Who flies first? – habitat‐specific phenological shifts of butterflies and orthopterans in the light of climate change: a case study from the south‐east Mediterranean

1. Insects undergo phenological change at different rates, showing no consistent trend between habitats, time periods, species or groups. Understanding how and why this variability occurs is crucial. 2. Phenological patterns of butterflies and Orthoptera were analysed using a novel approach of standardised major axis (SMA) analysis. It was investigated whether: (i) phenology (the mean date and duration of flight) of butterflies and Orthoptera changed from one survey (1998 and 1999 respectively) to another (2011), (ii) the rate at which phenology changed differed between taxa and (iii) phenological change was significantly different across habitat types (agriculture fields, grasslands, and forests). Using the 2011 dataset, we investigated relationships between habitat‐specific variables and species phenology. 3. For both groups, late‐emerging species had an advanced onset on the second survey while the duration showed no consistent trend for butterflies and did not change for Orthoptera. Although the rate at which phenology changed was consistent between the two groups, at the habitat level, a longer duration of flight period emerged for butterflies in agriculture fields while Orthoptera showed no differentiation in flight duration between habitats. We found an earlier emergence of butterflies in grasslands compared to forests, attributed to habitat‐specific temperature, whereas spatial variation in humidity had a significantly lower effect on butterflies' phenology in grasslands compared to forests. A gradual delay of butterfly appearances as the canopy cover increased was also found. 4. The utility of SMA analysis was demonstrated in phenological studies and evidence was detected that both habitat type and habitat‐specific variables refine species' phenological responses.     

Landscape heterogeneity at multiple spatial scales enhances spider species richness in an agricultural landscape

Landscape supplementation, which enhances densities of organisms by combination of different landscape elements, is likely common in heterogeneous landscapes, but its prevalence and effects on species richness have been little explored. Using grassland-dwelling spiders in an agricultural landscape, we postulated that richness and abundances of major constituent species are both highest in intermediate mixtures of forests and paddy fields, and that this effect derives from multi-scale landscape heterogeneity. We collected spiders in 35 grasslands in an agricultural landscape in Japan and determined how species richness and abundances of major species related to local and landscape factors across different spatial scales. We used a generalized linear model to fit data, created all possible combinations of variables at 15 spatial scales, and then explored the best models using Akaike's information criterion. Species richness showed a hump-shaped pattern in relation to surrounding forest cover, and the spatial scale determining this relationship was a 300–500-m radius around the study sites. Local variables were of minor importance for species richness. Abundances of major species also exhibited a hump-shaped pattern when plotted against forest cover. Thus, a combination of paddy fields and forests is important for enhancement of grassland spider species richness and abundance, suggesting habitat supplementation. The effective spatial scales determining abundances varied, ranging from 200 to >1000 m, probably representing different dispersal abilities. Landscape compositional heterogeneity at multiple spatial scales may be thus crucial for the maintenance of species diversity.     

Effects of land use and landscape patterns on Orthoptera communities in the Western Siberian forest steppe

Across Western Siberia, land use has changed substantially since the collapse of the Soviet Union in 1991: large cropland areas were abandoned and livestock numbers declined. In recent years these trends have partly been reversed, and an intensification of agricultural management has been observed that is still ongoing. We evaluated the impact of land use, as well as effects of landscape patterns and vegetation structure on Orthoptera communities and discuss them as drivers of community composition, species richness and abundance. We sampled Orthoptera using a box-quadrat on ancient grassland, ex-arable grassland (both including different management types: unmanaged, grazed and mown) and cereal fields. Landscape heterogeneity and composition strongly affected species richness and abundance of Orthoptera. Both were higher in grassland than in cropland, but did not differ significantly between ex-arable and ancient grasslands or different management practices. An Indicator Species Analysis revealed differentiation of Orthoptera communities between all management types. On croplands, the number of adult individuals and nymphs was influenced by the proportion of grassland in the surrounding landscape and tillage practices. Conservation tillage is most likely the key factor allowing Orthoptera to reproduce on croplands. After up to 24 years of succession, Orthoptera communities of ex-arable grasslands can be considered as completely recovered, as differences to ancient grasslands were minimal. Besides the continuation of low-intensity management, conservation strategies for this region should consider landscape composition and support habitat heterogeneity like ecotones with hemi-boreal forests in grassland-dominated landscapes.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Reprint of: Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Earthworms,spiders and bees as indicators of habitat quality and management in a low-input farming region—A whole farm approach

