Evaluating patterns of biodiversity in managed grasslands using spatial turnover metrics

Market and policy incentives that encourage agricultural intensification, such as incentives for bioenergy, may contribute to biodiversity decline when they encourage a large-scale conversion of native and semi-natural ecosystems to production fields. In order to appreciate the impact of these incentives on biodiversity, it is imperative to better understand how native and semi-natural ecosystems contribute to plant diversity and composition. We studied the five most common types of managed grasslands in Northeastern Kansas, a region undergoing agricultural intensification. We analyzed plant community data recorded at three spatial scales in 98 managed grassland sites, and examined patterns of diversity and composition of plant species and functional groups; and spatial turnover of species and functional groups measured at different spatial scales. We found reduced soil quality and plant diversity as well as a lack of scale-dependent community patterns on sites that were historically cultivated. Forage management practices (haying or grazing) altered plant diversity and composition in native grassland remnants but not in non-native grasslands. We identified several opportunities where changes to existing management practices could benefit both conservation and bioenergy production objectives. Recommendations to conserve biodiversity include increasing the use of hay management or other biomass collection on native grassland remnants and improving the Conservation Reserve Program by increasing enrollment, adding more native species to seed mixes, and incorporating a periodic biomass collection. We also found that using measures of spatial turnover in community composition added important insights in understanding the effects of management decisions on biodiversity.     

Spatial α-diversity patterns of diverse insect taxa in Northern China: Lessons for biodiversity conservation

Current approaches in terrestrial biodiversity conservation focus predominantly on plants and vertebrates. While these groups account for less than 4% of the estimated global species richness, it is commonly argued that especially the species richness in higher plants is a suitable indicator of overall biodiversity. We tested this assumption, investigating species richness and equitability patterns in three highly species-rich insect families and their links with the vegetation and other environmental factors. Vegetation surveys were combined with pitfall and light trapping to establish the α-diversity of ground beetles, geometrid moths and arctiid moths on 48 plots at varying altitudes between Beijing and the Inner Mongolian Plateau. Soil pH and nutrient status were also recorded. The α-diversity patterns in the three insect families were non-congruent, and links with phytodiversity were weak. The spatial α-diversity patterns in each of the three insect families were significantly correlated with the species density of individual plant families. These links varied between the three insect taxa and were mostly negative in moths. Furthermore, geometrid moth diversity decreased with increasing elevation and decreasing soil pH. Strongly diverging α-diversity patterns across different insect taxa illustrate that it is impossible to find a simple surrogate representing cross-taxon diversity for these highly diverse groups. Furthermore, phytodiversity and vegetation composition appear to play a limited role in governing insect diversity patterns. These results underline the significant risk that current plant-focused approaches in terrestrial biodiversity conservation are inadequate in addressing the conservation needs of the vast majority of species on earth.     

Grazing promotes dung beetle diversity in the xeric landscape of a Mexican Biosphere Reserve

We analyze the impact of grazing on dung beetle diversity at the Barranca de Metztitlán Biosphere Reserve, a xeric ecosystem in central Mexico with a long history of use by humans. We compared the community structure, as well as the alpha and beta diversity between two cover conditions (open and closed vegetation) that represent the impact of grazing within a habitat, and between habitat types (submountainous and crassicaule scrublands). From 576 samples we collected 75,605 dung beetles belonging to 20 taxa. While mean species richness and diversity were different between habitat types, cumulative species richness was not. The effects of grazing on vegetation structure influenced the cumulative species richness and diversity of dung beetles in the submountainous scrubland, where grazing has created land mosaics of a grassland matrix with scrubland patches. This was not the case in the crassicaule scrubland where the impact of grazing is not as evident. Beta diversity significantly responds to the effects of grazing on habitat conditions. We discuss the ecological factors that may promote these responses by landscape diversity components. We also identify the species that could act as useful indicators to monitor the effect of land management on biodiversity. Our results indicate cattle farming maintains a diversified land mosaic, and these areas support more diverse dung beetle ensembles than homogeneous areas of closed, shrubby vegetation cover. Thus, controlled grazing activity could certainly favour the conservation of dung beetle biodiversity and improve ecosystem functioning by maintaining dung decomposition rates.     

