Plants, insects and birds in semi-natural pastures in relation to local habitat and landscape factors

The preservation of remaining semi-natural grasslands in Europe has a high conservation priority. Previously, the effects of artificial fertilisation and grazing intensity on grassland animal and plant taxa have been extensively investigated. In contrast, little is known of the effects of tree and shrub cover within semi-natural grasslands and composition of habitats in the surrounding landscape on grassland taxa. We evaluated the effect that each of these factors has on species richness and community structure of vascular plants, butterflies, bumble bees, ground beetles, dung beetles and birds surveyed simultaneously in 31 semi-natural pastures in a farmland landscape in south-central Sweden. Partial correlation analyses showed that increasing proportion of the pasture area covered by shrubs and trees had a positive effect on species richness on most taxa. Furthermore, species richness of nectar seeking butterflies and bumble bees were negatively associated with grazing intensity as reflected by grass height. At the landscape level, species richness of all taxa decreased (butterflies and birds significantly so) with increasing proportion of urban elements in a 1-km² landscape area centred on each pasture, while the number of plant and bird species were lower in landscapes with large proportion of arable fields. Our results differed markedly depending on whether the focus was on species richness or community structure. Canonical correspondence analyses (CCA) showed that the abundance of most taxa was ordered along a gradient describing tree cover within pastures and proportion of arable fields in the landscape. However, subsets of grassland birds and vascular plants, respectively, showed markedly different distribution patterns along axis one of the CCA. In contrast to current conservation policy of semi-natural pastures in Sweden, our results strongly advise against using a single-taxon approach (i.e., grassland vascular plants) to design management and conservation actions in semi-natural pastures. Careful consideration of conservation values linked to the tree and shrub layers in grasslands should always precede decisions to remove trees and shrubs on the grounds of promoting richness of vascular plants confined to semi-natural grasslands. Finally, the importance of landscape composition for mobile organisms such as birds entails that management activities should focus on the wider countryside and not exclusively on single pastures.     

Different responses of plants and herbivore insects to a gradient of vegetation height: an indicator of the vertebrate grazing intensity and successional age

Empirical studies have suggested that species richness of grassland insects usually decreases under grazing management. By contrast, grazing has been shown to increase the species density and richness of vascular plants, especially on productive soils. In order to test the suggested differences in response to management between plants and insects, we simultaneously studied species richness of vascular plants and their insect herbivores, butterflies and moths, in 68 semi-natural grasslands with varying grazing intensity and frequency in SW Finland. Species richness of plants and insects was for the first time related to a quantitative measure of disturbance intensity and successional age, mean vegetation height, by using generalized additive models (GAM). The effects of soil nutrients on vegetation height were accounted for by using phosphorus concentration as a productivity surrogate.
The results showed that species richness of butterflies and moths peaked in taller vegetation compared with vascular plants, corresponding to a lower disturbance intensity and increasing time since abandonment. These patterns are discussed in the light of two hypotheses, the "structural diversity hypothesis" and "dynamic equilibrium model" of Huston, both suggesting a weaker disturbance tolerance of insects compared with plants. Butterflies and moths which are specialists in their larval host-plant use (monophagous and oligophagous species) preferred lower vegetation (higher disturbances) compared with generalists (polyphagous species), as predicted by Huston's model. This difference indicates a stronger relationship with plant species richness for specialist than for generalist butterflies and moths. Our results support the application of regionally and temporally varying grazing intensities in grassland conservation management.     

