Activity and satiation state in Pterostichus melanarius: an experiment in different agricultural habitats

1. A capture–recapture experiment was conducted to examine the effects of satiation state on the activity and foraging efficiency of the carabid beetle Pterostichus melanarius Ill., in four different habitats (a hedge, a transition zone between the hedge and the adjacent wheat crop, the margin of the wheat crop, and the centre of the wheat crop) in an agricultural ecosystem in France. 2. Prey biomass and density decreased from the centre of the hedge to the crop margin and to the centre of the crop, where prey density was low but prey biomass was moderate. Starved and satiated beetles were released in circular enclosures and recaptured using pitfall traps for 6 days. 3. Immediately after release, starved P. melanarius were significantly more active than satiated beetles, but this difference disappeared after the first day. At each recapture date, starved individuals had gained weight whereas satiated individuals had lost weight. The analysis of activity and weight variation at the end of the first day indicated that the centre of the crop seemed to be the most favourable habitat for starved individuals to find food rapidly, whereas the behaviour of satiated individuals was identical in all habitats. 4. This experiment shows that habitat use of P. melanarius is flexible. The results are discussed in the context of the classical view of foraging behaviour in carabid beetles. The activity and foraging efficiency in P. melanarius in different habitats can be explained as the result of the interaction between individual satiation state, prey availability in the habitat, and particular preferences of the species.     

Rapid Turnover and Edge Effects in Dung Beetle Assemblages (Scarabaeidae) at a Bolivian Neotropical Forest-Savanna Ecotone1

Habitat fragmentation and the widespread creation of habitat edges have recently stimulated interest in assessing the effects of ecotones on biodiversity. Ecotones, natural or anthropogenic, can greatly affect faunal movement, population dynamics, species interactions, and community structure. Few data exist, however, on insect community response to forest–savanna ecotones, a natural analog to anthropogenically cleared areas adjacent to forest. In this study, the abundance, total biomass, average individual biomass, and distribution of scarabaeine dung beetles were examined at a sharp tropical evergreen forest–savanna ecotone in Santa Cruz, Bolivia. The abundance, total biomass, and average individual biomass of dung beetles varied significantly across the forest, edge, and savanna habitats. Species richness (S_obs) also varied significantly across the three habitats, but statistical estimations of true species richness (S_est) did not. Habitat specificity of the dung beetles in this study was extremely high. Of the 50 most common species collected during the study, only 2 species were collected in both the forest and savanna habitats, signaling nearly complete community turnover in just a few meters. Strong edge effects were evidenced by the decline in abundance, total biomass, and species richness at the forest‐savanna boundary.     

Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats

Agricultural landscapes can be defined as mosaics of landscape elements which are affected by farming practices. Woodland habitats, even though they are managed, are amongst the most stable elements of agricultural landscapes and can play a key role in the maintenance of biodiversity. This study of the ground beetle (carabid) communities of woodlands and woody linear features in a Scottish agricultural landscape shows that these habitats contribute significantly to the overall landscape diversity of these beetles. Communities in woods and hedgerows display the same species diversity and are both characterized by the presence of forest species. The main factors constraining carabid communities in both environments are the grazing intensity and, to a lesser extent, the type of soil. Heavily grazed locations are characterized by the occurrence of grassland species while forest species are restricted to ungrazed locations. At the landscape scale, the distribution of the forest species is limited by spatial isolation, indicating that there are insufficient functional links between woodland habitats in the study area. Isolation could be compensated for either by a better control of grazing so that linear features can be used as dispersal corridors for forest carabids or by planting more linear features and woods in the area.     

