Status,future prospects,and management recommendations for alkaline fens in an agricultural landscape: A comprehensive survey

Due to agricultural intensification and cessation of traditional land use, alkaline fens of the Caricion davallianae alliance (EU-FFH 7230) are among the most endangered ecosystems in Europe. This study exhibits a vegetation analysis of these systems in Schleswig-Holstein (Northern Germany). We analyzed across three scales the effects of grazing and mowing on phytodiversity of core areas and recorded their adjacent vegetation to estimate the capability for habitat enlargement of fen species. Results revealed that species richness, evenness and number of endangered species varied insignificantly between mowing and grazing treatments, regardless of scale. The high proportion of fen species and Red-Book-listed species in core areas, along with a state-wide representation of only 2 ha of these vegetation types, underlines the need for further conservation measures. Floristic differences between grazed and mowed sites derived from the individual appearance of species within one treatment. Thus, to preserve the species pool of alkaline fens, both management strategies have to be considered. Moreover, as most small sedge reed species are low-productive and light-demanding, sufficient biomass removal of dominant tall-growing species is required. One cut per year in late summer, the traditional management, does not satisfy the requirements of target species, which is indicated by an increase of Phragmitetea species at larger plot sizes. Additionally, in grazed fens, vegetation adjacent to the core areas consisted mainly of more eutrophic wet grasslands, and, in mowed fens, mainly of reeds or woods. We recommend therefore several changes in current conservation measures that include an increase to two cuts per year in mowed fens, and stocking rates of at least 2 LU ha⁻¹ (summer grazing) or 0.7 LU ha⁻¹ (year-round grazing).     

Functional diversity analysis helps to identify filters affecting community assembly after fen restoration by top-soil removal and hay transfer

Top-soil removal followed by species introduction through hay transfer has appeared as a method to restore drained fens. This method addresses abiotic constraints by restoring hydrology and nutrient status, and biotic constraints by removing an unwanted seed bank and counteracting dispersal-limitation. Restoration works by altering environmental filters. Knowledge about the restoration actions effect on functional traits is necessary to understand which types of species may establish. In this study we analyse which factors in top-soil removal followed by hay transfer influence selection and composition of functional traits. Top-soil removal followed by hay transfer from reference sites was conducted at two sites in the Całowanie fen, 33 km SE of Warsaw, Poland. Species and abundance data were recorded for three consecutive years. These data, combined with data on functional traits were used to analyse the effect of the restoration actions on four functional diversity-indices and the community weighted mean of functional traits. Our results reveal a strong habitat filter in the restoration site that follows an elevation gradient. At low elevation this filter selects low values of autochory and specific leaf area and high values of clonal lateral spread, Ellenberg moisture values, and dispersal through hydrochory. The transferred hay differs in trait characteristics compared to the reference site vegetation by having species of higher specific leaf area, lower Ellenberg moisture value and lower dispersal by autochory and hydrochory. The result presented here has three important implications for fen restoration. First, the difference in trait-characteristics between the transferred hay and the reference site it was harvested from limits the restoration potential. Second, since for several fen species important functional traits are filtered along an elevation-gradient, careful planning regarding depth of top-soil removal is needed. Finally the results illustrate how a functional analysis can be used to detect environmental filters acting during ecological restoration.     

Shrub removal facilitates recovery of wetland species in a rewetted fen

Drainage and shrub expansion are the main threats to the biodiversity of fens and fen meadows, whereas rewetting and the removal of shrubby species are frequently applied restoration measures. We examine whether removal of shrubs enhances recovery of target species in a degraded fen subjected to moderate rewetting. The study was located in the drained fen Ca?owanie (central Poland), where remnants of open fen communities and willow-invaded fens exist in former turf-pits, surrounded by degraded meadows on dried peat. All these three habitat types were included in a monitoring grid, which covered an area of 2.2 ha. Within 55 quadrats of 20 m × 20 m we monitored occurrence of 52 species, i.e. two groups of target species (fen indicators and wet meadow indicators) and indicators of two failure scenarios (degraded fen indicators and eutrophic wetland indicators), during six years following shrub removal, rewetting and re-application of conservational mowing, using a 3-step ordinal abundance scale. NMDS ordination revealed a gradual convergence of shrub removal plots and reference plots. We noticed significant effects of year and habitat type on all indicator groups, but only fen indicators have shown a clear (increasing) trend within shrub removal plots. Degraded fen indicators (ruderal and opportunistic species) initially expanded on shrub removal plots, but this effect disappeared in the following years. We conclude that shrub removal enhances establishment of target species in a moderately drained and then rewetted fen and attribute this effect to lowered competition for light. However, given high costs of this method and long-lasting problems with shrub resprouts, we recommend applying shrub removal only to recently overgrown sites, which still retained high botanical diversity. Heavily degraded fen meadows did not react on the increase of moisture, which indicates that more advanced restoration measures, such as top soil removal are needed there.     

