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.     

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.     

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.     

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.

     

Grasshopper and butterfly local congruency in grassland remnants

Grasshoppers could be considered as appropriate ecological indicators for grasslands owing to their sensitive response to environmental features. However, if grasshoppers are a good ecological indicator, they must (i) also represent other taxa, and (ii) provide additional information over straight measurement of environmental variables. To assess this, we compared the congruence of species richness patterns of grasshoppers with butterflies and environmental variables in two areas with extensive ecological networks (ENs). ENs are landscape-scale remnants of corridors and nodes of natural habitat running throughout a transformed, usually agricultural, landscape. Species richness of grasshoppers and butterflies did not differ among reference and EN sites, but guild composition differed significantly. While ENs adequately conserved overall diversity of these two groups, they were utilized preferentially by small-sized grasshoppers and shrub and tree-feeding butterflies. Reference sites had significantly more graminivorous and intermediate-mobility grasshopper species, as well as more butterfly species with widespread distribution, herbaceous dicot feeders and those with no recorded association to forest edges. Nevertheless, grasshopper and butterfly species richness’ were highly correlated. These results were similar across geographic areas, despite the fact that the areas differed significantly in their overall richness and species composition. Although there were some specific significant correlations between environmental variables and diversity, none of the variables could adequately replace use of the insect assemblage for bioindication. We conclude that grasshopper species and guild richness are representative of the butterfly assemblage, and provide information which is not sufficiently clear when utilizing only environmental variables.     

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.     

Fruit‐feeding Butterfly Communities are Influenced by Restoration Age in Tropical Forests

Currently, a large‐scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit‐feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit‐feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.     

How does landscape context contribute to effects of habitat fragmentation on diversity and population density of butterflies?

Aim Studies on habitat fragmentation of insect communities mostly ignore the impact of the surrounding landscape matrix and treat all species equally. In our study, on habitat fragmentation and the importance of landscape context, we expected that habitat specialists are more affected by area and isolation, and habitat generalists more by landscape context. Location and methods The study was conducted in the vicinity of the city of Göttingen in Germany in the year 2000. We analysed butterfly communities by transect counts on thirty‐two calcareous grasslands differing in size (0.03–5.14 ha), isolation index (2100–86,000/edge‐to‐edge distance 55–1894 m), and landscape diversity (Shannon–Wiener: 0.09–1.56), which is correlated to percentage grassland in the landscape. Results A total of 15,185 butterfly specimens belonging to fifty‐four species are recorded. In multiple regression analysis, the number of habitat specialist (n = 20) and habitat generalist (n = 34) butterfly species increased with habitat area, but z‐values (slopes) of the species–area relationships for specialists (z = 0.399) were significantly steeper compared with generalists (z = 0.096). Generalists, but not specialists, showed a marginally significant increase with landscape diversity. Effects of landscape diversity were scale‐dependent and significant only at the smallest scale (landscape context within a 250 m radius around the habitat). Habitat isolation was not related to specialist and generalist species numbers. In multiple regression analysis the density of specialists increased significantly with habitat area, whereas generalist density increased only marginally. Habitat isolation and landscape diversity did not show any effects. Main conclusions Habitat area was the most important predictor of butterfly community structure and influenced habitat specialists more than habitat generalists. In contrast to our expectations, habitat isolation had no effect as most butterflies could cope with the degree of isolation in our study region. Landscape diversity appeared to be important for generalist butterflies only.     

Effects of forest degradation on butterfly communities in the Gwangneung Forest

This study was carried out to clarify the response of butterfly communities on forest degradation in the Gwangneung Forest, Korea. We monitored butterfly communities with varying degrees of human activities by conducting a line transect twice a month in 2011. A total of 70 species and 4676 individuals of butterflies were observed in four sites: natural forest (NR), two plantation forests, and the Korean National Arboretum (AR). The result on niche breadth, habitat breadth and habitat type of butterfly was not consistent with our predictions. Species richness of habitat type was only significantly different between NF and AR. Species diversity was significantly different among sites. Butterfly diversity associated with landscape patterns based on aerial photographs supported a mosaic concept. A forest management plan to conserve butterfly diversity in forests is necessary to maintain various habitats and to ensure that grasslands are protected.     

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.     

