The effectiveness of flower strips and hedgerows on pest control, pollination services and crop yield: a quantitative synthesis.

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.

Insectivorous birds disrupt biological control of cereal aphids.

Insect-feeding birds may interfere with trophic interactions in plant-insect food webs, which may be particularly important in agroecosystems. Here, we studied how Eurasian Tree Sparrows (Passer montanus) affect aphids and their predators in cereal fields using bird exclusion experiments. The Tree Sparrows fed their nestlings with aphid antagonists. Hoverflies and ladybird beetles accounted for 77% of the food for the nestlings during peak aphid density. When birds were excluded, densities of hoverfly larvae, which were the most abundant aphid predator group in the cereals, were 4% higher in wheat and 45% higher in oat, while aphid densities were 24% lower in wheat and 26% lower in oat. The demonstrated disruption of biological control by mesopredators through bird predation may be a common phenomenon in cropping systems characterized by small-sized and abundant pest species. Management of biotic interactions such as biological control needs a broad food-web perspective, even in simplified agroecosystems.

Forest defoliator pests alter carbon and nitrogen cycles.

Climate change may foster pest epidemics in forests, and thereby the fluxes of elements that are indicators of ecosystem functioning. We examined compounds of carbon (C) and nitrogen (N) in insect faeces, leaf litter, throughfall and analysed the soils of deciduous oak forests (Quercus petraea L.) that were heavily infested by the leaf herbivores winter moth (Operophtera brumata L.) and mottled umber (Erannis defoliaria L.). In infested forests, total net canopy-to-soil fluxes of C and N deriving from insect faeces, leaf litter and throughfall were 30- and 18-fold higher compared with uninfested oak forests, with 4333 kg C ha-1 and 319 kg N ha-1, respectively, during a pest outbreak over 3 years. In infested forests, C and N levels in soil solutions were enhanced and C/N ratios in humus layers were reduced indicating an extended canopy-to-soil element pathway compared with the non-infested forests. In a microcosm incubation experiment, soil treatments with insect faeces showed 16-fold higher fluxes of carbon dioxide and 10-fold higher fluxes of dissolved organic carbon compared with soil treatments without added insect faeces (control). Thus, the deposition of high rates of nitrogen and rapidly decomposable carbon compounds in the course of forest pest epidemics appears to stimulate soil microbial activity (i.e. heterotrophic respiration), and therefore, may represent an important mechanism by which climate change can initiate a carbon cycle feedback.

Farming practices change food web structures in cereal aphid-parasitoid-hyperparasitoid communities.

Agricultural intensification has been shown to result in a decline in biodiversity across many taxa, but the changes in community structure and species interactions remain little understood. We have analysed and compared the structure of feeding interactions for cereal aphids and their primary and secondary parasitoids in organically and conventionally managed winter wheat fields using quantitative food web metrics (interaction evenness, generality, vulnerability, link density). Despite little variation in the richness of each trophic group, food web structures between the two farming systems differed remarkably. In contrast to common expectations, aphids and primary parasitoids were characterized by (1) a higher evenness of interaction frequencies (interaction evenness) in conventional fields, which cascaded to interactions at the next trophic level, with (2) a higher interaction evenness, (3) a higher ratio of primary parasitoid taxa per secondary parasitoid (generality) and (4) a higher link density. Aphid communities in the organically managed fields almost exclusively consisted of a single ear-colonizing species, Sitobion avenae, while highly fertilized conventional fields were mainly infested by leaf-colonizing aphids that benefit from the nutritional status of winter wheat. In conclusion, agricultural intensification appears to foster the complexity of aphid-parasitoid food webs, thereby not supporting the general expectation on the importance of organic farming practices for species richness and food web complexity.

Agricultural intensification and cereal aphid-parasitoid-hyperparasitoid food webs: network complexity, temporal variability and parasitism rates.

