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.     

Differentiation of spring carabid beetle assemblages between semi‐natural habitats and adjoining winter wheat

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Heterogeneity from increasing crop types: effect on carabid beetles

We investigated the effect of crop type heterogeneity on carabid beetles in intensively managed agricultural landscapes. We compared different crops and habitat type configurations and compositions. In addition we assessed the effect of artificial fragmentation by paved roads and water channels in an agricultural landscape. Crop type heterogeneity negatively affected carabid assemblages. Spatial dissection by artificial structures had an even more negative effect on carabid diversity. The nested structure of the carabid assemblages among crop types was caused by a negative response to heterogeneity in the agricultural landscape. Even though crop types in an intensive agricultural landscape increased, the landscape may tend to fragment habitat. Although artificially fragmented habitats had more unstable carabid assemblages, heterogeneity from different crop types has also been shown to be a characteristic of fragmented landscapes. To produce a positive relationship between biodiversity and heterogeneity in an agricultural landscape, farming intensity should be reduced to allow for the conversion of highly productive lands into more natural habitats.     

Staphylinidae and Carabidae overwintering in wheat and sown wildflower areas of different age

Species richness and abundance of staphylinid and carabid beetles overwintering in winter wheat fields and 1- to 3-year-old wildflower areas were investigated during 2000/2001 on 16 study sites in Switzerland. Abundance and species richness of overwintering staphylinids significantly increased with successional age of the wildflower areas and were always higher in older wildflower areas than in winter wheat. A similar but less distinct pattern was observed for the abundance and species richness of carabid beetles. The influence of habitat parameters (vegetation cover, fine sand content, organic matter, pH, soil pore volume, surrounding landscape structure, habitat area) on the staphylinid and carabid assemblages based on the number of individuals per species and site was analysed using canonical correspondence analysis. Vegetation cover was the most significant parameter significantly characterizing both staphylinid and carabid assemblages. The amount of vegetation cover explained 15.7% of the variance, fine sand content accounted for 13.3% and surrounding landscape structure for 10.9% of the variance in the staphylinid assemblage. In the carabid assemblage, vegetation cover was the only significant factor, explaining 24.7% of the variance. This study showed for the first time that the significance of wildflower areas as a reservoir for hibernation for generalist predatory beetles increases with progressing successional age.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

Effects of landscape connectivity on the spatial distribution of insect diversity in agricultural mosaic landscapes

European agricultural landscapes are mosaics of intensively cultivated areas and semi-natural elements. Although comprising only a small fraction of the total area, semi-natural elements provide habitat for most of the landscape biodiversity. Agricultural intensification has increasingly fragmented semi-natural elements and species numbers are in decline. Insights into the effects of landscape structure on species’ distributions within and among semi-natural habitats are needed to conserve biodiversity in agricultural landscapes more effectively. We investigated the landscape- and habitat-specific diversity partitions of wild bees, true bugs, and carabid beetles in two differently structured agricultural landscapes in Switzerland. In each landscape, we partitioned the total species diversity (γ) into its additive components within (_P) and among patches (β_P) and among habitats (β_H). In the landscape characterized by a patchy, isolated distribution of habitat elements, among-patch diversity (β_P) explained 44% of the total species richness (γ) and was significantly higher than expected under a random distribution of samples among habitat patches; in the landscape with higher habitat connectivity, among-patch diversity (β_P) comprised 32% of the total species richness (γ) and did not differ from the random expectation. Habitat-specific within-patch contributions to species richness were similarly low across habitat types (_P=23–24%) in the patchy landscape, whereas in the more connected landscape within-patch partitions tended to be higher and differed among habitat types (_P=22–38%). Functionally different groups of bees, true bugs, and carabids also responded differently to landscape structure in a manner that was consistent with known differences in resource specialization and dispersal ability. Differences in diversity partitions among landscapes and taxa indicate the need for flexible conservation strategies. Conservation of habitat-specific diversity may require more habitat patches in landscapes that have lower habitat connectivity and low within-patch diversity (_P) than in landscapes with higher within-patch diversity (_P).     