The benefits of low input farming on biodiversity and ecosystem services are already well-established, however most of these studies focus only on the focal field scales. We aimed to study whether these benefits exist at the whole farm scale, to find the main environmental driving effects on biodiversity at the whole farm scale in farms of different grassland grazing intensity, applying three well-known species diversity indicator groups of different ecological traits.Edaphic (earthworms), epigeic (spiders) and flying (bees) taxa were sampled in each identified habitat type within 18 low-input farms in Central Hungary, 2010. The number of habitat types, the number of grassland plots, the cumulative area of grasslands and habitat type had an effect on the species richness and abundance of spiders, while grassland grazing intensity influenced the species richness of bees. Both bees and spiders were sensitive to vegetation and weather conditions, resulting in more bees on flower-rich farms and those having higher temperature; and more spiders on farms with more heterogeneous vegetation structure and in low-wind areas. Relatively few earthworms were found in the whole study, and their abundance was not influenced by any of the farm composition and management variables.We conclude that local field management (grazing intensity of grassland patches) can have a farm scale effect, detectable on species diversity indicators that have high dispersal ability and strong connection to grasslands as important foraging sites (bees). However, other farmland biota (spiders) is also strongly determined by farmland composition and habitat diversity, therefore the maintenance of a mosaic within-farm habitat structure is strongly recommended. The application of earthworms as farmland composition or management indicators is strongly restricted because of their special needs of soil conditions.     

A test of concordance in community structure between leafhoppers and grasslands in the central Tien Shan Mountains

Leafhoppers and related Auchenorrhynchous Hemiptera (AH) are among the most diverse grassland herbivores, and many have been linked inexorably to grassland vegetation through diet and shelter for millions of years. Are AH–plant communities in natural grasslands tightly integrated, how does the interaction differ across major ecological gradients, and do habitat or environmental factors explain the most variance in AH community structure? These questions have implications for the conservation of biodiversity and in evaluating effects of a warming climate. Using grasslands of the central Tien Shan Mountains as a natural laboratory, we examine whether AH species assemblages are concordant with vegetation in terms of community structure using closely associated species-level samples. Data were recorded from a nearly 3000-m elevation gradient crossing four arid and three montane grassland vegetation classes. We found elements of AH–plant community classification and structure to be closely correlated except for at the arid–montane habitat transition where a small group of widespread AH species were significant indicators for vegetation classes in both major grassland types. AH species richness and abundance are positively correlated with plant species density and percent cover and, correspondingly, peak at mid-elevations in association with montane grasslands. While overall elevation (and covariate mean annual temperature) explains the most variance in AH species assemblages, the sum total of habitat factors explain more variance than environmental factors when arid and montane grasslands were examined separately, but environmental factors are co-equal with habitat factors when the grassland types are combined. Unexplained variance in the AH community assemblages, attributable to individualistic species responses to environmental and habitat factors, slightly exceeds the total accounted for by the model.     

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.     

Cape ground squirrels as ecosystem engineers: modifying habitat for plants,small mammals and beetles in Namib Desert grasslands

Burrowing and foraging of semi‐fossorial rodents can affect species distribution and composition. Ground squirrels dig large burrow systems for refuge from predators and temperature extremes. Burrowing and foraging around burrows by squirrels may affect habitat and resource distributions for other organisms. We examined the impact of Cape ground squirrels (Xerus inauris) on vegetation, small mammals and beetles during winter and summer in grasslands on the edge of the Namib Desert. At each burrow system and paired control site without burrows, we estimated plant cover and height using quadrats (N = 8 paired sites), small mammal abundance and species richness using mark‐recapture techniques (N = 8 paired sites) and beetle abundance and species richness using pitfall traps (N = 6 paired sites, winter only). Squirrel burrowing and foraging activities resulted in lower plant cover and height, higher small mammal abundance and lower beetle abundance and species richness. Squirrels also reduced more plant cover in winter compared to summer, but had no effect on small mammal species richness. Furthermore, plant cover and height were higher in summer, whereas small mammal abundance and species richness were higher in winter. Our results suggest that Cape ground squirrels are important ecosystem engineers that influence plant and animal communities in the Namib Desert grasslands.     