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.     

Relationship between light availability and species richness during fen grassland succession

The complex relation between standing crop and species richness in herbaceous plant communities is still one of the most intensively discussed topics in vegetation ecology. In this study, we focus on the extent to which light availability is able to explain this relationship in fen grasslands. We analyse changes of light availability (measured as relative irradiance, RI), standing crop and species richness during fen grassland succession. Our study include communities representing differences in drainage intensity and nutrient status (‘land use intensification sere’) and successional stages developing after abandonment (‘abandonment seres’). Both in the land use intensification sere and in the abandonment seres, we recorded an increase in standing crop reaching highest values of approx. 800 g m^‐2. RI, species richness and number of small‐growing species declined in the successional seres. RI was highest in stands of low‐productive communities, where light compensation points (5% RI) were already attained at soil surface and light saturation points (30% RI) at soil surface or at 15 cm height. Horizontal heterogeneity of RI at soil surface and at 30 cm height decreased in the abandonment seres, but not at 30 cm height in the land use intensification sere. Furthermore, we detected a signiñcant negative relation between standing crop and RI. Species richness declined with increasing standing crop and thus consequently also with decreasing RI. This result points out that light competition might be of importance for this pattern. The number of small‐growing species decreased exponentially with an increase in standing crop (and decrease in RI). It can be assumed that low light availability negatively affects small‐growing species at standing crop values higher than 400–500 g m². As the maintenance of small‐growing species and the improvement of habitat conditions for their establishment are important aims of nature conservation in fen grasslands, management strategies should be orientated at maintaining standing crops that are not higher than these values.     

Convergence in plant traits between species within grassland communities simplifies their monitoring

Plant trait measurement is a very powerful and promising method for assessing the effects of land use change on ecosystem behavior in grasslands, but it is very time-consuming. Hence we pose the following questions for simplifying diagnosis and monitoring: (i) are plant traits (PTs) similar between plant life forms (PLF: grasses, rosettes, upright forbs, legumes) within a plant community? (ii) is it possible to define the main plant community characteristics by measuring traits on one PLF or a limited number of dominant species?Six PTs known for their ability to characterize the capacity of species to exploit resource-rich or -poor environments and for their competitive dominance in response to disturbance (specific leaf area (SLA), leaf dry matter content (LDMC), plant height (H), C and N contents, flowering time) were measured on the species of 18 plant communities located in Central Pyrenees. The experiment combined 2 fertility levels and 3 defoliation regimes (cutting, grazing). Comparisons were made between the weighted values at community, PLF and two dominant species levels. Regression analysis shows that there were significant correlations between grasses and rosettes for 4 PTs. For H, N and C:N ratio, data for both grass and rosette PLFs were close to the bisecting line. The largest difference in the intercept was observed for LDMC. On the basis of plant traits weighted for all the species, plant communities were ranked in similar ways for SLA and H (Spearman r > 0.93; p < 0.001) and to a lesser extent for LDMC (r = 0.72; p < 0.001). Convergence in weighted plant traits for different PLFs within a plant community mean that in the studied grasslands, defoliation regime and nutrient availability act as strong filters that impose, at least at PLF level, very similar PFTs. This determines a specific local community structure and composition. An application of this result in managed grasslands is the possibility of focusing on one PLF or a limited number of species for vegetation diagnosis and monitoring.     

Woodland regeneration on grazed former arable land: A question of tolerance,defence or protection?

