The role of linear mown firebreaks in conserving butterfly diversity: Effects of adjacent vegetation and management

Semi‐natural grasslands are vital for maintaining grassland butterflies in Japan, as well as in Europe. However, severe decline in these grassland environments has recently attracted attention to linear grasslands, such as firebreaks and power‐line corridors, as alternative habitats for grassland insects. We surveyed butterflies in an abandoned grassland and nearby linear mown firebreaks adjacent to different vegetation at the northern foot of Mt. Fuji, central Japan, over 5 successive years, particularly focusing on species on the 2012 Japanese Red List of Threatened Species (“red‐list” species). We found that the firebreaks were consistently higher in species richness and abundance of butterflies than the long‐term abandoned grassland, and that species composition differed among the firebreaks depending on conditions of the adjacent vegetation. The firebreaks surrounded by forests were mainly utilized by forest and edge species, whereas the firebreaks adjacent to the grassland were essential for conserving red‐list grassland species. Thus, only the mown firebreaks adjacent to the grassland were regarded as a high‐quality alternative habitat for many grassland butterflies, but the area was limited. Therefore, creating heterogeneity in the abandoned grassland by infrequent mowing could help conserve grassland butterflies, including red‐list species.     

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 effects on butterfly assemblages in an agricultural region

We examined the butterfly fauna at 62 sites in southeastern Sweden within a region exhibiting high variation in the landscape surrounding the studied grasslands. The landscape varied from an intensively-managed agricultural landscape with a large amount of open fields to a landscape with a high amount of deciduous forest/semi-natural grassland. We made 12 179 observations of 57 species of butterflies. The amount of neighbouring deciduous forest/semi-natural grassland, with >25% tree and bush cover, was the most important environmental factor explaining the variation in the butterfly assemblages. Landscape analyses at three different spatial scales showed that the variation in butterfly assemblages could be explained only at the largest scale (radius 5000 m) and not at the smaller ones (radii 500 and 2000 m).
Logistic regressions were used to predict presence/absence of butterfly species. Our study indicated that there may be critical thresholds for the amount of habitat at the landscape scale for several butterfly species as well as for species richness. For Melitaea athalia , there was a sharp increase in occupancy probability between 3 and 10% deciduous forests/semi-natural grasslands at the 5000-m scale. For 12 other species, the value for 50% probability of occurrence varied between 2 and 12% deciduous forest/semi-natural grassland. Species which had high occupancy probabilities in landscapes with a low amount of surrounding deciduous forests/semi-natural grasslands were significantly more mobile than others.
Our study highlights the importance of applying a landscape perspective in conservation management, and that single-patch management might fail in maintaining a diverse butterfly assemblage.     

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.     

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.

     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

The method of conserving herbaceous grassland specialists through silvicultural activities under deer browsing pressure

Early-successional herbs such as grassland specialists in semi-natural grasslands are decreasing because the intensity of using semi-natural grassland is decreasing for the decrease of human population or the usage of fossil fuel as energy source. Deer browsing also decreases grassland specialists. Silvicultural activities such as clearcutting, mowing of understory vegetation, and deer countermeasures may provide a suitable habitat for such grassland specialists. We aimed to clarify the method of conserving grassland specialists through silvicultural activities. We recorded all plant species that emerged in each quadrat, and measured their maximum heights within 80 quadrats that were established in four types of treatment sites in Yanagidaira, Japan. The four treatment sites were clearcut, fenced, and mowed (hereafter “fence site”); clearcut and mowed with tubed saplings (hereafter “tube site”); clearcut without deer countermeasures and mowing (hereafter “open site”); and not clearcut forest (hereafter “forest”). The total number of species and the number of species that occurred only in one treatment site of the four sites were highest in fence sites among the treatments. Those of tube and open sites were intermediate, whereas those of forest were the lowest. Grassland specialist forbs dominated in fence site, and grassland specialist graminoids and unpalatable grassland specialist forbs dominated in tube and open sites. The presence probabilities and maximum heights of grassland specialist forbs were significantly increased by deer exclusion, but those of grassland specialist graminoids were rarely affected by deer exclusion. Establishment of deer-proof fences in clearcut sites is effective in conserving grassland specialist forbs.     

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.     

