmental conditions within a local agricultural landscape.     

The impact of forest encroachment after agricultural land abandonment on passerine bird communities: The case of Greece

Agricultural land abandonment is one of the main drivers of land use change, leading to various responses of farmland ecological communities. In an effort to better understand the effect of agricultural land abandonment on passerine bird communities, we sampled 20 randomly selected sites [1 km × 1 km] in remote Greek mountains, reflecting an abandonment gradient, in terms of forest encroachment. We sampled 169 plots using the point count method of fixed distance (47 passerine species), and we investigated bird diversity and community structure turnover along the gradient. We found that grazing intensity has a beneficial effect hampering forest encroachment that follows progressively land abandonment. Habitat composition changes gradually with forests developing at the expense of open meadows and heterogeneous grasslands. Forest encroachment has a significant negative effect on bird diversity and species richness, affecting in particular typical farmland and Mediterranean shrubland species. Birds form five distinct ecological clusters after land abandonment: species mostly found in pinewoods and cavity-dwelling species; species that prefer open forests forest edges or ecotones; species that prefer shrubland or open habitats with scattered woody vegetation; Mediterranean farmland birds that prefer semi-open habitats with hedges and/or woodlots; and, generalist forest-dwelling or shrubland species. We extracted a set of 22 species to represent the above ecological communities, as a new monitoring tool for agricultural land use change and conservation. We suggest that the maintenance of rural mosaics should be included in the priorities of agricultural policy for farmland bird diversity conservation.     

Avian fossil assemblages at the onset of the LGM in the eastern Alps: A palaecological contribution from the Rio Secco Cave (Italy)

The avian fossil assemblages from the late Pleistocene deposits of the Rio Secco Cave (north-eastern Italy) is presented herein. We studied the layers that date back to the end of MIS3 and the beginning of MIS2, which also contain evidence of Gravettian frequentation dated to 33.5–30 ka cal BP. The systematic analysis revealed the presence of 18 species and other supraspecific taxa that supported palaeoenvironmental reconstructions. Taxa indicate that, at the onset of LGM, site surroundings were characterised by conifer or mixed forests, open grasslands, slow-flowing water bodies and mountain meadows with rocky outcrops, as indicated by Lagopus muta. Today, this environment is found above the tree line (beyond 1500–2000 m) and cannot be detected near the site, located at 580 m asl. Noteworthy, is also the finding of the second Italian late Pleistocene fossil record of Picus canus.     

Plants as indicators of snow layer duration in southern Norwegian mountains

The main aim of this paper was to study the responses of mountain plants in relation to the time of snowmelt. Three mountain areas situated along an oceanic–continental gradient were selected as study sites, and the sample plots ranged from 182 m below to 473 m above the climatic forest limit. In total, 185 quadrats (2 m × 2 m), stratified to include only oligotrophic and mesotrophic mountain vegetation types, were selected to represent a topographic range along altitudinal gradients. In each quadrat, the percentage groundcover of the species was recorded. From the beginning of April until July 2004, snow thickness was monitored, and the Julian day when the snow had completely melted was determined for all plots. The relationship between species abundances and Julian day of snowmelt were analysed by two different numerical methods: (1) relative values for species optimum and tolerance were given by Detrended Canonical Correspondence Analysis (DCCA) with Julian day of snowmelt as the constraining variable. (2) Species responses were modelled by Generalized Linear Models (GLM). For species with significant unimodal responses, optimum and tolerance were calculated. For species with significant linear models, different species response models were identified by the regression intercepts. One hundred and twenty six species (taxa) were tested, and 103 evidenced statistically significant (p < 0.05) distribution responses. Several common alpine plants had a distribution that appeared to be independent of snow. On the basis of the results of the numerical methods, the species were separated into nine Snow Indicator (SI) classes, as a parallel to the Ellenberg indicator values. The species’ SI values were used to calculate weighted average SI values to examine the relationships between previously described plant communities and vegetation transects which experience different snow conditions.     

The significance of habitat continuity and current management on the compositional and functional diversity of grasslands in the uplands of Lower Saxony,Germany

