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.     

Species diversity of remnant calcareous grasslands in south eastern Germany depends on litter cover and landscape structure

Species diversity depends on, often interfering, multiple ecological drivers. Comprehensive approaches are hence needed to understand the mechanisms determining species diversity. In this study, we analysed the impact of vegetation structure, soil properties and fragmentation on the plant species diversity of remnant calcareous grasslands, therefore, in a comparative approach.We determined plant species diversity of 18 calcareous grasslands in south eastern Germany including all species and grassland specialists separately. Furthermore, we analysed the spatial structure of the grasslands as a result of fragmentation during the last 150 years (habitat area, distance to the nearest calcareous grassland and connectivity in 1830 and 2013). We also collected data concerning the vegetation structure (height of the vegetation, cover of bare soil, grass and litter) and the soil properties (content of phosphorous and potassium, ratio of carbon and nitrogen) of the grassland patches. Data were analysed using Bayesian multiple regressions.We observed a habitat loss of nearly 80% and increasing isolation between grasslands since 1830. In the Bayesian multiple regressions the species diversity of the studied grasslands depended negatively on cover of litter and to a lower degree on the distance to the nearest calcareous grassland in 2013, whereas soil properties had no significant impact.Our study supports the observation that vegetation structure, which strongly depends on land use, is often more important for the species richness of calcareous grasslands than fragmentation or soil properties. Even small and isolated grasslands may, therefore, contribute significantly to the conservation of species diversity, when they are still grazed.     

Fragmentation in calcareous grasslands: species specialization matters

Habitat fragmentation resulting from anthropogenic land-use change may negatively affect both biodiversity and ecosystem structure and function. However, susceptibility to fragmentation varies between species and may be influenced by for instance specialization, functional traits and trophic level. We examined how total and specialist species richness, species composition and functional trait composition at two trophic levels (vascular plants and sap-feeding hoppers) vary with habitat fragmentation (patch size and connectivity) in dry calcareous grasslands in southeast Norway. We found that fragmentation affected plant and hopper species composition both totally and of habitat specialists, but with a net species loss only for the specialists, indicating greater susceptibility of specialized species. Reductions in patch size and increasing isolation negatively affected plant specialists with different sets of traits, effectively reducing the number of species with trait combinations suitable to persist in small and isolated patches. Fragmentation influenced trait composition of the total hopper community, but not of habitat specialists. A lesser degree of habitat association could explain why hoppers, despite belonging to a higher tropic level, seemed to be less susceptible to fragmentation than plants. Nonetheless, our study shows that habitat fragmentation affects both species richness, species composition and trait composition of plants and hoppers, indicating that fragmentation leads not only to a loss of species, but also alters dominance hierarchies and the functionality of grassland communities.     

After the hotspots are gone: Land use history and grassland plant species diversity in a strongly transformed agricultural landscape

Question: We asked how landscape configuration and present management influence plant species richness and abundance of habitat specialists in grasslands in a ‘modern’(much exploited and transformed) agricultural Swedish landscape. Location: Selaön, south‐eastern Sweden (59°24’ N, 17°10’ E). Methods: Present and past (150 and 50 years ago) landscape pattern was analysed in a 25 km² area. Species richness was investigated in 63 different grassland patches; grazed and abandoned semi‐natural grasslands, and grazed ex‐arable fields. Influence of landscape variables; area, past and present grassland connectivity, present management on total species richness, density and abundance of 25 grassland specialists was analysed. Results: Semi‐natural grasslands (permanent unfertilised pastures or meadows formed by traditional agricultural methods) had declined from 60% 150 years ago to 5% today. There was a significant decline in species richness and density in abandoned semi‐natural grasslands. Total species richness was influenced by present management, size and connectivity to present and past grassland pattern. Landscape variables did not influence species density in grazed semi‐natural grassland suggesting that maintained grazing management makes grassland patches independent of landscape context. The abundance of 16 grassland specialists was mainly influenced by management and to some extent also by landscape variables. Conclusion: Although species richness pattern reflect management and to some extent landscape variables, the response of individual species may be idiosyncratic. The historical signal from past landscapes is weak on present‐day species richness in highly transformed, agricultural landscapes. Generalizations of historical legacies on species diversity in grasslands should consider also highly transformed landscapes and not only landscapes with a high amount of diversity hotspots left.     