Impact of rubber plantation on carabid beetle communities and species distribution in a changing tropical landscape (southern Yunnan, China)

Carabid beetles (Coleoptera: Carabidae) have widely been used to assess biodiversity values of different habitats in cultivated landscapes, but rarely in the humid tropics. This study aimed to investigate effects of land use change on the carabid assemblages in a tributary valley of the Mekong River in tropical southern Yunnan, China. The study area includes habitats of traditional land use systems (rice production and shifting cultivation successions) and was dominated by natural forests until about 30?years ago. Since then, large areas of forest have been, and still are, successively transformed into commercial rubber monoculture plantations. In total, 102 species of Carabidae (including Cicindelinae) were recorded from 13 sites over different seasons, using pitfall traps, Malaise traps and aerial collectors in trees. Cluster analysis and indicator species analysis showed that three types of habitat (rice field fallows, early natural successions and natural forest) possess a degree of uniqueness in species composition. Non-metric multidimensional scaling revealed that the environmental factors explaining 80% of the total variation in carabid assemblage composition are the degree of vegetational openness of a habitat and its plant species diversity. Rice field fallows had significantly higher numbers of species and individuals than any other type of habitat and are probably dominated by species originating from other regions. Carabid assemblages of young rubber plantations (5 and 8?years) were quantitatively similar to those of forests, but without species of significant indicator value. With increasing plantation age (20 and 40?years), the number of carabid species decreased. Increasing age and a further spatial expansion of rubber plantations at the expense of forest areas will have negative impacts on the native forest carabid assemblages with strongest effects on forest specialists and rare species.     

Effects of narrow roads on the movement of carabid beetles (Coleoptera,Carabidae) in Nopporo Forest Park,Hokkaido

Forest-dwelling carabid beetles that have no flight ability were studied using mark-recapture methods in late-June to mid-October 2007. This study was done to determine the effects of narrow roads in Nopporo Forest Park, Hokkaido on carabid beetle movement and habitat use. The investigation was conducted at four sites: one site was an abandoned grassy road with a width of 3.5 m, two sites were gravel roads with widths of 3.5 and 4.5 m, and another site was an asphalt-paved road with a width of 4.5 m. A total of 3,580 individuals from six species of carabid beetles were collected using dry pit-fall traps, and recapture rates ranged from 6.1 to 36.2%. All examined roads acted as barriers against the movement of Leptocarabus arboreus ishikarinus. All roads, except the abandoned grassy road, acted as a barrier against Carabus granulatus yezoensis movement. Forest–roadside verge comparisons demonstrated that some carabid beetles avoid even narrow roadside verges. Harmful effects increase with increasing road width and both paved roads and narrow roads negatively affect the movement of carabid beetles inhabiting the bordering forest. Therefore, forest specialist beetles are influenced by a barrier effect that starts at the forest road verge, and this barrier effect may be exacerbated by vehicular traffic. Therefore, these barrier effects on carabid beetles should be considered when planning and implementing road construction and maintenance in forests.     

Distribution of carabid beetles (Coleoptera: Carabidae) across a forest-grassland ecotone in southwestern China

This paper studied the occurrence of carabid beetles (Coleoptera: Carabidae) in the forest edge, the adjacent forest interior, and the surrounding grassland in southwestern China. Beetles were collected with pitfall traps along five replicated transects. Forest species rarely penetrated into the grassland from the forest interior, and the grassland specialists were not found in the forest interior. The forest edge hosted additional species from the adjacent grassland that increased its overall species richness. Nearly all forest species (23 of 24 species) and grassland species (13 of 15 species) can be found in the forest edge. Carabids of the forest edge were more similar to those of the forest interior than to those of the grassland by ordination and cluster analysis. Based on the specificity and fidelity, carabids can be distinguished into five species groups: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analysis showed that canopy cover and/or shrub cover were the most important factors in determining the richness, abundance, and diversity of carabids. The forest edge may serve as a transition zone for dispersal and re-colonization of carabid beetles from adjacent habitats and therefore is important for natural conservation.     

The role of various meadow margin types in shaping carabid and staphylinid beetle assemblages (Coleoptera: Carabidae,Staphylinidae) in meadow dominated landscapes