Effects of restoration with cattle grazing on plant species composition and richness of semi-natural grasslands

Restoration of semi-natural grasslands by cattle grazing is among the most practical options for reversing the decline of northern European floristic diversity, but no studies on this subject are available. In this work the success of restoration of abandoned, privately owned mesic semi-natural grasslands by farmers receiving support from the EU agri-environmental support scheme was studied in southwestern Finland. Three kinds of grasslands were compared: old (continuously cattle grazed), new (cattle grazing restarted 3–8 years ago) and abandoned pastures (grazing terminated >10 years ago). Plant species composition of the three pasture types was floristically different in multivariate analyses (non-metric multidimensional scaling). Total species richness, richness of grassland plants, indicator plants and rare plants were highest in old and lowest in abandoned pastures in all studied spatial scales (0.25–0.8 ha, 1 and 0.01 m²). The results were congruent with different scales and species list definitions, suggesting that species density scale (1 m²) can be used as a partial surrogate for large scale species richness. Species richness of new pastures was 20% higher on 0.25–0.8 ha, 40–50% higher on 1 m² and 30% higher on the 0.01 m² scale compared to abandoned grasslands. Rare species showed insignificant response to resumed grazing. Despite problems in management quality, this study showed promising results of restoration of abandoned grasslands by cattle grazing on private farms. However, populations of several rare grassland plants may not recover with present cattle grazing regimes. Management regulations in the agri-environmental support scheme need to be defined more precisely for successful restoration.     

Railway Embankments as New Habitat for Pollinators in an Agricultural Landscape

Pollinating insect populations, essential for maintaining wild plant diversity and agricultural productivity, rely on (semi)natural habitats. An increasing human population is encroaching upon and deteriorating pollinator habitats. Thus the population persistence of pollinating insects and their associated ecosystem services may depend upon on man-made novel habitats; however, their importance for ecosystem services is barely understood. We tested if man-made infrastructure (railway embankments) in an agricultural landscape establishes novel habitats that support large populations of pollinators (bees, butterflies, hoverflies) when compared to typical habitats for these insects, i.e., semi-natural grasslands. We also identified key environmental factors affecting the species richness and abundance of pollinators on embankments. Species richness and abundance of bees and butterflies were higher for railway embankments than for grasslands. The occurrence of bare (non-vegetated) ground on embankments positively affected bee species richness and abundance, but negatively affected butterfly populations. Species richness and abundance of butterflies positively depended on species richness of native plants on embankments, whereas bee species richness was positively affected by species richness of non-native flowering plants. The density of shrubs on embankments negatively affected the number of bee species and their abundance. Bee and hoverfly species richness were positively related to wood cover in a landscape surrounding embankments. This is the first study showing that railway embankments constitute valuable habitat for the conservation of pollinators in farmland. Specific conservation strategies involving embankments should focus on preventing habitat deterioration due to encroachment of dense shrubs and maintaining grassland vegetation with patches of bare ground.     

Regional hotspots of butterfly diversity in a protected area: Are they indicators of unique assemblages and areas with more species of conservation concern?

The implementation of biodiversity studies to set conservation priorities assumes that the different criteria used in prioritisation are correlated. Based on butterfly data, we examined whether high total diversity (species density and richness), unique species assemblages, and high numbers of different categories of Species of European Conservation Concern (SPEC) were co-located in the same habitats in the Picos de Europa National Park, Spain. Species density (number of species per sample) declined with elevation and was significantly higher in Genista shrublands on southern slopes (Southern Genista shrubland). We found unique species assemblages for Southern Genista shrublands, heathlands, mown meadows and subalpine meadows, whereas Genista shrubland on northern slopes (Northern Genista shrubland) and pastures were occupied by more habitat generalist species. We also found an effect of elevation on species composition. Species with global distributions concentrated in Europe (SPECs 4b) tended to be associated with Southern Genista shrubland, whereas species with global distributions restricted to Europe (SPECs 4a) were associated with higher elevations. Thus, we suggest a higher conservation value for butterfly assemblages located both on Southern Genista shrublands and at high elevations as they contain unique assemblages with a higher number of SPECs. By comparing our data with a previous study on carabid beetles, we found a low coincidence between butterflies and beetles for habitats with high species diversity and unique assemblages. Our study suggests that preserving areas in the landscape supporting higher butterfly diversity only partially preserves those species of greater conservation concern, and that butterflies cannot be used as a reliable indicator of other priority taxa for conservation, such as carabid beetles.     