宁夏香山荒漠草原区生物多样性与环境特征研究

采用大样地、典型固定样地调查方法,对宁夏香山荒漠草原区生物多样性与环境特征进行了研究。结果表明,研究区所调查151种植物分属39科、95属;植物群落分布体现了山地荒漠草原的特征,多数植物群落类型分布在山地阴坡;群落多样性动态受时间、地形影响较大,在土壤属性中受土壤含水量影响较大;灌丛植被在该山地生态系统中处于重要地位,是生态功能维持的主要因素;昆虫群落符合干旱地区草地植物群落特征,大型动物的稀有也提示对该地区进行生物多样性保护日益紧迫。     

Regrowth and revegetation in temperate Australia presents a conservation challenge for reptile fauna in agricultural landscapes

Secondary vegetation, associated with changes in land use, presents a conservation issue in the preservation of biological diversity in agricultural landscapes. We examine the interactive effect of eucalypt regrowth and rock habitat on reptile species richness and assemblage structure in fragmented agricultural landscapes in south-eastern Australia. Zoogeographic and geomorphic factors influenced species richness and community composition. Saxicolous and arboreal species were less abundant in grassy woodland regrowth, whereas Bassian and fossorial species responded positively to forest regrowth (and tree plantings). Regrowth with rock habitat had higher reptile richness, and more old growth-associated taxa, than regrowth without rock habitat. Thus, the presence of saxicolous habitat can reduce the time required for regrowth to attain a climax community structure and elements of old growth fauna. However, in the absence of vegetation management, secondary vegetation can reduce habitat suitability for a broad range of reptiles. In agricultural landscapes historically supporting savannah-like vegetation, habitat manipulation may be necessary to maintain reptile diversity. Furthermore, tree plantings in temperate agricultural landscapes should aim to restore historical vegetation composition and structure, thereby reflecting the habitat requirements of extant species and facilitating evolutionary processes. In grassy woodland ecosystems, this may involve heterogeneous plantings which emulate natural levels of canopy cover and solar penetration. Maintaining biodiversity in agricultural landscapes will therefore involve managing trade-offs to preserve areas of dense regeneration for regrowth-dependant fauna, while at the same time, creating open-canopy environments to enhance habitat for ectothermic organisms.     

Grazing management of calcareous grasslands and its implications for the conservation of beetle communities

Calcareous grasslands are an important habitat for floral and faunal communities in the UK and Europe. Declines due to changes in management, scrub invasion and agricultural improvement have left much of the remnants of this habitat in a degraded and fragmented state. Grazing, by cattle or sheep, is one of the main management practices used to maintain and improve the floral and faunal quality of calcareous grassland. The long-term impacts of different grazing regimes, however, are poorly understood, particularly in terms of the invertebrate communities. This study contrasted the impacts of recently introduced and long-term sheep or cattle grazing on beetle communities present on one of the largest areas of calcareous grassland in Europe, the Salisbury Plain military training Area, UK. No effects of grazing management on beetle abundance, species richness or evenness were found, but plant diversity and overall percentage cover of grasses did influence beetle diversity. Proportions of the total number of individuals and overall species richness within beetle guilds (predatory, phytophagous, flower/seed feeders, root feeders and foliage feeders) were strongly influenced by both the duration and type of grazing animal. At the species level, beetle community structure showed significant differences between ungrazed, long-term cattle and long-term sheep grazing treatments. Changes in plant community structure were found to influence beetle community structure. The significance of these results is discussed in terms of the long-term impacts of grazing on beetle community structure, and the benefits of different grazing regimes for the conservation management of calcareous grasslands.     

Spatial congruence between biodiversity and ecosystem services in South Africa

Ecosystems services sustain humans all over the world. The unsustainable use of ecosystem services around the world has led to widespread degradation which now threatens human health and livelihoods. Although the maintenance of ecosystem services is often used to justify biodiversity conservation actions, it is still unclear how ecosystem services relate to different aspects of biodiversity and to what extent the conservation of biodiversity will ensure the provision of services. The aim of this study was to find out whether biodiversity priorities, biomes, species richness and vegetation diversity hotspots co-occur in space with ecosystem services. The distribution of the ranges and hotspots of five ecosystem services (surface water supply, water flow regulation, carbon storage, soil accumulation, and soil retention) was assessed in South African biomes. Coincidence, overlap, and correlation analyses were used to assess spatial congruence between ecosystem services and species richness (plants and animals) and vegetation diversity hotspots. The grassland and savanna biomes contained significant amounts of all five ecosystem services. There was moderate overlap and a generally positive but low correlation between ecosystem services hotspots and species richness and vegetation diversity hotspots. Species richness was mostly higher in the hotspots of water flow regulation and soil accumulation than would be expected by chance. The water services showed varying levels of congruence with species richness hotspots and vegetation diversity hotspot. These results indicate that actions taken to conserve biodiversity in South Africa will also protect certain ecosystem services and ecosystem services can be used to strengthen biodiversity conservation in some instances.     

