Grassland Butterfly Fauna under Traditional Animal Husbandry: Contrasts in Diversity in Mown Meadows and Grazed Pastures

Semi-natural grasslands resulting from traditional land use practices (mowing and grazing) are severely endangered throughout Europe due to the intensification of agriculture. The ecological impact of mowing and grazing on grassland butterflies was studied in eight mown meadows and eight grazed pastures under traditional animal husbandry in NW Russia and adjacent Finland. Transect count data over 3 years (1997–1999) covered a total of 48 species and 5742 individuals. The butterfly fauna was rather similar under both management forms; species richness, diversity and total abundance did not differ significantly between meadows and pastures, yet meadows were preferred by more species. In both groups, the most abundant species were Aphantopus hyperantus, Pieris napi and Thymelicus lineola. Of 37 species observed as a minimum of five individuals, Polyommatus amandus, Ochlodes sylvanus and A. hyperantus showed a significant preference for mowing management. According to the ordination, butterfly communities were affected more by the origin and age of the grassland than the present management method. Landscape factors (meadow or pasture surrounded by forests or open environments), the abundance of nectar plants and the intensity of tilling were the most important factors differentiating older grasslands from the younger ones evolved from old Finnish hay fields cultivated prior to the 1940s.     

Impact of logging on the richness and diversity of forest butterflies in a tropical dry forest in Thailand

Studies on the impact of logging on tropical forest butterflies have been almost exclusively conducted in moist forest habitats. This study considers the impacts of small-scale logging on butterfly communities at three sites of varying disturbance intensity in a tropical dry forest in western Thailand. Butterfly species richness was similar at all sites, but the abundance of butterflies and diversity of the butterfly community decreased with increased logging disturbance. The recorded decrease in diversity at the relatively large sampling scale used lends further support to the hypothesis that disturbance effects are scale dependent. Species abundance data for butterflies fitted a log-normal distribution at all sites, but also a log-series distribution at the two disturbed sites. These analyses suggest a more complex butterfly community at the undisturbed site, but also that log-series and log-normal distributions may not to be sufficiently sensitive to be useful indicators of community changes following logging. Community ordination separates both the butterfly species and transect samples into three distinct regions corresponding to the three study locations. Ordination axes are correlated with tree density, understorey cover and understorey plant richness. Species with the smallest geographic ranges tend to be the least abundant and occurred most frequently in the undisturbed site. The observed diverging responses to disturbance among butterfly families diminishes the value of butterfly communities as biodiversity indicators, and forest managers should perhaps focus on restricted range species or of groups of recognized sensitive species for this purpose.     

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.     

Haying and grazing effects on the butterfly communities of two Mediterranean-area grasslands

Grasslands are usually the most suitable environment for butterflies, but have been also traditionally used for productive activities. This paper compares the impact of mowing and grazing on butterfly biodiversity in two S Italian (Campania) grasslands, at Campo Somma (CS) and Pianoro di Prada (PP) located at an identical altitude of 850 m. These grasslands have an area of approximately 6 ha each and are at a 3 km distance from each other. They share similar climate and are both surrounded by woods, mainly of sweet chestnut. CS is managed for haying and is mown once a year, in June. PP is used for sheep and cattle grazing. Weekly transects were made from April to September in 2008 and 2009, during the butterfly flight activity. We analysed data from eight monthly transects by Kruskal–Wallis and Mann–Whitney tests. We observed 45 butterfly species at PP, and 28 at CS. Diversity indexes were significantly higher at PP, whereas evenness was similar. The monthly comparisons of species richness showed that, except for April, PP values were always significantly higher. Species abundance was significantly higher at PP in June, July and August. In all months, except in April, the Shannon–Wiener index was also significantly higher at PP, as was Simpson’s index in May, June and September. Dominance index differed significantly only in May, June and September, whereas evenness was never significantly different between the two grasslands. These data show that, as concerns butterflies, the impact of mowing was much stronger than that of grazing.     

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

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

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.     

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.     

不同放牧强度对荒漠草原蝗虫群落的影响

采用无底样框法对四子王旗荒漠草原不同放牧强度下的蝗虫群落进行了取样调查,比较了不同放牧强度下荒漠草原蝗虫群落的丰富度、多样性和均匀度指数及群落中蝗虫种类和数量随季节的动态变化,并对蝗虫种群分布与植物群落特征进行了相关性分析。结果表明,在不同放牧强度下蝗虫的群落结构及时间动态均存在一定差异,说明放牧活动对蝗虫群落结构有明显影响,其中,轻度放牧和重度放牧样地中蝗虫种群总数显著高于其他样地。不同种类的蝗虫群落与放牧强度之间关系表现复杂。     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

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.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Impacts of grazing intensity and increased precipitation on a grasshopper assemblage (Orthoptera: Acrididae) in a meadow steppe

