Plant Trait Assembly Affects Superiority of Grazer's Foraging Strategies in Species-Rich Grasslands

Background

Current plant – herbivore interaction models and experiments with mammalian herbivores grazing plant monocultures show the superiority of a maximizing forage quality strategy (MFQ) over a maximizing intake strategy (MI). However, there is a lack of evidence whether grazers comply with the model predictions under field conditions.

Methodology/Findings

We assessed diet selection of sheep (Ovis aries) using plant functional traits in productive mesic vs. low-productivity dry species-rich grasslands dominated by resource-exploitative vs. resource-conservative species respectively. Each grassland type was studied in two replicates for two years. We investigated the first grazing cycle in a set of 288 plots with a diameter of 30 cm, i.e. the size of sheep feeding station. In mesic grasslands, high plot defoliation was associated with community weighted means of leaf traits referring to high forage quality, i.e. low leaf dry matter content (LDMC) and high specific leaf area (SLA), with a high proportion of legumes and the most with high community weighted mean of forage indicator value. In contrast in dry grasslands, high community weighted mean of canopy height, an estimate of forage quantity, was the best predictor of plot defoliation. Similar differences in selection on forage quality vs. quantity were detected within plots. Sheep selected plants with higher forage indicator values than the plot specific community weighted mean of forage indicator value in mesic grasslands whereas taller plants were selected in dry grasslands. However, at this scale sheep avoided legumes and plants with higher SLA, preferred plants with higher LDMC while grazing plants with higher forage indicator values in mesic grasslands.

Conclusions

Our findings indicate that MFQ appears superior over MI only in habitats with a predominance of resource-exploitative species. Furthermore, plant functional traits (LDMC, SLA, nitrogen fixer) seem to be helpful correlates of forage quality only at the community level.     

Correlations among anatomical,morphological, chemical and agronomic characteristics of leaf blades in Panicum maximum genotypes

Tropical forage grasses present high growth rates and biomass yields, partly due to its C₄ photosynthetic pathway. Considering this, the anatomy of the grasses related to morphology and chemical composition of leaf blades may influence consumption and digestibility, thus interfering in the forage quality. Agronomic, morphological, anatomical and chemical characteristics of leaves of nine Panicum maximum genotypes were evaluated in Brazil, to verify if these characteristics and the associations among them may influence the quality of the leaf blades. A randomized complete blocks design was used with nine treatments and three replications. Three evaluation harvests were done in the rainy season and one in the dry season to evaluate forage yields and quality. Quality was determined through NIRS – Near Infrared Spectrometer. One day prior to each harvest, four leaf blades per plot were harvested for morphological and anatomical evaluations. Results were subjected to analysis of variance and mean comparison by Tukey test, and to simple linear and canonical correlations by SAS. Leaf width was positively correlated with mesophyll. The parenchyma bundle sheath was associated with leaf area and specific leaf area. Neutral detergent fibre was positively correlated with the parenchyma bundle sheath area and specific leaf area. The specific leaf area was negatively correlated with in vitro organic matter digestibility. Morphological differences among P. maximum genotypes did not interfere in biomass accumulation. Considering this, leaf width may be a supplementary tool, that may be used in the early phases of the process of genotype selection, for discriminating qualitatively promising high yielding materials.     

Biomass production potential of grasslands in the oak savanna region of Minnesota, USA