Successional trajectories of soil nematode and plant communities in a chronosequence of ex-arable lands

Conversion of arable land into semi-natural grassland or heath land is a common practice for restoring and conserving plant diversity. However, little is known about the effectiveness of land conversion for restoring and conserving taxonomic and functional diversity in the soil. We studied soil nematode community development in a chronosequence of abandoned fields and related this to plant community development. The taxonomic and functional composition of the soil nematode community was analyzed to detect changes in soil food web structure, using semi-natural sites and theoretical plant and soil communities as references.While plant communities clearly developed towards the semi-natural references, there was less direction in succession of nematode taxa. The number of fungal feeding nematodes increased after land abandonment. Numbers of omni-carnivorous nematodes expanded only during the first years, after which there were no substantial changes for the next three decades. Plant communities on the ex-arable fields developed towards the theoretical reference plant associations Galio hercynici-Festucetum ovinae and Genisto anglicae-Callunetum. Nematode communities developed away from a theoretical community indicative of arable land, but there was no clear development towards a theoretical (semi-)natural reference. Our results show that restoration and conservation of plant communities is of limited indicative value for developments belowground: successful restoration of plant diversity does not necessarily imply successful restoration of belowground diversity. Assessing the impact of conservation measures on restoring soil biodiversity requires information on belowground community composition of (semi-)natural areas in order to establish proper references for restoration sites.     

Invertebrate diversity and national responsibility for species conservation across Europe - A multi-taxon approach

Invertebrates play important roles in ecosystem functioning, but with the decline in biodiversity worldwide, it has become increasingly evident that our knowledge of invertebrate diversity distributions and of how to adequately integrate many invertebrates into conservation strategies is insufficient. We analyzed congruence between patterns of total as well as of endemic species richness of twelve invertebrate taxa across Europe and compared these patterns to those of so far predominantly studied vertebrates and plants. Coarse-scale centers of high collective diversity were identified, all in southern European regions. However, the strength of covariation between diversity patterns strongly depended on the respective taxa, with ground beetles emerging as one of the invertebrate taxa significantly reflecting endemism and total species richness distributions of many invertebrates, vertebrates and plants. Plants and ants showed similarly strong cross-taxon correlations. In contrast, correlations of vertebrates with invertebrates were in part less strong, questioning whether conservation strategies based on the former adequately account for diversity of invertebrates as a whole. Moreover, as the potential impact of environmental factors on diversity distributions varies among taxa, future climate change might affect relationships between richness patterns of different taxa. Our findings draw attention to the necessity of incorporating invertebrates, represented for instance by the highly diverse ground beetles, into national and international conservation decisions. They also highlight the importance of individual countries in maintaining European biodiversity and the responsibility of these countries for specific invertebrate taxa within international conservation frameworks, emphasizing the need for adequate consideration of invertebrates in the implementation of conservation actions.     

Effects of vineyard management on biodiversity at three trophic levels

In annual crops, where disturbance intensities are generally high, numerous investigations demonstrated beneficial effects of organic and other extensive farming practices on biodiversity. It is however unclear if organic farming has the same beneficial effect in perennial crops (e.g. vineyards) because of a generally reduced background disturbance level. Moreover, the impact of farming practices on biodiversity may differ between trophic levels. We tested the effect of farming practices differing in disturbance intensity on the abundance, species richness and community composition of three trophic levels (plants, grasshoppers and spiders) in Swiss vineyards. Organic and conventional vineyards were compared, and within the latter different options of weed control (mowing vs. mulching) and pest management (hand spraying vs. helicopter spraying) were investigated. In contrast to findings in annual crops, organic farming promoted neither diversity nor abundance at any trophic level. Grasshopper diversity was even significantly lower in organic compared to conventional vineyards. Weed control as well as pest management differed in their effects, depending on the trophic level. Within conventionally managed vineyards, spider and grasshopper diversity were enhanced by mulching, while plants benefited from the lower disturbance of mowing. The greater disturbance of fungicide spraying by hand than by helicopter was disadvantageous for grasshoppers, but did not affect spiders and plants. Thus, different taxa respond differently not only to an increase in disturbance but also to different types of disturbance and local site conditions. We conclude that disturbance in organically farmed vineyards appears to be too low to be beneficial for biodiversity.     

