Effects of Restoration on Plant Species Richness and Composition in Scandinavian Semi-Natural Grasslands

Plant species richness in rural landscapes of northern Europe has been positively influenced by traditional management for millennia. Owing to abandonment of these practices, the number of species‐rich semi‐natural grasslands has decreased, and remaining habitats suffer from deterioration, fragmentation, and plant species decline. To prevent further extinctions, restoration efforts have increased during the last decades, by reintroducing grazing in former semi‐natural grasslands. To assess the ecological factors that might influence the outcome of such restorations, we made a survey of semi‐natural grasslands in Sweden that have been restored during the last decade. We investigated how plant species richness, species density, species composition, and abundance of 10 species that are indicators of grazing are affected by (1) the size of the restored site, (2) the time between abandonment of grazing and restoration, (3) the time elapsed since restoration, and (4) the abundance of trees and shrubs at the restored site. Only two factors, abundance of trees and shrubs and time since restoration, were positively associated with total species richness and species density per meter square at restored sites. Variation in species composition among restored sites was not related to any of the investigated factors. Species composition was relatively similar among sites, except in mesic/wet grasslands. The investigated factors had small effects on the abundance of the grazing‐indicator species. Only Campanula rotundifolia responded to restoration with increasing abundance and may thus be a suitable indicator of improved habitat quality. In conclusion, positive effects on species richness may appear relatively soon after restoration, but rare, short‐lived species are still absent. Therefore, remnant populations in surrounding areas may be important in fully recreating former species richness and composition.     

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.     

Passive restoration contributes to bird conservation in Brazilian Pampa grasslands

The global decline of biodiversity makes it important to find affordable ways to conserve and restore habitats. Restoration is useful for conserving native grasslands, with passive restoration defined as either natural colonization or unassisted recovery. Grasslands in southeastern South America have been transformed into croplands and impacted by other human activities. We describe the first assessment of passive restoration as a management tool to conserve birds in the Pampa grasslands of Brazil. We compared bird species richness using coverage‐based rarefaction and extrapolation, applying PERMANOVA for composition, and the abundance of bird communities between sites undergoing passive restoration (PR) and sites with native grasslands (NG). We employed fitted generalized linear mixed models (GLMM) to quantify relationships between bird occurrence and vegetation structure and cover. We recorded 61 species of birds during our study (45 in PR and 46 in NG) and 762 individuals (333 in PR and 429 in NG). Of these species, 15 were restricted to PR and 16 to NG. Grassland specialists and threatened species were found in both PR and NG, and only vegetation height differed between PR and NG. We detected eight species of conservation concern, including three recorded only in PR, three only in NG, and two in both PR and NG. The absence of marked differences in species richness and composition of bird communities between passive‐restoration and native grasslands in our study suggests that grasslands in the process of passive restoration can provide habitat for many species of grassland birds and that passive restoration is an appropriate management tool for biodiversity conservation in Brazilian grasslands.     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

The effects of grassland management and aspect on Orthoptera diversity and abundance: site conditions are as important as management

Calcareous grasslands represent local hotspots of biodiversity in large parts of Central and Northern Europe. They support a great number of rare species which are adapted to these xerothermic habitats. Due to massive changes in land use, calcareous grasslands have become a rare habitat type and their conservation has been given a high priority in the habitats directive of the European Union. It is well known that grassland management may affect biodiversity substantially. However, the quality of calcareous grasslands is also influenced by abiotic conditions, such as aspect (i.e. sun exposure), which affects the local mesoclimate. We studied the combined effects of aspect and grassland management on Orthoptera diversity on 16 sites in Central Germany, in an unbalanced crossed design with three factors: aspect, management type and management intensity. For both response variables (diversity and abundance) we obtained a similar pattern. South-facing pastures maintained a greater diversity than north-facing pastures, but both had a greater diversity than extensively used meadows. Intensively used meadows maintained the lowest diversity and abundances. A multivariate analysis revealed that the abundance of rare Orthoptera species correlated with bare ground cover and forb cover, both of which were greatest at south-facing pastures. Our results suggest that grazing is a more suitable management for maintaining a high biodiversity in calcareous grasslands than mowing. Moreover, the mesoclimate (in this studied measured by its surrogate: aspect) is a crucial factor determining species richness and needs to be considered in reserve planning.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Butterfly responses to environmental factors in fragmented calcareous grasslands