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

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

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

Tracking the effects of one century of habitat loss and fragmentation on calcareous grassland butterfly communities

Habitat loss and fragmentation are known to reduce patch sizes and increase their isolation, consequently leading to modifications in species richness and community structure. Calcareous grasslands are among the richest ecosystems in Europe for insect species. About 10% (1,150 ha) of the total area of a calcareous ridge region (Calestienne, Belgium) and its butterfly community was analysed over a timeframe of about 100 years. Since 1905 to present day (2005), the Calestienne region has undergone both calcareous grassland loss and fragmentation: not only did calcareous grassland size decrease and isolation increase, but also, the number of calcareous grassland patches within the landscape increased until 1965, and subsequently decreased, clearly reflecting the effects of fragmentation. These processes have had a profound effect on the butterfly community: extinction and rarefaction affected significantly more often specialist species, which means that generalist species are more and more overrepresented. This ecological drift, i.e. the replacement of specialists by generalists in species assemblages is likely to be a general effect of habitat loss and fragmentation on natural communities.     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

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.     

Nitrogen deposition causes widespread loss of species richness in British habitats

We use national scale data to test the hypothesis that nitrogen (N) deposition is strongly negatively correlated with plant species richness in a wide range of ecosystem types. Vegetation plots from a national ecological surveillance programme were drawn from heathland, acid, calcareous and mesotrophic grassland habitats. Mean species number and mean plant traits were calculated for each plot and related to atmospheric N deposition. There was a significant reduction in species richness with N deposition in acid grassland and heathland even after fitting covarying factors. In acid grassland and heathland, evidence from trait changes suggested that acidification rather than increased fertility was responsible for species loss. In contrast, calcareous grassland showed evidence of eutrophication in response to increasing N deposition. Loss of species richness from chronic N deposition is apparent in infertile grasslands and heathland. Mechanisms associated with loss of species richness differ between habitats so mitigation of N deposition should be targeted to habitat type.     

Trait-Specific Responses of Wild Bee Communities to Landscape Composition,Configuration and Local Factors

Land-use intensification and loss of semi-natural habitats have induced a severe decline of bee diversity in agricultural landscapes. Semi-natural habitats like calcareous grasslands are among the most important bee habitats in central Europe, but they are threatened by decreasing habitat area and quality, and by homogenization of the surrounding landscape affecting both landscape composition and configuration. In this study we tested the importance of habitat area, quality and connectivity as well as landscape composition and configuration on wild bees in calcareous grasslands. We made detailed trait-specific analyses as bees with different traits might differ in their response to the tested factors. Species richness and abundance of wild bees were surveyed on 23 calcareous grassland patches in Southern Germany with independent gradients in local and landscape factors. Total wild bee richness was positively affected by complex landscape configuration, large habitat area and high habitat quality (i.e. steep slopes). Cuckoo bee richness was positively affected by complex landscape configuration and large habitat area whereas habitat specialists were only affected by the local factors habitat area and habitat quality. Small social generalists were positively influenced by habitat area whereas large social generalists (bumblebees) were positively affected by landscape composition (high percentage of semi-natural habitats). Our results emphasize a strong dependence of habitat specialists on local habitat characteristics, whereas cuckoo bees and bumblebees are more likely affected by the surrounding landscape. We conclude that a combination of large high-quality patches and heterogeneous landscapes maintains high bee species richness and communities with diverse trait composition. Such diverse communities might stabilize pollination services provided to crops and wild plants on local and landscape scales.     