Landscape elements as potential barriers and corridors for bees, wasps and parasitoids

Habitat loss and fragmentation in agricultural landscapes lead to severe declines of abundance and richness of many insect species in the remaining isolated semi-natural habitats. We analysed possible barrier effects of large hedges and corridor effects of narrow grass strips that were hypothesized to affect foraging and dispersal of hymenopterans. We selected calcareous grasslands in the vicinity of Göttingen (Germany), which harbour high Hymenoptera diversity and are starting points for foraging and dispersal in the landscape. We installed pan traps to sample bees (i) on the grasslands; (ii) on grassland edges behind adjacent hedges (potential barriers) and without hedges; (iii) on grass strips in 100 m distance to the grassland, which were connected or unconnected to the grassland; and (iv) unconnected (isolated) grass strips in 300 m and 750 m distance to test for corridor and isolation effects on abundance and species richness of foraging wild bees. Additionally we provided trap nests for bees, wasps and their parasitoids on the grasslands and the strips. Species abundance and richness declined with increasing isolation from grasslands for foraging solitary bees, trap-nesting bees, wasps and parasitoids, but not for foraging bumblebees. Hedges did not confine movement of foraging bees. We found no mitigating effects of (100 m) corridor strips on any of the observed groups. We conclude that conservation of semi-natural habitats as sources of bee and wasp diversity is important and that grass strips act as sinks rather than corridors when high quality patches are nearby.     

Soil community changes during secondary succession to naturalized grasslands

Succession to a naturalized grassland from former agricultural land and pastures is accompanied by changes in plant biodiversity and in the soil community. These changes are the result of a reduction or elimination of management, fertilizer applications and of grazing by large herbivores. We review soil biology studies on agricultural land that are in various successional stages towards naturalized grasslands, where interactions between plant species composition changes and the soil ecology affect each other. In many chronosequence studies, the soil microbial community tends to shift towards a less bacterial, and more fungal dominated food web energy channel following a reduction in fertilizer inputs and grazing intensity. Whereas changes in microarthropod communities are obscured, nematode trophic functional group (ecological guild) changes respond to both plant and soil community changes. There are opportunities to further study the feedback interactions between roots and soil organisms in grasslands. A better understanding of the molecular feedback mechanisms would be beneficial in long-term grassland management.     

东北平原典型玉米种植区农业景观植物多样性研究——以昌图县为例

非耕作生境是农业景观中的重要组成部分,为生物提供食物和栖息地,对生物多样性保护具有重要意义。本文以辽宁省昌图县为例,选取了20个非耕作生境比例为0~50%的网格(1 km?1 km)为样区,采用典型样地法对调查区域内农业景观中的林地、田间路、草地、沟渠、果园5类主要非耕作生境中的植物群落进行调查,探讨不同非耕作生境类型和比例中植物多样性及其与景观异质性之间的关系。研究结果表明,随着非耕作生境比例增加,植物物种多样性和均匀度先增加后减小,在20%~30%时达到峰值;非耕作生境比例在0~40%范围内时,植物物种丰富度同样呈现先增加后减小,且在20%~30%时达到峰值;非耕作生境比例高于40%时,植物物种丰富度再次出现峰值且高于20%~30%。随着非耕作生境比例的增加,农业景观香浓多样性、香浓均匀度指数和景观斑块形状复杂程度逐渐增加,蔓延度恰好相反。不同生境中重要值最高的均为禾本科植物,但植物种不同;优势种优势度为果园沟渠草地田间路林地。果园、草地和沟渠的物种多样性和均匀度指数显著高于林地和田间路,丰富度指数为草地林地沟渠果园田间路,且草地、果园和田间路的不同调查区域间植物群落指数存在较大的变异性。草地、果园、沟渠、田间路和林地中景观异质性与物种多样性相关性顺次降低。研究结论表明,东北平原典型玉米种植区农业景观中,最适宜非耕作生境比例为20%~30%;非耕作生境中林地、沟渠对物种多样性维持、湿生植物多样性保护具有积极作用;不同生境干扰类型和方式不同,景观异质性与植物物种多样性相关性程度也不同。在今后农业景观生物多样性保护中应综合考虑非耕作生境的类型和比例,干扰的类型和方式等多种因素对生物多样性、生态服务及农业产量的影响。     