Although there is much research showing a strong negative effect of habitat fragmentation and deterioration on the viability of different insect populations and on species richness, the effect of fragmentation is modified by other local and landscape factors. One of the most substantial gaps in knowledge is whether species are similar in their response to the same environmental factors and if their response mirrors response of the entire community. From the conservation point of view this knowledge is of primary importance in planning conservation actions, yet these studies are rare. In this paper we test the relative effects of habitat patch and landscape characteristics on butterflies inhabiting calcareous grasslands in southern Poland. Butterfly species richness and abundance were positively affected by patch size and wind shelter. In the case of species richness there was also a positive effect of plant species richness. Butterfly diversity was enhanced in wind sheltered patches, and commonness (non-rarity) enhanced by distance to buildings and by shorter vegetation. Multivariate analysis suggested differences in the responses of individual species to the examined environmental variables, with some species more responsive to patch size and shelter and others to sward height. The conservation of butterfly communities requires sensible and complex management to ensure high habitat diversity. The most important challenge for future studies on calcareous grasslands is to formulate a model of management that guarantees high species richness and conservation of each individual species.     

Plant species richness in calcareous grasslands as affected by dispersability in space and time

Abstract. Species richness in calcareous grassland is discussed against the background of historical dispersal processes associated with traditional land-use management such as grazing, but also the artificial establishment by hayseed. Important vectors in the traditionally man-made landscape were sheep and cattle or other livestock such as goats. Calcareous grasslands were not only connected to each other but also to other habitats such as villages, forests, arable fields and heathlands by these vectors which could cover large distances (e.g. transhumance shepherding), which is not the case in the current man-made landscape. Species richness after restoration management of abandoned and afforested calcareous grasslands was predicted by using characters of dispersability in space and time. These were the persistence of the species in the vegetation and the diaspore bank after abandonment or afforestation and the dispersal capacity through wind and sheep. The results reveal that reintroduction of sheep grazing is necessary to reestablish the original species richness. The first validation of the prediction of the succession on clear-cut sites and a comparison with data of species composition in abandoned quarries and the surroundings made it obvious that a species' own dispersal capacity in space is very low except for some well wind-dispersed species. Therefore, it is recommended to include and to simulate dispersal processes in future management to be able to restore the original species richness.     

Dark diversity in dry calcareous grasslands is determined by dispersal ability and stress‐tolerance

Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat‐specific regional species pool may be absent from local communities and represent the ‘dark diversity’ of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat‐specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing). We analysed a total of 1223 vegetation plots (1 × 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co‐occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S‐strategy type scores were used to characterise low stress‐tolerance. Levels of small‐scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions. We conclude that, even if temperate grasslands have high levels of small‐scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress‐tolerance.     

Process‐based long‐term evaluation of an ecological network of calcareous grasslands connected by sheep herding

This paper evaluates the long‐term effect of an ecological network of calcareous grasslands, a habitat type that experienced dramatic habitat loss and fragmentation during the 20th century, on species richness of habitat specialist plants. Calcareous grasslands are of special conservation concern as the habitat type with the highest diversity in plant and invertebrate species in central Europe. A baseline survey in 1989 established complete vascular plant species lists for all 62 previously abandoned calcareous grassland patches in the study area and assessed the presence of 48 habitat specialist plant species. An ecological network was initiated in 1989 to reconnect these patches with existing grazed pastures (core areas) through large flock sheep herding where feasible, as sheep are thought to be the primary dispersal vectors for calcareous grassland plants. An evaluation survey in 2009 showed significant increase in species richness of habitat specialist plants in patches reconnected by sheep herding, indicating successful colonizations by habitat specialist plants, while ungrazed patches showed no significant change. Observed increase in species richness between 1989 and 2009 was related to connectivity by sheep herding and the presence of a diversity of structural elements providing microsites for establishment. Baseline species richness of the patches, which had been abandoned since at least 1960, was associated with patch area, supporting the effect of ecological drift, and with vegetation type, which suggests that delays in extinction may be related to site factors governing the strength of competition with later seral species. The implementation of this ecological network represents a long‐term ‘natural experiment’ with baseline data, manipulation, and evaluation of hypothesized effects on a clearly defined target variable. It thus provides much needed empirical evidence that species loss in fragmented calcareous grassland communities can be counteracted by restoring functional connectivity among remnant patches.     

