Cork-oak woodlands as key-habitats for biodiversity conservation in Mediterranean landscapes: a case study using rove and ground beetles (Coleoptera: Staphylinidae, Carabidae)

Land-use intensification in Mediterranean agro-forest systems became a pressure on biodiversity, concerning particularly the woodland sensitive species. In 2001, the effects of a land-use gradient from old-growth cork-oak forest to a homogeneous agricultural area were assessed using rove beetles as indicators in a Mediterranean landscape. The aim was to find which species were negatively affected by land-use intensification at the landscape level and whether they benefited from cork-oak patches occurring along the land-use gradient. A total of 3,196 rove beetles from 88 taxa were sampled from all landscape types. Agricultural area recorded significantly higher numbers of abundance and species richness in relation to the cork-oak mosaics, i.e. the old-growth forest and the managed agro-forest landscapes (montados). Moreover, 70% of rove beetle indicator species common enough to be tested by IndVal displayed their highest indicator value for agriculture, showing a lower number of woodland indicators in comparison to ground beetles. Nevertheless, one rove beetle taxon was considered a specialist of closed woodland mosaics while no specialist ground beetle was found for that landscape typology. Some rare rove beetle species were also important in typifying diversity patterns of old-growth cork-oak forests. Hence, future management in Mediterranean landscapes should take into account not only indicator species common enough to be tested by IndVal, but also rare and endemic species. Considering the added value of cork-oak woodland cover for sensitive rove and ground beetle diversity, the strengthening of cork-oak woodland connectivity seems to be a crucial management that is required in agricultural Mediterranean landscapes.     

Introduced and native ground beetle assemblages (Coleoptera: Carabidae) along a successional gradient in an urban landscape

According to the intermediate disturbance hypothesis (IDH), species diversity should be higher at sites with intermediate levels of disturbance. We tested this hypothesis using ground beetles (Coleoptera: Carabidae) collected in pitfall traps from sites that varied in time since last disturbance. This successional gradient was embedded in an urban landscape near Montreal, Quebec. We predicted that diversity in young forests and old fields would be higher than in agricultural fields and old forests. Fifty-five species (2932 individuals) were found in 2003 and 46 species (2207 individuals) in 2004. In both years, species richness was highest from traps placed in agricultural fields. We collected nine introduced species; these had higher catch rates than the native species in both years (64.8% of total catch). When introduced species were removed from the Nonmetric Multidimensional Scaling ordination analysis, the assemblages from agricultural fields were less distinct compared to those of the other habitats, suggesting the introduced fauna is important in structuring carabid assemblages from the agricultural fields. Introduced species may play a significant role in the community composition of ground beetles in urban landscapes, and their influence may be the cause of the lack of support found for the IDH.     

Dung Beetle Community and Functions along a Habitat-Disturbance Gradient in the Amazon: A Rapid Assessment of Ecological Functions Associated to Biodiversity

Although there is increasing interest in the effects of habitat disturbance on community attributes and the potential consequences for ecosystem functioning, objective approaches linking biodiversity loss to functional loss are uncommon. The objectives of this study were to implement simultaneous assessment of community attributes (richness, abundance and biomass, each calculated for total-beetle assemblages as well as small- and large-beetle assemblages) and three ecological functions of dung beetles (dung removal, soil perturbation and secondary seed dispersal), to compare the effects of habitat disturbance on both sets of response variables, and their relations. We studied dung beetle community attributes and functions in five land-use systems representing a disturbance gradient in the Brazilian Amazon: primary forest, secondary forest, agroforestry, agriculture and pasture. All response variables were affected negatively by the intensification of habitat disturbance regimes, but community attributes and ecological functions did not follow the same pattern of decline. A hierarchical partitioning analysis showed that, although all community attributes had a significant effect on the three ecological functions (except the abundance of small beetles on all three ecological functions and the biomass of small beetles on secondary dispersal of large seed mimics), species richness and abundance of large beetles were the community attributes with the highest explanatory value. Our results show the importance of measuring ecological function empirically instead of deducing it from community metrics.     