Low congruence between the diversity of Waxcap (Hygrocybe spp.) fungi and vascular plants in semi-natural grasslands

The diversity of fungi in semi-natural grasslands is poorly known, partly due to difficulties in species identification in the field but also because there are few specialists available. Diversity assessments of grassland fungi would be facilitated if a potential surrogate group for fungal diversity could be identified. The aim of this study was to assess whether plant diversity patterns in semi-natural grasslands are congruent with diversity patterns of Waxcap (Hygrocybe spp.) fungi. Waxcaps, together with several other groups of fungi (e.g. the genera Entoloma, Dermoloma, Camarophyllopsis, and the families Clavariaceae and Geoglossaceae) have habitat requirements similar to many plants typical for semi-natural grasslands and they are all threatened by ceased management and nutrient enrichment. Diversity data from 31 semi-natural grasslands in southern Sweden were used to examine if there is congruence in species richness, nestedness, β-diversity and species composition between plants and Hygrocybe fungi. Species richness of Hygrocybe was significantly positively correlated with plant richness, although the relationship was not strong (r²=0.14). Both plant and Hygrocybe species composition was significantly nested, i.e. species-poor sites contain a subset of species from species-rich sites, which suggests that rare species mostly occur in the species-rich sites. A species similarity analysis between the grassland sites showed that there is low overlap between species composition of plants and Hygrocybe, indicating that conservation decisions based solely on plants may not fulfil the requirements of the Hygrocybe species. The conclusion is that there is low congruence between plant and Hygrocybe species diversity, making plants a poor surrogate group for Hygrocybe fungi, and probably also for other grassland fungi.     

Species richness in agroecosystems: the effect of landscape, habitat and farm management

It has been suggested that biodiversity in agroecosystems depends on both landscape heterogeneity and farm management, but at the same time, studies of biodiversity in relation to both landscape variables and farm management are rare. We investigated the species richness of plants, butterflies, carabids, rove beetles and the diversity of spiders in cereal fields, leys (grass and clover crop) and semi-natural pastures at 16 farms in Central East Sweden. The farms were divided into eight pairs of one conventional and one organic farm to enable us to separate the effects of landscape and farm management on biodiversity. The pairing was based on land use, location, and landscape features. Species richness of different taxonomic groups was generally not correlated. There were no differences in species richness between the farming systems, except for carabids that had higher numbers of species on conventional farms. The species richness generally increased with landscape heterogeneity on a farm scale. Habitat type had a major effect on the species richness for most groups, with most species found in pastures and leys. The correlations between species richness and landscape variables on a farm scale, and not on a scale of multiple farms, identify farmers as the important decision-maker in conservation issues for these taxonomic groups. We discuss the role of species richness of pests' natural enemies for biological control and conservation strategies of the more common species in the agricultural landscape.     

How can we preserve and restore species richness of pollinating insects on agricultural land?

During recent decades, concern about the loss of biodiversity on agricultural land has increased, and semi‐natural grasslands have been highlighted as critical habitats. Temperate European agricultural landscapes require distinct and appropriate management to prevent further impoverishment of the flora and fauna. This is especially urgent for pollinating insects that provide important ecosystem services. Our aim was to examine how species richness of three important groups of pollinating insects; solitary bees, butterflies and burnet moths are related to different farm characteristics, and if there are any differences between these three groups. A further aim was to test if red‐listed species are related to any farm characteristics. Species richness of solitary bees, butterflies and burnets was measured on all semi‐natural grasslands at 16 farms in a forest‐dominated area of 50 km² in southern Sweden, using systematic transect walks in April to September 2003 (only butterflies and burnets) and 2005. Species richness of solitary bees and butterflies was intercorrelated, both before and after controlling for the area of semi‐natural grassland. Species richness of solitary bees increased with the area of semi‐natural grassland. After controlling for the effect of the area of semi‐natural grassland species richness was strongly positively related with the density of the plant Knautia arvensis and negatively related with the proportion of grazed grassland. The results were similar for solitary bees and butterflies. The number of red‐listed solitary bees was positively related to the proportion of meadows with late harvest (after mid‐July) and decreased with increased farm isolation. The number of burnet species (all red‐listed) was positively related to vegetation height, flower density and the proportion of meadows with late harvest on a farm. Areas with a high density of K. arvensis and with traditional hay‐meadow with late harvest present, harbour most species. Promoting traditional hay‐meadows, late extensive grazing and the herb K. arvensis, people managing agricultural biodiversity can encompass high species richness of pollinating insects and support red‐listed species. Further, we suggest that the density of K. arvensis at a farm can be used as a biodiversity indicator, at least for pollinating insects.     