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

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

Response of plant functional traits during the restoration of calcareous grasslands from forest stands

In this survey, we studied the response of plant functional traits to calcareous grassland restoration in the Calestienne region, Southern Belgium (restoration protocol: forest clear-cutting followed by grazing at all sites). We considered traits related to dispersal, establishment, and persistence that integrate the main challenges of plants to re-establish and survive in restored areas. Functional traits were compiled from databases and compared among (i) pre-restoration and young restoration forests; (ii) restoration areas of different ages; and (iii) old restorations and reference grasslands. The following questions were addressed: (i) What is the early response (2–4 years) in terms of plant functional trait following one restorative clear-cut event? (ii) What plants functional trait responses occur from restorative management (i.e., sheep and goat grazing)? (iii) Which differences still persist between the oldest restored parcels (10–15 years), and the historical reference grasslands? Forest clear-cuts induced several changes among functional traits, including decreased mean seed mass and certain vegetative traits (i.e., decreased phanerophytes, branching species; and increased short lifespan species i.e., annuals and biennials). During restorative management, clonal, epizoochorous and autumn germinating species were favored. Despite numerous other changes during this phase, many differences remained compared to reference grasslands. In particular, geophytes, mycorrhizal and evergreen species abundance were not approaching reference grassland values. The observed pattern helped to draw inferences on the possible mechanisms operating under vegetation recovery following restorative forest clear-cut and subsequent management were identified and described in this study. Results indicated grazing was an important factor, which increased epizoochorous species, and autumn germinating taxa that filled niches in vegetation opened by summer grazing animals. Finally, differences between old restoration and reference grasslands emphasized that management should focus on reduction in soil fertility, and geophyte rhizomatous grasses. Long-term monitoring is vital to assess if management plans are effective in the complete restoration of species functional trait assemblages.     

Threatened species in a vanishing habitat: plants and invertebrates in calcareous grasslands in the Swiss Jura mountains

We examined the diversity of vascular plants, butterflies, grasshoppers, gastropods and carabid beetles in three calcareous grasslands in the northwestern Swiss Jura mountains, a habitat type that has decreased dramatically during the recent decades. As many as 58 species (ca 22%) of the 266 collected are listed as threatened in northern Switzerland. The number of threatened species ranged from 27 to 49 per site, and 26 species occurred in only one of the three grasslands. The species richness of butterflies and vascular plants correlated positively among the grasslands, while the species richness of other groups did not covary. An index of complementarity indicated that the species compositions (including non-threatened species, and spiders and oribatid mites) of the groups varied greatly among the grasslands. In addition, herbivorous groups were more widely distributed than predators among the three grasslands. Due to this variation in species composition none of the three sites c an substitute for the others, if the biodiversity of these grasslands is to be maintained. Furthermore, the taxonomic groups studied are poor indicators of each others' diversity. Consequently, we support the shopping basket approach to conservation evaluation, i.e. measuring species richness, species composition and complementarity of several groups instead of just one.     

Asymmetric carabid beetle spillover between calcareous grasslands and coniferous forests

Central European calcareous grasslands are considered biodiversity hotspots, but are severely threatened by the change in land-use and by habitat fragmentation. Coniferous forests are typical adjacent habitats to calcareous grasslands, as abandoned calcareous grasslands are often afforested or develop into coniferous forests by succession. To investigate spillover between calcareous grasslands and coniferous forests, a total of 144 pitfall traps for carabid beetles were placed at three different distances (1, 5, 20 m) from the edge in both habitats at eight locations from April to late August. We found that both habitats had a distinct species assemblage and a decrease in spillover with increasing distance from the habitat edge into the adjacent habitat. Calcareous grasslands were more affected by spillover from the adjacent coniferous forests than vice versa because more forest specialists penetrated into calcareous grasslands than grassland specialists penetrated into coniferous forests. We conclude that spillover into small and isolated habitats can severely change species assemblages, which has to be considered in conservation measures. The protection of large sites with small edge-interior ratios can reduce negative effects on species assemblages in endangered calcareous grasslands.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