Biomass harvested from grasslands formerly used for forage production or set aside for conservation has been identified as a potential source of bioenergy feedstocks. Our objective was to characterize yield and chemical composition of biomass harvested from existing grasslands in the oak savanna region of Minnesota and to determine whether aggregated soil properties and grassland type influence biomass yield and feedstock properties. The influence of soil type and dominant functional plant groups on biomass yield, theoretical ethanol yield, and mineral, ash, and lignocellulosic concentration was measured on biomass harvested from 32 grassland sites. Soils with high productivity ratings, as measured by the Minnesota Crop Productivity Index, produced 36 % more biomass than lower quality soils. Grasslands dominated by warm-season species produced 18 % more biomass than those dominated by cool-season species, when measured after senescence during the late-fall harvest time. Biomass harvested from sites dominated by cool-season grasses had higher N, Mg and Cl concentrations than those dominated by warm-season grasses, suggesting that such grasslands could have lower efficiency in thermochemical conversion processes and that repeated harvesting from such grasslands could remove nutrients from the systems. In addition, glucose and xylose concentrations were slightly higher in biomass from sites dominated by warm-season grasses, which resulted in an estimated additional 12 L?Mg^?1 of ethanol over those dominated by cool-season grasses. Combined with the greater yields, warm-season grasslands could produce an additional 376 L?ha^?1 year^?1.     

Grazing Lawns in Terai Grasslands,Royal Bardia National Park,Nepal1

We compared the community structure, nutritive quality, and aboveground biomass of grazing lawns (patches of shortgrass communities) to neighboring grasslands in the Terai of western Nepal. Grazing lawns differed from the adjacent grasslands in species composition and community structure. Species diversity and species richness were higher on grazing lawns (H = 1.60, S = 20.93) than the grasslands (H = 0.97, S = 8.97). Fencing that excluded grazers for 150 days made areas of grazing lawns indistinguishable from neighboring grasslands in terms of plant height and biomass. Growing shoots of forage from grazing lawns had higher digestibility, crude protein, and sodium than forage from the grasslands. Grazing lawns appear to be maintained by continuous grazing and are enriched by deposition of urine, dung, and by certain plant species not found in the the adjacent grasslands.     

Nutritive value of papyrus (Cyperus papyrus,Cyperaceae), a tropical emergent macrophyte

Papyrus (Cyperus papyrus, Cyperaceae) covers large areas in tropical African wetlands. Analysis of its nutritive value has revealed that crude protein is higher in umbels than culms. Ruminai dry matter digestibility of papyrus is, however, higher in culms than umbels. Both the crude protein and ruminai dry matter digestibility decrease with increasing age of the plant. Values for crude protein and ruminai dry matter digestibility are similar to those reported for the range grasses that constitute the greatest percentage of forage in East Africa. In general, papyrus has some grazing potential and could be used as fodder especially in the dry season when other forage is scarce and of low nutritive value.     

Isolation, characterization and fibre degradation potential of anaerobic rumen fungi from cattle

A total of 20 fungal cultures were isolated from the rumen of cattle fed a high fibre-containing diet. All of the isolates showed polycentric growth patterns and were identified as different strains of Orpinomyces and Anaeromyces. Enzyme assays of most of the isolates showed the highest carboxymethylcellulase (CMCase) and xylanase activities after 96 h of growth and highest avicelase activity after 120 h. Among all enzymes tested, xylanase activity was the highest, followed by CMCase and avicelase. The results of the in vitro fibre digestibility and rumen fermentation analyses revealed that the addition of fungal cultures significantly increased acetate, in vitro dry matter digestibility, partition factor values and microbial biomass synthesis levels. Overall, Orpinomyces spp. were found to be the better enzyme producers and fibre degraders than Anaeromyces spp.     

Effect of fencing and grazing on a Kobresia-dominated meadow in the Qinghai-Tibetan Plateau

Grazing is one of the most important factors influencing community structure and productivity in natural grasslands. Fencing to exclude grazers is one of the main management practices used to protect grasslands. Can fencing improve grassland community status by restraining grazing? We conducted a field community study and indoor soil analyses to determine the long-term effects of fencing and grazing on the above-ground community and soil in a Kobresia-dominated meadow in the Qinghai-Tibetan Plateau, NW China. Our results showed that fencing significantly improved above-ground vegetation productivity but reduced plant density and species diversity. Long-term fencing favored the improvement of forage grass functional groups and restrained the development noxious weed functional groups. There were significant positive effects of fencing on below-ground organic matter, total nitrogen, available nitrogen, total phosphorus and available phosphorus. The productivity of grazed meadow showed a weak decrease over time. There were long-term decreasing trends for plant density both in fenced and grazed meadows. Our study suggests that grazing can be considered as a useful management practice to improve species diversity and plant density in long-term fenced grasslands and that periodic grazing and fencing is beneficial in grassland management.     