Remnant grassland habitats as source communities for plant diversification in agricultural landscapes

Lately there has been a shift in Sweden from grazing species-rich semi-natural grasslands towards grazing ex-arable fields in the modern agricultural landscape. These fields normally contain a fraction of the plant species richness compared to semi-natural grasslands. However, small remnant habitats have been suggested as important for plant species diversity and conservation as they may function as refugia for grassland specialists in fragmented and highly modified agricultural landscapes. In this study, we examined whether plant communities on small remnant habitats, i.e. midfield islets, can function as sources for grassland species to disperse out into surrounding grazed ex-fields (former arable fields). We examined species richness and grassland specialists (species favoured by grazing) and their ability to colonize fields after 5 and 11 years of grazing. The fields that had been grazed for a shorter time were fairly species-poor with few grassland specialists. A longer period of grazing had a positive effect on total and small-scale species diversity in both islets and fields. Species composition became more similar with time, and the number of grassland specialists in both habitats increased. We found that grassland specialists dispersed step-wise into the fields, and the number of grassland specialists decreased with distance from the source. Our study suggests that remnant habitats, such as midfield islets, do function as a source community for grassland specialists and enhance diversification of grassland species when grazing is introduced. For long-term conservation of plant species, incorporating small refugia into larger grazing complexes may thus enhance species richness.     

Grassland management influences spatial patterns of soil microbial communities

Soil micro-organisms play a vital role in grassland ecosystem functioning but little is known about the effects of grassland management on spatial patterns of soil microbial communities. We compared plant species composition with terminal restriction fragment length polymorphism (T-RFLP) fingerprints of soil bacterial and fungal communities in unimproved, restored and improved wet grasslands. We assessed community composition of soil micro-organisms at distances ranging from 0.01 m to 100 m and determined taxa–area relationships from field- to landscape level. We show that land management type influenced bacterial but not fungal community composition. However, extensive grassland management to restore aboveground diversity affected spatial patterns of soil fungi. We found distinct distance–decay and small-scale aggregation of fungal populations in extensively managed grasslands restored from former arable use. There were no clear spatial patterns in bacterial communities at the field-scale. However, at the landscape level there was a moderate increase in bacterial taxa and a strong increase in fungal taxa with the number of sites sampled. Our results suggest that grassland management affects soil microbial communities at multiple scales; the observed small-scale variation may facilitate plant species coexistence and should be taken into account in field studies of soil microbial communities.     

Small-scale plant species richness and evenness in semi-natural grasslands respond differently to habitat fragmentation

The study explores whether small-scale species diversity, species evenness and species richness in semi-natural grassland communities are similarly associated with present management regime and/or present and historical landscape context (percentage of different land-cover types in the surroundings). Species diversity, evenness and richness were recorded within 441 50 × 50 cm grassland plots in 4.5 × 4.5 km agricultural landscape on Öland, Sweden. Recent and historical land-cover maps (years 2004, 1959, 1938, 1835, and 1800) were used to characterize the present and past landscape context of the sampled vegetation plots. Partial regression and simultaneous autoregressive models were used to explore the relationships between species diversity measures (Shannon diversity, richness and evenness) and different explanatory variables while accounting for spatial autocorrelation in the data. The results indicated that species richness was relatively sensitive to grassland isolation, while the response of species evenness to isolation was characterized by a degree of inertia. Because the richness and evenness components of species diversity may respond differently to habitat fragmentation, we suggest that monitoring projects and empirical studies that focus on changes in biodiversity in semi-natural grasslands should include the assessment of species evenness - as a complement to the assessment of species richness. In addition, our results indicated that the development and persistence of a species-rich and even grassland vegetation was favoured in areas that have historically (in the 19th century) been surrounded by grasslands. Information on landscape history should, whenever possible, be incorporated into the planning of strategies for grassland conservation.     