Agricultural intensification (AI) is currently a major driver of biodiversity loss and related ecosystem functioning decline. However, spatio-temporal changes in community structure induced by AI, and their relation to ecosystem functioning, remain largely unexplored. Here, we analysed 16 quantitative cereal aphid-parasitoid and parasitoid-hyperparasitoid food webs, replicated four times during the season, under contrasting AI regimes (organic farming in complex landscapes vs. conventional farming in simple landscapes). High AI increased food web complexity but also temporal variability in aphid-parasitoid food webs and in the dominant parasitoid species identity. Enhanced complexity and variability appeared to be controlled bottom-up by changes in aphid dominance structure and evenness. Contrary to the common expectations of positive biodiversity-ecosystem functioning relationships, community complexity (food-web complexity, species richness and evenness) was negatively related to primary parasitism rates. However, this relationship was positive for secondary parasitoids. Despite differences in community structures among different trophic levels, ecosystem services (parasitism rates) and disservices (aphid abundances and hyperparasitism rates) were always higher in fields with low AI. Hence, community structure and ecosystem functioning appear to be differently influenced by AI, and change differently over time and among trophic levels. In conclusion, intensified agriculture can support diverse albeit highly variable parasitoid-host communities, but ecosystem functioning might not be easy to predict from observed changes in community structure and composition.

Landscape moderation of biodiversity patterns and processes - eight hypotheses.

Understanding how landscape characteristics affect biodiversity patterns and ecological processes at local and landscape scales is critical for mitigating effects of global environmental change. In this review, we use knowledge gained from human-modified landscapes to suggest eight hypotheses, which we hope will encourage more systematic research on the role of landscape composition and configuration in determining the structure of ecological communities, ecosystem functioning and services. We organize the eight hypotheses under four overarching themes. Section A: 'landscape moderation of biodiversity patterns' includes (1) the landscape species pool hypothesis-the size of the landscape-wide species pool moderates local (alpha) biodiversity, and (2) the dominance of beta diversity hypothesis-landscape-moderated dissimilarity of local communities determines landscape-wide biodiversity and overrides negative local effects of habitat fragmentation on biodiversity. Section B: 'landscape moderation of population dynamics' includes (3) the cross-habitat spillover hypothesis-landscape-moderated spillover of energy, resources and organisms across habitats, including between managed and natural ecosystems, influences landscape-wide community structure and associated processes and (4) the landscape-moderated concentration and dilution hypothesis-spatial and temporal changes in landscape composition can cause transient concentration or dilution of populations with functional consequences. Section C: 'landscape moderation of functional trait selection' includes (5) the landscape-moderated functional trait selection hypothesis-landscape moderation of species trait selection shapes the functional role and trajectory of community assembly, and (6) the landscape-moderated insurance hypothesis-landscape complexity provides spatial and temporal insurance, i.e. high resilience and stability of ecological processes in changing environments. Section D: 'landscape constraints on conservation management' includes (7) the intermediate landscape-complexity hypothesis-landscape-moderated effectiveness of local conservation management is highest in structurally simple, rather than in cleared (i.e. extremely simplified) or in complex landscapes, and (8) the landscape-moderated biodiversity versus ecosystem service management hypothesis-landscape-moderated biodiversity conservation to optimize functional diversity and related ecosystem services will not protect endangered species. Shifting our research focus from local to landscape-moderated effects on biodiversity will be critical to developing solutions for future biodiversity and ecosystem service management.

The relationship between agricultural intensification and biological control: experimental tests across Europe.

Agricultural intensification can affect biodiversity and related ecosystem services such as biological control, but large-scale experimental evidence is missing. We examined aphid pest populations in cereal fields under experimentally reduced densities of (1) ground-dwelling predators (-G), (2) vegetation-dwelling predators and parasitoids (-V), (3) a combination of (1) and (2) (-G-V), compared with open-fields (control), in contrasting landscapes with low vs. high levels of agricultural intensification (AI), and in five European regions. Aphid populations were 28%, 97%, and 199% higher in -G, -V, and -G-V treatments, respectively, compared to the open fields, indicating synergistic effects of both natural-enemy groups. Enhanced parasitoid: host and predator: prey ratios were related to reduced aphid population density and population growth. The relative importance of parasitoids and vegetation-dwelling predators greatly differed among European regions, and agricultural intensification affected biological control and aphid density only in some regions. This shows a changing role of species group identity in diverse enemy communities and a need to consider region-specific landscape management.

Agricultural intensification and biodiversity partitioning in European landscapes comparing plants, carabids, and birds.