Effects of vegetation structure and floristic diversity on detritivore,herbivore and predatory invertebrates within calcareous grasslands

Calcareous grasslands in Europe have shown wide scale declines in their extent and quality as a result of modern agricultural practices, increased atmospheric eutrophication and lack of management. In addition to being a key habitat for specialist plants, calcareous grasslands are also important for many threatened invertebrates. In this UK based study, we investigated the impact of military vehicle activity, floral species richness and vegetation structure on assemblages of detritivore, herbivore and predatory invertebrates. We also consider the impact that disturbance by military vehicle activity on the proportion of invertebrate species capable of flight, a surrogate for dispersal ability. Sward height was negatively correlated with detritivore, herbivore and predator species richness. Herbivores species richness was positively correlated with both forb and grass species richness. Spatial variation in the number of plant species was negatively correlated with herbivore species richness. Those sites most heavily disturbed by military vehicle activity supported the lowest proportions of flightless invertebrates. Successful management for calcareous grassland invertebrates should aim to maintain short swards with high floristic diversity, in terms of both the forbs and grasses. It should be noted, however, that these findings refer to principally surface rather than sward active invertebrates. While disturbance associated with military vehicle activity was not found to affect invertebrate species richness, it has negative consequences for the structure of invertebrate assemblages by selecting against invertebrates with low dispersal ability. To support invertebrate diversity in calcareous grasslands we emphasise the need for variety in the timing and type of management applied to promote heterogeneity in sward structure.     

Size and heterogeneity rather than landscape context determine plant species richness in semi‐natural grasslands

Question: Is plant diversity in fragmented semi‐natural grasslands related to present and historical landscape context? Location: Southern Sweden. Methods: Plant diversity was described at 30 semi‐natural grassland sites in terms of total and specialist plant species richness at the site and species density at different scales (0.5–10 m²). These measures are commonly used to assess conservation value of semi‐natural grasslands. Landscape context was measured as contemporary connectivity to other semi‐natural grasslands, historical connectivity 50 years ago, amount of linear elements potentially suitable for dispersal (road verges, power line clearings), and amount of forest (inverse of the openness of the landscape). Results: The diversity measures were generally correlated with each other, implying that species richness in a subset of the grassland can predict the total richness. Plant species density at three scales (0.5 m², 10 m² and total) was related to the landscape context using an information theoretic approach. Results showed that total species richness increased with increased size of grasslands, contrary to earlier diversity studies in semi‐natural grasslands. Larger grasslands were more heterogeneous than smaller grasslands, and this is a likely reason for the species‐area relationship. Heterogeneity was also of high importance at the smaller scales (0.5 m², 10 m²). With increased amount of forest, total species richness increased but species density on 10 m² decreased. There was no influence of connectivity in either the contemporary or the historical landscape, contrary to previous studies. Conclusions: Grassland size and heterogeneity are of greater importance for plant diversity in semi‐natural grassland, than grassland connectivity in the landscape.     

Remnant habitats for grassland species in an abandoned Swedish agricultural landscape

Questions: Which factors influence the persistence of vascular grassland plants in long‐abandoned (at least 50 yr) arable fields and meadows? What might be the implications of current levels of species richness on abandoned arable fields and meadows for future restoration? Location: Forested highlands of Kilsbergen, south central Sweden. Methods: The abundance of all vascular plant species was investigated in three habitat types: former arable fields, hay meadows and outlands (pastures) at 27 farms, abandoned for either approximately 50 yr or 90 yr. Time since abandonment, tree cover, soil depth, degree of soil podsol development, size of the infield area and two measures of connectivity were used as predictors for species richness and species composition. Results: Former outland had denser tree cover, fewer species and fewer grassland species than former arable fields and hay meadows, irrespective of time since abandonment. Former hay meadows and arable fields with a longer time since abandonment were less rich in species, more wooded and had greater podsolization than meadows and fields abandoned at a later stage. Species richness was higher in hay meadows and arable fields at farms with larger infield area and deeper soils compared with farms with smaller infield area and shallower soils. The greatest richness of species and most open habitat were former arable fields at larger farms abandoned 50 yr before the study. Former arable fields had the highest number of grassland species. Conclusion: After 50 yr of abandonment, former arable fields were the most important remnant habitats for grassland species and may be a more promising target for restoration than formerly managed grasslands.     