Management intensity at field and landscape levels affects the structure of generalist predator communities

Agricultural intensification is recognised as a major driver of biodiversity loss in human-modified landscapes. Several agro-environmental measures at different spatial scales have been suggested to mitigate the negative impact of intensification on biodiversity and ecosystem services. The effect of these measures on the functional structure of service-providing communities remains, however, largely unexplored. Using two distinct landscape designs, we examined how the management options of organic farming at the field scale and crop diversification at the landscape level affect the taxonomic and functional structure of generalist predator communities and how these effects vary along a landscape complexity gradient. Organic farming as well as landscapes with longer and more diversified crop rotations enhanced the activity-density of spiders and rove beetles, but not the species richness or evenness. Our results indicate that the two management options affected the functional composition of communities, as they primarily enhanced the activity-density of functionally similar species. The two management options increased the functional similarity between spider species in regards to hunting mode and habitat preference. Organic farming enhanced the functional similarity of rove beetles. Management options at field and landscape levels were generally more important predictors of community structure when compared to landscape complexity. Our study highlights the importance of considering the functional composition of generalist predators in order to understand how agro-environmental measures at various scales shape community assemblages and ecosystem functioning in agricultural landscapes.     

Comparative analysis of two edaphic zoocoenoses (Oribatid mites and Carabid beetles) in five habitats of the ‘Pietraporciana’ and ‘Lucciolabella’ Nature Reserves (Orcia Valley,central Italy)

Oribatid mite and Carabid beetle communities were investigated at five sites in the ‘Pietraporciana’ and ‘Lucciolabella’ Nature Reserves (central Italy). In this part of southern Tuscany many attempts have been made to encourage the regeneration of native habitats and to preserve existing ones. Human-induced changes in the original forest landscape have had a direct impact on mite and carabid populations. Significant differences in species diversity and abundance among different sites were revealed throughout the sampling period. Species richness, abundance and diversity of oribatid mites decrease from woodland sites to open habitats where evenness was high. There is an inverse trend between the number of species and richness of carabid beetles and those of oribatid mites. Canonical correspondence analysis (CCA) of oribatid and carabid beetle compositions discriminated the sites, demonstrating how even small areas with different vegetation, composition, structure, environment and microclimate were characterised by distinct edaphic populations.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

Inter‐assemblage facilitation: the functional diversity of cavity‐producing beetles drives the size diversity of cavity‐nesting bees

Inter‐specific interactions are important drivers and maintainers of biodiversity. Compared to trophic and competitive interactions, the role of non‐trophic facilitation among species has received less attention. Cavity‐nesting bees nest in old beetle borings in dead wood, with restricted diameters corresponding to the body size of the bee species. The aim of this study was to test the hypothesis that the functional diversity of cavity‐producing wood boring beetles ‐ in terms of cavity diameters ‐ drives the size diversity of cavity‐nesting bees. The invertebrate communities were sampled in 30 sites, located in forested landscapes along an elevational gradient. We regressed the species richness and abundance of cavity nesting bees against the species richness and abundance of wood boring beetles, non‐wood boring beetles and elevation. The proportion of cavity nesting bees in bee species assemblage was regressed against the species richness and abundance of wood boring beetles. We also tested the relationships between the size diversity of cavity nesting bees and wood boring beetles. The species richness and abundance of cavity nesting bees increased with the species richness and abundance of wood boring beetles. No such relationship was found for non‐wood boring beetles. The abundance of wood boring beetles was also related to an increased proportion of cavity nesting bee individuals. Moreover, the size diversity of cavity‐nesting bees increased with the functional diversity of wood boring beetles. Specifically, the mean and dispersion of bee body sizes increased with the functional dispersion of large wood boring beetles. The positive relationships between cavity producing bees and cavity nesting bees suggest that non‐trophic facilitative interactions between species assemblages play important roles in organizing bee species assemblages. Considering a community‐wide approach may therefore be required if we are to successfully understand and conserve wild bee species assemblages in forested landscapes.     

Arthropod Diversity and Community Structure in Relation to Land Use in the Mekong Delta, Vietnam

Declining biodiversity in agro-ecosystems, caused by intensification of production or expansion of monocultures, is associated with the emergence of agricultural pests. Understanding how land-use and management control crop-associated biodiversity is, therefore, one of the key steps towards the prediction and maintenance of natural pest-control. Here we report on relationships between land-use variables and arthropod community attributes (for example, species diversity, abundance and guild structure) across a diversification gradient in a rice-dominated landscape in the Mekong delta, Vietnam. We show that rice habitats contained the most diverse arthropod communities, compared with other uncultivated and cultivated land-use types. In addition, arthropod species density and Simpson’s diversity in flower, vegetable and fruit habitats was positively related to rice cover in the local landscape. However, across the landscape as a whole, reduction in heterogeneity and the amount of uncultivated cover was associated, generally, with a loss of diversity. Furthermore, arthropod species density in tillering and flowering stages of rice was positively related to crop and vegetation richness, respectively, in the local landscape. Differential effects on feeding guilds were also observed in rice-associated communities with the proportional abundance of predators increasing and the proportional abundance of detritivores decreasing with increased landscape rice cover. Thus, we identify a range of rather complex, sometimes contradictory patterns concerning the impact of rice cover and landscape heterogeneity on arthropod community attributes. Importantly, we conclude that that land-use change associated with expansion of monoculture rice need not automatically impact diversity and functioning of the arthropod community.     