Less intensively managed semi-natural habitats, e.g., field and meadow margins like hedgerows, are thought to be crucial landscape components for maintaining biodiversity in highly disturbed and intensively managed agricultural landscapes. In this study, we focused on the effects of three meadow margin types on activity-density, species richness and species composition of carabid and staphylinid beetles recorded by pitfall traps in Central European landscapes dominated by intensively managed meadows. Carabid activity-density was significantly higher in meadows than in meadow margins and within meadow margins their activity-density increased from grassy meadow margins via shrubby ones to woody meadow margins. We found that recorded species richness of both carabid and staphylinid beetles was not significantly affected by habitat identity (meadow margin or neighbouring meadow) and meadow margin type. Recorded species composition of both investigated taxa was significantly affected by habitat identity and interaction between habitat identity and meadow margin type (i.e. it differed between particular meadow margin types). Assemblages inhabiting various meadow margin types were more dissimilar between each other than assemblages from neighbouring meadows. Meadow margins within grassland dominated landscapes maintain local species richness by hosting different species from those living in surrounding meadows. Dissimilarity of carabid and staphylinid assemblages from meadows neighbouring both sides of particular meadow margin did not differ between meadow margin types. Our results indicate that semi-natural habitats play an important role in maintaining biodiversity not only in agricultural landscapes dominated by arable fields, but also in those dominated by meadows.     

The impact of hedge-forest connectivity and microhabitat conditions on spider and carabid beetle assemblages in agricultural landscapes

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.     

Landscape level processes driving carabid crop assemblage in dynamic farmlands

Landscape heterogeneity has been shown to be a major factor in the maintenance of biodiversity and associated services in agricultural landscapes. Farmlands are mosaics of fields with various crop types and farming practices. Crop phenology creates asynchrony between fields sown and harvested in different periods (winter vs. spring crops). The present study was conducted to examine the influence of such spatio-temporal heterogeneity on biodiversity, with the hypothesis that it would lead to spatio-temporal redistribution (shifting) of species. Species richness and activity-density of carabid beetles in winter cereal (winter) and maize (spring) crops were compared across 20 landscapes distributed along a double gradient of relative area and spatial configuration of winter and spring crops. Maize fields were sampled in spring and late summer for comparison over time. The response of carabid species richness to landscape heterogeneity was weak in spring, but maize field richness benefited from adjacencies with woody habitat, in late summer. In spring, increased length of interfaces between winter and spring crops lowered carabid activity-density in winter cereal fields, suggesting that maize fields acted as sinks. Interfaces between woody habitats and crops increased activity-density in both crop types. We found no evidence of spatio-temporal complementation, but different species benefited from winter cereals and maize in spring and late summer, increasing overall diversity. These findings confirm the role of adjacencies between woody and cultivated habitats in the conservation of abundant carabid assemblage in winter cereals and maize. We conclude that between-field population movement occurs, and advocate for better consideration of farmland heterogeneity in future research.     

Relevance of exotic pine plantations as a surrogate habitat for ground beetles (Carabidae) where native forest is rare

Plantation forests are of increasing importance worldwide for wood and fibre production, and in some areas they are the only forest cover. Here we investigate the potential role of exotic plantations in supporting native forest-dwelling carabid beetles in regions that have experienced extensive deforestation. On the Canterbury Plains of New Zealand, more than 99% of the previous native forest cover has been lost, and today exotic pine (Pinus radiata) plantations are the only forest habitat of substantial area. Carabids were caught with pitfall traps in native kanuka (Kunzea ericoides) forest remnants and in a neighbouring pine plantation, grassland and gorse (Ulex europaeus) shrubland. A total of 2,700 individuals were caught, with significantly greater abundance in traps in young pine, grassland and gorse habitats than in kanuka and older pine. Rarefied species richness was greatest in kanuka, a habitat that supported two forest specialist species not present in other habitat types. A critically endangered species was found only in the exotic plantation forest, which also acts as a surrogate habitat for most carabids associated with kanuka forest. The few remaining native forest patches are of critical importance to conservation on the Canterbury Plains, but in the absence of larger native forest areas plantation forests are more valuable for carabid conservation than the exotic grassland that dominates the region.     

New approach for evaluating habitat stability using scarce records for both historical and contemporary specimens: a case study using Carabidae specimen records

Natural history collections, such as specimen records, are crucial resources for conservation and habitat management. However, these data are usually scarce compared to physical environmental data (e.g., digital terrain maps) that we often have little species data and a lot of physical environmental data with which to evaluate habitats. In this paper, we propose a method for evaluating habitat stability using scarce natural history records and abundant physical environmental data. We used both historical and contemporary specimen records of carabid beetles (areas in which records of the same species were recorded during both periods) and evaluated the attributes of these areas using terrain characteristics. We found two common terrain characteristics among the occupied areas: large total river length and low variation in elevation. These terrain characteristics suggest that habitats of carabid species have been conserved in disturbed and wet environments for a long time. These results are consistent with the ecological characteristics of carabid beetles. Our study shows that scarce natural history collections, combined with ingenuity, can be useful for evaluating habitats.     

