Importance of boreal grasslands in Sweden for butterfly diversity and effects of local and landscape habitat factors

A widespread decline in biodiversity in agro-ecosystems has been reported for several groups of organisms in Western Europe. The butterfly fauna was studied in 60 selected semi-natural grasslands in a coniferous-dominated boreal landscape in south-eastern Sweden. The aim was to investigate how butterfly assemblages were affected by the amount of semi-natural grasslands in the surrounding landscape. Furthermore, we wanted to determine if semi-natural grasslands in boreal landscapes harboured species otherwise declining in other parts of Europe. For each study site, the amounts of semi-natural grasslands in the landscape within radii of 500, 2,000 and 5,000 m were studied. Nine local habitat factors were also recorded. Only the amount of semi-natural grasslands within a 5,000 m radius could explain a significant part of the variation in butterfly composition, but there was no clear relationship between the amount of semi-natural grassland and butterfly diversity. Instead, this study showed that local habitat quality was very important for butterfly diversity at individual sites. Flower abundance, sward height and herb composition were the most important local factors. Patches surrounded by a small amount of semi-natural grasslands had high butterfly diversity, contrary to expectations. This may be explained by the fact that forest habitat provides a matrix with several features suitable for butterflies. The butterfly fauna was rich in species representative of low-productivity grasslands, species that are declining in other countries in Western Europe.     

The Influence of Landscape Grain Size on Butterfly Diversity in Grasslands

The relationship between butterfly diversity and both habitat and landscape variables was studied in two areas of southern Sweden. The habitat quality of the grasslands was similar in the two study areas but the landscape pattern differed in grain size and amount of grassland and forest. Using a transect survey method, a total of 3341 butterflies were observed and 30 taxa identified. We found that both habitat and landscape variables influenced the butterfly diversity of the investigated grasslands. Species composition differed markedly between the two study areas. A study area with a fine-grained landscape pattern, a high cover of semi-natural grassland and many forest edges had twice as many butterfly species but half the number of individuals compared with a coarser-grained study area with larger grasslands widely spread in a matrix of arable fields. The results of our study indicate that both habitat quality and landscape pattern have to be considered when developing conservation strategies for grassland butterflies.     

The value of semi-natural grasslands for the conservation of carabid beetles in long-term managed forested landscapes

Species rich semi-natural grasslands are disappearing across Europe, affecting invertebrate diversity negatively. In NW Spain, the recent abandonment of traditional farming practices and the gradual decrease in grazing pressures are reducing the number and extent of montane grasslands. In this context, we investigated the composition of carabid beetle (Coleoptera, Carabidae) assemblages that inhabit semi-natural grasslands situated in long-term managed oak and beech forested landscapes. According to their spatial arrangement, the studied grasslands were classified into: (1) interior or gap grasslands (small and completely surrounded by continuous forest) and (2) exterior grasslands (large and connected to a variety of habitat types). Our results indicate that, within each forested landscape, the gap and exterior grasslands harboured particular carabid assemblages (i.e. exclusive or abundantly collected species), which were also distinct from the surrounding forest carabid fauna. Dissimilarities between gap and exterior grasslands in each landscape suggest great carabid diversity at the regional scale. We also detected species-specific responses as several carabids were mainly associated with gap or exterior grasslands. Consequently, in highly modified forested landscapes, semi-natural grassland remnants may constitute great value for the protection of the carabid fauna. Specifically, we recommend conservation strategies that preserve variety in grassland features and maintain proper management activities to prevent the loss of specialised species and a decrease in regional carabid diversity.     

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.     

Pattern of land mosaics affecting butterfly assemblage at Mt Ikoma, Osaka, Japan

We studied the effects of habitat mosaics on butterfly assemblage on multiple spatial scales: landscape, landscape element, local habitat, and microhabitat, based on the transect counts conducted along a 3.84 km route. The transect route, including 21 local habitats, passed through two distinct areas: 1.65 km of a secondary deciduous Quercus forest and the grove of a shrine in Hiraoka, and 2.19 km of a mosaic of secondary deciduous Quercus forest, grassland, and farmland in Narukawa. The diversity of the landscape elements and species richness were higher in Narukawa than in Hiraoka; the landscape mosaic enhanced the species richness in Narukawa. However, the diversity indices and specialist species (univoltine tree feeder) were decreased in this mosaic landscape. The species richness at local habitats was also increased by the mosaic of microhabitats, such as the herbaceous layer, glade, and mantle in the local habitats, whereas it was decreased by an abundant shrub layer. The ratios of species richness to abundance in the local habitats were lower than expected based on random sampling from the total of Hiraoka and Narukawa. This means that local assemblages were non-random samples from an assemblage on the landscape or regional scale, and were made up by the process of habitat selection of butterfly species in the assemblages on the landscape or regional scale. For conservation of butterfly assemblages, we recommend that woodlands should be kept without fragmentation, but with glades or small grasslands, and with clearance of the shrub layer along the path.     