Ecological restoration of native woodlands and wooded pastures on former agricultural land is an important topic in modern conservation practice. The introduction of large herbivores is increasingly used to achieve these aims. We investigated how grazing, resistance traits of plants (concerning herbivory) and associational resistance interact and affect the establishment pattern of woody species on abandoned arable land (N-Belgium, W-Europe). In these early successional tree assemblages, we tested whether grazing increased or decreased spatial heterogeneity, which is supposed to be a crucial factor for biodiversity.With repeated measurements, 2-3 and 5-7 years after the cessation of agricultural use, we sampled 87 grazed and 56 ungrazed plots (314 m²) in 14 sampling areas (former arable land) on nutrient rich, (sandy) loam soils. We recorded established tree frequencies, related to grazing, time, resistance traits and unpalatable/spiny vegetation cover in the herb and low shrub layer. We investigated horizontal and vertical heterogeneity using variances in establishment frequencies and variances in frequencies of trees that were able to grow beyond the browse line, respectively.We found massive colonisation of grazing tolerant and resistant woody species in early successional stages. Grazing decreased frequencies and height of the most abundant tolerant species (mainly Salix caprea L.). After 5-7 years, frequencies of defensive and tolerant species were equal, but the former (mainly Betula pendula Roth) were able to grow beyond the browse line. When the cover of unpalatable/spiny vegetation was high enough (>60% of plot size), it also provided suitable nurse sites for tolerant species to grow out. In early assemblages, grazing increased horizontal and vertical heterogeneity, resulting in intermediate successional stages. In the long-term, the mechanism of associational resistance will also allow non-resistant and tolerant species to grow beyond the browse line and promote forest succession and the order of species establishment and replacement.     

Rapid restoration of a species-rich ecosystem assessed from soil and vegetation indicators: The case of calcareous grasslands restored from forest stands

Calcareous grasslands have long been recognized as biodiversity hotspots in Europe. However, in recent decades these ecosystems have seen rapid decline. In Belgium, more than 100 ha of calcareous grasslands have been restored from oak coppices and pine forests since the 1990s. The aim of the present study was to provide a quantitative assessment of the success of these restoration efforts, using two sets of indicators: one related to soil conditions, the other related to vascular plant communities. Soil conditions were evaluated by comparing soil samples from pre-restoration forest stands, restored grasslands (3-age classes: 2–4 years; 5–8 years, and 10–15 years) and reference grasslands. The analysis revealed no significant differences in soil N, P, and K contents between pre-restoration forests and restored and reference grasslands. We observed a decrease in the mineralization rate indicators in both pre-restoration forests and recent grassland restorations, which was resorbed in older restorations. Floristic surveys revealed that plant species composition of older restorations was most like reference grasslands. However, some differences in species composition persisted after 15 years. Moreover, a few rare species did not colonize restored grasslands despite a close seed source. Non-recolonization by a set of species expected on calcareous grasslands may be due to dispersal limitation and higher cover by native invasive grasses in restored parcels. These results were discussed in term of implications for management.     

Measuring the Short‐term Success of Grassland Restoration: The Use of Habitat Affinity Indices in Ecological Restoration

Agricultural intensification threatens grasslands worldwide and the restoration of grasslands from arable lands can at least partially counter this threat. We studied grassland restoration by following early successional changes of arthropod assemblages (spiders, Araneae; true bugs, Heteroptera; orthopterans, Orthoptera; and ground beetles, Carabidae) on 1‐ and 2‐year‐old restorations using arable lands and native grasslands as two ends of the succession timescale. To examine the changes in species composition among the habitat types, we used habitat affinity indices based on fidelity and/or specificity of the species. We found that the number of species did not differ between habitat types, while species composition changed markedly with time. Species richness was thus not adequate to detect favorable changes after grassland restoration. Habitat affinity indices, on the other hand, were useful to detect compositional changes caused by the increasing numbers of species characteristic of target grasslands as early as the second year after restoration. Habitat affinity indices are easy‐to‐use, easy‐to‐interpret measures of restoration success; therefore, we recommend their use as measures complementary to species richness and simple similarity. Our results show that sowing low‐diversity seed mixture followed by mowing and grazing can be particularly successful in grassland restoration in time periods as short as 2 years.     

Life-history traits and habitat preferences of colonizing plant species in long-term spontaneous succession in abandoned gravel–sand pits

The following question was asked: Species of what traits and ecological demands are the best and the least successful colonizers in the course of succession in disused gravel–sand pits?Abandoned gravel–sand pits (36) were studied in the Czech Republic. Together 224 phytosociological relevés were recorded in 5 m×5 m plots in all available seral stages ranging in age from 1 to 75 years. Complete lists of species occurring in (semi-)natural habitats were surveyed up to a distance of 100 m from each relevé. The colonization success of each species was expressed by an index between 0 and 1 which was obtained as the ratio of the number of relevés with species present in a pit/the number of relevés with the species occurrence in their surroundings. Species characteristics were taken from available databases. Data were elaborated by ordination analysis and regression tree analysis. Generally, the best colonizers of gravel–sand pits were hydrophytes with the ability to vegetatively reproduce. In particular, at the beginning of succession, anemochorous species played the most important role. Stress tolerant species, possessing light diaspores, were typical of nitrogen poor and acidic habitats. Later, sciophilous and nitrophilous species of mesic habitats increased in importance. They have heavier diaspores, and are mostly phanerophytes or geophytes, often with the ability to vegetatively reproduce as well as exhibiting higher demand for pH. Plant functional traits were recognized as powerful tools to predict the colonization success of plants available in the local species pool. They may help predict vegetation succession in various human-disturbed sites and thus be used in various restoration programmes.     