松嫩平原不同生境蝴蝶群落多样性及其保护建议

随着气候变化加剧和人类活动影响，生物多样性变化及其保护逐渐受到广泛关注。蝴蝶作为开花植物的传粉媒介和生态环境监测及评价的关键指示者，其多样性变化能够在一定程度上反映生境状况，因此，有必要清晰认识不同生境中的蝴蝶多样性变化。为明确松嫩平原蝴蝶资源和不同生境的群落多样性差异，采用样线法于2016年5月-2018年8月对松嫩平原的割草草地、湿地、农田、放牧利用草地及恢复草地共五种生境类型进行调查研究。结果发现，调查共记录蝴蝶5108头，隶属于6科21属26种，其中牧女珍眼蝶（Coenonympha amaryllis）和红珠灰蝶（Plebejus argyrognomon）为优势种类，分别占蝴蝶个体总数的25.61%和31.66%，且在五种生境类型中均有分布。不同生境类型中，蝴蝶群落的物种丰富度指数和均匀度指数无明显差异，而恢复草地生境的蝴蝶群落Shannon-Wiener多样性指数较高，优势度指数较低。农田生境中的蝴蝶个体数量较少，且群落组成与其他四种生境之间均具有显著差异。五种生境类型中的蝴蝶数量和多样性均呈现一定的月动态和年动态变化趋势。除湿地和农田外，其余三种生境中蝴蝶物种和个体数量从5月到8月均持续升高。四种生境的蝴蝶物种数量、个体数量（除农田外）在2018年均出现明显下降趋势。物种丰富度指数等指标的月动态和年动态在不同生境类型间存在较大差异。这些结果表明，生境类型和人类活动与蝴蝶多样性变化关系密切，表现为单一生境中蝴蝶多样性较低，复杂生境有利于保护蝴蝶多样性。本研究有助于厘清松嫩平原蝴蝶资源的基础数据，并为该地区蝴蝶多样性保护和利用及评估该区域生态环境提供一定理论支撑。     

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.     

Low-Intensity Agricultural Landscapes in Transylvania Support High Butterfly Diversity: Implications for Conservation

European farmland biodiversity is declining due to land use changes towards agricultural intensification or abandonment. Some Eastern European farming systems have sustained traditional forms of use, resulting in high levels of biodiversity. However, global markets and international policies now imply rapid and major changes to these systems. To effectively protect farmland biodiversity, understanding landscape features which underpin species diversity is crucial. Focusing on butterflies, we addressed this question for a cultural-historic landscape in Southern Transylvania, Romania. Following a natural experiment, we randomly selected 120 survey sites in farmland, 60 each in grassland and arable land. We surveyed butterfly species richness and abundance by walking transects with four repeats in summer 2012. We analysed species composition using Detrended Correspondence Analysis. We modelled species richness, richness of functional groups, and abundance of selected species in response to topography, woody vegetation cover and heterogeneity at three spatial scales, using generalised linear mixed effects models. Species composition widely overlapped in grassland and arable land. Composition changed along gradients of heterogeneity at local and context scales, and of woody vegetation cover at context and landscape scales. The effect of local heterogeneity on species richness was positive in arable land, but negative in grassland. Plant species richness, and structural and topographic conditions at multiple scales explained species richness, richness of functional groups and species abundances. Our study revealed high conservation value of both grassland and arable land in low-intensity Eastern European farmland. Besides grassland, also heterogeneous arable land provides important habitat for butterflies. While butterfly diversity in arable land benefits from heterogeneity by small-scale structures, grasslands should be protected from fragmentation to provide sufficiently large areas for butterflies. These findings have important implications for EU agricultural and conservation policy. Most importantly, conservation management needs to consider entire landscapes, and implement appropriate measures at multiple spatial scales.     

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.     