Invertebrate conservation at the interface between the grassland matrix and natural Afromontane forest fragments

Diversity patterns of amphipods, carabid beetles and ants were investigated in five naturally-fragmented Afromontane forest remnants, and in the surrounding grassland matrix. Forests were architecturally similar. In contrast, grasslands surrounding these forests are subject to great differences in anthropogenic impacts. Consequently, transition from forest to grassland ranged from being abrupt (heavy disturbance) to gradual (little disturbance). Significantly different mean numbers of carabid individuals and species were captured between sites and multivariate analyses showed clear separation in carabid assemblage-structure with level of disturbance. Carabids were, furthermore, significantly more diverse in forests, compared to grasslands. Ants, however, were equally species rich between sites but were significantly more abundant and species rich in grasslands than forests. Amphipods, represented here by a single species, Talistroides africana, was significantly less abundant at highly disturbed sites and significantly more abundant in forests than grasslands. Results support the hypothesis that the dynamics of remnants are influenced by their surrounding landscape. Here, the dynamics of amphipods and carabids (predominantly forest taxa) were influenced by different disturbance regimes in grasslands surrounding these forests. Epigaeic ants, a predominantly grassland taxon here, also showed significant differences in assemblage-composition between sites with varying disturbance. Conserving Afromontane grasslands should be of prime concern because this will include the protection of forest/grassland ecotones and forest remnants.     

Contrasting trends of butterfly species preferring semi-natural grasslands,field margins and forest edges in northern Europe

Indicator classifications help us to focus on the most relevant groups of species in monitoring the effects of land use changes on biodiversity. We studied changes in distribution area of 74 butterfly species preferring one of the three common habitats of boreal agricultural landscapes: semi-natural grasslands (35 species), arable field margins (7) and forest edges (32). Using extensive atlas data from four time periods during the last 50 years in Finland, we quantified trends in the occupancy of the species in 10 km grid squares, and classified them into four classes: declining (23), stable (17), increasing (27) and fluctuating (7) species. Trends among the species favouring three habitats were different: 60% of the species of semi-natural grasslands had declined, whereas 86% of the species typical of open field margins had increased. An increase also predominated in species associated with forest edges. Declining and increasing species differed in three ecological characteristics: increasing species were more mobile, utilized a wider range of habitats and, based on their larval host plants, lived in more eutrophic habitats than declining species. Species overwintering as adults showed more positive trends in occupancy than species overwintering as eggs, larvae or pupae. Observed trends in occupancy are in good agreement with long-term changes in land use and habitat availability in Finland: a long-continued decrease in the area of semi-natural grasslands and an increased amount of open forest edges and clearings due to modern forestry during the past 50 years.     

High value of short rotation coppice plantations for phytodiversity in rural landscapes

The demand for wood from short rotation coppice (SRC) plantations as a renewable energy source is currently increasing and could affect biodiversity in agricultural areas. The objective was to evaluate the contribution of SRC plantations to phytodiversity in agricultural landscapes assessed as species richness, species–area relationships, Shannon indices, detrended correspondence analysis on species composition, Sørensen similarities, habitat preference proportions, and species proportions found in only one land use. Vegetation surveys were conducted on 12 willow (Salix spp.) and three poplar (Populus spp.) coppice sites as well as on surrounding arable lands, grasslands and forests in central Sweden and northern Germany. SRC plantations were richer in plant species (mean: 30 species per 100 m²) than arable land (10), coniferous forests (13) and mixed forests in Germany (12). Comparing SRC plantations with other land uses, we found lowest similarities in species composition with arable lands, coniferous forests and German mixed forests and highest similarities with marginal grassland strips, grasslands and Swedish mixed forests. Similarity depended on the SRC tree cover: at increased tree cover, SRC plantations became less similar to grasslands but more similar to forests. The SRC plantations were composed of a mixture of grassland (33%), ruderal (24%) and woodland (15%) species. Species abundance in SRC plantations was more heterogeneous than in arable lands. We conclude that SRC plantations form novel habitats leading to different plant species composition compared to conventional land uses. Their landscape‐scale value for phytodiversity changes depending on harvest cycles and over time. As a structural landscape element, SRC plantations contribute positively to phytodiversity in rural areas, especially in land use mosaics where these plantations are admixed to other land uses with dissimilar plant species composition such as arable land, coniferous forest and, at the German sites, also mixed forest.     