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.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

Past and present disturbances influence biodiversity value of subtropical grassland ecological networks

The effect of transforming natural habitat to commercial land uses can be mitigated by implementing ecological networks (ENs) in production landscapes. However, EN ability to conserve biodiversity is potentially influenced by past and present disturbances, and spatial configuration of habitat patches. Here, we investigated how plant assemblages of a subtropical EN in a timber production landscape in South Africa respond to disturbance history (i.e. natural vs. recovering after removal of Pinus spp.), mowing in firebreaks (i.e. 30 m wide grassy strips intended to stop spread of fires in plantations), and three design parameters: corridor width, distance to protected area boundary, and context i.e. proportion eucalypt compartments within 500 m radius. Reference sites were in the adjacent iSimangaliso Wetland Park, a protected area (PA). Past disturbances and the current practice of mowing in firebreaks significantly influenced plant species composition, but not plant species richness. Plant dominance was greatest in mowed firebreaks, but similar between natural and recovering grassland. None of the design variables affected plant species richness, or composition. However, dominance was significantly and negatively correlated with corridor width, and positively correlated with context. Hence, sites are ordered in terms of their biodiversity value from greatest to lowest: natural areas > recovering areas > firebreaks. Based on these results, natural grassland in wide, well-connected corridors should have greatest conservation priority. We recommend that past disturbances, and current management practices should be taken into account and integrated into the design of future ENs. Such an approach is necessary for effective biodiversity conservation in production landscapes.     

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.     

Butterfly and grasshopper diversity patterns in humid Mediterranean grasslands: the roles of disturbance and environmental factors

The present paper studies butterfly, grasshopper and vascular plant communities in ten seasonally flooded grasslands with different anthropogenic disturbance regimes (NW Greece). Disturbance intensity was assessed on the basis of disturbance frequency and type (grazing, mowing, trampling, constructions). The distribution patterns of butterflies are regulated by humidity and elevation (Redundancy Analysis). Elevation, flower-heads abundance, low disturbance intensity and plant species richness predict grasshopper species richness well, while the latter together with humidity predict plant species richness (Generalized Linear Models). Chorthippus lacustris, a critically endangered endemic grasshopper species, is positively associated with humid microhabitats with high flower-heads abundance. An indicator value procedure reveals four butterfly species as being typical species for habitats with a pronounced character of hedgerows and tree lines. Conservation management of grassland butterflies should focus on the maintenance of the humid character of the humid grasslands as well as on the maintenance of hedgerows and tree lines. The reduction of human-induced disturbance towards occasional grazing and mowing seems to benefit both butterfly and grasshopper communities. Finally, we suggest the use of grasshoppers as surrogates for vascular plants and vice versa, given their congruent species richness patterns.     

Extensively managed or abandoned urban green spaces and their habitat potential for butterflies

Urban green spaces are often subject to intensive management and therefore only provide habitat for very few plant and animal species. Many studies have already proven that less intensive management of near-natural grassland habitats can increase species diversity, for example of butterflies. However, there is still a lack of knowledge as to what extent less intensive management of urban green spaces can promote urban butterfly diversity. Therefore, we examined parameters that determine the number of butterfly species in less maintained or even abandoned urban green spaces. We expected butterfly species richness and observation numbers to be affected by different management regimes. Butterfly surveys were carried out on 56 sites in the city of Hannover, Germany, in three consecutive years. The sites were subject to different management regimes. We explored parameters such as area size, the diversity of nectar plant species and habitat diversity for each site. A total of 34 species were recorded, with a minimum of 0 and a maximum of 17 species per site and year. The extreme drought and heat in 2018 clearly influenced the results of our study and this was reflected in massive decreases of observation numbers for some species. Less intensive management showed preferable effects in comparison to intensive management, but area size and nectar plants were more important factors influencing butterfly species richness and number of observations. Our results suggest that site parameters such as area size, nectar plant species richness or number of habitat types had stronger influences on butterflies than variations of extensive management. We conclude that less intensive urban green space management for good butterfly habitats is not a question of grazing, mowing or temporary abandonment, but instead must aim to increase nectar plant and habitat diversity.     

Relationships between the age of northern Kantou plain (central Japan) coppice woods used for production of Japanese forest mushroom logs and butterfly assemblage structure

To discern mechanisms maintaining the diversity of grassland and forest butterflies in coppice woods managed for the production of Japanese forest mushroom logs, we investigated the butterfly fauna in cut-over land tracts shortly after felling and 5 year later, and in forest stands 10, 15, and 25 year after felling (here, we use the term “forests” when referring to the chronosequence of these treed stands). Butterfly species richness and diversity (H′) and the densities of individuals were highest in cut-over lands 5 year after clear-cutting, followed by 25-year-old forest stands. In forests, the richness and densities of forest butterfly species were higher than were those of grassland species. Among forest stands of different ages, forest butterfly species’ richness and the densities of individuals were highest in 25-year-old woods nearing felling time. Some forest butterfly species were observed only in forests. The species richness and densities of grassland butterflies were much higher in cut-over lands 0 and 5 year post felling than in forests; grassland species were rarely found in stands ≥10 year old. Thus, cut-over lands seem to function as temporary habitats for grassland species. Furthermore, the number of forest butterfly species was the same in cut-over lands 5 year after felling and in 25-year-old forest stands; the densities of forest butterfly species was higher in these cut-over lands than in the forest stands. Forest butterfly species living on cut-over land 5 year post felling sipped flower nectar, laid eggs on host plants, and practiced territorial behaviour involved in mate finding. Hence, these cut-over lands functioned as important habitats for various developmental stages of forest butterflies. In conclusion, traditional coppicing in woods for production of Japanese forest mushroom logs is very important for the maintenance of diversity in grassland and forest butterfly species.     