Community composition of butterflies and bumblebees in fallows: niche breadth and dispersal capacity modify responses to fallow type and landscape

Fallows (i.e. fields temporarily taken out of production) provide important habitat for flower-visiting insects in intensively cultivated agricultural landscapes. Cost-efficiency of fallowing schemes could be enhanced through improved understanding of key characteristics of fallows and surrounding landscape that determine community composition and provide support for species of conservation concern. Impacts of fallow characteristics and landscape structure on the species composition of butterflies and bumblebees were studied in two types of perennial fallows in boreal farmland. To understand species’ responses to environmental factors from a conservation perspective, community composition was examined with respect to two species traits—niche breadth and dispersal capacity. Whereas overall species composition of butterflies and bumblebees was strongly affected by forest cover in the surrounding landscape, the studied species traits were most related to fallow type and the cover of perennial grasslands. Habitat breadth of butterflies was narrowest in long-term grassland fallows in landscapes with high grassland cover. Dispersal capacity of butterflies was also lowest in grassland-rich landscapes. Diet breadth of bumblebees was narrower in long-term grassland fallows than in short-term fallows. The results confirm that the diversity of butterflies and bumblebees can be enhanced by establishing and managing fallows both in open and forested landscapes. For conservation of habitat specialists and less mobile species, retention of long-term fallows in grassland-rich landscapes is apparently the best option. The results provide no justification for exempting forested regions or farms with high grassland cover from the ecological focus area requirement under the European Union’s current agricultural policy.     

Missing the target? A critical view on butterfly conservation efforts on calcareous grasslands in south-western Germany

Butterflies are strongly declining on grassland habitats of Central Europe. Therefore, the success of conservation measures on high quality grassland habitats is controversially discussed. We compared the changes in butterfly diversity and community structure on six managed calcareous grasslands with eight unmanaged vineyard fallows. We obtained strong losses of species diversity and remarkable shifts of community compositions on both habitat types. However, the changes on vineyard fallows were only slightly more severe but more stochastic than on the calcareous grasslands. The shifts in community composition with respect to functional species traits were rather similar between the two different grassland types so that complex butterfly communities evolved into generalist-dominated ones. Connectivity was higher among vineyard fallows than among calcareous grasslands. Consequently, conservation measures on calcareous grasslands only partly achieved their goal to maintain the high species diversity and functional complexity still observed in the 1970s. The negative impacts of eutrophication and monotonisation of the landscape as well as climate change are affecting all habitats, independently from management concepts. Therefore, management on conservation sites can buffer against these effects, but is not sufficient for a full compensation.     

Quality of remnant indigenous grassland linkages for adult butterflies (Lepidoptera) in an afforested African landscape

Retention of interconnected, remnant grassland linkages is proposed here to reduce the adverse effects of alien pine afforestation in Afromontane grasslands. Adult butterflies were sampled at 38 grassland sites, representing increasing levels of disturbance both within the afforested area and outside it. Butterfly species richness and abundance in the lesser disturbed grassland remnants within the afforested area were similar to those of the surrounding natural grasslands. In contrast, butterfly species richness, but not necessarily abundance, decreased significantly in the highly disturbed sites, both in the grassland linkages and outside. Although some highly disturbed sites were relatively rich in species, most were visited by geographically widespread and vagile species. In contrast, wide, relatively undisturbed grassland linkages, as well as grasslands outside, were important for localised, sedentary and local endemic butterfly species. Nectar plants, especially the alien Verbena bonariensis, were the most significant variable explaining local butterfly distribution. In addition, tall grasses, hills, topographical landmarks, thermoregulatory sites, shelter and water features were also vital for particular species. It did not matter how deep the grassland linkages were situated inside the afforested area, as long as they were made up of good habitat. Retention of wide, quality grassland linkages are a way forward to maximise biodiversity alongside agroforestry.     

长江三峡库区蝶类群落的等级多样性指数

在三峡库区蝶类物种多样性调查基础上,从等级多样性角度进行分析,结果表明,蝴蝶属级,科级和等级多样性指数在灌丛生境较高,森林、农田和草地生境多样性较低;非参数检验和方差分析结果表明4种生境在属级和等级多样性上差异显著,灌丛生境和其他生境差异显著。海拔500-1500m高度,蝴蝶的生物多样性相对较高,是库区蝴蝶最重要的生存地带。属级和科级多样性指数的变异系数在森林植被型样本间最大,前者有4个小生境在100％以上,后者有6个小生境超过100%;农田科的变异系数有1个小生境超过100%;草地的属和科变异系数均是1个小生境超过100%;灌丛则全部低于100%。这些数值,反映了三峡库区的形成对蝴蝶的生存及分布的产生了很大的影响,进一步证明库区蝶类生存环境的多样性和破碎化;其中,森林对蝴蝶生存的影响是最复杂的;通过蝴蝶做环境质量评估时,灌丛可作为首选考察对象。     

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.     

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.