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.     

Local and landscape effects on butterfly density in northern Idaho grasslands and forests

Understanding butterfly response to landscape context can inform conservation management and planning. We tested whether local-scale resources (host and nectar plants, canopy cover) or landscape context, measured at two scales, better explained the densities of four butterfly species. The density of Coenonympha tullia, which has host plants strongly associated with grassland habitats, was positively correlated with the amount of grassland in 0.5- and 1-km radius landscapes and only occurred in forests when they bordered grasslands. For the other species, Celastrina ladon, Cupido amyntula, and Vanessa cardui, local-scale resources better explained butterfly densities, emphasizing the importance of local habitat quality for butterflies. These three species also used host plants that were distributed more heterogeneously within and among habitat types. Our findings demonstrate the importance of host plant spatial distributions when determining the scale at which butterfly density relates to resources, and we recommend that both these distributions and landscape context be evaluated when developing butterfly monitoring programs, managing for species of concern, or modeling potential habitat.     

Comparison of butterfly communities and abundances between marginal grasslands and conservation lands in the eastern Great Plains

The alteration and fragmentation of native tallgrass prairie in the Midwestern United States has created a need to identify other land types with the ability to support grassland butterfly species. This study examines butterfly usage of marginal grasslands, which consist of semi-natural grasslands existing within in a larger agricultural matrix, compared to grasslands managed for conservation of prairie species. Using generalized linear mixed models we analyzed how land purpose (marginal vs. conservation grasslands) affected butterfly abundance. We found grassland butterfly species to be significantly more common on conservation grasslands, whereas generalist species were significantly more common on marginal grasslands. Results of ordination analyses indicated that while many species used both types of habitats, butterfly species assemblages were distinct between habitat types and that edge to interior ratio and the floristic quality index of sites were important habitat characteristics driving this distinction. Within conservation grasslands we examined the relationship between butterfly abundance and the planting diversity used in restoring each site. We found higher diversity restorations hosted more individuals of butterflies considered habitat generalists, as well as species considered to be of conservation concern.     

Grassland area determines beetle assemblage dissimilarity from surrounding floodplain forest

Patch size is known to affect biodiversity in fragmented landscapes, but is usually examined in systems where the surrounding matrix habitat is unfavourable. We examined beetle diversity in a floodplain ecosystem that is characterised by naturally occurring grassland patches within a dominant matrix of contrasting yet habitable forest. We asked whether differences in the beetle assemblage between grassland and forest vegetation depended on the area of the grassland patch, which is a function of its flooding frequency and duration: smaller grasslands tend to be higher on the floodplain and are flooded less often and for shorter periods than larger grasslands. We found a negative relationship between grassland area and beetle abundance and species richness, and a positive relationship between grassland area and compositional dissimilarity from the surrounding forest. As expected, we found an overall difference in composition between forest and grassland assemblages, with five beetle species more common in the grasslands. Our study indicates that floodplain grasslands not only support beetle assemblages that are distinct from the surrounding forest, but that assemblages from the larger grasslands are compositionally more distinct than those from smaller grasslands. A likely cause of this pattern is the reduced edge effects and greater environmental contrast between forest and large grasslands that may be exposed to greater variation in local climate. Ongoing changes to flood regimes and potential encroachment of forest plants may decrease grassland area in the future, which may reduce spatial heterogeneity in the insect community in this unique floodplain ecosystem.     

Woody elements benefit bird diversity to a larger extent than semi-natural grasslands in cereal-dominated landscapes

Increasing landscape complexity can mitigate negative effects of agricultural intensification on biodiversity by offering resources complementary to those provided in arable fields. In particular, grazed semi-natural grasslands and woody elements support farmland birds, but little is known about their relative effects on bird diversity and community composition. In addition, the relative importance of local habitat versus landscape composition remains unclear. We investigated how the presence of semi-natural grasslands, the number of woody elements and the composition of the wider agricultural landscape affect bird species richness, true diversity (exponential Shannon diversity) and species composition. Bird communities were surveyed four times on 16 paired transects of 250 m each with 8 transects placed between a crop field and a semi-natural grassland and 8 transects between two crop fields with no semi-natural grasslands in the vicinity. The number of woody elements around transects was selected as an important predictor in all models, having a positive effect on species richness and true diversity, while the local presence of semi-natural grasslands was not selected in the best models. However, species richness and true diversity increased with increasing cover of ley and semi-natural grasslands, whereas species composition was modified by the coverage of winter wheat at the landscape scale. Furthermore, bird species richness, true diversity and species composition differed between sampling dates. As bird diversity benefited from woody elements, rather than from the local presence of semi-natural grasslands as such, it is important to maintain woody structures in farmland. However, the positive effect of grassland at the landscape scale highlights the importance of habitat variability at multiple scales. Because species richness and true diversity were affected by different landscape components compared to species composition, a mosaic of land-use types is needed to achieve multiple conservation goals across agricultural landscapes.     