Heat shock increases the reliability of a temperate calcareous grassland seed bank study

Reliable assessment of the density and species richness of the viable seeds in the soil is essential to estimate the probability of successful restoration of a particular plant community. Since temperate calcareous grasslands contain several thermophilous species typical of fire prone habitats, heat shock can be expected to break dormancy of target species for calcareous grassland restoration. We tested the effect of heat shock on the estimated density and species richness of seeds in soil samples. Heat shock treatments between 70 and 110°C resulted in both a higher estimated seed density and estimated species richness, in particular of specialist calcareous grassland species. Moreover, 25 species germinated exclusively from the heat-treated samples. This indicates that several species would have been missed by handling soil seed bank samples following the standard germination protocol, resulting in an underestimation of the restoration potential.     

Land use history and site location are more important for grassland species richness than local soil properties

Lately there has been a shift in Sweden from grazing species‐rich semi‐natural grasslands towards grazing ex‐arable fields in the modern agricultural landscape. Grazing ex‐arable fields contain a fraction of the plant species richness confined to semi‐natural grasslands. Still, they have been suggested as potential target sites for re‐creation of semi‐natural grasslands. We asked to what extent does fine‐scale variation in soil conditions, management history and site location effect local plant diversity in grazed ex‐arable fields. We examined local soil conditions such as texture, pH, organic carbon, nitrogen (N) and extractable phosphate (P) and effects on plant richness in ten pairs of grazed ex‐fields and neighbouring semi‐natural grasslands in different rural landscapes. Each grassland pair where in the same paddock. A multivariate test showed that site location and land use history explained more of differences in species richness than local soil property variables. Plant species richness was positively associated to grazed ex‐fields with low pH, low N and P levels. Sites with high plant richness in semi‐natural grasslands also had more species in the adjacent grazed ex‐fields, compared to sites neighbouring less species‐rich semi‐natural grasslands. Although both soil properties and species richness were different in grazed ex‐fields compared to semi‐natural grassland, the site location within a landscape, and vicinity to species‐rich grasslands, can override effects of soil properties. In conclusion, if properly located, ex‐arable fields may be an important habitat to maintain plant diversity at larger spatio‐temporal scales and should considered as potential sites for grassland restoration.     

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.     

Does natural reforestation represent a potential threat to dung beetle diversity in the Alps?

Traditional agro-pastoral practices are in decline over much of the Alps, resulting in the complete elimination of livestock grazing in some areas. Natural reforestation following pastoral abandonment may represent a significant threat to alpine biodiversity, especially that associated with open habitats. This study presents the first assessment of the potential effects of natural reforestation on dung beetles by exploring the relationships between the beetle community (abundance, diversity, species turnover and assemblage structure) and the vegetation stages of ecological succession following pastoral abandonment. A hierarchical sampling design was used in the montane belt of the Sessera Valley (north-western Italian Alps). Dung beetles were sampled across 16 sampling sites set in four habitat types corresponding to four different successional stages (pasture, shrub, pioneer forest and beech forest) at two altitudinal levels. The two habitats at the extremes of the ecological succession, i.e. pasture and beech forest, had the greatest effect on the structure of local dung beetle assemblages. Overall, dung beetle abundance was greater in beech forest, whereas species richness, Shannon diversity and taxonomic diversity were significantly higher in pasture, hence suggesting this latter habitat can be considered as a key conservation habitat. Forests and pastures shared a lower number of species than the other pairs of habitats (i.e. species turnover between these two habitats was the highest). The two intermediate seral stages, i.e. shrub and pioneer forest, showed low dung beetle abundance and diversity values. Local dung beetle assemblages were also dependent on season and altitude; early-arriving species were typical of pastures of high elevation, whereas late-arriving species were typical of beech forests. It is likely that grazing in the Alps will continue to decrease in the future leading to replacement of open habitats by forest. This study suggests therefore that, at least in the montane belt, reforestation may have potentially profound and negative effects on dung beetle diversity. Maintaining traditional pastoral activities appears to be the most promising approach to preserve open habitats and adjacent beech forests, resulting in the conservation of species of both habitats.     

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.