Changes in agricultural management drive the diversity of Burkholderia species isolated from soil on PCAT medium

In order to assess the diversity of culturable Burkholderia populations in rhizosphere and bulk soil and to evaluate how different agricultural management regimes and land use history affect this diversity, four treatments were evaluated: permanent grassland; grassland converted into maize monoculture; arable land and arable land converted into grassland. Burkholderia isolates obtained on PCAT medium were grouped in 47 clusters using 16S ribosomal RNA gene based PCR-DGGE combined with BOX genomic fingerprinting (DGGE-BOX). The distribution of the isolates in the DGGE-BOX clusters was used to calculate the Shannon diversity index per treatment. Interestingly, we observed that the Burkholderia diversity was affected by changes in the agricultural management, since the highest diversity was observed in permanent grassland and in continuous arable land. In addition, the diversity tended to be higher in the rhizosphere than in the corresponding bulk soil. The use of species abundance models indicated that rhizosphere communities had more even distributions than communities collected from the bulk soil. Identification of isolates revealed that only 2% of these belonged to the B. cepacia complex and that the majority was assigned to either (1) new Burkholderia species or (2) Burkholderia species that had originally been isolated from soil. Isolates classified as B. hospita, B. caledonica and Burkholderia sp. ‘LMG 22934’ and ‘LMG 22936’ were found mainly in the arable land, while isolates belonging to Burkholderia sp. ‘LMG 22929’ and B. phytofirmans were associated with the grassland area. Another potentially new Burkholderia species, ‘LMG 22932’, was found in both areas, in close association with the maize rhizosphere.     

Temporary conservation for urban biodiversity

Urban habitats, particularly wastelands and brownfields, maintain rich biodiversity and offer habitat for many species, even rare and endangered taxa. However, such habitats are also under socio-economic pressures due to redevelopment for housing and industrial uses. In order to maintain urban biodiversity, it is currently unknown how much open area must be preserved and whether conservation is possible without complete exclusion from economic development. In this study, we applied a simulation model based on species distribution models for plants, grasshoppers, and leafhoppers to investigate planning options for urban conservation with special focus on business areas. Altogether, we modelled the occurrence of 81 species of the urban species pool and analysed settings of different proportions of open sites, different habitat turnover times, and different lot sizes. Our simulations demonstrated that dynamic land use supports urban biodiversity in terms of species richness and rarity. Setting aside brownfields before redevelopment for a period of on average 15 years supported the highest conservation value. Consequently, we recommend integrating the concept of ‘temporary conservation’ into urban planning for industrial and business areas. This concept requires habitat to be destroyed by redeveloping brownfield sites to built-up sites, but simultaneously creating new open spaces due to abandonment of urban land uses at other locations. This maintains a spatio-temporal mosaic of different successional stages ranging from pioneer to pre-forest communities.     

Fifty years of change in Central European grassland vegetation: Large losses in species richness and animal-pollinated plants

There is growing concern that biodiversity loss in European agricultural landscapes is having negative effects on functional trait diversity. Long-term studies examining vegetation changes from the period before agricultural industrialisation are however rare. Here, we ask how management intensification and increased nutrient input initiated in the 1950/1960s have altered grassland plant community composition, species diversity and functional trait composition using comprehensive datasets from five floodplain regions (plus one protected reference region) in northern Germany. Sites with available historical relevés and vegetation maps (1950/1960s, 1990s) were resampled in 2008 to facilitate the analysis of a period spanning four to five decades.Plant community composition changed tremendously in all study regions during the 50 year period, which was related to increasing Ellenberg indicator values for nutrient availability. Species richness at the plot-level fell by 30–50% over the period, and losses in functional diversity were equally large. A non-formal comparison with the results from the protected reference study region indicates that the changes may mostly be attributable to local nutrient input rather than to supra-regional climate change. Our results indicate a consistent trend towards much more species-poor communities dominated by mow-tolerant, N-demanding competitive grasses, whereas species with more ruderal strategies, species flowering early in the season and, in particular, insect-pollinated herbs have all decreased. The substantial loss of nectar-producing grassland herbs is likely to have negative effects on the abundance of pollinating insects, with consequences for the grassland animal communities. This highlights the growing need for adequate grassland management schemes with low N input to preserve high-nature-value grassland.