Recreating semi-natural grasslands: A comparison of four methods

Semi-natural grasslands and their species and populations are declining rapidly throughout Europe, bringing about a need for successful vegetation recreation methods. To maintain biodiversity and ecological services of semi-natural grasslands, we need more knowledge on the relative performance of different recreation methods. In a replicated experiment in western Norway, we evaluated two hay transfer methods (hard or light raking of local hay), sowing of local seeds and natural regeneration for recreating semi-natural grassland in a road verge. We compared treated trial plots with their respective donor plots (where hay and seeds were harvested) for three successive years by evaluating vegetation cover, species richness and species transfer rates, and vegetation dynamics analysed by Bray–Curtis compositional dissimilarity (BC) and GNMDS (Global Non-Metric Multidimensional Scaling) ordination. Vegetation cover at the trial site exceeded that of donor sites in three years. Transfer rates of common species were high for seed sowing and both hay transfer procedures. Species composition in trial plots for all three treatments became significantly more similar to donor plots, but was still relatively dissimilar after three years. Natural regeneration showed a different temporal pattern and also had a higher successional rate. The species composition of the other treatments followed the same trajectory toward the donor sites as revealed by GNMDS. We found relatively small differences between the two hay transfer methods and seed sowing. Transfer of local hay therefore appears to be a successful method of establishing local species when recreating semi-natural grasslands, and is generally cheaper than using commercial local seed mixtures.     

Can grassland plant communities be preserved on road and railway verges?

Abstract. Semi‐natural grasslands are a threatened biotope type in many countries. Typical grassland plant species are adapted to continuous grazing or mowing that keeps the environment open. With the decline in grassland area, these species are ever more reliant on alternative habitats such as road verges. To find out whether plant communities comparable to those of semi‐natural grasslands can be maintained on modern road and railway sides, the vegetation of 92 grasslands and 90 road and railway verges was studied. These biotope types were compared with each other according to their community structure, total number of species, number of grassland species and restricted‐range diversity. Further study of the vegetation of road and railway verges was carried out in order to identify the treatments and environments which are most likely to support diverse plant communities. The species number and the restricted‐range diversity proved to be higher next to roads and railways than on grasslands. Grassland species were, however, most abundant on grasslands. Furthermore, the community structure of these biotope types was totally divergent. In their present state, road and railway verges are not a substitute for semi‐natural grasslands. Nevertheless, the occurrence of grassland species in verges may be enhanced by a suitable mowing regime, by giving up the use of de‐icing salt and herbicides and by allowing natural establishment of vegetation on the verges.     

Species introduction in restoration projects – Evaluation of different techniques for the establishment of semi-natural grasslands in Central and Northwestern Europe

During recent decades, many studies have shown that the successful restoration of species-rich grasslands is often seed-limited because of depleted seed banks and limited seed dispersal in modern fragmented landscapes. In Europe, commercial seed mixtures, which are widely used for restoration measures, mostly consist of species and varieties of non-local provenance. The regional biodiversity of a given landscape, however, can be preserved only when seeds or plants of local provenance are used in restoration projects. Furthermore, the transfer of suitable target species of local provenance can strongly enhance restoration success.We review and evaluate the success of currently used near-natural methods for the introduction of target plant species (e.g. seeding of site-specific seed mixtures, transfer of fresh seed-containing hay, vacuum harvesting, transfer of turves or seed-containing soil) on restoration sites, ranging from dry and mesic meadows to floodplain grasslands and fens. Own data combined with literature findings show species establishment rates during the initial phase as well as the persistence of target species during long-term vegetation development on restoration sites.In conclusion, our review indicates that seed limitation can be overcome successfully by most of the reviewed measures for species introduction. The establishment of species-rich grasslands is most successful when seeds, seed-containing plant material or soil are spread on bare soil of ex-arable fields after tilling or topsoil removal, or on raw soils, e.g. in mined areas. In species-poor grasslands without soil disturbance and on older ex-arable fields with dense weed vegetation, final transfer rates were the lowest. For future restoration projects, suitable measures have to be chosen carefully from case to case as they differ considerably in costs and logistic effort. Long-term prospects for restored grassland are especially good when management can be incorporated in agricultural systems.     