Effect of land-use on the earthworm assemblages in semi-deciduous forests of Central-West Ivory Coast

In this study, the impact of forest disturbance on earthworm assemblages was assessed using monoliths dug out at 5 m intervals along a gradient of land-use intensification. The land-use types comprised primary forest (as a baseline), secondary forest, tree plantation, fallow, perennial and annual crop. Forest disturbance resulted in a significant decrease in soil organic carbon and pH, while earthworm abundance and biomass increased along the gradient of disturbance. Surprisingly, anthropogenic disturbances in semi-deciduous forest areas have not led to the disappearance of native species to the benefit of exotic species as revealed in former studies. As a result, in this study land-use change had no impact on species richness at the landscape level, even if at local scales, recurrent Chromolaena odorata fallows, multispecies tree plantations and 4 year-old teak plantations hosted the highest average species richness. Multiple regression analyses performed between earthworm communities and environmental variables showed that soil organic carbon and pH are potential indicators of earthworm abundance change.     

Carabid beetle (Coleoptera: Carabidae) richness and functional traits in relation to differently managed grasslands in the Alps

Species richness, composition, and functional traits of carabid beetle assemblages (Coleoptera: Carabidae) were studied in relation to different grassland management. Carabid beetles were sampled during the summers 2008 and 2009 by 165 traps located in 11 sites in the central-eastern Italian Alps. Using mixed effect models to account for potential spatial bias, we found that mown grasslands had significantly more species, a lower proportion of wingless species and a lower proportion of species with long larval development than grazed and natural grasslands. Within grazed and mown grasslands, neither cattle density nor number of cuts had any significant effect neither on species richness nor on any of the traits. The influence of grassland management can be summarised as follows: (1) grazing does not change community structure and functional traits compared to natural grasslands; (2) mowing negatively affects the carabid beetle assemblages; (3) the intensity of grazing and of cutting may not affect the structure of species assemblages of ground beetles. Our results support the hypothesis that agroecosystem practices in alpine grasslands influence carabid beetle communities. Specifically, the species with traits typical of undisturbed habitats (low dispersal abilities and long larval development) are more sensitive to perturbations (e.g. cutting). Our suggestion for agricultural and environmental planning and for conservation schemes is that the preservation of natural grasslands (e.g. forest gaps) and the implementation of grazing should be promoted during the planning of agroecosystem mosaics.     

High-resolution distribution of bumblebees (Bombus spp.) in a mountain area marked by agricultural decline

Since the 1980s, bumblebee species have declined in Europe, partly because of agricultural intensification. Yet little is known about the potential consequences of agricultural decline on bumblebees. In most mountainous areas, agricultural decline from rural exodus is acute and alters landscapes as much as intensive farming. Our study aims at providing a quantitative assessment of agricultural decline through its impact on landscapes, and at characterising bumblebee assemblages associated with land-use types of mountain regions. The studied area (6.2 km²) belongs to the Eyne’s valley in the French Pyrenees, known to host the exceptional number of 33 bumblebee species of the 45 found in continental France. We compare aerial photographs from 1953 and 2000 to quantify agricultural decline. We cross a bumblebee database (2849 observations) with land-use types interpreted from aerial photographs from 2000. Comparison of land-use maps from 1953 and 2000 reveals a strong progression of woodland and urbanised areas, and a decline of agricultural land (pastures and crops), except for hayfields. Spatial correlations between low altitude agro-pastoral structure and the occurrence of bumblebee species shows that bumblebee specific richness is highest in agro-pastoral land-uses (pastures and hayfields) and in the ski area, and poorest in woodland and urbanised areas. Urbanisation and agricultural decline, through increased woodland areas, could lead to a loss of bumblebee diversity in the future. To preserve high bumblebee richness, it is crucial to design measures to maintain open land habitats and the landscape’s spatial heterogeneity through agro-pastoral practices.     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

Landscape-scale crop diversity interacts with local management to determine ground beetle diversity