Small-scale patterns of species richness in Swedish semi-natural grasslands: the effects of community species pools

Temperate semi-natural grasslands are known for their high plant species richness at small spatial scales. We examined the variation in small-scale species richness in a sample of 63 sites from Swedish semi-natural grasslands, located as fragments in the modem landscape dominated by forest and agricultural land. Data were obtained from two spatial scales at each site. 1 dm² and 4 m². Using an analysis based on a Monte Carlo simulation, we found support for the species-pool hypothesis: a high species richness at the I dm- scale was associated with high species richness at the 4 m2 scale. The conclusion from this pattern analysis would, however, be considerably strengthened if we could reduce the likelihood that other mechanisms than sampling from species pools of unequal size influence the pattern of small-scale species richness. Additional analyses were made in order to identify such mechanisms. We examined whether four putative key traits: seed size, seed production, plant size and reproductive allocation were different among species at comparatively species-rich vs species-poor I dm' plots. We found only a little evidence for such differences. There was a weak tendency that species in the plots with high species richness possessed larger (and fewer) seeds than species from species-poor plots. Our results are congruent with the main prediction of the species pool model: variation in small-scale species richness (1 dm²- scale) is basically a result of sampling from unequally sized community species pools (4 m² scale). Variation in species richness between the 4 nr semi-natural grassland "patches" may thus be sought for among mechanisms operating al larger spatial scales than 4 m². We briefly discuss such mechanisms, based on other studies performed in the same study area.     

Asymmetric dispersal and survival indicate population sources for grassland butterflies in agricultural landscapes

We tested the hypothesis that populations in small habitat fragments remaining in agricultural landscapes are maintained by repeated immigration, using three grassland butterflies ( Aphantopus hyperantus , Coenonympha pamphilus and Maniola jurtina ). Transect counts in 12 matched sets of semi-natural pastures, and linear habitat elements proximate and isolated from the pastures showed that population densities of M.  jurtina and C.  pamphilus were significantly higher in pastures and in linear habitats adjacent to these than in isolated linear elements. A mark-recapture study in a 2×2 km landscape indicated that individuals of all three species are able to reach even the isolated linear elements situated at least 1 km from the grasslands. For two of the species, A.  hyperantus and C.  pamphilus , analysis of the mark-recapture data revealed higher daily local survival rates in the semi-natural pastures and more individuals dispersing from pastures to linear habitat elements. The proportion of old compared to young individuals of C.  pamphilus and M.  jurtina were significantly higher in linear elements than in semi-natural pastures, which suggests that butterflies emerging in pastures subsequently dispersed to the linear elements. In combination, these results suggest that semi-natural pastures act as population sources, from which adult butterflies disperse to surrounding linear elements. Hence, preservation of the remaining fragments of semi-natural grassland is necessary to keep the present butterfly abundance in the surrounding agricultural landscape.     

Adjacent woodlands rather than habitat connectivity influence grassland plant,carabid and bird assemblages in farmland landscapes

One response to biodiversity decline is the definition of ecological networks that extend beyond protected areas and promote connectivity in human-dominated landscapes. In farmland, landscape ecological research has focused more on wooded than open habitat networks. In our study, we assessed the influence of permanent grassland connectivity, described by grassland amount and spatial configuration, on grassland biodiversity. We selected permanent grasslands in livestock farming areas of north-western France, which were sampled for plants, carabids and birds. At two spatial scales we tested the effects of amount and configuration of grasslands, wooded habitats and crops on richness and abundance of total assemblages and species ecological groups. Grassland connectivity had no significant effects on total richness or abundance of any taxonomic group, regardless of habitat affinity or dispersal ability. The amount of wooded habitat and length of wooded edges at the 200 m scale positively influenced forest and generalist animal groups as well as grassland plant species, in particular animal-dispersed species. However, for animal groups such as open habitat carabids or farmland bird specialists, the same wooded habitats negatively influenced richness and abundance at the 500 m scale. The scale and direction of biodiversity responses to landscape context were therefore similar among taxonomic groups, but opposite for habitat affinity groups. We conclude that while grassland connectivity is unlikely to contribute positively to biodiversity, increasing or maintaining wooded elements near grasslands would be a worthwhile conservation goal. However, the requirements of open farmland animal species groups must be considered, for which such action may be deleterious.     