Positive versus negative environmental impacts of tree encroachment in South Africa

Woody plant encroachment in grasslands is a worldwide phenomenon. Despite many studies, the consequences of woody plant encroachment on sub-canopy vegetation and soil properties are still unclear. To better understand the impacts of trees on grassland properties we examined the following questions using a mountainous sub-tropical grassland of South Africa encroached by an indigenous tree, Acacia sieberiana as a case study: (1) Do trees increase sub-canopy herbaceous diversity, quality and biomass and soil nitrogen content? (2) Do large trees have a stronger effect than medium-sized trees on grass and soil properties? (3) Does the impact of trees change with the presence of livestock and position of trees in a catena? We studied grass and non-graminoid species diversity and biomass, grass quality and soil properties during the wet season of 2009. Nitrogen in grass leaves, soil cation exchange capacity and calcium and magnesium ion concentrations in the soil increased under tall Acacia versus open areas. Medium-sized Acacia decreased the gross energy content, digestibility and neutral detergent fibre of grasses but increased the species richness of non-graminoids. Tall and medium Acacia trees were associated with the presence of Senecio inaequidens, an indigenous species that is toxic to horses and cattle. The presence of livestock resulted in a decrease in herbaceous root biomass and an increase in soil carbon and leaf biomass of grass under Acacia. Tree position in the catena did not modify the impact of trees on the herbaceous layer and soil properties. For management of livestock we recommend retaining tall Acacia trees and partially removing medium-sized Acacia trees because the latter had negative effects on grass quality.     

Chemical composition of the water fern,salvinia molesta,and its potential as feed source for ruminants

The chemical composition of the now widespread tropical aquatic fern Salvinia molestawas analysed with regard to its suitability as a source of forage for ruminants. Three different stages of plant growth collected in Kerala, India, were air dried, brought to Germany, and used for Weende analysis, detergent fibre analysis, and the determination of gross energy, amino acids, tannins, and minerals. The concentration of 12.4% crude protein in sand-corrected dry matter (DM) and of some other nutrients in Salviniais comparable to that in conventional forage. However, the high content of crude ash (17.3% in DM) and of lignin (13.7%) and the presence of tannins (0.93%) may reduce acceptance as well as digestibility and therefore restrict the use of Salviniaas a potential feed source for ruminants.     

Changes in light- and nitrogen-use and in aboveground biomass allocation patterns along productivity gradients in grasslands

Light- and nitrogen-use change was examined along productivity gradients in natural grasslands at Laelatu, western Estonia, both at community level and in most abundant species. Aboveground biomass (M) ranged from 341 to 503 g m^?2 in wet (W) and from 248 to 682 g m^?2 in dry (D) community. Aboveground leaf area ratio (aLAR) decreased with rising M in D site, while it increased in W site. In a high-aLAR W community (significantly higher compared to D), adjustment of leaf morphology through an increase in specific leaf area is responsible for an increase in aLAR with rising productivity. In low-aLAR stand, by contrast, adjustment of biomass allocation due to decrease in aboveground leaf mass fraction is primarily responsible for the tendency of aLAR to decline. In conclusion, a decrease in aLAR is not a universal response to increasing M. We hypothesise that there exists an optimum of light acquisition efficiency (Φ_M) along a productivity gradient independent of community type. Aboveground nitrogen-use efficiency (aNUE) decreased in high-aLAR, W community with increasing M, while in low-aLAR, D site, there was no relationship along a gradient, although aNUE increased along six plots dominated by graminoids. A trade-off was established between leaf nitrogen content per unit leaf area (N _A) and aLAR.     