Crop diversification and maintenance of semi-natural habitats (grasslands and field boundaries) are suggested to enhance farmland biodiversity, but the relative importance of these factors remains poorly known. We evaluated how crop diversity and availability of semi-natural grasslands at a landscape-scale interacted with local farming management (three management types from low to high intensity: ley < winter wheat < sugar beet) in their effect on ground beetle assemblages in southern Sweden. Ground beetle diversity increased with crop diversity either independently of local management (Simpson species diversity), or only in the less intensively managed habitats (rarefied species richness). While ground beetle diversity in leys tended to increase with field boundary length, no such relationship was observed in winter wheat or sugar beet fields. In contrast, the landscape proportions of leys and semi-natural grasslands did not affect ground beetle species richness and diversity. We conclude that (a) semi-natural grasslands and leys may not function as source habitats at a landscape-scale if they comprise a low proportion of the total land-use, while (b) increasing crop diversity is correlated to ground beetle richness and diversity in agricultural landscapes dominated by arable land. The beneficial effect of landscape-scale crop diversification on farmland biodiversity may depend on the general level of agricultural intensity of a region.     

Termite diversity along a land use intensification gradient in a semi-arid savanna

Disturbance, particularly agricultural expansion is one of the major threats to the biodiversity and ecological functions of tropical and sub-tropical ecosystems. In this regard, we examined changes in the species richness, abundance, and diversity of termites across different disturbance treatments in a sub-tropical semi-arid savanna in south eastern Zimbabwe. Nine transects (100?×?2 m) representing three habitat disturbance treatments (primary woodland; grazing area; agricultural field) were sampled for termites using a rapid biodiversity assessment protocol. Termites were more abundant and species-rich in primary woodland and grazing area than in the agricultural field. Twelve termite species from three sub-families were present, with Microtermes sp. constituting 35% of the identified termite species. Termite feeding group structure differed significantly among land-use types, and of all termites present, wood-feeding termites were the most abundant while soil-feeders were rare in the agricultural field. In conclusion the observed pattern in termite species richness and relative abundance indicates that termites are very resilient to natural disturbance and might actually benefit from some natural disturbances like they did in the grazing area of this study, but they are not resilient to extreme anthropogenic disturbance. Although there was no notable difference in termite species richness and relative abundance between agricultural field and primary woodland, the pattern observed across the three sites may be potential support for the IDH suggesting that intermediate levels of physical disturbance intensity influence the structure and functioning of termite assemblages in semi-arid savanna.     

Does Land-Use Intensification Decrease Plant Phylogenetic Diversity in Local Grasslands?

Phylogenetic diversity (PD) has been successfully used as a complement to classical measures of biological diversity such as species richness or functional diversity. By considering the phylogenetic history of species, PD broadly summarizes the trait space within a community. This covers amongst others complex physiological or biochemical traits that are often not considered in estimates of functional diversity, but may be important for the understanding of community assembly and the relationship between diversity and ecosystem functions. In this study we analyzed the relationship between PD of plant communities and land-use intensification in 150 local grassland plots in three regions in Germany. Specifically we asked whether PD decreases with land-use intensification and if so, whether the relationship is robust across different regions. Overall, we found that species richness decreased along land-use gradients the results however differed for common and rare species assemblages. PD only weakly decreased with increasing land-use intensity. The strength of the relationship thereby varied among regions and PD metrics used. From our results we suggest that there is no general relationship between PD and land-use intensification probably due to lack of phylogenetic conservatism in land-use sensitive traits. Nevertheless, we suggest that depending on specific regional idiosyncrasies the consideration of PD as a complement to other measures of diversity can be useful.     

From natural woodlands to cultivated land: diversity of fruit‐feeding butterflies and beetles in the mid‐Zambezi valley,northern Zimbabwe

We tested the effect of cultivation on butterfly (Nymphalidae: Charaxes) and beetle (Coleoptera: Scarabaeidae: Cetoniinae) species richness and abundance along a cultivation intensification gradient. Results showed significant differences in species richness and abundance between natural woodlands and cultivated landscapes with larger differences in areas of high cultivation intensity. The results indicate that natural woodland clearing for cultivation purposes has negative impacts on arthropod diversity, a situation more severe in highly intensified cultivated areas. Our results imply that mosaics of different land‐use units, each in a different phase of clearance‐cultivation‐abandonment‐recovery‐clearance cycle could counter the negative effects of cultivation intensity on arthropod diversity.     