Habitat complementarity and butterfly traits are essential considerations when mitigating the effects of exotic plantation forestry

Habitat loss threatens insect diversity globally. However, complementary vegetation types in remaining habitat increases opportunities for species survival. We assess the extent to which indigenous forest patches moderate the impact of exotic commercial afforestation on grassland butterflies. Butterflies were sampled in grassland along uncorrelated gradients of landscape-scale indigenous forest and plantation cover, while controlling for variation in local vegetation composition. We separately assessed responses by butterfly groups differing in habitat preference, larval diet, and mobility. There was no effect of landscape- or local-scale variables on species richness, but there was a strong interactive effect of forest and plantation cover on butterfly assemblage structure. The effect varied according to species traits. When forest cover was high, assemblages did not differ at different levels of plantation cover. However, plantation cover significantly influenced assemblage structure when forest cover was low. Grassland with limited forest cover in the protected area supported unique assemblages with high frequency of less mobile, specialized species with herbaceous larval host plants, whereas grassland with low forest cover near plantations had a prevalence of mobile, generalist species. A positive association between forest cover and butterflies with woody larval host plants suggests that indigenous forest patches improved the suitability of fragmented grassland for a subset of butterflies, emphasising the value of natural heterogeneity in transformed areas. However, certain butterfly traits associated with large, open grassland were under-represented in grassland between plantations, underscoring the importance of open areas in the broader landscape to conserve the full diversity of species.

     

Estimating the consequences of land-use changes on butterfly diversity in a marginal agricultural landscape in Sweden

Abandonment of farming with the resultant increase in forest cover is one of the major threats to semi-natural grasslands in marginal agricultural areas. In Sweden, the loss of semi-natural grassland is a serious nature conservation problem since it is one of the most species-rich habitats. In this study, the consequences of grassland abandonment and afforestation on butterfly diversity and butterfly dispersal costs are estimated and used to compare three different future land-use scenarios for a marginal agricultural landscape in Sweden. Based on previous butterfly surveys on grasslands in the area, a relationship between land-use type and butterfly diversity was established. By comparing land-use maps of different scenarios, the number of suitable habitat patches and total suitable habitat patch area with low, medium and high butterfly diversity could be estimated. To obtain an indication of possible fragmentation effects, a least-cost analysis was used to compare travel costs of the butterflies between suitable habitat patches for the different scenarios. The results show that different land-use scenarios affect butterfly diversity and travel costs differently. In the extreme case scenario of cessation of full-time farming and a reduction in part-time farming, nearly all valuable butterfly habitats will vanish, since the most species-rich habitats lie in the periphery of the settlement and are expected to be abandoned and afforested first. If, on the other hand, grassland management is less reduced the effect of abandonment on butterflies depends very much on which areas continue to be managed. To preserve the most important grasslands for butterflies an active management strategy for the whole study area would be needed. While it seems relatively easy to identify the areas most important to conserve from a butterfly diversity perspective, it will be more difficult to find an optimal spatial solution that also minimises dispersal costs for butterflies.     

Chronosequential change in a butterfly community after clear-cutting of deciduous forests in a cool temperate region of central Japan

Transect counts of butterflies were conducted in the northern part of Ibaraki, central Japan, from 1997 to 2001 at 11 census sites, composed of successive stages of deciduous forest development: grassland (one site, early abandoned stage); cutover land (one site, 1–5 years after clear‐cutting); secondary forests (very young (two sites, 6–9 years), middle (two sites, 16–22 years) and old (two sites, 47–51 years)) and old‐growth natural forests (three sites, ≥124 years old). A total of 86 species and 8858 individual butterflies were recorded by 29 sets (406 times) of transect counts. The species richness (number of species), abundance (number of individuals) and diversity indices (Shannon–Wiener H′ and Simpson's 1–λ) of butterflies were high in the early stages (grassland, cutover land and very young secondary forests) of secondary succession. Typical natural forest species increased with the progress of succession, and the old secondary forests and old‐growth natural forests had similar species composition. In contrast, most of the typical natural grassland species were recorded only in the grassland site. In the cutover land site, the number of individuals of grassland species considerably decreased from the first to the second year; furthermore, only one typical natural grassland species was recorded. Thus, the suitable stage for grassland butterfly species lasts for only 1–2 years after clear‐cutting. Old secondary forests (approximately>50 years old) would be able to give refuge to the forest butterfly species, including typical natural forest species. Based on the results, a practical, forestry‐based plan to conserve butterfly diversity was proposed.     

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.