The Restoration of Phytophagous Beetles in Species‐Rich Chalk Grasslands

This study focuses on the restoration of chalk grasslands over a 6‐year period and tests the efficacy of two management practices, hay spreading and soil disturbance, in promoting this process for phytophagous beetles. Restoration success for the beetles, measured as similarity to target species–rich chalk grassland, was not found to be influenced by either management practice. In contrast, restoration success for the plants did increase in response to hay spreading management. Although the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonized, few beetle species colonized as soon as their host plants became established. Morphological characteristics and feeding habits of 27 phytophagous beetle species were therefore tested to identify factors that limited their colonization and persistence. The lag time between host plant establishment and colonization was greatest for flightless beetles. Beetles with foliage‐feeding larvae both colonized at slower rates than seed‐, stem‐, or root‐feeding species and persisted within the swards for shorter periods. Although the use of hay spreading may benefit plant communities during chalk grassland restoration, it did not directly benefit phytophagous beetles. Without techniques for overcoming colonization limitation for invertebrate taxa, short‐term success of restoration may be limited to the plants only.     

Restoration of species‐rich grasslands by transfer of local plant material and its impact on species diversity and genetic variation—Findings of a practical restoration project in southeastern Germany

Restoration of species‐rich grasslands is a key issue of conservation. The transfer of seed‐containing local plant material is a proven technique to restore species‐rich grassland, since it potentially allows to establish genetically variable and locally adapted populations. In our study, we tested how the transfer of local plant material affected the species diversity and composition of restored grasslands and the genetic variation of the typical grassland plant species Knautia arvensis and Plantago lanceolata.For our study, we selected fifteen study sites in southeastern Germany. We analyzed species diversity and composition and used molecular markers to investigate genetic variation within and among populations of the study species from grasslands that served as source sites for restoration and grasslands, which were restored by transfer of green hay and threshed local plant material.The results revealed no significant differences in species diversity and composition between grasslands at source and restoration sites. Levels of genetic variation within populations of the study species Knautia arvensis and Plantago lanceolata were comparable at source and restoration sites and genetic variation among populations at source and their corresponding restoration sites were only marginal different.Our study suggests that the transfer of local plant material is a restoration approach highly suited to preserve the composition of species‐rich grasslands and the natural genetic pattern of typical grassland plant species.     

Fitness in Naturally Occurring and Restored Populations of a Grassland Plant Lychnis flos‐cuculi in a Swiss Agricultural Landscape

Wildflower seed mixtures are widely used for restoration of grasslands. However, the genetic and fitness consequences of using seed mixes have not been fully evaluated. Here, we studied the role of genetic diversity, origin (commercial regional seed mixtures, natural populations), and environmental conditions for the fitness of a grassland species Lychnis flos‐cuculi. First, we examined the relationship between genetic diversity, environmental parameters, and fitness in sown and natural populations of this species in a Swiss agricultural landscape. Second, we established an experiment in the study area and in an experimental garden to study the implications of local adaptation for plant fitness. Third, to examine the response of plants to different soil properties, we conducted an experiment in climate chambers, where we grew plants from sown and natural populations of L. flos‐cuculi as well as from seed suppliers on soils with different nutrient and moisture content. We detected no significant effect of genetic diversity on the fitness of sown and natural populations. There was no clear indication that plants from natural populations were better adapted to local environment than plants from sown populations or seed suppliers. However, plants of natural origin invested more into generative reproduction than plants from sown populations or seed suppliers. Furthermore, in the climate chamber, plants originating from natural populations tended to flower earlier. Our results indicate that using nonlocal seeds for habitat recreation may influence restoration success even if the seeds originate from the same seed zone as the restored site.     

Grassland restoration on former croplands in Europe: an assessment of applicability of techniques and costs

Grasslands used to be vital landscape elements throughout Europe. Nowadays, the area of grasslands is dramatically reduced, especially in industrial countries. Grassland restoration is widely applied to increase the naturalness of the landscape and preserve biodiversity. We reviewed the most frequently used restoration techniques (spontaneous succession, sowing seed mixtures, transfer of plant material, topsoil removal and transfer) and techniques used to improve species richness (planting, grazing and mowing) to recover natural-like grasslands from ex-arable lands. We focus on the usefulness of methods in restoring biodiversity, their practical feasibility and costs. We conclude that the success of each technique depends on the site conditions, history, availability of propagules and/or donor sites, and on the budget and time available for restoration. Spontaneous succession can be an option for restoration when no rapid result is expected, and is likely to lead to the target in areas with high availability of propagules. Sowing low-diversity seed mixtures is recommended when we aim at to create basic grassland vegetation in large areas and/or in a short time. The compilation of high-diversity seed mixtures for large sites is rather difficult and expensive; thus, it may be applied rather on smaller areas. We recommend combining the two kinds of seed sowing methods by sowing low-diversity mixtures in a large area and high-diversity mixtures in small blocks to create species-rich source patches for the spontaneous colonization of nearby areas. When proper local hay sources are available, the restoration with plant material transfer can be a fast and effective method for restoration.     