Farmland carabid beetle communities at multiple levels of spatial scale

Carabid beetle communities were studied at three levels of spatial scale, i.e. within fields (three sets of traps, mean distance 15 m), between fields (five fields within 1 km from each other) and between patches of farmland (four study areas 4-8 km from each other). We compared carabid assemblages sampled on five crop types in each study area, i.e. ley. set-aside, spring cereal, potato, and sugar beet. Because of small sample sizes, only the community composition was studied within fields with a DCA ordination, but the differences were small. Analyses of species richness, activity density, community composition as revealed by DCA ordinations, dominance structure, diversity, and evenness showed that carabid communities varied significantly among patches of farmland but not between fields with different crops within the patches. Only the communities of potato fields were found lo resemble each other. Only five carabid beetle species showed a preference for crops, and twelve species favoured some study areas. We conclude that the effect of spatial scale should be emphasised in further studies as it seems that carabid beetles may be very localised even in agricultural habitats.     

Effects of dung-pad conditions and density on coprophagous beetle assemblages in a Mediterranean rangeland

Dung beetles highly depend on the ephemeral microhabitat dung which is food resource and larval habitat at the same time. Environmental conditions surrounding a dung pad, such as vegetation structure, have an impact on dung beetle assemblages. We investigated the influence of dung conditions and surrounding habitat characteristics on Mediterranean dung beetle assemblages in a permanently grazed landscape in northern Sardinia. We sampled the dung beetle assemblages of donkey and horse dung in three different vegetation types and assessed species richness and abundance of dung beetles. Species richness was determined by dung and surrounding habitat conditions, whereas abundance was solely affected by dung conditions. However, species richness and abundance decreased with increasing dung density. The effect of dung density on species richness varied depending on vegetation type, with dry grassland exhibiting the highest number of dung beetles species at high dung density. Species composition in dung pads was influenced by abiotic factors with dwellers being negatively affected by increasing dung-pad temperature. Our results underline the importance of diverse vegetation, particularly with respect to the complexity of vegetation which interrelates with the microclimate. Furthermore, our findings illustrate the negative effect of high dung densities on dung beetle assemblages, suggesting that the degree of the intensity of use by grazing animals is important when considering measures for the conservation of dung beetles.     

Habitat structure and neighbor linear features influence more carabid functional diversity in olive groves than the farming system

The effects of land-use management and environmental features at different scales on carabid beetle diversity and trait structure were assessed across olive groves in northeastern Portugal. We selected organic and integrated olive groves that were distinct in terms of specific management practices, local linear features and landscape configurations. Besides the management intensification levels, differences in carabid diversity and community traits were mainly due to local habitat and ecological linear structures at a finer spatial scale. Carabid community traits related to disturbance, namely traits of body size and species dispersal ability, responded to land-use intensity and particular olive grove features were influencing diversity patterns. Within the olive grove patches, larger and brachypterous species were associated to plots with more dense vegetation cover while macropterous and small-sized species were more associated to open areas. Also, larger carabid species benefitted from higher patch size heterogeneity within the landscape mosaics. Our findings indicate that the effects of farming system is contingent on the specific management practices, local and linear features present in agroecosystems such as olive groves. Particularly, the influence of local features on carabid diversity patterns and community traits linked to dispersal and movement may be crucial in maintaining pest control at a landscape scale.     

Nutrient concentrations and fibre contents of plant community biomass reflect species richness patterns along a broad range of land-use intensities among agricultural grasslands

Understanding changes in biodiversity in agricultural landscapes in relation to land-use type and intensity is a major issue in current ecological research. In this context nutrient enrichment has been identified as a key mechanism inducing species loss in Central European grassland ecosystems. At the same time, insights into the linkage between agricultural land use and plant nutrient status are largely missing. So far, studies on the relationship between chemical composition of plant community biomass and biodiversity have mainly been restricted to wetlands and all these studies neglected the effects of land use. Therefore, we analyzed aboveground biomass of 145 grassland plots covering a gradient of land-use intensities in three regions across Germany. In particular, we explored relationships between vascular plant species richness and nutrient concentrations as well as fibre contents (neutral and acid detergent fibre and lignin) in the aboveground community biomass.We found the concentrations of several nutrients in the biomass to be closely linked to plant species richness and land use. Whereas phosphorus concentrations increased with land-use intensity and decreased with plant species richness, nitrogen and potassium concentrations showed less clear patterns. Fibre fractions were negatively related to nutrient concentrations in biomass, but hardly to land-use measures and species richness. Only high lignin contents were positively associated with species richness of grasslands. The N:P ratio was strongly positively related to species richness and even more so to the number of endangered plant species, indicating a higher persistence of endangered species under P (co-)limited conditions. Therefore, we stress the importance of low P supply for species-rich grasslands and suggest the N:P ratio in community biomass to be a useful proxy of the conservation value of agriculturally used grasslands.