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

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

Vegetation structure and photosynthesis respond rapidly to restoration in young coastal fens

Young coastal fens are rare ecosystems in the first stages of peatland succession. Their drainage compromises their successional development toward future carbon (C) reservoirs. We present the first study on the success of hydrological restoration of young fens. We carried out vegetation surveys at six young fens that represent undrained, drained, and restored management categories in the Finnish land uplift coast before and after restoration. We measured plant level carbon dioxide (CO₂) assimilation and chlorophyll fluorescence (Fv/Fm) from 17 most common plant species present at the sites. Within 5 years of restoration, the vegetation composition of restored sites had started to move toward the undrained baseline. The cover of sedges increased the most in response to restoration, while the cover of deciduous shrubs decreased the most. The rapid response indicates high resilience and low resistance of young fen ecosystems toward changes in hydrology. Forbs had higher photosynthetic and respiration rates than sedges, deciduous shrubs, and grasses, whereas rates were lowest for evergreen shrubs and mosses. The impact of management category on CO₂ assimilation was an indirect consequence that occurred through changes in plant species composition: Increase in sedge cover following restoration also increased the potential photosynthetic capacity of the ecosystem. Synthesis and applications. Restoration of forestry drained young fens is a promising method for safeguarding them and bringing back their function as C reservoirs. However, their low resistance to water table draw down introduces a risk that regeneration may be partially hindered by the heavy drainage in the surrounding landscape. Therefore, restoration success is best safeguarded by managing the whole catchments instead of carrying out small‐scale projects.     

Modelling biomass of mountainous grasslands by including a species composition map

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

Community assembly and biomass production in regularly and never weeded experimental grasslands

We studied the natural colonisation of new species in experimental grasslands varying in plant species richness (from 1 to 60) and plant functional group richness (from 1 to 4) in either regularly or never weeded subplots during the first 3 years after establishment. Sown species established successfully, with no differences in species richness or their relative abundances between the regularly and never weeded subplots during the study period. Aboveground biomass of sown species increased with increasing sown species richness in both treatments. While a positive relationship between sown species richness and total aboveground biomass (including colonising species) existed in the 2nd year after sowing in the regularly and never weeded subplots, this positive relationship decayed in the 3rd year in the never weeded subplots because of a higher biomass of colonising species in species-poor mixtures. Total aboveground biomass varied independently of total species richness 3 years after sowing in both treatments. Jaccard similarity of coloniser species composition between regularly and never weeded subplots decreased from the 2nd to the 3rd year, indicating a divergence in coloniser species composition. Coloniser immigration and turnover rates were higher in regularly weeded subplots, confirming that weeding counteracts species saturation and increases the chance that new colonisers would establish. Although our study shows that low diversity plant communities are unstable and converge to higher levels of biodiversity, the effects of initially sown species on community composition persisted 3 years after sowing even when allowing for succession, suggesting that colonising species mainly filled empty niche space.     

Proximity of valuable habitats affects succession patterns in abandoned quarries

The study tested the hypothesis that the composition of vegetation formed during primary succession in basalt quarries is affected by the distance to, and area of, conservation-valuable biotopes of surrounding xerophilous grasslands. The successional vegetation was recorded in 270 relevés collected in 34 quarries in the area of Ceske Stredohori Hills, Czech Republic. We used detrended correspondence analysis to visualise the relationship between successional vegetation, ages of individual sites, and distances to the closest xerophilous grasslands. Subsequent regression analyses of fidelities of individual reléves to the grassland alliances Festucion valesiacae and Allyso-Festucion pallentis corroborated the view that the probability of development of valuable habitats within the quarries decreased with distance to the closest grassland sites, and increased with their area. It also increased with successional age, but this effect was suppressed if quarry identity was considered as covariable in the regressions. Our results show that the valuable biotopes would eventually develop in quarries situated less than 100 m from adjoining xerophilous grasslands. We advocate that quarry operators pay attention to conservation management of biotopes that surround excavation sites, because maintaining valuable vegetation in the vicinity will eventually reduce costs of post-excavation restoration.     