There is a growing concern that land use intensification is having negative effects on semi-natural grasslands and that it leads to a general loss of biodiversity among all types of formerly extensively managed grasslands of poor to medium nutrient richness. Since the 1950s, many Central European uplands have been subject to an increase in grassland cover as a result of changes in land use practices. Using such a landscape in Lower Saxony, Germany, as a model region, we assessed environmental factors that control grassland diversity, including plant community composition, species richness and pollination trait composition. In 2007, 189 vegetation sampling sites were randomly distributed among grasslands covering some 394 ha within a 2500 ha study area. Plant communities were classified using TWINSPAN and the effects of environmental factors (soil, topography, current management and habitat continuity) were analysed by canonical correspondence analysis and regression analysis reducing for the effects of spatial autocorrelation by using principal coordinates of neighbour matrices.We found a wide range of six species-poor (<15 plant spp.) to extremely species-rich (>27 spp.) grassland types under mesic to dry site conditions, including sown, Cynosurion, Arrhenatherion and semi-natural grasslands. Grassland community composition was best explained by soil factors and species richness and pollination type composition by combined effects of current management and habitat continuity. During the 1950/60s, the extent of grassland area within the studied landscape rapidly increased to more than double its previous extent, and in 2007, grasslands comprised 16%. Natura 2000 grassland types comprised 1% of the surveyed site and medium-rich, high-nature-value grasslands a further 5%. While the number of wind-pollinated plant species was equal among all grassland types, there was a parallel decline in insect-pollinated plants and overall median species richness in the grassland communities along a gradient of increasing land use intensity (mowing, nutrient supply). Moreover, insect-pollinated plants occurring in intensively managed grasslands were found to additionally have the ability for self-pollination. Species-rich grasslands – including semi-natural grasslands and a semi-improved, species-rich Arrhenatherion community – occurred exclusively on old sites (with >100 years of habitat continuity) that had been used for traditional sheep grazing (environmental contracting). Medium-rich Arrhenatherion grasslands were established primarily on less productive, formerly arable fields (<30 years). We conclude that conservation efforts should focus on extant species-rich grassland types and should aim to implement traditional land use practices such as sheep grazing. Additional restoration efforts should focus on establishing new grasslands on less productive sites in the proximate surroundings of species-rich grasslands to facilitate seed dispersal, but nitrogen deposition should be buffered where appropriate. These measures would enhance the interaction between nature reserves and agricultural grasslands and thus improve the ecological quality of grasslands at the landscape scale.     

Assessing the impact of land abandonment, nitrogen enrichment and fairy-ring fungi on plant diversity of Mediterranean grasslands

Semi-natural calcareous grasslands are of great conservation interest because of their high species richness, but they are threatened by land abandonment and nitrogen eutrophication. These plant communities evolved as a result of a long history of human activity, which generated and maintained these habitats by extensive grazing and mowing. Calcareous grasslands are listed as a priority for conservation in the EC Habitats Directive. However, the effects of different management regimes, nitrogen enrichment, and soil-borne pathogens on plant species diversity are less clear for grasslands of the Mediterranean Basin, compared to meadows in Northern and Central Europe. In this study, we assessed the impact of land abandonment, nitrogen enrichment, and fairy-ring fungi on species diversity in semi-natural grasslands found in the Mediterranean Basin by comparing the available literature with findings from recent studies carried out in Central Italy. In a series of field experiments, the cutting of abandoned grassland consistently reduced the living biomass of the dominant perennial grasses, such as Brachypodium rupestre and Bromus erectus, and promoted a rapid increase in species richness and diversity by allowing the establishment of rare species. There was a similar, but less effective, restoration of species diversity and composition in mowed grassland after litter removal. We also show that nitrogen enrichment at levels comparable to atmospheric deposition depresses species diversity, which also hampers the positive effects of litter removal. Our findings are consistent with previous results achieved in Northern and Central Europe, which however, mainly focused on grasslands with intermediate to high primary productivity levels. The limited availability of data from low-productivity, drought-prone Mediterranean grasslands requires further studies to assess the impact of land abandonment and nitrogen eutrophication in such ecosystems. Finally, we discuss the role of fairy-ring fungi in the maintenance of plant diversity in species-rich grassland. We show that fairy-ring fungi (e.g. Agaricus campestris) critically affect the spatial distribution and diversity of coexisting plant species. By killing the dominant perennial herbs, these radially growing plant pathogens produce empty niches for rare, short-lived species, thus affecting the vegetation pattern. Overall, our results are of interest for environmental managers, as they provide guidelines for the restoration of abandoned areas and the conservation of these species-rich habitats.     

Altitude modifies species richness–nutrient indicator value relationships in a country-wide survey of grassland vegetation

Nutrient enrichment is a threat to botanical diversity in Europe, and its assessment is part of biodiversity monitoring schemes. In Switzerland, this is done by calculating the average nutrient (N) indicator value of the vegetation based on a country-wide systematic vegetation survey. However, it is questionable whether N values indicate eutrophication and resulting species loss equally well across an entire country, which includes wide topographic gradients and distinct biogeographic regions. Here we analyze vascular plant species lists from 415 grassland plots (10 m²) between 365 and 2770 m a.s.l. throughout Switzerland to investigate how the relationship between N value and species richness differs with altitude and among regions. The N value strongly decreased with altitude (piecewise regression: r² = 0.77), particularly between 800 and 2000 m a.s.l., where this decrease was related to a decreasing proportion of fertilized grasslands. In the alpine belt, lower N values were associated with a greater frequency of acidic soils and a restricted species pool. Vascular plant species richness was maximal at intermediate altitude (piecewise regression: r² = 0.33) and intermediate N value (polynomial regression: r² = 0.46). When analyzed separately by altitudinal belt, the relationship between species richness and N value was negative in the lowlands and montane belt but unimodal in the subalpine belt. In the alpine belt, soil pH (R indicator values) explained most of the variation in species richness. Two indices of between-plot diversity (floristic dissimilarity and the contribution of individual plots to total species richness) were negatively related to N values from the lowlands to the subalpine belt but not in the alpine belt. All relationships differed little among the biogeographic regions of Switzerland, but they might be modified by changes in management and by the expansion of common lowland species into mountain grasslands.     

Direct and indirect effects of land‐use intensification on ant communities in temperate grasslands

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species.