Top‐soil translocation as a technique in the re‐creation of species‐rich meadows

Abstract. In France, most civil engineering and excavation projects are at present accompanied by compensatory measures with the aim of preserving biodiversity. In order to avoid the destruction of a habitat of high conservation interest in NE France, harbouring two legally protected plant species, an experiment of soil translocation was conducted on an area of 1 ha. The donor site was an extensively managed mesophilic meadow and the receiving site was a neighbouring arable land. The vegetation of the translocated meadow was described 8 and 17 months after soil translocation, and compared (1) with vegetation resulting from more classical restoration techniques tested on the arable land (natural regeneration and seed mixture sowing) and (2) with the soil seed bank and vegetation previously present on the donor site. Results showed that the soil translocation technique permitted the development of many meadow species, including two legally protected species, and few ruderal species despite a large area of bare ground. This technique seems effective in terms of number and abundance of meadow species compared to natural regeneration and commercial seed sowing. In the case of the two classical methods, species richness was lower and only widespread species were present. Topsoil translocation provides a good compensatory method to avoid habitat and species destruction. However, the study should be continued, with the aim of assessing the longer term development and stabilization of the vegetation of the translocated meadow.     

Evaluation of Initial Restoration Measures during the Restoration of Calcareous Grasslands on Former Arable Fields

In 1993, experiments on the restoration of calcareous grasslands on ex‐arable fields were started in order to provide new habitats for species of a small nature reserve with ancient grasslands north of Munich (Germany). The effects of diaspore transfer by the application of seed‐containing hay on vegetation establishment were studied on restoration fields with and without topsoil removal for 5 years. The aim of the study was to assess plant diversity for the evaluation of restoration success by different methods including determination of species with viable seeds in the hay by germination tests, phenological investigations on hay‐transfer source sites at the time of harvest, and vegetation analyses on the restoration sites. Total seed content of the hay and the number and composition of plant species with viable seeds were affected by the time of harvesting and differed between a site which had been used as arable field until 1959 and ancient grassland sites. Nevertheless, the number of established hay‐transfer species showed only few differences between restoration fields. The proportion of species transferred to restoration fields in relation to the number of species with viable seeds in the hay was between 69 and 89%. Five years after the hay transfer, the proportion of the established species was still between 58 and 76%. Up to now, topsoil removal had no significant effect on the number of established hay‐transfer species. After triple hay application the absolute number of transferred grassland species was higher than on sites with single hay application, but restoration efficiency was lower because many of the species with viable seeds in the hay did not establish. In general, our results showed that the transfer of autochthonous hay is a successful method to overcome dispersal limitation in restoration projects.     

Grassland restoration after afforestation: No direction home?

The coastal grasslands in north‐eastern South Africa are a severely threatened vegetation type rich in plant species, particularly forbs. Many of the forbs have underground storage organs which allow them to resprout rapidly after fires. A significant portion of this land was placed under commercial pine afforestation in the 1950s. The pine plantations have since been removed starting 17 years ago and restored to grasslands within a conservation area. We assessed the effects of plantations on grassland plant diversity and functional trait composition by sampling 64 circular quadrats of 5 m radius distributed equally in restored versus natural grasslands. The difference in plant diversity was dramatic with the natural grassland supporting 221 species of which 163 were forbs compared with 144 and only 73 forb species in restored grasslands. Major differences in species composition were recorded, especially for forb species. Natural grasslands were dominated by resprouters (130 species) but these were rare in the restored grasslands (36 species). Differences in plant species response to fire were also evident for the two grassland states. In contrast to coastal forest restoration studies in the same area which have shown near linear increases in woody species with time, restored grasslands showed no increase in forb species richness with increasing time since clear‐felling of pines. Our results indicate that current methods for restoring these grasslands are inadequate and that restoring grasslands may be a lot harder than previously thought. Considerable effort should be made in conserving what is left of natural grasslands.     

Restoration of Seminatural Grasslands: What is the Impact on Ants?