Species abundance patterns of plants in Swedish semi-natural pastures

This investigation is based on a species inventory m semi-natural pastures, remnants of an "old" traditional landscape, in Sweden The aim was to examine relationships between species richness and pasture area, neighbour pasture area, distance to nearest pasture, pasture heterogeneity and fertilization Species occurrences were also investigated individually for the same parameters Pasture area, neighbour pasture area and distance to nearest pasture were not found to influence species richness Habitat heterogeneity and indications that pastures had been influenced by fertilization were however associated with both species richness and species occurrences Heterogeneity had only positive effects whereas the effects of fertilization were only negative Species regional abundance pattern tended to be bimodal, as predicted by the core-satellite hypothesis No correlation between abundance and plant height, or seed size was found, as would have been expected if competitive dominance effects determined species regional abundance There was a close to significant tendency that pasture area was positively associated with occurrences of species with features favouring a high seed dispersal capacity We suggest that the previous history of pasture distribution and management may be of vital importance for present day species abundance pattern, even though this remains obscure in the recent landscape     

The effect of local anthropogenic habitat heterogeneity on assemblages of carabids (Coleoptera,Caraboidea) endemic to the Alps

We assessed the response of a guild of endemic medium and large-bodied carabid species to the local (fine-grained) habitat heterogeneity induced by man. The study was conducted by means of trapping (without killing and preservative agents) and radio-tracking methods (to collect data on the spatial ecology of the endangered species Carabus olympiae Sella). Habitat differentiation induced by man (a mosaic of beech forests, alpen rose shrubberies and pastures) affected species richness and abundance of the guild, which were significantly higher in forests and/or shrubberies than in pastures. Most species preferred forests and shrubberies, with the only exception of Carabus concolor Fabricius, which preferred open areas. Species distribution was also affected by stone density and cattle grazing intensity. In particular, petrophilous species peaked on the roadside because of the high stone density there (stones amassed during road construction), while species abundances significantly lowered in overgrazed pastures. The compactness of the paths made by C. olympiae individuals (evaluated through a tortuosity index) was significantly higher in beech forests than in alpen rose shrubberies, indicating movements in the latter habitat type were bound by the spatial distribution of shrubs, which imposes a limit on path tortuosity. Our findings suggest that local ground beetle species diversity strongly depends on small-scale anthropogenic variables (namely habitat type, stone density and grazing intensity), and that habitat modifications (namely from forest to shrubbery) may significantly affect species movement patterns. General and local conservation management suggestions are given in conformity with these results.     

Scale-dependent effects of land use on plant species richness of mountain grassland in the European Alps