Carabids of differently aged reforested pinewoods and a natural pine forest in a historically modified landscape

We studied the responses of carabid beetles to Pinus sylvestris ageing in four plantations aged 2–3, 10–12, 40–45 and 80 years, and one mature natural pine forest – in NW Spain. In 2003, a total of 16,866 carabid individuals (50 species) were collected using pitfall traps. Overall carabid abundance and species richness differed significantly among the five age classes with the highest values occurring in the youngest age class. The carabid assemblages of the initial stages of the ageing sequence differed considerably from the later stages that were relatively similar to each other – a consequence of differences in species dominance at specific age classes. Yet, the carabid assemblage of the 80-year-old stage did not approach that of the natural pine forest, but, unexpectedly, the 40–45-year-old age class did. Canopy cover and soil pH were the main environmental variables that affected the carabid distribution. We also detected species-level responses to the ageing process, with open habitat species more abundantly collected at the youngest stages and forest specialists at the older stages. We conclude that carabid beetles responded predictably to the pine plantation ageing process, and that although pine forests in NW Spain do not seem to provide habitat for unique species, they act as secondary habitats for forest specialist species.     

Dispersal of a carabid beetle in farmland is driven by habitat‐specific motility and preference at habitat interfaces

Carabid beetles are common predators of pest insects and weed seeds in agricultural systems. Understanding their dispersal across farmland is important for designing farms and landscapes that support pest and weed biological control. Little is known, however, about the effect of farmland habitat discontinuities on dispersal behaviour and the resulting redistribution of these beetles. We released 1,985 well‐fed and 1,680 food‐deprived individuals of the predatory carabid beetle Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on a farm in Wageningen, The Netherlands. We recaptured 23.6% of those beetles over a period of 23 days in 2010. The farmland comprised agricultural fields with various crop species and tillage, separated by strips of perennial vegetation. We developed discrete Fokker‐Planck diffusion models to describe dispersal based on motility (m² day^?1) and preferential behaviour at habitat interfaces. We used model selection and Akaike’s information criterion to determine whether movement patterns were driven by variation in motility between habitats, preferential behaviour at habitat interfaces, or both. Model selection revealed differences in motility among habitats and gave strong support for preferential behaviour at habitat interfaces. Behaviour at interfaces between crop and perennial vegetation was asymmetric, with beetles preferentially moving towards the crop. Furthermore, beetles had lower motility in perennial strips than in arable fields. Also between arable habitats movement was asymmetric, with beetles preferentially moving towards the habitat in which motility was lowest. Neither crop type nor tillage explained differences in motility between crop habitats. Recapture data representing dispersal patterns of beetles were best described by a model that accounted for differences in motility between farmland habitats and preferential behaviour at habitat interfaces. Motility in farmland and behaviour at interfaces can also be estimated for other organisms and farmland habitats to support design of farmland conducive to natural pest suppression. Landscape design for early recruitment of carabids into arable fields should take into account the quantity and quality of resource habitats in the landscape, their proximity to crop fields, movement rates, and the possibility of movement responses at interfaces between landscape elements.     

Effects of urbanization on ground beetle assemblages (Coleoptera, Carabidae) of grassland habitats in western Canada