Landscape-scale connectivity and fragment size determine species composition of grassland fragments

As a consequence of agricultural intensification and habitat fragmentation since the mid-20th century, biological diversity has declined considerably throughout the world, particularly in Europe. We assessed how habitat and landscape-scale heterogeneity, such as variation in fragment size (small vs. large) and landscape configuration (measured as connectivity index), affect plant and arthropod diversity. We focused on arthropods with different feeding behaviour and mobility, spiders (predators, moderate dispersal), true bugs (mainly herbivores and omnivores with moderate dispersal), wild bees (pollinators with good dispersal abilities), and wasps (pollinators, omnivores with good dispersal abilities). We studied 60 dry grassland fragments in the same region (Hungarian Great Plain); 30 fragments were represented by the grassland component of forest-steppe stands, and 30 were situated on burial mounds (kurgans). Forest-steppes are mosaics of dry grasslands with small forests in a matrix of plantation forests. Kurgans are ancient burial mounds with moderately disturbed grasslands surrounded by agricultural fields. The size of fragments ranged between 0.16–6.88 ha (small: 0.16–0.48 ha, large: 0.93–6.88 ha) for forest-steppes and 0.01–0.44 ha (small: 0.01–0.10 ha and large: 0.20–0.44 ha) for kurgans. Fragments also represented an isolation gradient from almost cleared and homogenous landscapes, to landscapes with relatively high compositional heterogeneity. Fragment size, connectivity, and their interaction affected specialist and generalist species abundances of forest-steppes and kurgans. Large fragments had higher species richness of ground-dwelling spiders, and the effect of connectivity was more strongly positive for specialist arthropods and more strongly negative for generalists in large than in small fragments. However, we also found a strong positive impact of connectivity for generalist plants in small kurgans in contrast to larger ones. We conclude that besides the well-known effect of enhancing habitat quality, increasing connectivity between fragments by restoring natural and semi-natural habitat patches would help to maintain grassland biodiversity.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

Relative importance of habitat area, connectivity, management and local factors for vascular plants: spring ephemerals in boreal semi-natural grasslands

Using species and environmental data from an extensive grassland area in south-western Finland, we investigated the effect of patch area and connectivity, management and local habitat variables on the occurrence of spring-flowering vascular plants and their richness in boreal agricultural landscapes. Generalized linear models (GLM) and variation partitioning were used to study the explanatory power of the three groups of variables and their combined contributions on the richness and occurrence of six spring-flowering plant species. Generalized additive models (GAMs) and associated cross-validation tests were used to evaluate the predictability of the species occurrence and richness patterns. Present-day grassland patch area and connectivity were important predictors for occurrence and richness of the studied plant species. In addition, local habitat factors, especially radiation, accounted for major fractions of occurrence patterns of the studied species. Hybrid models including variables from all three variable groups had higher explanatory power and predictive capability than partial models. However, performance of the separate single-species models varied considerably between the six study species. Exclusion of radiation or connectivity from the hybrid models decreased their predictive performance, suggesting that these factors are of particular importance for grassland plant species at their northern range margins. When developing conservation and management planning for grassland plant species in Northern Europe, attention should be paid to well-connected networks of grassland patches including large, steeply-sloped patches with a favorable microclimate.     

The significance of habitat continuity and current management on the compositional and functional diversity of grasslands in the uplands of Lower Saxony,Germany