The number of species‐rich seminatural grasslands in Northern Europe has decreased significantly due to the abandonment of traditional land use practices. To preserve these habitats, an increasing number of abandoned and overgrown grasslands have been restored by cutting down trees and shrubs and reintroducing grazing. These practices are considered a useful tool to recover the species richness of vascular plants, but their impact on other taxa is hardly known. Here we studied ants as one important group of grassland insects. We investigated (1) the effects of restoration of nongrazed and afforested seminatural grasslands, compared to continuously managed reference sites; and (2) the modulating impacts of habitat characteristics and time elapsed since restoration. We found a total of 27 ant species, 11 of these were characteristic of open habitats and seven characteristic of forests. Neither species richness per site nor the number of open‐habitat species, nor the number of forest species differed between restored and reference sites. Yet, within the restored sites, the total species richness and the number of open‐habitat species was positively related to the time since restoration and the percentage of bare rock. High frequencies of most open‐habitat species were associated with low vegetation, older restored sites, and reference sites. Most forest species showed their highest frequencies in tree‐ and shrub‐dominated habitat. We conclude that restoration efforts have been successful in terms of retrieving species richness. A regular and moderate grazing regime subsequent to the restoration is suggested in order to support a high abundance of open‐habitat species.     

Successful restoration of abandoned terraced vineyards and grasslands in Southern Switzerland

Extensively managed semi-natural grasslands represent species-rich habitats and therefore play a key role for the maintenance of biodiversity in agricultural areas. In marginal and poorly accessible areas, the traditional management of grassland is frequently abandoned, which leads to the spread of forest. In Southern Switzerland, terraced vineyards (a special grassland type) and terraced grasslands are part of the cultural heritage and local biodiversity hotspots. Yet, many of them are overgrown by forest. In the past years, several abandoned terraced vineyards and grasslands have been restored by removing the forest, rebuilding the walls and re-introducing the traditional management. We examined restoration success by assessing plant species richness, diversity and species composition in both the aboveground vegetation and soil seed bank in (1) restored, (2) abandoned for 25–50 years, and (3) permanently used areas of six terraced vineyards and six terraced grasslands. Plant species richness and diversity were reduced and species composition altered in the aboveground vegetation of abandoned vineyards and grasslands compared to the permanently used and restored ones. However, species richness, Shannon-diversity and species composition of the aboveground vegetation did not differ between restored and permanently used areas, indicating a successful restoration of the vegetation 10–15 years after restoration. In abandoned vineyards, species richness of plants emerging from the soil seed bank was slightly higher than in permanently used and restored vineyards. No difference in seedling species richness was found between abandoned, permanently used and restored terraced grasslands. Our results showed that the soil seed bank played a minor role for the re-establishment of the above-ground vegetation. We assume that the large species pool in the surroundings and the presence of dispersal vectors are essential for the successful passive restoration of abandoned grassland in this region.     

Large‐Scale Restoration of Dry Grasslands on Ex‐Arable Land Using a Regional Seed Mixture: Establishment of Target Species

Large‐scale (circa 500 ha) restoration of species‐rich dry grasslands was conducted using a high‐diversity regional seed mixture in the White Carpathians Protected Landscape Area and Biosphere Reserve, Czech Republic, Central Europe. After sowing, the restored grasslands were regularly mown. Vegetation was analyzed at sites restored 1–12 years ago and compared with that of ancient, extremely species‐rich grasslands nearby. Nearly all (98%) sown target species successfully established and nearly half of unsown target species established spontaneously, partly dependent on distance to the ancient grasslands. Early mowing in the first half of June appeared to support species diversity and broad‐leaved forbs at the expense of competitive grasses. Using a regional seed mixture appeared to be an effective way of restoring dry grasslands and is especially recommended in the proximity of still existing ancient grasslands where spontaneous establishment of unsown target species may reinforce the success of restoration more easily.     

Strengths and potential pitfalls of hay transfer for ecological restoration revealed by RAD‐seq analysis in floodplain Arabis species