Dynamic effects of ground-layer plant communities on beetles in a fragmented farming landscape

Vegetation effects on arthropods are well recognized, but it is unclear how different vegetation attributes might influence arthropod assemblages across mixed-agricultural landscapes. Understanding how plant communities influence arthropods under different habitat and seasonal contexts can identify vegetation management options for arthropod biodiversity. We examined relationships between vegetation structure, plant species richness and plant species composition, and the diversity and composition of beetles in different habitats and time periods. We asked: (1) What is the relative importance of plant species richness, vegetation structure and plant composition in explaining beetle species richness, activity-density and composition? (2) How do plant-beetle relationships vary between different habitats over time? We sampled beetles using pitfall traps and surveyed vegetation in three habitats (woodland, farmland, their edges) during peak crop growth in spring and post-harvest in summer. Plant composition better predicted beetle composition than vegetation structure. Both plant richness and vegetation structure significantly and positively affected beetle activity-density. The influence of all vegetation attributes often varied in strength and direction between habitats and seasons for all trophic groups. The variable nature of plant-beetle relationships suggests that vegetation management could be targeted at specific habitats and time periods to maximize positive outcomes for beetle diversity. In particular, management that promotes plant richness at edges, and promotes herbaceous cover during summer, can support beetle diversity. Conserving ground cover in all habitats may improve activity-density of all beetle trophic groups. The impacts of existing weed control strategies in Australian crop margins on arthropod biodiversity require further study.     

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.     

Influence of abandonment on syrphid assemblages in mountainous meadows

Grasslands are often managed with different intensities in the European Alps. Studies have shown that a medium management can benefit plant and animal biodiversity in these ecosystems. However, in recent decades abandonment of extensively managed meadows is an ongoing global challenge. Syrphids (hoverflies) have been recognized as a threatened group due to environmental drivers, and these managed grasslands provide preferred habitats for them. Yet, at what extent syrphids respond to grassland management strategies is little known. We investigated whether abandonment of managed mountain meadows (mown once a year, no fertilizer use) affects syrphid abundance, richness and species assemblages. Moreover, the influence of plant richness, flower frequency and surrounding landscape on syrphids were assessed. Four managed meadows were compared with four abandoned meadows in a mountainous region in Austria. Adult syrphids were sampled over two consecutive years (June and August 2015–2016) by sweep netting using line transects and observation plots. Syrphid abundance was significantly higher in managed than abandoned meadows and increased with increasing plant richness and flower frequency across management types. Management and sampling time also significantly affected syrphid species assemblages. Syrphid richness and abundance were not influenced by surrounding landscape. We conclude that both abandoned and managed meadows provide important habitat types for syrphids. Therefore, efforts should be made to maintain both managed and abandoned meadows in order to protect syrphid species within mountainous landscapes.     

Effects of fen management and habitat parameters on staphylinid beetle (Coleoptera: Staphylinidae) assemblages in north-eastern Germany

Because fens have undergone dramatic declines in recent decades, an important question is which management regimes and habitat parameters are most effective in preserving fen biodiversity. The aim of the present study was to assess the effects of five different management regimes (intensive grassland, moist meadows, summer harvested sites, winter harvested sites, fallows) on staphylinid beetle assemblages. During the study period 5,989 individuals from 92 staphylinid beetle species were recorded. Species richness and abundance were highest on intensive grassland and fallows, and water level and vegetation height had significant impacts on the abundance of staphylinids. On winter-harvested sites species richness and abundance were lowest. In general, staphylinid beetles did not show pronounced variation among management regimes, while the environmental factors water level, vegetation height and top soil mineralisation seemed to have a larger structuring impact. The number of threatened species and the conservation index were highest on summer-harvested sites and fallows, representing fairly well-preserved fens. We conclude that summer harvest of reed beds or no management at all appears to be most beneficial for the conservation of staphylinid beetles.     

Ground beetle (Coleoptera: Carabidae) assemblages associated with a satoyama landscape in Japan: the effects of soil moisture,weed height,and distance from woodlands

Agricultural landscapes generally include not only crop fields but also semi-natural habitats. In Japan, such a mixed rural landscape is called “satoyama.” Although ground beetles are potential predators of pests, the environmental factors that determine their distribution in Japanese rural landscapes have not been fully elucidated. To understand the effects of distance from woodland edges, soil moisture, and weed height on assemblages of carabid beetles, we examined the number of adult beetles in pitfall traps placed in a satoyama landscape in the lowlands of western Honshu, Japan. Our results show that the carabid species could be largely differentiated into woodland, intermediate, and open-land species. The “intermediate species” group includes species that depend on woodland or woodland edges for at least part of their life cycles. Paddy fields must have long provided semi-natural habitats that complement those in natural grasslands and wetlands for open-land beetles that prefer wet conditions. Weeds can also increase the abundance of some intermediate and woodland species; thus, the arrangement of such landscape elements as woodlands and paddies can determine the species richness and abundance of ground beetles in agricultural fields.     