Molecular breeding of transgenic perennial ryegrass (<Emphasis Type="Italic">Lolium perenne</Emphasis> L.) with altered fructan biosynthesis through the expression of fructosyltransferases

Perennial ryegrass is the most important forage grass used in temperate agriculture. Transgenic perennial ryegrass events with altered fructan biosynthesis have the potential not only to increase animal production by improving digestibility of the grasses, but also to increase pasture intake by the animal due to lower neutral detergent fibre (NDF) concentrations. Transgenic perennial ryegrass plants were shown to have increased (P?<?0.05) in fructan concentrations of leaf blades in transgenic T₀ events and have thus an increase in water-soluble carbohydrate and in vivo dry matter digestibility concentrations as well as a decrease in the NDF concentration within the plant in spring and summer. These changes in nutritive value have led to an increase in metabolisable energy of up to 1.7 MJ ME·kg DM^?1 in selected T₀ events compared to FLp418-20 during spring and summer, with no differences in autumn or winter. The field evaluation of these events and the further development of these events using molecular breeding technologies are described in this paper.     

Effects of grazing on plant species diversity and pasture quality in subalpine grasslands in the eastern Pyrenees (Andorra): Implications for conservation

Mountain areas boast high levels of biodiversity due to the wide variety in topography, climates and traditional land-use practices they embrace. Recent changes in the mountain agro-ecosystems of Europe have led to a decline in these traditional practices (above all grazing) and have had negative effects on biodiversity. In this study we evaluated the effects of grazing pressure on plant species diversity and also focused on the common and rare species diversity, forage quality and forage production in the Festuca airoides, Festuca eskia, Festuca gautieri, Festuca paniculata and Nardus stricta grassland communities. These analyses could be of great importance in identifying the pastoral management practices that are most consistent with the conservation of plant diversity in the Andorran subalpine and alpine grasslands. Data were collected at 113 sampling sites exposed to either high or low grazing pressures. In all, 222 vegetation line-transects were established from which samples were collected and analysed using the NIRS method. In addition the lignin, protein and fibre fractions were obtained in order to evaluate the quality of the forage. In the Andorran summer pastures, the Festuca gautieri calcareous grasslands had the greatest plant species diversity, while the Festuca eskia, Festuca airoides and Nardus stricta grasslands had the lowest plant species diversity and a lack of common species due to their less homogeneous spatial structure. The highest quality forage corresponds to the Festuca gautieri and Nardus stricta grasslands, while the Festuca eskia grasslands have the lowest quality forage. Although high grazing pressure can lead to greater species diversity, it also tends to diminish forage quality and/or energy content by increasing the lignin content. Thus, domestic livestock is a fundamental element in these grasslands and an increase in the grazing pressure of the different types of livestock (cattle, sheep and horses) is necessary since a more balanced distribution of grazers between the grassland types – rather than an increase in grazers – will not be sufficient to maintain plant biodiversity. Likewise, there is a need to re-establish the practice of transhumance, a traditional pastoral practice based on the movement of livestock between winter lowland pastures and upland summer pastures.     

Distribution,density and impact of invasive plants in Lake Nakuru National Park,Kenya

Invasive plants have invaded swathes of grasslands in Lake Nakuru National Park thus necessitating the Park management to institute measures to control them. Despite this, information on the status and impact of invasive plants in these grasslands is lacking. Six grassland types were identified and assigned random numbers. Five study sites were then randomly selected from each grassland type. The Point‐Centre Quarter method was used to determine the distribution and density of seven major invasive plants, whilst their impact was assessed by comparing biomass and crude protein content of important forage grasses in invaded and noninvaded grasslands. Results show that the distribution of invasive plants did not differ among the grasslands (F_{5, 30} = 1.47, P = 0.229) but their densities differed among the plants (F_{6, 30} = 20.99, P < 0.001). The mean biomass in invaded and noninvaded grasslands (22 g m^?2 versus 37.2 g m^?2) was significantly different (t‐test, P < 0.05), while crude protein content of grasses in invaded (7.73, % DM) and noninvaded (9.3, % DM) patches was not significantly different (t‐test; P > 0.05). Our results suggest that invasive plants lower grass production and hence might reduce food availability to grazers.     