No general edge effects for invertebrates at Afromontane forest/grassland ecotones

The Afromontane region of South Africa is characterised by numerous small, remnant forests in a grassland matrix. The edges, or ecotones between forests and grasslands are usually sharp (typically just over a few metres) and are mainly maintained by both natural and, more recently, anthropogenic fires. We investigated epigaeic amphipod, carabid and ant distribution patterns across Afromontane forest/grassland ecotones and found little evidence to support the biological edge effect. Five of the fifty-two sampled species however, did increase significantly in abundance at the ecotone. Among these was a very distinct edge species, the amphipod Talistroides africana. Overall, carabids were more abundant and species rich in forests while for ants it was in the grasslands. Ants and carabids were both more abundant and species rich in spring and summer than in autumn and winter. More interestingly, the abundance and species richness patterns across the ecotone did not change with the passing of the seasons. We argue that a conservation strategy for the Afromontane forest patches must also incorporate the surrounding grassland. The grassland habitat is often perceived as less valuable than forest and, as a consequence, is subject to many anthropogenic disturbances such as fragmentation, cattle grazing and afforestation. Protecting grasslands around forest patches not only conserves the rich ant diversity, but also conserves the biota in the forests and at the edges, and would therefore be more meaningful in terms of the overall conservation of Afromontane biodiversity.     

Spider communities in urban green patches and their relation to local and landscape traits

Urbanization and urban landscape characteristics greatly alter plant and animal species richness and abundances in negative and positive directions. Spiders are top predators, often considered to be sensitive to habitat alteration. Studies in urban environments frequently focus on ground-dwelling spiders or on spiders in built structures, leaving aside foliage spiders. Effects of habitat, landscape type and structure and local characteristics on spider species composition, richness and relative abundance were evaluated in urban green patches in a temperate city of South America. We also assess whether Salticidae could be an indicator group for the broader spider community in the urban environment. Spiders were sampled with a G-VAC (aspirator) in urban green patches in Córdoba city, Argentina, in urban, suburban and exurban habitats (18 sites; six per habitat) and local and landscape traits were assessed. Overall, the exurban was richer than the urban habitat, however, at the site level Salticidae richness and abundance (but not the total spider assemblage) were significantly lower in urban sites. Species composition moderately differed between urban and exurban sites. Results indicate that on urban green spaces a low impervious surface cover, a coverage of trees, herbaceous vegetation and a vertical structure of vegetation at least up to 1 m in height contribute to higher richness and abundance of spiders, Salticidae being more sensitive than the overall spider community to local effects. In addition, Salticidae richness can predict 74% of the total spider richness recorded and may be used as spider diversity bio-indicators in this climatic region.     

What determines Orthoptera species distribution and richness in temperate semi-natural dry grassland remnants?

Wide-spread fragmentation and isolation of habitats with high nature conservation value lends increasing importance to a better understanding of the factors which determine species richness in isolated habitat patches. Using data of one of the most abundant invertebrate groups in grasslands, Orthoptera, we analysed how species richness and distribution in 60 isolated semi-natural grassland remnants in Austria were affected by five environmental variables (altitude, habitat and land use diversity within each patch, habitat diversity of areas adjacent to each patch, patch size), and related to diversity of their main food source, i.e. vascular plants. We found a significant positive correlation between Orthoptera and vascular plant species richness, with threatened Orthoptera species having the lowest correlation coefficients. Life form diversity of plants was only moderately positively correlated with Orthoptera species richness. Habitat diversity within and adjacent to the grassland patch had by far the highest loadings on the first two axes of the principal component analysis, which jointly explained 99?% of the variance, and proved to be significant for total, threatened and not threatened Orthoptera, as well as for the two Orthoptera orders occurring in Central Europe (Caelifera, Ensifera). Additionally, the distribution of the majority of those 14 Orthoptera species analysed individually was mainly correlated with habitat diversity within and adjacent to the grassland patch. However, the distribution of a significant proportion of species was associated with other factors: five species were closely related to on-site land use diversity and patch size, and the distribution of three Ensifera species was not significantly correlated to any of the explanatory variables. We conclude that a surrogate taxa approach, i.e. the use of well-known taxonomic groups (e.g. vascular plants), may indeed deliver good results for capturing total, but less so for threatened, Orthoptera species richness in semi-natural grassland remnants. Small scale habitat diversity may be crucial to allow for the co-existence of a large number of Orthoptera species and has to be taken equally into account as patch size in nature conservation.     

Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

Direct and indirect effects of ski run management on alpine Orthoptera

Alpine landscapes are heavily influenced by ski run management, which can have severe impacts on alpine biodiversity. To assess these impacts on alpine Orthoptera, we compared species richness and species abundance in 41 plot pairs on ski runs and adjacent off-slope control plots in three ski resorts in Austria and Germany. A mixed modelling approach was used to assess the impacts of ski run preparation, artificial snow-making and environmental variables such as altitude, cover of dwarf shrubs and the application of fertilizer. Ski run plots showed a significantly lower species richness and number of individuals than control plots. Moreover, artificial snow led to a further decrease in species number. Hierarchical variance partitioning revealed that Orthoptera community composition is best predicted by environmental variables indirectly related to ski run management (fertilization, cover of dwarf shrubs) and to altitude. Only one out of five species significantly decreased in abundance after artificial snow-making. Other species were more sensitive to fertilizing and altitude. Dwarf shrubs were negatively associated with ski run management but positively associated with abundance of three species and species richness. Our data provide evidence for both direct and indirect consequences of ski runs and artificial snow-making on alpine Orthoptera. Overall, Orthoptera communities are suitable indicators for human-induced changes in alpine environments. In particular, a shift towards generalist species such as Chorthippus parallelus along with a decrease in typical alpine species gives cause for concern as this implies a homogenisation of biodiversity owing to ski run management.     

Measuring the Short‐term Success of Grassland Restoration: The Use of Habitat Affinity Indices in Ecological Restoration

Agricultural intensification threatens grasslands worldwide and the restoration of grasslands from arable lands can at least partially counter this threat. We studied grassland restoration by following early successional changes of arthropod assemblages (spiders, Araneae; true bugs, Heteroptera; orthopterans, Orthoptera; and ground beetles, Carabidae) on 1‐ and 2‐year‐old restorations using arable lands and native grasslands as two ends of the succession timescale. To examine the changes in species composition among the habitat types, we used habitat affinity indices based on fidelity and/or specificity of the species. We found that the number of species did not differ between habitat types, while species composition changed markedly with time. Species richness was thus not adequate to detect favorable changes after grassland restoration. Habitat affinity indices, on the other hand, were useful to detect compositional changes caused by the increasing numbers of species characteristic of target grasslands as early as the second year after restoration. Habitat affinity indices are easy‐to‐use, easy‐to‐interpret measures of restoration success; therefore, we recommend their use as measures complementary to species richness and simple similarity. Our results show that sowing low‐diversity seed mixture followed by mowing and grazing can be particularly successful in grassland restoration in time periods as short as 2 years.     

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.     

The effects of grassland management and aspect on Orthoptera diversity and abundance: site conditions are as important as management

Calcareous grasslands represent local hotspots of biodiversity in large parts of Central and Northern Europe. They support a great number of rare species which are adapted to these xerothermic habitats. Due to massive changes in land use, calcareous grasslands have become a rare habitat type and their conservation has been given a high priority in the habitats directive of the European Union. It is well known that grassland management may affect biodiversity substantially. However, the quality of calcareous grasslands is also influenced by abiotic conditions, such as aspect (i.e. sun exposure), which affects the local mesoclimate. We studied the combined effects of aspect and grassland management on Orthoptera diversity on 16 sites in Central Germany, in an unbalanced crossed design with three factors: aspect, management type and management intensity. For both response variables (diversity and abundance) we obtained a similar pattern. South-facing pastures maintained a greater diversity than north-facing pastures, but both had a greater diversity than extensively used meadows. Intensively used meadows maintained the lowest diversity and abundances. A multivariate analysis revealed that the abundance of rare Orthoptera species correlated with bare ground cover and forb cover, both of which were greatest at south-facing pastures. Our results suggest that grazing is a more suitable management for maintaining a high biodiversity in calcareous grasslands than mowing. Moreover, the mesoclimate (in this studied measured by its surrogate: aspect) is a crucial factor determining species richness and needs to be considered in reserve planning.