Effects of agricultural intensification (AI) on biodiversity are often assessed on the plot scale, although processes determining diversity also operate on larger spatial scales. Here, we analyzed the diversity of vascular plants, carabid beetles, and birds in agricultural landscapes in cereal crop fields at the field (n = 1350), farm (n = 270), and European-region (n = 9) scale. We partitioned diversity into its additive components alpha, beta, and gamma, and assessed the relative contribution of beta diversity to total species richness at each spatial scale. AI was determined using pesticide and fertilizer inputs, as well as tillage operations and categorized into low, medium, and high levels. As AI was not significantly related to landscape complexity, we could disentangle potential AI effects on local vs. landscape community homogenization. AI negatively affected the species richness of plants and birds, but not carabid beetles, at all spatial scales. Hence, local AI was closely correlated to beta diversity on larger scales up to the farm and region level, and thereby was an indicator of farm- and region-wide biodiversity losses. At the scale of farms (12.83-20.52%) and regions (68.34-80.18%), beta diversity accounted for the major part of the total species richness for all three taxa, indicating great dissimilarity in environmental conditions on larger spatial scales. For plants, relative importance of alpha diversity decreased with AI, while relative importance of beta diversity on the farm scale increased with AI for carabids and birds. Hence, and in contrast to our expectations, AI does not necessarily homogenize local communities, presumably due to the heterogeneity of farming practices. In conclusion, a more detailed understanding of AI effects on diversity patterns of various taxa and at multiple spatial scales would contribute to more efficient agri-environmental schemes in agroecosystems.

Food web structure and biocontrol in a four-trophic level system across a landscape complexity gradient.

Decline in landscape complexity owing to agricultural intensification may affect biodiversity, food web complexity and associated ecological processes such as biological control, but such relationships are poorly understood. Here, we analysed food webs of cereal aphids, their primary parasitoids and hyperparasitoids in 18 agricultural landscapes differing in structural complexity (42-93% arable land). Despite little variation in the richness of each trophic group, we found considerable changes in trophic link properties across the landscape complexity gradient. Unexpectedly, aphid-parasitoid food webs exhibited a lower complexity (lower linkage density, interaction diversity and generality) in structurally complex landscapes, which was related to the dominance of one aphid species in complex landscapes. Nevertheless, primary parasitism, as well as hyperparasitism, was higher in complex landscapes, with primary parasitism reaching levels for potentially successful biological control. In conclusion, landscape complexity appeared to foster higher parasitism rates, but simpler food webs, thereby casting doubt on the general importance of food web complexity for ecosystem functioning.

Crop-noncrop spillover: arable fields affect trophic interactions on wild plants in surrounding habitats.

Ecosystem processes in agricultural landscapes are often triggered by resource availability in crop and noncrop habitats. We investigated how oilseed rape (OSR; Brassica napus, Brassicaceae) affects noncrop plants in managed systems and semi-natural habitat, using trophic interactions among wild mustard (Sinapis arvensis, Brassicaceae), rape pollen beetles (Meligethes aeneus, Nitidulidae) and their parasitoids (Tersilochus heterocerus, Ichneumonidae). We exposed wild mustard as phytometer plants in two cropland habitat types (wheat field, field margin) and three noncrop habitat types (fallow, grassland, wood margin) across eight landscapes along a gradient from simple to complex (quantified as % arable land). Both landscape and local factors affected the abundance of rape pollen beetles and parasitoids. Rape pollen beetle infestation and parasitism rates on these plants were lower in noncrop habitats and higher in wheat fields and field margins, whereas beetles and parasitoids responded differently to landscape scale parameters. We found the hypothesized spillover from OSR crop onto wild plants in surrounding habitats only for parasitoids, but not for pollen beetles. Parasitism rates were not related to landscape simplification, but benefited from increasing proportions of OSR. In contrast, rape pollen beetles benefited from simple landscape structures, presumably due to multi-annual population build-ups resulting from long-term OSR planting (as part of the crop rotation). In conclusion, we showed that spillover from cropland affects parasitism rates on related wild plants outside cropland, which has not been shown so far, but can be expected to be a widespread effect shaping noncrop food webs.

Changes in herbivore control in arable fields by detrital subsidies depend on predator species and vary in space.

Prey from the decomposer subsystem may help sustain predator populations in arable fields. Adding organic residues to agricultural systems may therefore enhance pest control. We investigated whether resource addition (maize mulch) strengthens aboveground trophic cascades in winter wheat fields. Evaluating the flux of the maize-borne carbon into the food web after 9 months via stable isotope analysis allowed differentiating between prey in predator diets originating from the above- and belowground subsystems. Furthermore, we recorded aphid populations in predator-reduced and control plots of no-mulch and mulch addition treatments. All analyzed soil dwelling species incorporated maize-borne carbon. In contrast, only 2 out of 13 aboveground predator species incorporated maize carbon, suggesting that these 2 predators forage on prey from the above- and belowground systems. Supporting this conclusion, densities of these two predator species were increased in the mulch addition fields. Nitrogen isotope signatures suggested that these generalist predators in part fed on Collembola thereby benefiting indirectly from detrital resources. Increased density of these two predator species was associated by increased aphid control but the identity of predators responsible for aphid control varied in space. One of the three wheat fields studied even lacked aphid control despite of mulch-mediated increased density of generalist predators. The results suggest that detrital subsidies quickly enter belowground food webs but only a few aboveground predator species include prey out of the decomposer system into their diet. Variation in the identity of predator species benefiting from detrital resources between sites suggest that, depending on locality, different predator species are subsidised by prey out of the decomposer system and that these predators contribute to aphid control. Therefore, by engineering the decomposer subsystem via detrital subsidies, biological control by generalist predators may be strengthened.