We pitfall-trapped carabid beetles across an urban-rural gradient in central Alberta, Canada to assess the relative contributions of landscape structure and habitat quality in explaining variation in composition of assemblages. The gradient ranged from the "urban" zone in the city of Edmonton, through a "suburban" zone just outside the city, into a "rural" zone 25–60 km east of the city. We trapped beetles at three replicated sites for each of two habitats (unmanaged grasslands and highly managed graveyards) in each of the three zones. Most of the 24 species (3162 individuals) captured were commonly associated with open habitats. Pterostichus melanarius , an introduced species, made up 80% of the total catch. Carabid assemblages of grasslands and graveyards were distinct, with lower species richness and lower catch rates of native, introduced and brachypterous carabids in graveyards. Graveyard assemblages also had proportionally fewer brachypterous carabid individuals (32 vs 52%) and species (20 vs 36%), suggesting that populations recruit locally from elsewhere after disturbances. Native carabids were least abundant in the urban areas. Brachypterous carabids were similarly abundant across the gradient, but more occurred in the urban zone than elsewhere (32 vs 5–10%). Samples from graveyards were characterized by lower beta diversity (variation among samples) than those from grasslands, but beta diversity was more-or-less equally variable across the gradient within habitats. The amount of unmanaged grassland habitat and the degree of urbanization significantly influence carabid assemblages, and therefore are relevant to management and conservation of urban grassland areas.     

Invasion and retreat: shifting assemblages of dung beetles amidst changing agricultural landscapes in central Peru

This study examines the effects of deforestation, and the habitat value of coffee and regenerated forest for tropical dung beetles, a functionally significant insect group. Pitfall trapping was conducted at 22 sites in a montane region of central Peru during April and November/December of 2002. Sites included primary and secondary forest, shade-coffee, regenerated forest and open farms (mainly with banana, yuca, and corn). Ordination techniques indicated that beetle assemblages in forests, regenerated forest and coffee were relatively similar. However, assemblage compositions in forested areas differed even at similar altitudes under the influence of biogeographical factors, and the assemblages at disturbed sites (farms/coffee) were influenced by beetle dispersal from adjacent forests. During dry months, when beetle activity is low, communities at all habitat types tended to converge because fewer unique species were recorded in forests at that time and habitat/season generalists were dominant. Preliminary results also indicate that beetles in shady crops such as bananas responded to plant growth: as the banana canopy closed-in, producing more shade, open-habitat specialists retreated and forest/shade specialists invaded the sites. Chronosequence data at two of the sites demonstrate the rapid and dramatic changes in species richness and assemblage composition caused by deforestation. As forests become increasingly fragmented, and open farms continue to expand, dung beetles will become more restricted to the remaining fragments and reserves. In the mosaic landscape studied here, shade crops, like coffee, act as habitat and corridors for many dung beetle species. Small farm size and the consequent magnitude of edge effects, likely contributed to beetle movement between habitat types and determined the apparent generalist nature of many of the dung beetle species in this study.     

Movement patterns of selected insect groups between natural forest,open land and rubber plantation in a tropical landscape (southern Yunnan,SW China)

In many regions of tropical Asia, the expansion of rubber monoculture plantations is conducted by replacement of natural forest areas and strongly affects biodiversity and movement patterns of wild species, including insects. Against this background, we conducted a study on selected insect groups (longhorn beetles, bark beetles, wild bees and hoverflies) along transects between rainforest patches, open uncultivated land and rubber plantation habitats in a region of Xishuangbanna (southern Yunnan, China), with the objectives to identify (a) movement directions and patterns of selected insect groups based on their abundances in modified Malaise traps in the different habitats, and (b) the role of remaining natural rainforest patches and rubber plantations, respectively, for insect diversity maintenance and conservation. The highest total numbers of species and individuals of bark beetles, longhorn beetles and wild bees were recorded from the natural forest edge compared to open land and rubber plantation edge. This result clearly indicates that the natural forest plays an important role in maintenance of these three insect groups. However, the highest number of hoverfly species and individuals was recorded from the open land sites, indicating the most relevant habitat type for this group of species. Overall, the lowest species and individual numbers were recorded from the rubber plantation edge, indicating the unsuitability of this habitat type for all insect groups considered. The distribution of species and individuals in the opposite trap sides along the transect indicates that longhorn beetles, bark beetles and wild bees show not only movements from the forest to the surrounding habitats, but also return back after encountering the unsuitable rubber plantation habitat. Bark beetle composition showed the relatively highest similarity between all trap sites and opposite trap sides among the insect groups considered, indicating a higher movement activity than the other groups. The four insect groups considered in this study show different movement modes between the forest, open land and rubber plantation, which are not the same for all taxa. Except for hoverflies, the natural forest was found to be the most important habitat for the maintenance of species diversity in the different land use types of the study area.     