Achieving high intraspecific genetic diversity is a critical goal in ecological restoration as it increases the adaptive potential and long‐term resilience of populations. Thus, we investigated genetic diversity within and between pristine sites in a fossil floodplain and compared it to sites restored by hay transfer between 1997 and 2014. RAD‐seq genotyping revealed that the stenoecious floodplain species Arabis nemorensis is co‐occurring with individuals that, based on ploidy, ITS‐sequencing and morphology, probably belong to the close relative Arabis sagittata, which has a documented preference for dry calcareous grasslands but has not been reported in floodplain meadows. We show that hay transfer maintains genetic diversity for both species. Additionally, in A. sagittata, transfer from multiple genetically isolated pristine sites resulted in restored sites with increased diversity and admixed local genotypes. In A. nemorensis, transfer did not create novel admixture dynamics because genetic diversity between pristine sites was less differentiated. Thus, the effects of hay transfer on genetic diversity also depend on the genetic make‐up of the donor communities of each species, especially when local material is mixed. Our results demonstrate the efficiency of hay transfer for habitat restoration and emphasize the importance of prerestoration characterization of microgeographic patterns of intraspecific diversity of the community to guarantee that restoration practices reach their goal, that is maximize the adaptive potential of the entire restored plant community. Overlooking these patterns may alter the balance between species in the community. Additionally, our comparison of summary statistics obtained from de novo‐ and reference‐based RAD‐seq pipelines shows that the genomic impact of restoration can be reliably monitored in species lacking prior genomic knowledge.     

Mass effects,clonality, and phenology but not seed traits predict species success in colonizing restored grasslands

Preserved grasslands commonly host species‐rich plant communities but a large part of the grasslands were plowed up in the past. Their restoration often requires a long time and initial restoration measures might trigger ecosystem recovery, which is then followed by spontaneous colonization. We evaluate the establishment success of target grassland species, which were not sown but established spontaneously in the restored grasslands of Bílé Karpaty Mts., SE Czech Republic. According to their key functional traits and incidence in the landscape (mass effect; acquired from the results of a grid mapping project in the region), we examined the frequency of species and their mean cover in 82 restored grasslands. The best predictor of species frequency in the grasslands was their mass effect, followed by a high capacity for clonal growth and late phenology. Seed dispersal traits (seed mass, terminal velocity, epizoochory ranking index) and plant height had no significant effect. Specific leaf area was positively correlated with mass effect. Species having a high cover in the restored grasslands had a high capacity for clonal growth. In the preparation of seed mixtures, we should therefore consider that nonclonal species relying on regeneration from seeds will be generally less able to reproduce and should be promoted by artificial sowing. At the same time, species common in the landscape, which spread well clonally, and those with a late phenology, might be expected to colonize restored meadows on their own, so that sowing them is not necessary.     

Problems, Approaches, and Results in Restoration of Dutch Calcareous Grassland During the Last 30 Years

This paper is based on research of the restoration of species‐rich calcareous grasslands in The Netherlands, over the last 30 years. Chalk grassland is a semi‐natural vegetation with a high density of species at a small scale. This type of vegetation was once widespread in Western Europe as common grazing land, mainly for flocks of sheep for which the main function was dung production. In some regions of Central Europe, these grasslands were also used for hay production. The dung was used to maintain arable field production at a reasonable level. In the chalk district in the southernmost part of The Netherlands some 25 sites of this vegetation, varying in area from 0.05–4.5 ha, are still present. Chalk grassland completely lost its significance for modern agricultural production after the wide application of artificial fertilizer following World War II. This grassland has a high conservation value both for plants and animal species, of which a large number of species are exclusively restricted to this biotope. When conservation activities started at a large scale in the early 1960s, three different types of restoration activities could be distinguished: (1) restoration of fertilized sites; (2) restoration of abandoned grasslands; and (3) recreation of chalk grassland on former arable fields. The main aim of the restoration attempt is to create and/or improve sustainable conditions for both plant and animal species characteristic of the chalk grassland ecosystem. In the process of restoration, several phases of different activities can be distinguished: (1) pre‐restoration phase, during which information of the land use history is collected and, based on these data, clear restoration goals are established; (2) initial restoration phase, during which effects of former, non‐conservational land use has to be undone in order to stimulate germination and establishment of target species originating from soil seed bank and species pool; (3) consolidation phase, including the introduction and continuation of a regular management system for sustainable conservation; and (4) long‐term conservation strategy, including measures to prevent disturbance from the outside and genetic erosion and extinction of locally endangered plant populations.     