Beta diversity at different spatial scales: plant communities in organic and conventional agriculture.

Biodiversity studies that guide agricultural subsidy policy have generally compared farming systems at a single spatial scale: the field. However, diversity patterns vary across spatial scales. Here, we examined the effects of farming system (organic vs. conventional) and position in the field (edge vs. center) on plant species richness in wheat fields at three spatial scales. We quantified alpha-, beta-, and gamma-diversity at the microscale in 800 plots, at the mesoscale in 40 fields, and at the macroscale in three regions using the additive partitioning approach, and evaluated the relative contribution of beta-diversity at each spatial scale to total observed species richness. We found that alpha-, beta-, and gamma-diversity were higher in organic than conventional fields and higher at the field edge than in the field center at all spatial scales. In both farming systems, beta-diversity at the meso- and macroscale explained most of the overall species richness (up to 37% and 25%, respectively), indicating considerable differences in community composition among fields and regions due to environmental heterogeneity. The spatial scale at which beta-diversity contributed the most to overall species richness differed between rare and common species. Total richness of rare species (present in < or = 5% of total samples) was mainly explained by differences in community composition at the meso- and macroscale (up to 27% and 48%, respectively), but only in organic fields. Total richness of common species (present in > or = 25% of total samples) was explained by differences in community composition at the micro- and mesoscale (up to 29% and 47%, respectively), i.e., among plots and fields, independent of farming system. Our results show that organic farming made the greatest contribution to total species richness at the meso (among fields) and macro (among regions) scale due to environmental heterogeneity. Hence, agri-environment schemes should exploit this large-scale contribution of beta-diversity by tailoring schemes at regional scales to maximize dissimilarity between conservation areas using geographic information systems rather than focusing entirely at the classical local-field scale, which is the current practice.

The landscape context of cereal aphid-parasitoid interactions.

Analyses at multiple spatial scales may show how important ecosystem services such as biological control are determined by processes acting on the landscape scale. We examined cereal aphid-parasitoid interactions in wheat fields in agricultural landscapes differing in structural complexity (32-100% arable land). Complex landscapes were associated with increased aphid mortality resulting from parasitism, but also with higher aphid colonization, thereby counterbalancing possible biological control by parasitoids and lastly resulting in similar aphid densities across landscapes. Thus, undisturbed perennial habitats appeared to enhance both pests and natural enemies. Analyses at multiple spatial scales (landscape sectors of 0.5-6 km diameter) showed that correlations between parasitism and percentage of arable land were significant at scales of 0.5-2 km, whereas aphid densities responded to percentage of arable land at scales of 1-6 km diameter. Hence, the higher trophic level populations appeared to be determined by smaller landscape sectors owing to dispersal limitation, showing the 'functional spatial scale' for species-specific landscape management.

Relative importance of predators and parasitoids for cereal aphid control.

Field experiments with manipulations of natural enemies of plant-feeding insects may show how a diverse enemy group ensures an important ecosystem function such as naturally occurring biological pest control. We studied cereal aphid populations in winter wheat under experimentally reduced densities of: (i) ground-dwelling generalist predators (mostly spiders, carabid and staphylinid beetles); (ii) flying predators (coccinellid beetles, syrphid flies, gall midges, etc.) and parasitoids (aphidiid wasps), and a combination of (i) and (ii), compared with open controls. Aphid populations were 18% higher at reduced densities of ground-dwelling predators, 70% higher when flying predators and parasitoids were removed, and 172% higher on the removal of both enemy groups. Parasitoid wasps probably had the strongest effect, as flying predators occurred only in negligible densities. The great importance of parasitism is a new finding for aphid control in cereal fields. In conclusion, a more detailed knowledge of the mechanisms of natural pest control would help to develop environmentally sound crop management with reduced pesticide applications.