Traditionally managed mountain grasslands in the Alps are species‐rich ecosystems that developed during centuries of livestock grazing. However, changes in land use including fertilisation of well accessible pastures and gradual abandonment of remote sites are increasingly threatening this diversity. In five regions of the Swiss and French Alps we assessed the relationship between land use, soil resource availability, cover of the unpalatable species Veratrum album, species richness and vegetation composition of mountain grasslands across four spatial scales ranging from 1 to 1000 m². Mean species richness and the increase in the number of species with increasing area were lower in intensively grazed, fertilised pastures than in traditional pastures or in abandoned pastures. Species composition of abandoned pastures differed from that of the other management types. Plant species richness was influenced by different factors at different spatial scales. At the 1 m² scale, plant species richness was negatively related to soil nitrate and influenced by the cover of V. album, depending on land use: species richness and cover of V. album were negatively correlated in abandoned pastures, but positively correlated in fertilised grasslands. At the 1000 m² scale, a negative effect of fertilization on richness was evident. These results indicate that at small scales species richness in mountain grasslands is determined by competition for light, which should be more important if nutrient availability is high, and by positive and negative interactions with unpalatable plants. In contrast, species richness at the large scale appears to be mainly influenced by land use. This result emphasizes the importance of studying such inter‐relationships at multiple scales. Our study further suggests that the maintenance of the traditional land use scheme is crucial for the conservation of plant species richness of mountain pastures as both intensification and abandonment changed species composition and reduced plant species diversity.     

The effect of horse, cattle and sheep grazing on the diversity and abundance of land snails in nutrient-poor calcareous grasslands

Livestock grazing is a common management practise in semi-natural grasslands in Central Europe. Different types of livestock (horses, cattle, sheep) and grazing intensity are known to affect the richness and composition of plant species. However, knowledge of grazing-dependent effects on invertebrates is limited. We examined the influence of horse, cattle and sheep grazing on the richness, abundance and composition of land snail species in 21 calcareous nutrient-poor grassland areas in the northwestern Jura Mountains, Switzerland. Grazing by different livestock species did not affect the species richness, abundance and species composition of land snails. Furthermore, the number of open-land species and the ratio of large- to small-sized snail species or individuals did not differ among the three pasture types. However, independent of livestock species, grazing intensity negatively influenced the snail fauna. Snail species richness, abundance and number of Red list species decreased with increasing grazing intensity. Grazing intensity also affected the occurrence of individual snail species (Truncatellina cylindrica, Cecilioides acicula, Candidula unifasciata and Trichia plebeia). To preserve the snail fauna in nutrient-poor grasslands, pastures can be stocked with horses, cattle or sheep. However, both maximum stocking rate (number of livestock units per hectare) and grazing duration (number of grazing days per year) must be carefully defined for the proper management of the pastures.     

Is there a benefit of excluding sheep from pastures at flowering peak on flower-visiting insect diversity?

Permanent grasslands are an important habitat for insect communities, including pollinator populations which are in pan-European decline. Here, we investigated the benefits of temporarily excluding sheep from pastures on butterfly (Lepidoptera: Rhopalocera and Zygaenidae) and bumblebee (Hymenoptera: Bombidae) communities in two semi-natural grasslands differing in soil fertility and surrounding landscapes. We compared the impact of continuous grazing against rotational grazing (RG) at the same stocking rate but in which a subplot was excluded from the rotation during the main flowering period. We predicted that the diversity of flower-visiting insect community would be improved by RG due to the preservation of flower-rich patches and the maintenance of sward heterogeneity. Benefits of RG management were mainly evidenced on bumblebee density and species richness, with some additional effects on local density of butterflies during the subplot-exclusion period. Temporarily excluding sheep from pastures during peak flowering could thus offer an opportunity to preserve the diversity of flower-visiting insects, in spite of weaker benefits than could have been expected from previous surveys with cattle.     

Can re-establishment of cattle grazing restore bryophyte diversity in abandoned mesic semi-natural grasslands?