1. Grasshoppers are dominant herbivores in grassland ecosystems, and many studies have examined how grazing by large herbivores and precipitation patterns individually influence the dynamics of grassland grasshopper assemblages, but their combined effects are largely unknown. 2. In this study, grazing intensities (ungrazed, moderate, and heavy) were manipulated and precipitation (ambient and increased amount of rainfall) altered in a field experiment to test the effects of grazing and altered precipitation on a grasshopper community in a meadow steppe in northeastern China. 3. It was found that grasshopper species richness did not change according to different grazing intensities under ambient precipitation, but was significantly higher (by 38.1%) in moderate grazing intensities under increased precipitation. Grasshopper abundance increased considerably with increasing grazing intensities in ambient precipitation treatments; however, grasshopper abundance in heavy grazing intensities was significantly lower (by 32.9%) than in the other two grazing intensities under increased precipitation. Moreover, the responses of grasshopper abundance to grazing under altered precipitation were species‐specific. 4. Grazing effects on grasshopper species diversity were mediated through the species richness and biomass of grasses (food resources), but the effects on grasshopper abundance were mediated through plant height (vegetation structure) under altered precipitation. 5. These results suggest that appropriate grazing by large herbivores would be considered as beneficial management practices for maintaining grasshopper diversity and abundance under conditions of increased precipitation in grassland ecosystems. Additionally, greater attention should be paid to the population dynamics of different grasshopper species to better understand the responses of grasslands to grazing and altered precipitation.     

Habitat type plays a greater role than livestock grazing in structuring shrubsteppe plant–pollinator communities

Livestock grazing is a widespread grassland disturbance and can negatively impact biodiversity. Pollinators constitute a vital component of grassland ecosystems, but the impact of grazing on pollinator diversity has seldom been evaluated in North America. We assessed vegetation structure, and pollinator and flowering plant abundance, richness, diversity, and community composition in four pairs of spring-grazed/ungrazed sites in south-central British Columbia, Canada. We also investigated whether pollinator or floral communities differed between the two threatened shrubsteppe habitat types we sampled—antelope-brush and big sagebrush shrubsteppe. Pan-trapping surveys captured 5907 bees, flies, beetles, wasps and butterflies constituting 253 species. We found that the percent cover of shrubs and bare soil increased with grazing, while the height of grasses and forbs decreased. In contrast, pollinator and flowering plant abundance, richness, diversity, and community composition were not significantly affected by grazing. Flowering plant and pollinator community composition did differ significantly between shrubsteppe habitats. Our results indicate that grasslands in North America, when managed responsibly, can maintain pollinator and flowering plant diversity under grazing pressure. The continued effort of land managers to balance ecological integrity and economic viability will be important for the conservation of grassland pollination systems.     

Ants and plants as indicators of biodiversity,ecosystem services,and conservation value in constructed grasslands

Grasslands are constructed for soil and wildlife conservation in agricultural landscapes across Europe and North America. Constructed grasslands may mitigate habitat loss for grassland-dependent animals and enhance ecosystem services that are important to agriculture. The responses of animal species richness and abundance to grassland habitat quality are often highly variable, however, and monitoring of multiple taxa is often not feasible. We evaluated whether multiple animal taxa responded to variation in constructed grassland habitats of southwest Ohio, USA, in ways that could be predicted from indicators based on quality assessment indices, Simpson diversity, and the species richness of ants and plants. The quality assessment indices included a widely used Floristic Quality Assessment (FQA) index, and a new Ant Quality Assessment (AntQA) index, both based on habitat specificity and species traits. The ant and plant indicators were used as predictor variables in separate general linear models of four target taxa—bees, beetles, butterflies and birds—with response variables of overall species richness and abundance, and subsets of taxa that included the abundance of ecosystem-service providers and grassland-associated species. Plant Simpson diversity was the best-fitting predictor variable in models of overall bee and beetle abundance, and the abundance of bees classified as ecosystem-service (ES) providers. FQA and plant richness were the best predictors of overall butterfly species richness and abundance. Ant species richness was the best predictor of overall bird species richness and abundance as well as the abundance of ES birds, while the AntQA index was the best predictor for the abundance of grassland bird and butterfly species. Thus, plant Simpson diversity and ant species richness were the most effective indicators for complementary components of grassland animal communities, whereas quality assessment indices were less robust as indicators and require more knowledge on the habitat specificity of individual ant and plant species.     

Butterfly Community Composition Across a Successional Gradient in a Human‐disturbed Afro‐tropical Rain Forest

Knowledge of the recovery of insect communities after forest disturbance in tropical Africa is very limited. Here, fruit‐feeding butterflies in a tropical rain forest at Kibale National Park, Uganda, were used as a model system to uncover how, and how fast, insect communities recover after forest disturbance. We trapped butterflies monthly along a successional gradient for one year. Traps were placed in intact primary forest compartments, heavily logged forest compartments with and without arboricide treatment approximately 43 years ago, and in conifer‐clearcut compartments, ranging from 9 to 19 years of age. The species richness, total abundance, diversity, dominance, and similarity of the community composition of butterflies in the eight compartments were compared with uni‐ and multivariate statistics. A total of 16,728 individuals representing 88 species were trapped during the study. Butterfly species richness, abundance, and diversity did not show an increasing trend along the successional gradient but species richness and abundance peaked at intermediate stages. There was monthly variation in species richness, abundance, diversity and composition. Butterfly community structure differed significantly among the eight successional stages and only a marginal directional change along the successional gradient emerged. The greatest number of indicator species and intact forest interior specialists were found in one of the primary forests. Our results show that forest disturbance has a long‐term impact on the recovery of butterfly species composition, emphasizing the value of intact primary forests for butterfly conservation.     