There is a growing concern that land use intensification is having negative effects on semi-natural grasslands and that it leads to a general loss of biodiversity among all types of formerly extensively managed grasslands of poor to medium nutrient richness. Since the 1950s, many Central European uplands have been subject to an increase in grassland cover as a result of changes in land use practices. Using such a landscape in Lower Saxony, Germany, as a model region, we assessed environmental factors that control grassland diversity, including plant community composition, species richness and pollination trait composition. In 2007, 189 vegetation sampling sites were randomly distributed among grasslands covering some 394 ha within a 2500 ha study area. Plant communities were classified using TWINSPAN and the effects of environmental factors (soil, topography, current management and habitat continuity) were analysed by canonical correspondence analysis and regression analysis reducing for the effects of spatial autocorrelation by using principal coordinates of neighbour matrices.We found a wide range of six species-poor (<15 plant spp.) to extremely species-rich (>27 spp.) grassland types under mesic to dry site conditions, including sown, Cynosurion, Arrhenatherion and semi-natural grasslands. Grassland community composition was best explained by soil factors and species richness and pollination type composition by combined effects of current management and habitat continuity. During the 1950/60s, the extent of grassland area within the studied landscape rapidly increased to more than double its previous extent, and in 2007, grasslands comprised 16%. Natura 2000 grassland types comprised 1% of the surveyed site and medium-rich, high-nature-value grasslands a further 5%. While the number of wind-pollinated plant species was equal among all grassland types, there was a parallel decline in insect-pollinated plants and overall median species richness in the grassland communities along a gradient of increasing land use intensity (mowing, nutrient supply). Moreover, insect-pollinated plants occurring in intensively managed grasslands were found to additionally have the ability for self-pollination. Species-rich grasslands – including semi-natural grasslands and a semi-improved, species-rich Arrhenatherion community – occurred exclusively on old sites (with >100 years of habitat continuity) that had been used for traditional sheep grazing (environmental contracting). Medium-rich Arrhenatherion grasslands were established primarily on less productive, formerly arable fields (<30 years). We conclude that conservation efforts should focus on extant species-rich grassland types and should aim to implement traditional land use practices such as sheep grazing. Additional restoration efforts should focus on establishing new grasslands on less productive sites in the proximate surroundings of species-rich grasslands to facilitate seed dispersal, but nitrogen deposition should be buffered where appropriate. These measures would enhance the interaction between nature reserves and agricultural grasslands and thus improve the ecological quality of grasslands at the landscape scale.     

Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands

Metapopulation theory predicts that species richness and total population density of habitat specialists increase with increasing area and regional connectivity of the habitat. To test these predictions, we examined the relative contributions of habitat patch area, connectivity of the regional habitat network and local habitat quality to species richness and total density of butterflies and day-active moths inhabiting semi-natural grasslands. We studied butterflies and moths in 48 replicate landscapes situated in southwest Finland, including a focal patch and the surrounding network of other semi-natural grasslands within a radius of 1.5 km from the focal patch. By applying the method of hierarchical partitioning, which can distinguish between independent and joint contributions of individual explanatory variables, we observed that variables of the local habitat quality (e.g. mean vegetation height and nectar plant abundance) generally showed the highest independent effect on species richness and total density of butterflies and moths. Habitat area did not show a significant independent contribution to species richness and total density of butterflies and moths. The effect of habitat connectivity was observed only for total density of the declining butterflies and moths. These observations indicate that the local habitat quality is of foremost importance in explaining variation in species richness and total density of butterflies and moths. In addition, declining butterflies and moths have larger populations in well-connected networks of semi-natural grasslands. Our results suggest that, while it is crucial to maintain high-quality habitats by management, with limited resources it would be appropriate to concentrate grassland management and restoration to areas with well-connected grassland networks in which the declining species currently have their strongest populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Landscape structure and management regime as indicators of calcareous grassland habitat condition and species diversity

This study investigates the importance of spatial landscape characteristics and habitat management on the condition of calcareous grassland in the North Down Natural Area, Kent UK. We used a digitised map of the study area containing shapefiles of all the habitats including 82 patches of calcareous grassland together with management information for each patch and data on the presence and abundance of a range of calcareous grassland indicator plant species. We defined habitat condition by presence of indicator species and used classification trees to generate models with rules for predicting habitat condition from the landscape spatial characteristics and management information. We also applied the same method to investigate the factors affecting presence or diversity of three ecological groups of positive indicator species and dominance of a negative indicator species. All the models except one showed good classification accuracy and high kappa statistic. Favourable habitat condition was predicted by presence of different types of grazing management, presence of woodland around patches of calcareous grassland and shape complexity. These results indicate that calcareous grassland in favourable condition is management-dependent but also located in less intensively managed landscapes. Unfavourable habitat condition was predicted by threat factors such as lack of management and high incidence of arable or improved grassland around patches of calcareous grassland, indicating nutrient enrichment and habitat degradation. Some of these factors also predicted high diversity of the different ecological species groups. The value of this method for predicting habitat condition and species diversity from baseline ecological data for conservation monitoring at the landscape level is emphasised.