Changes of agricultural practices have led to decline of semi-natural habitats sustained by traditional animal husbandry in many European regions. The abandonment of semi-natural pastures leads to increase of vascular plant biomass and subsequent decline of weak competitors such as bryophytes. Re-establishing traditional animal husbandry may potentially restore biodiversity but the success of such measures remains insufficiently known. In this study, we asked if re-establishing cattle grazing on previously abandoned grasslands will restore their bryophyte communities. The effect of cattle grazing on bryophyte communities of mesic semi-natural grasslands was studied in south-western Finland in a comparison of (i) continuously grazed pastures, (ii) previously abandoned pastures where grazing was re-established during 1990s, and (iii) abandoned pastures, where grazing had ceased during late 1960s to early 1980s. The average cover, species richness, species density and species diversity of bryophytes were significantly higher in the continuously grazed than in the abandoned grasslands. Ordination analyses revealed clear differences also in community structure between the management classes. Re-established grasslands were ecologically heterogeneous and situated in between the continuously grazed and abandoned grasslands in all characteristics, indicating variable effect of the restoration measure. Seventeen bryophyte species were recognized as significant indicators of the three grassland classes, four of which could be used as indicators of valuable grassland habitats. Although there was variation in the consequences of re-introduction of grazing, the results give evidence of positive effect of grazing on regaining bryophyte diversity of abandoned grasslands.     

Do protected areas in urban and rural landscapes differ in species diversity?

Previous studies from Central Europe and North America showed that species richness is higher in urban than in rural landscapes. Do protected areas, which can be found in both city and countryside, reflect this species richness pattern? The impact of urban land-use might reduce conservation success and necessitate special management strategies. We compared species richness and species spatial turnover of selected animal and plant taxa (carabids, butterflies, snails, birds, lichens, mosses, vascular plants) in 30 protected areas in the city of Halle and 56 protected areas in the adjacent rural district of Saalkreis (Central Germany). Species were mapped by experienced biologists within a systematic species inventory. We corrected species numbers for the effects of landscape structure (e.g. size, shape and distance of habitats) which might influence species diversity beyond urbanisation effects. Butterflies, birds and lichens had significantly higher species numbers in the rural protected areas. Species spatial turnover was higher among urban areas than among rural areas or pairs of urban and rural areas for most taxa. Diversity in all taxa depended on the size of a protected area. We discussed these patterns in the context of the general urban-rural species diversity patterns. Our results indicate an increasing isolation of species assemblages with urbanisation and highlight that space for protected areas is even more limited in urban than rural areas. An effective conservation of urban species diversity should include both typical urban and semi-natural habitats to cover the full range of species living in cities.     

Effect of cultivating croplands and grazing in arid grassland habitats on the conservation of melitaeine butterflies in a mountainous area in Northern China

In the study area (Yanjiaping Village, Hebei Province, China), grazing extensity varies at different locations, small and discontinuous croplands are imbedded in some arid grassland, which are habitats for the melitaeine butterflies, Euphydryas aurinia and Melitaea phoebe. These two species of butterfilies coexist in this area, in which grazing and cultivation are the main disturbances. Grazing and cultivation have a reciprocal effect on E. aurinia, rather than M. phoebe. We observed that E. aurinia preferred to occupy patches with moderate grazing and imbedded with small and discontinuous croplands, where E. aurinia also has high population density. The percentage of E. aurinia larval groups in the ribbings was significantly higher than that of M. phoebe, whereas larvae of both species tended to increase in recent years. Our data also showed that the population density and the patch occupancy rate of both E. aurinia and M. phoebe were the highest under moderate grazing. It indicates that cultivation of small and discontinuous croplands within the patch has a significant effect on the population density of both species of melitaeine butterflies. Thus, to artificially create or maintain semi-natural habitats, complemented by moderate grazing, might be an ecological strategy to conserve melitaeine butterflies effectively. Considering the distinct impacts of cultivation and grazing on the population distribution and dynamics of the two different species, human disturbance in the mountainous area might be strategically involved in proposing conservation plans for the target species in the future.     

Floristic diversity and the richness of locally endangered plant species of semi-natural grasslands under different management practices,southern Kyushu,Japan

Background: The rapid decline of semi-natural grasslands in Japan threatens many relic and endemic plant species. There is insufficient knowledge on how the impacts of land-use changes and management of grasslands have been affecting grassland ecosystems and what conservation measures may be taken to conserve as much of the existing plant diversity as possible.