Impact of chicory inclusion in a cereal-based diet on digestibility,organ size and faecal microbiota in growing pigs

A total of 30 7-week-old pigs were used to evaluate the effects of chicory inclusion on digestibility, digestive organ size and faecal microbiota. Five diets were formulated: a cereal-based control diet and four diets with inclusion of 80 and 160 g/kg chicory forage (CF80 and CF160), 80 g/kg chicory root (CR80) and a mix of 80 g/kg forage and 80 g/kg chicory root (CFR). Generally, the pigs showed a high growth rate and feed intake, and no differences between the different diets were observed. The coefficients of total tract apparent digestibility (CTTAD) of energy, organic matter and CP did not differ between the control and CF80, whereas they were impaired in diet CF160. The CTTAD of non-starch polysaccharides and especially the uronic acids were higher (P < 0.05) with chicory inclusion, with highest (P < 0.05) values for diet CF160. Coliform counts were lower and lactobacilli : coliform ratio was higher (P < 0.05) in diet CFR than in the control. Global microbial composition was investigated by terminal restriction fragment length polymorphism combined with cloning and sequencing. Analysis of gut microbiota pattern revealed two major clusters where diet CF160 differed from the control and CR80 diet. Chicory forage diets were correlated with an increased relative abundance of one species related to Prevotella and decreased abundance of two other species related to Prevotella. For diet CFR, the relative abundance of Lactobacillus johnsonii was higher than in the other diets. This study shows that both chicory forage and root can be used as fibre sources in pig nutrition and that they modulate the composition of the gut microbiota differently.     

贝加尔针茅响应降水变化敏感指标及关键阈值

降水是影响中国北方草原群落结构和生产力的关键环境因子,弄清植物响应降水变化敏感指标,确定关键阈值,可为草原的科学管理与制定应对气候变化措施提供依据。以贝加尔针茅为材料,基于不同降水的模拟实验,采用正态总体统计容忍区间及容忍限的估算方法,对测定的叶面积、株高、叶绿素和叶N含量等14个植物生理生态指标响应降水变化的敏感性及其阈值进行了研究。结果表明,植株总叶面积、水势(或叶绿素含量)和地上生物量可分别从形态、生理和生物量积累三方面较好地指示贝加尔针茅对降水变化的敏感性,受到胁迫时的降水量(6、7和8月3个月份总降水量)阈值分别是283、276(276)mm和280 mm。研究结果可为客观辨识贝加尔针茅草原干旱的发生发展与监测预警提供参考。     

Root traits and soil properties in harvested perennial grassland,annual wheat,and never-tilled annual wheat

Background and aims

Root functional traits are determinants of soil carbon storage; plant productivity; and ecosystem properties. However, few studies look at both annual and perennial roots, soil properties, and productivity in the context of field scale agricultural systems.

Methods

In Long Term and Conversion studies in North Central Kansas, USA; root biomass and length, soil carbon and nitrogen, microbial biomass, nematode and micro-arthropod communities were measured to a depth of one meter in paired perennial grassland and cropland wheat sites as well as a grassland site that had been converted to cropland using no tillage five years prior.

Results

In the Long Term Study root biomass was three to seven times greater (9.4 Mg ha^?1 and 2.5 Mg ha^?1 in May), and root length two times greater (52.5 km m^?2 and 24.0 km m^?2 in May) in perennial grassland than in cropland. Soil organic carbon and microbial biomass carbon were larger, numbers of Orbatid mites greater (2084 vs 730 mites m^?2), and nematode communities more structured (Structure Index 67 vs 59) in perennial grassland versus annual cropland. Improved soil physical and biological properties in perennial grasslands were significantly correlated with larger, deeper root systems. In the Conversion Study root length and biomass, microbial biomass carbon, mite abundance and nematode community structure differed at some but not all dates and depths. Isotope analysis showed that five years after no-till conversion old perennial roots remained in soils of annual wheat fields and that all soil fractions except coarse particulate organic matter were derived from C₄ plants.