Grazing response patterns indicate isolation of semi‐natural European grasslands

Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species‐rich semi‐natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on‐going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades. It is a largely unresolved question to what degree current temperate grassland communities already reflect a decline of regional processes such as longer distance seed dispersal. Answering this question is challenging since it requires both a mechanistic approach to community dynamics and a sufficient data basis that allows identifying general patterns. Here, we present results of a local individual‐ and trait‐based community model that was initialized with plant functional types (PFTs) derived from an extensive empirical data set of species‐rich grasslands within the ‘Biodiversity Exploratories’ in Germany. Driving model processes included above‐ and belowground competition, dynamic resource allocation to shoots and roots, clonal growth, grazing, and local seed dispersal. To test for the impact of regional processes we also simulated seed input from a regional species pool. Model output, with and without regional seed input, was compared with empirical community response patterns along a grazing gradient. Simulated response patterns of changes in PFT richness, Shannon diversity, and biomass production matched observed grazing response patterns surprisingly well if only local processes were considered. Already low levels of additional regional seed input led to stronger deviations from empirical community pattern. While these findings cannot rule out that regional processes other than those considered in the modeling study potentially play a role in shaping the local grassland communities, our comparison indicates that European grasslands are largely isolated, i.e. local mechanisms explain observed community patterns to a large extent.     

Grassland connectivity by motor vehicles and grazing livestock

In addition to habitat loss and fragmentation, agricultural change has led to a change in seed dispersal processes in the rural landscape through a loss of structural and functional connectivity. Here, human‐mediated dispersal vectors are prevalent, and we explored whether the loss of connectivity via free‐ranging livestock could be mitigated by the increase in roads and motor vehicles. We found that structurally, 39% of all valuable semi‐natural grassland habitats in southern Sweden are adjacent to public road verges, which in the rural landscape are often considered to be suitable habitat for grassland species. Additionally, by collecting mud attached to cars and farming machinery and manure from livestock (cattle, horse, sheep) grazing semi‐natural grassland pasture, we found that motor vehicles are also capable seed dispersers. A similar number of species were dispersed by both vectors, although the composition of samples was quite different. Motor vehicles dispersed more grassland specialists than invasive species, although in much lower abundances than did grazing livestock. Despite these differences, motor vehicles were found to be able to disperse species with the same kinds of dispersal traits as livestock. A high number of seeds, species and specialists in manure samples means that greater movement of livestock is desirable to increase functional grassland connectivity. However, effective management could improve the suitability of roadsides as grassland corridors and increase the availability of seeds for long‐distance human‐mediated dispersal via cars and tractors. Our results suggest that in many rural landscapes, connectivity by road networks could help mediate habitat loss and fragmentation of grasslands. However, such effects can be context dependent, and the connectivity provided by roads could have serious negative consequences in other regions.     

Restoration of upland hay meadows over an 11‐year chronosequence: an evaluation of the success of green hay transfer

Grassland restoration has become a key tool in addressing the drastic losses of seminatural grassland since the mid‐twentieth century. This study examined the restoration by green hay transfer of upland hay meadows, a particularly scarce and vulnerable habitat, over an 11‐year chronosequence. The community composition of 18 restoration meadows was compared with that of donor reference sites in two study areas in the Pennine region of northern England. The study investigated: differences in community composition between donor and restoration meadows; transfer of upland hay meadow target species; and the effect of time and isolation from neighboring meadows on the community composition of the restoration meadows. Results showed that restoration meadows differed from donor meadows in that some target species were easily transferred whilst others were not found in the restoration meadows, or were at low levels of cover. Time had a significant effect on the community composition of the restoration meadows, but the similarity between restoration sites and donor sites did not increase with time, and the effect of isolation was not significant. The study showed that the green hay transfer method increases botanical diversity and is an important first step in meadow restoration. However, further restoration activity, such as seed addition, is likely to be required if restoration sites are to resemble closely the reference donor sites.     

Scale‐dependence of vegetation‐environment relationships in semi‐natural grasslands

Questions: Which environmental and management factors determine plant species composition in semi‐natural grasslands within a local study area? Are vegetation and explanatory factors scale‐dependent? Location: Semi‐natural grasslands in Lærdal, Sognog Fjordane County, western Norway. Methods: We recorded plant species composition and explanatory variables in six grassland sites using a hierarchically nested sampling design with three levels: plots randomly placed within blocks selected within sites. We evaluated vegetation‐environment relationships at all three levels by means of DCA ordination and split‐plot GLM analyses. Results: The most important complex gradient determining variation in grassland species composition showed a broad‐scale relationship with management. Soil moisture conditions were related to vegetation variation on block scale, whereas element concentrations in the soil were significantly related to variation in species composition on all spatial scales. Our results show that vegetation‐environment relationships are dependent on the scale of observation. We suggest that scale‐related (and therefore methodological) issues may explain the wide range of vegetation‐environment relationships reported in the literature, for semi‐natural grassland in particular but also for other ecosystems. Conclusions: Interpretation of the variation in species composition of semi‐natural grasslands requires consideration of the spatial scales on which important environmental variables vary.