Aim: We assessed the existing management regimes for their suitability for conserving Red Data Book (RDB) species.

Methods: We conducted our study in four districts of Kushima, Kyushu, south-west Japan, with different land-use histories. We compared species richness, plant density and abundance in six grassland types: regularly burnt, regularly mown, paddy levee, roadside, landslip and wetland communities in a total of 289 1 m x 1 m quadrats, recorded in172 grassland patches. Species richness plant density and abundance were analysed with special reference to RDB species under different land use history.

Results: Species richness of grasslands did not differ across different land use histories, yet our analysis showed that the reduced area of grasslands markedly affected the density of RDB species. Grassland types differed in their ability to support RDB species: regularly burnt grasslands were the richest in RDB species and poorest in alien species, followed by regularly mown grasslands, paddy levees, landslip, wetland and roadside communities.

Conclusions: Traditional management regimes, such as regular burning or mowing of grasslands have the best potential for conserving RDB species, and thus should be part of conservation management practices of semi-natural grasslands.     

Alternative management on fens: Response of vegetation to grazing and mowing

Abstract. The impact of cattle grazing on the vegetation of calcareous fens was compared to the effects of traditional autumn mowing in southern Germany. Vegetation composition was studied in adjacent pairs of fen meadows and pastures with similar environmental conditions and biomass production. Vegetation data were analysed with respect to species richness, species composition and response of species traits to disturbance, including morphology, defence mechanisms, clonal growth form and generative reproduction. Species richness was significantly reduced by grazing, but the percentage of typical fen species or Red Data Book species was not affected by land use type. Detrended Corrspondence Analysis indicated that species composition could best be explained in terms of a land use gradient. Species traits showed a clear trend in their response to land use type. Grazing favoured grasses and small forbs. Only a few species with defence mechanisms against foraging were more frequent or abundant on pastures. Many other species with defence mechanisms, however, did not have an advantage on pastures. Flowering and seed dispersal traits did not respond significantly to grazing or mowing. Species with fast spreading stem derived clonal organs were favoured on pastures, whereas all other clonal growth forms and, particularly, non‐clonal species were more abundant on meadows. More indicator species of wet soil conditions and species adapted to flooding were found on pastures. Grazing can be recommended as an alternative land use to mowing in contrast to abandonment, but a reduction in species richness and changes in species composition and species traits may occur.     

Importance of semi-natural habitats for the conservation of butterfly communities in landscapes dominated by pine plantations

While the area of plantation forests continues to increase worldwide, their contribution to the conservation of biodiversity is still controversial. There is a particular concern on the central role played by natural habitat remnants embedded within the plantation matrix in conserving species-rich insect communities. We surveyed butterflies in maritime pine plantation landscapes in south-western France in 83 plots belonging to seven habitat types (five successional stages of pine stands, native deciduous woodlands and herbaceous firebreaks). The effect of plot, habitat and landscape attributes on butterfly species richness, community composition and individual species were analysed with a General Linear Model (GLM), partial Canonical Correspondence Analysis (CCA) and the IndVal method. The most important factors determining butterfly diversity and community composition were the presence of semi-natural habitats (deciduous woodlands and firebreaks) at the landscape scale and the composition of understorey vegetation at the plot scale. Pure effects of plot variables explained the largest part of community variation (12.8%), but landscape factors explained an additional, independent part (6.7%). Firebreaks were characterized by a higher species richness and both firebreaks and deciduous woodlands harboured species not or rarely found in pine stands. Despite the forest-dominated landscape, typical forest butterflies were rare and mainly found in the deciduous woodlands. Threatened species, such as Coenonympha oedippus and Euphydryas aurinia, were found in pine stands and in firebreaks, but were more abundant in the latter. In the studied plantation forest, the conservation of butterflies depends mainly on the preservation of semi-natural habitats, an adequate understorey management and the maintenance of soil moisture levels.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.