Landscape-scale crop diversity interacts with local management to determine ground beetle diversity

Crop diversification and maintenance of semi-natural habitats (grasslands and field boundaries) are suggested to enhance farmland biodiversity, but the relative importance of these factors remains poorly known. We evaluated how crop diversity and availability of semi-natural grasslands at a landscape-scale interacted with local farming management (three management types from low to high intensity: ley < winter wheat < sugar beet) in their effect on ground beetle assemblages in southern Sweden. Ground beetle diversity increased with crop diversity either independently of local management (Simpson species diversity), or only in the less intensively managed habitats (rarefied species richness). While ground beetle diversity in leys tended to increase with field boundary length, no such relationship was observed in winter wheat or sugar beet fields. In contrast, the landscape proportions of leys and semi-natural grasslands did not affect ground beetle species richness and diversity. We conclude that (a) semi-natural grasslands and leys may not function as source habitats at a landscape-scale if they comprise a low proportion of the total land-use, while (b) increasing crop diversity is correlated to ground beetle richness and diversity in agricultural landscapes dominated by arable land. The beneficial effect of landscape-scale crop diversification on farmland biodiversity may depend on the general level of agricultural intensity of a region.     

Management intensity at field and landscape levels affects the structure of generalist predator communities

Agricultural intensification is recognised as a major driver of biodiversity loss in human-modified landscapes. Several agro-environmental measures at different spatial scales have been suggested to mitigate the negative impact of intensification on biodiversity and ecosystem services. The effect of these measures on the functional structure of service-providing communities remains, however, largely unexplored. Using two distinct landscape designs, we examined how the management options of organic farming at the field scale and crop diversification at the landscape level affect the taxonomic and functional structure of generalist predator communities and how these effects vary along a landscape complexity gradient. Organic farming as well as landscapes with longer and more diversified crop rotations enhanced the activity-density of spiders and rove beetles, but not the species richness or evenness. Our results indicate that the two management options affected the functional composition of communities, as they primarily enhanced the activity-density of functionally similar species. The two management options increased the functional similarity between spider species in regards to hunting mode and habitat preference. Organic farming enhanced the functional similarity of rove beetles. Management options at field and landscape levels were generally more important predictors of community structure when compared to landscape complexity. Our study highlights the importance of considering the functional composition of generalist predators in order to understand how agro-environmental measures at various scales shape community assemblages and ecosystem functioning in agricultural landscapes.     

The value of semi-natural grasslands for the conservation of carabid beetles in long-term managed forested landscapes

Species rich semi-natural grasslands are disappearing across Europe, affecting invertebrate diversity negatively. In NW Spain, the recent abandonment of traditional farming practices and the gradual decrease in grazing pressures are reducing the number and extent of montane grasslands. In this context, we investigated the composition of carabid beetle (Coleoptera, Carabidae) assemblages that inhabit semi-natural grasslands situated in long-term managed oak and beech forested landscapes. According to their spatial arrangement, the studied grasslands were classified into: (1) interior or gap grasslands (small and completely surrounded by continuous forest) and (2) exterior grasslands (large and connected to a variety of habitat types). Our results indicate that, within each forested landscape, the gap and exterior grasslands harboured particular carabid assemblages (i.e. exclusive or abundantly collected species), which were also distinct from the surrounding forest carabid fauna. Dissimilarities between gap and exterior grasslands in each landscape suggest great carabid diversity at the regional scale. We also detected species-specific responses as several carabids were mainly associated with gap or exterior grasslands. Consequently, in highly modified forested landscapes, semi-natural grassland remnants may constitute great value for the protection of the carabid fauna. Specifically, we recommend conservation strategies that preserve variety in grassland features and maintain proper management activities to prevent the loss of specialised species and a decrease in regional carabid diversity.     

Extinction debt in naturally contracting mountain meadows in the Pacific Northwest,USA: varying responses of plants and feeding guilds of nocturnal moths

Fire suppression and climate change are leading to habitat fragmentation in temperate montane meadows across the globe, raising concerns about biodiversity loss. Restoration strategies may depend on the rate and nature of species response to habitat loss. We examined the effects of habitat loss and fragmentation on plants and nocturnal moths in natural montane meadows in the western Cascades, Oregon, USA, using generalized additive mixed models, non-metric multidimensional scaling, and multiple response permutation procedure. Historic (1949) rather than current (2005) meadow size explained species richness of herbaceous plants and herb-feeding moths and meadow plant community structure, indicating that loss of meadow species may be delayed by many decades following loss of meadow habitat, resulting in an extinction debt. In contrast, abundance of herb-feeding moths and species richness and abundance of woody plant-feeding moths were related to recent meadow configuration: as meadows are invaded by woody plants, abundance of meadow species declines, and woody plants and associated moths increase. Despite decades of fire suppression and climate change, montane meadows in many temperate mountain landscapes may still be amenable to restoration.     

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.     