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.     

Parasitism rate of Plebejus argyrognomon (Lepidoptera: Lycaenidae) under different levels of mowing management

Plebejus argyrognomon is an endangered grassland butterfly species in Japan. In this study we focused on the parasitism rate of tachinid flies under different mowing management to obtain knowledge of this possible top-down effect that has been largely ignored. We established three mowing treatments differing in the intensity of mowing, as represented by vegetation height. The results showed that the parasitism rate of P. argyrognomon increased with vegetation height; the parasitism rate in blocks subjected to the highest mowing intensity was 8%, whereas in blocks subjected to the lowest mowing intensity, the rate was 42%. We suggest that clarifying interspecific interactions is important for understanding how habitat management affects grassland-inhabiting butterflies in general.     

Conserving butterflies in fragmented plantation forests: are edge and interior habitats equally important?

Edge effects are increasing in forest-dominated landscapes worldwide, due to increased fragmentation by other land uses. Understanding how species respond to edges is therefore critical to define adequate conservation measures. We compared the relative importance of interior and edge habitats for butterflies in a landscape composed of even-aged pine plantations interspersed with semi-natural habitats. Butterfly assemblages were surveyed simultaneously at the edge and the interior of 68 patches belonging to four main habitat types: herbaceous firebreaks, clearcuts and young pine stands, older pine stands, and deciduous woodlands. Butterfly species richness was higher at edges than in interior habitats, especially for pine stands. Assemblage composition differed significantly between edge and interior habitats, except for firebreaks. Of the 23 most abundant butterfly species, seven were significantly more abundant in one or all edge habitat types, five in interior habitats, and 11 species showed no edge-interior preference. Modelling the presence of individual species in edge habitats revealed the importance of habitat variables such as the abundance of nectar and host-plants, but also of the abundance of the same species in the adjacent interior habitat. Moreover, our results suggest that most species use several, different habitat types to find supplementary or complementary resources, including micro-climatic refuges to escape hot temperatures during summer. The use of adjacent edge and interior habitats by butterflies is probably a key process in such mosaic landscapes and underlines the importance of landscape heterogeneity for butterfly conservation.     

Evaluating butterflies as surrogates for birds and plants in semi-natural grassland buffers

Semi-natural grasslands can support diverse faunal and floral communities, including grassland birds, beneficial insects, and native wildflowers. Monitoring biodiversity of this type of ecosystem is important to assess abundance and richness of grassland-associated species, evaluate success of establishing grasslands, and to assess overall ecosystem health. We tested butterflies as surrogates for birds and plants to assess establishment success of semi-natural grassland buffers in north-central Mississippi using Spearman rank correlation (Spearman’s ρ). Disturbance and grassland butterfly guilds were generally not suitable surrogates for grassland bird metrics, non-grassland bird metrics, or nest density metrics. Butterflies did have consistent positive correlations with plant species richness and forb metrics, as well as consistent negative correlations with grass metrics, but these correlations were generally smaller than what is considered suitable to serve as surrogates. In general, butterflies were not suitable surrogates for birds or plants in semi-natural grassland buffers.     

Importance of local habitat conditions and past and present habitat connectivity for the species richness of grassland plants and butterflies in power line clearings

Agricultural intensification has caused drastic declines in the area and species richness of semi-natural grasslands across Europe. Novel habitats, such as power line clearings, provide alternative habitats and niches for grassland species, and might therefore mitigate these declines. However, it is not fully understood which environmental factors determine the occurrence of grassland species in the clearings. Identifying the most important drivers for grassland species occurrence would help understand the value of the clearings for grassland conservation and target enhanced management into clearings with most potential as grassland habitat. We studied the effects of local environmental conditions, and past and present connectivity to semi-natural grasslands, on the species richness of grassland plants and butterflies in 43 power line clearings in Finland. The results of generalized linear models and hierarchical partitioning showed that increasing time since clear-cut and amount of clearing residue decreased the species richness of both species groups, while the cover of mesic habitats increased it. However, the two species groups showed also divergent responses. Present-day local environmental conditions appeared to be the sole driver of grassland butterfly species richness, whereas the richness of grassland plants was related both to current conditions and historical connectivity to grasslands in 1870–1880s. This suggests the presence of an extinction debt in the studied grassland plant communities, emphasizing the need for enhanced management to increase suitable grassland habitat in the clearings. This would diminish the potential future losses of grassland plant species in the clearings and create valuable habitat for grassland butterflies as well.