Conclusions

Significant correlation between larger, longer roots in grasslands compared to annual croplands and improved soil biological, physical and chemical properties suggest that perennial roots are an important factor allowing perennial grasslands to maintain productivity and soil quality with few inputs. Perennial roots may persist and continue to influence soil properties long after conversion to annual systems.     

Short-term regrowth responses of four steppe grassland species to grazing intensity, water and nitrogen in Inner Mongolia

Inner Mongolia steppe grasslands are widely used for livestock farming and the regrowth ability of grassland species is therefore strongly influenced not only by water and nutrient availability but also quite heavily by grazing. However, little is known on how grazing, water and nitrogen interactively affect the dominant C3 species (Leymus chinensis, Stipa grandis, Agropyron cristatum) and the C4 species (Cleistogenes squarrosa). Therefore in the 2007 and 2008 growing seasons, a field experiment was carried out to test the hypothesis that under different grazing intensities the dominant species show different short-term regrowth response to simultaneous variation in the availability of water and nitrogen. Single factor and interaction effects of the addition of water (rainfed vs. simulated wet-year) and nitrogen (0 or 25 kg N ha^?1) were analysed along a gradient of four grazing intensities (ungrazed, lightly, moderately and heavily grazed) after one month of grazing exclusion. Water and nitrogen addition affected short-term regrowth of all species in a similar way whereas species responded differently to grazing. Simulated wet-year water availability consistently resulted in higher standing biomass, relative growth rate and cellulase digestible organic matter yield. Supplementary nitrogen promoted standing biomass and crude protein concentration. The nutritive value of all species’ standing biomass showed a similar increase with more intensive grazing. However, heavy grazing led to a clear shift in the relative biomass of the species, i.e. mainly a promotion of the C4 grass, C. squarrosa. In contrast to our hypothesis, there were no differences among species in their response to water or nitrogen addition, whereas, heavy grazing induced the expected species-specific response. Our results suggest that heavy grazing rather than nitrogen or water determine short-term shifts in species composition of the investigated steppe ecosystem. Furthermore, differences in the species-specific growth response to grazing may increase the proportion of the C4 grass C. squarrosa in steppe communities, whereas higher availability of nitrogen and water may lead to higher forage biomass and nutritive value of all investigated species but in short-term cannot compensate for the grazing induced changes in species composition.     

内蒙古草甸草原与典型草原地下生物量与生产力季节动态及其碳库潜力

地下根系是草原生态系统的重要组成部分,其生物量及其净生产力对地下碳库具有直接与间接作用,分析地下生物量季节动态与周转对深入揭示草原生态系统碳库动态及其固碳速率与潜力具有重要意义。应用钻土芯法对不同利用方式或管理措施下内蒙古草甸草原、典型草原地下生物量动态及其与温度、降水的相关性研究表明:草甸草原和典型草原地上生物量季节动态均为单峰型曲线,与上月降水显著正相关(P0.05),但地下生物量季节动态表现为草甸草原呈"S"型曲线,典型草原则是双峰型曲线,与温度、降水相关性均不显著(P0.05);两种草原根冠比和地下生物量垂直分布均为指数函数曲线,根茎型草原地下生物量集中在土壤0—5 cm,丛生型草原地下生物量集中于土壤5—10 cm,根冠比值在生长旺季(7—8月份)最小。草甸草原地下净生产力及碳储量范围分别为2167—2953 g m-2a-1和975—1329 gC m-2a-1,典型草原为2342—3333 g m-2a-1和1054—1450 gC m-2a-1,地下净生产力及其碳储量约为地上净生产力及其碳储量的10倍,具有较大的年固碳能力,且相对稳定;地下净生产力与地上净生产力呈显著负相关性(P0.05);地下生物量碳库是地上生物量碳库的10倍左右,适度放牧可增加地下生产力,但长期过度放牧显著降低其地下生物量与生产力,并使其垂直分布趋向于浅层化。     

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.