Organic versus conventional management in an apple orchard: effects of fertilization and tree-row management on ground-dwelling predaceous arthropods

1 Organic and conventional management of apple orchards may have a different effect on arthropod communities.
2 We conducted a 3-year study to assess the effect of two strategies of fertilizer treatment (organic versus chemical) and three tree-row management systems (straw mulching, tillage and herbicide) on activity-density and biodiversity of epigeic predators. Ground beetles (Carabidae), rove beetles (Staphylinidae), ants (Formicidae) and spiders (Araneae) were sampled monthly with pitfall traps in the same apple orchard during 2003, 2004 and 2005.
3 A total of 4978 individuals were collected. Carabids (56.8% of the total catches) were the most abundant taxonomic group, followed by spiders (20.7%), ants (14.8%) and rove beetles (7.7%). Tree-row management had a greater influence on predator catches than fertilizer treatment. Total predator catches were lower under the mulch. Mulching also reduced carabid abundance, but increased staphylinid catches.
4 Tree row management also had a significant effect on biodiversity parameters. Species richness did not significantly differ among treatments for ants, spiders or the total catches, but was higher on herbicide-treated plots for carabids and on mulched plots for staphylinids. Shannon–Wiener's diversity index was significantly greater in the mulched and herbicide treated plots for total predators and carabids. For staphylinids, this index was significantly greater on the mulched plots. Fertilizer application strategy only influenced the species richness of rove beetles, which was greater in the chemically-treated plots.
5 The results showed that a change from conventional to organic fertilizer treatment of apple trees may be performed without differential effects on predator activity-density or biodiversity. However, a change from herbicide treatment to mulching or mechanical weed control may be significant, depending on the taxonomic group.     

Species Richness and Assemblages in Landscapes of Different Farming Intensity – Time to Revise Conservation Strategies?

Worldwide conservation goals to protect biodiversity emphasize the need to rethink which objectives are most suitable for different landscapes. Comparing two different Swedish farming landscapes, we used survey data on birds and vascular plants to test whether landscapes with large, intensively managed farms had lower richness and diversity of the two taxa than landscapes with less intensively managed small farms, and if they differed in species composition. Landscapes with large intensively managed farms did not have lower richness than smaller low intensively managed farms. The landscape types were also similar in that they had few red listed species, normally targeted in conservation. Differences in species composition demonstrate that by having both types of agricultural landscapes regional diversity is increased, which is seldom captured in the objectives for agro-environmental policies. Thus we argue that focus on species richness or red listed species would miss the actual diversity found in the two landscape types. Biodiversity conservation, especially in production landscapes, would therefore benefit from a hierarchy of local to regional objectives with explicit targets in terms of which aspects of biodiversity to focus on.     

Influence of within‐field position and adjoining habitat on carabid beetle assemblages in winter wheat

• 1 The influence of within‐field position and adjoining habitat on carabid beetles was studied in 20 winter wheat fields in ten different Swiss agricultural landscapes. In each landscape, two winter wheat fields (one with adjoining sown wildflower area and one with adjoining grassy margin) were investigated.
• 2 Carabid beetles were caught in pitfall traps 3 and 30 m from the edge in each of the 20 wheat fields. Significantly more individuals were found in the centres (30‐m position) than at the edges (3‐m position). Conversely, species richness was significantly higher at the field edges than in the centres.
• 3 Of the ten most abundant species, Poecilus cupreus, Agonum muelleri and Pterostichus melanarius were significantly more abundant in the field centres than at the edges. Harpalus rufipes was significantly more abundant in the fields adjoining sown wildflower areas than in the fields adjoining grassy margins.
• 4 In conclusion, the response of carabid beetles to within‐field position and adjoining habitats was species specific. This needs to be taken into account in habitat management for biodiversity conservation and pest control.

     

Assessing human impact on fen biodiversity: effects of different management regimes on butterfly, grasshopper, and carabid beetle assemblages

Fens and concomitantly the associated species assemblages have undergone dramatic declines in recent decades. Given that many remaining or restored fens are currently lying fallow, an important question is which management regime, if any, is most effective in preserving fen biodiversity. Against this background we here investigate the effects of five management regimes (intensive grassland, moist meadows, summer harvested sites, winter harvested sites, fallows) on three insect taxa (butterflies, grasshoppers, carabid beetles) in riverine fens in north-eastern Germany. Butterflies and grasshoppers showed highest species numbers, diversity and numbers of threatened species on moist meadows and were detrimentally affected by high vegetation. In contrast, ground dwelling carabid beetles were less strongly affected by different management regimes, but responded very sensitive to drainage. Winter harvested sites and fallows seemed to be particularly beneficial to carabid beetles. Overall, drained, intensive grassland was not suitable for preserving fen-specific communities, while extensively managed moist meadows harboured overall the highest numbers of threatened species. We conclude that in terms of management maintaining high water tables is most important. Furthermore, some management seems necessary to maintain stands with lower vegetation turf for associated species. However, care is needed to also maintain fallows and to consider the specific demands of some highly endangered, management-sensitive species.