Effects of land-use change on productivity depend on small-scale plant species diversity

Understanding the interplay between land-use change, species diversity and ecosystem function is critical for the prediction of global change impacts on ecosystem services. Biodiversity experiments with artificial species assemblages have shown that community-scale species richness may affect ecosystem productivity and spatial stability. However, the use of synthetic communities with controlled levels of species density for biodiversity experiments has been criticised and their relevance for natural communities has been questioned. Here, we use a land-use change experiment to investigate the biodiversity effects on production within managed, upland grasslands. We examine species diversity and productivity at both the small plant-neighbourhood scale (14×14 cm) and the field scale (15 m×25 m) for two land-use trajectories under field conditions: intensification through fertilisation, and extensification through the cessation of mowing. Both intensification and extensification were associated with a decrease in species number, but the magnitude of this decrease was greater at the small scale. Extensification was associated with a decrease in small-scale productivity whereas intensification had no significant effect on small-scale productivity. Effects of land-use treatments on biomass production were mediated by variation in small-scale species number; species number showed a significant positive relationship with small-scale productivity within each land-use treatment. Furthermore, species number was associated with a decrease in the variance of small-scale green biomass. In contrast, no species diversity effects were found on productivity at the field scale. Instead, field-scale species diversity decreased with increase in the total above-ground biomass (green biomass+litter). This study demonstrates that biodiversity effects can be observed under field conditions at the small scale and may play an important role for ecosystem functioning and stability even in low-diversity plant communities.     

The quality of flower-based ecosystem services in field margins and road verges from human and insect pollinator perspectives

We contrasted traditionally used indicators of service provision quality, such as overall species richness and growth form composition, to three more specific functional properties: functional diversity, functional intensity, and functional stability. We defined flower colour as a functional trait perceived differently by humans and insect pollinators, and used user specific colour richness, flower size, and species richness within colour group as indicators of these three properties. We asked (1) do field margins and road verges provide flower-based ecosystem services with the quality of permanent grasslands; and (2) do traditional and detailed functional indicators of service provision quality agree on the service quality ranking of habitats?In an agricultural landscape of central and south-eastern Estonia (115 ÿ 95 km area, centroid 26°49⿲43⿳ and 58°54⿲49⿳) we sampled 87 field margins and 111 road verges as linear grassland-substitution habitats, and 84 permanent grasslands to scale their service quality.Linear habitats generally provided service of lower quality than permanent grasslands, but detailed indicators showed less evident contrast among habitat types than the overall species richness and stronger contrast than the proportion of forbs. Detailed indices, however, had strong seasonal dynamics to take into account. Vegetation in the first year field margins had greater colour richness (functional diversity) and species richness within colour groups (functional stability), but the smallest flower size (functional intensity), in contrast to road verges. By the third year of succession, field margins had become more similar to road verges. Indication of service provision quality differed between humans and pollinators, but their estimates were correlated across habitats.We showed that (1) combinations of specific service quality indicators provide more adequate information than generalized richness or growth form system, and (2) single grassland surrogate habitat type is an insufficient service providing substitute for permanent grasslands, although a mosaic of these habitats might be more efficient. Therefore, remnant fragments of semi-natural grasslands should receive top priority attention for conservation and restoration, particularly in agriculture dominated landscapes.     

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.     

Congruence among vascular plants and butterflies in the evaluation of grassland restoration success

Using surrogate taxa in the monitoring of restoration success is strongly advocated but currently under debate. Are butterfly and vascular plant communities good surrogates for each other in monitoring the restoration of calcareous grassland grassland? The research was conducted in a small area in Tuscany (central Italy), which was the focus of a restoration project. We sampled vascular plants using 35 plots (1 m²) and butterflies using 20 transects (100 m in length) belonging to four treatment types: control scrubland, restored ex-arable, restored grassland and reference grassland. A mixed model nested ANOVA showed that reference grassland, 3 years after restoration, had the highest number of vascular plant species and the highest number of butterfly species. The different treatments were found to be distinct from a compositional point of view: pRDA showed that the “treatment” factor, independent of spatial variables, accounted for 22.1 and 21.9% of the total variance in the plant community and butterfly community, respectively (p < 0.01). The butterfly community was significantly correlated with the entire plant community data set and with the phanerophyte community (i.e. shrubs); larval host plants were significantly correlated with the composition of non-migratory butterflies. The results indicated that butterfly and vascular plant communities were congruent, suggesting the use of community composition of different taxa as a valuable tool for the evaluation of management success.