Taxon specific response of carabids (Coleoptera,Carabidae) and other soil invertebrate taxa on invasive plant Amorpha fruticosa in wetlands

False indigo (Amorpha fruticosa L.) is an invasive exotic plant introduced to Europe in the early eighteenth century. Its spread has been rapid, particularly in disturbed wetland habitats, where it forms dense impermeable monospecific stands and modifies habitat conditions. The impact of A. fruticosa on native plant communities has been well analyzed, however knowledge concerning the possible effects on soil invertebrates and particularly carabid beetles is completely lacking. This study analyzed the impact of an A. fruticosa invasion on carabid beetles and other soil invertebrates. Soil fauna was sampled by pitfall traps at natural habitats, initially colonized by A. fruticosa, and habitats largely invaded by A. fruticosa. In total 2,613 carabid beetles belonging to 50 species and 72,166 soil invertebrates were collected. The invasion of A. fruticosa strongly affected the carabid beetle species composition, which clearly differed between all studied sites. Widespread euritopic carabid beetle species showed positive responses to A. fruticosa invasion, while the activity density of open habitat species strongly declined. Mean individual biomass was significantly higher at invaded sites due to increased incidence of large carabids (genus Carabus Linné, 1758). Carabid beetle activity density and abundance of soil invertebrates were considerably higher at invaded sites than in natural sites. Conversely, the impact of A. fruticosa on carabid beetle species richness and diversity was less pronounced, most likely due to immigration from adjacent habitats. Changes in carabid beetle species composition and abundance of soil invertebrates were most likely due to changes in vegetation structure and microclimate. The results suggest that A. fruticosa invasion considerably affected carabid beetles, an insect group that is only indirectly related to plant composition. Therefore, severe future changes can be expected in invertebrate groups that are closely related to plant composition, since A. fruticosa cannot be completely removed from the habitat and covers relatively large areas.     

Dung beetle assemblages in primary forest and disturbed habitats in a tropical dry forest landscape in western Mexico

Dung beetles (Scarabaeinae) are conspicuous components in most terrestrial ecosystems, performing important ecological functions and services. Being sensitive to several types of disturbance, they have been successfully used as indicators of habitat change. Dung beetle communities in tropical rainforests have been well studied, but considerably less information is available for tropical dry forests. In this study I sampled dung beetles in two undisturbed habitats, deciduous forest and semideciduous forest, and two disturbed habitats, secondary forest and open area habitat, in the Chamela-Cuixmala region of western Mexico. Dung beetle species with high indicator value for each habitat were identified. Beetle abundance, observed species richness and estimated species richness were similar in the three forest habitats, but significantly lower in the open area habitat. A more detailed analysis of species-specific abundances in the three forest habitats revealed some differences. Transects of one of the undisturbed habitats, the deciduous forest, were more similar to the non-adjacent transects of disturbed secondary forest, than to the adjacent undisturbed semideciduous forest transects. Unlike studies in other tropical sites that have found a decrease in equitability in Scarabaeinae assemblages between undisturbed forest and disturbed habitat (particularly open habitats), in the Chamela-Cuixmala region all four habitats showed similar low equitability in community structure, with two or three very dominant species.     

Asymmetric carabid beetle spillover between calcareous grasslands and coniferous forests

Central European calcareous grasslands are considered biodiversity hotspots, but are severely threatened by the change in land-use and by habitat fragmentation. Coniferous forests are typical adjacent habitats to calcareous grasslands, as abandoned calcareous grasslands are often afforested or develop into coniferous forests by succession. To investigate spillover between calcareous grasslands and coniferous forests, a total of 144 pitfall traps for carabid beetles were placed at three different distances (1, 5, 20 m) from the edge in both habitats at eight locations from April to late August. We found that both habitats had a distinct species assemblage and a decrease in spillover with increasing distance from the habitat edge into the adjacent habitat. Calcareous grasslands were more affected by spillover from the adjacent coniferous forests than vice versa because more forest specialists penetrated into calcareous grasslands than grassland specialists penetrated into coniferous forests. We conclude that spillover into small and isolated habitats can severely change species assemblages, which has to be considered in conservation measures. The protection of large sites with small edge-interior ratios can reduce negative effects on species assemblages in endangered calcareous grasslands.