区域变动尺度上不同生境类型及环境因子对西双版纳蝴蝶多样性的影响

【目的】生境类型和环境因子对物种分布和维持具有重要的影响。本研究通过分析不同生境类型对蝴蝶群落多样性及其群落结构影响的差异,以及环境因子对蝴蝶物种丰富度和多度的影响,为区域变动尺度蝴蝶多样性维持机制的研究奠定基础。【方法】于2019年8月和10月,在西双版纳地区采用样线法,调查了天然林、次生林、复合生境、人工林和农田5种生境中蝴蝶的物种,分析了蝴蝶群落多样性、群落结构相似性及物种丰富度和多度与环境因子的关系。【结果】2019年从西双版纳共采集蝴蝶2 226头,隶属于11科98属175种,在西双版纳州级尺度上蝴蝶物种丰富度高于县域尺度。在西双版纳州级尺度上,蝴蝶的物种丰富度和多度在5种生境间存在显著差异,而在县域尺度上,物种丰富度、多度和Chao 1物种丰富度估计值没有一致性规律。群落结构相似性结果显示,在西双版纳州级尺度上,蝴蝶群落结构在不同生境类型间存在极显著差异,在县域尺度上,仅勐腊区域蝴蝶群落结构在不同生境类型间存在显著差异。蝴蝶物种丰富度和多度不仅受到生境类型的影响,还受到温度、年均降水和海拔的影响。【结论】本研究结果表明,在区域变动尺度上,生境类型对西双版纳蝴蝶的多样性的影响较大,而温度、年均降水和海拔是维持蝴蝶物种多样性的重要因素。这些发现对当前人类导致的生境丧失和气候变化时代生物多样性的保护具有重要意义。     

Significant decrease in local butterfly community during the last 15 years in a calcareous hill of the middle Korea

Recently the significant decreases of species richness and abundance among terrestrial animals including butterflies are reported due to habitat change, overexploitation, and global warming. We compared the butterfly species composition and abundance from 1999 and 2014–2015 in a calcareous hill site of the middle part of Korea using a line transect method. There was a significant decrease in the number of individuals (abundance) and the number of species (richness) from 1999 to 2014–2015. This decrease was more prevalent among northern species than southern species, and the local extinct species were more prevalent among northern species, showing the influence of global warming on butterfly assemblages. However, no impact of habitat change was observed because of maintenance of the grasslands, which is caused by the dry soils of the calcareous region.     

Effects of managing semi-natural grassland buffers on butterflies

Butterflies are important components of biodiversity in grassland ecosystems and provide ecosystem services such as pollination. Although agricultural intensification has led to a scarcity of native grassland habitats within most agricultural landscapes of North America, fragmented remnants and semi-natural habitats may support diverse communities, including butterflies, as long as vital resources such as host plants are available. The United States Department of Agriculture’s (USDA) Conservation Reserve Program practice CP33 Habitat Buffers for Upland Birds (USDA 2004) provides semi-natural grassland habitat in agricultural landscapes, but a knowledge gap exists about impacts of prescribed disturbance (e.g. burning or disking) on butterflies. We monitored butterfly habitat and butterfly communities on experimentally manipulated CP33 grassland buffers in Clay County, Mississippi from 2007 to 2009. Disturbance guild butterfly species richness did not differ among treatments. However, disturbance guild abundance was positively affected by disking in both the first and second growing seasons following disking, and the magnitude of this response varied between years. Effects of burning on disturbance guild abundance did not differ from the control treatment. There were no treatment differences for grassland guild butterfly abundance and species richness suggesting that periodic disturbance does not unduly impact grassland-associated butterflies in the southeastern US. Our results support current USDA practice standards that require periodic disturbance during the 10-year contract, but restrict the disturbance to 1/3 or 1/4 of grassland buffer area in a given year.     

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

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

Butterfly community structure in fragmented habitats

We analysed effects of habitat fragmentation on the diversity, abundance, and life history traits of butterflies on 33 calcareous grasslands. Diversity of butterflies was positively correlated with habitat area (as was plant diversity), but not with habitat isolation. In contrast to expectations, butterfly densities of polyphagous and oligophagous species declined with habitat area whereas densities of monophagous species increased. The z -values, i.e. the slope of species–area relationships, increased with food plant specialization, from 0.07 in polyphagous, 0.11 in oligophagous, 0.16 in strongly oligophagous to 0.22 in monophagous species, and were 0.14 in plant species. Significant z -values were not only found for total species richness, based on a sample size adjusted to fragment area ( z  = 0.12), but also for the local density of butterfly species richness, based on equal sample size across all habitat fragments ( z  = 0.06). To our knowledge, this is the first study to show differential responses of monophagous, oligophagous and polyphagous species to area with respect to species richness and population density.