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The idea that noncrop habitat enhances pest control and represents a win-win opportunity to conserve biodiversity and bolster yields has emerged as an agroecological paradigm. However, while noncrop habitat in landscapes surrounding farms sometimes benefits pest predators, natural enemy responses remain heterogeneous across studies and effects on pests are inconclusive. The observed heterogeneity in species responses to noncrop habitat may be biological in origin or could result from variation in how habitat and biocontrol are measured. Here, we use a pest-control database encompassing 132 studies and 6,759 sites worldwide to model natural enemy and pest abundances, predation rates, and crop damage as a function of landscape composition. Our results showed that although landscape composition explained significant variation within studies, pest and enemy abundances, predation rates, crop damage, and yields each exhibited different responses across studies, sometimes increasing and sometimes decreasing in landscapes with more noncrop habitat but overall showing no consistent trend. Thus, models that used landscape-composition variables to predict pest-control dynamics demonstrated little potential to explain variation across studies, though prediction did improve when comparing studies with similar crop and landscape features. Overall, our work shows that surrounding noncrop habitat does not consistently improve pest management, meaning habitat conservation may bolster production in some systems and depress yields in others. Future efforts to develop tools that inform farmers when habitat conservation truly represents a win-win would benefit from increased understanding of how landscape effects are modulated by local farm management and the biology of pests and their enemies., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

Cover cropping is a common practice in U.S. Midwest carrot production for soil conservation, and may affect soil ecology and plant-parasitic nematodes-to which carrots are very susceptible. This study assessed the impact of cover crops-oats ( Avena sativa ), radish ( Raphanus sativus ) cv. Defender, rape ( Brassica napus ) cv. Dwarf Essex, and a mixture of oats and radish-on plant-parasitic nematodes and soil ecology based on the nematode community in Michigan carrot production systems. Research was conducted at two field sites where cover crops were grown in Fall 2014 preceding Summer 2015 carrot production. At Site 1, root-lesion ( Pratylenchus penetrans ) and stunt ( Tylenchorhynchus sp.) nematodes were present at low population densities (less than 25 nematodes/100 cm 3 soil), but were not significantly affected ( P > 0.05) by cover crops. At Site 2, P. penetrans population densities were increased ( P ≤ 0.05) by 'Defender' radish compared to other cover crops or fallow control during cover crop growth and midseason carrot production. At both sites, there were few short-term impacts of cover cropping on soil ecology based on the nematode community. At Site 1, only at carrot harvest, radish-oats mixture and 'Dwarf Essex' rape alone enriched the soil food web based on the enrichment index ( P ≤ 0.05) while rape and radish increased structure index values. At Site 2, bacterivore abundance was increased by oats or radish cover crops compared to control, but only during carrot production. In general, cover crops did not affect the nematode community until nearly a year after cover crop growth suggesting that changes in the soil community following cover cropping may be gradual.

The strength and prevalence of trophic cascades, defined as positive, indirect effects of natural enemies (predatory and parasitic arthropods) on plants, is highly variable in agroecosystems. This variation may in part be due to the spatial or landscape context in which hese trophic cascades occur. In 2011 and 2012, we conducted a natural enemy exclusion experiment in soybean fields along a gradient of landscape composition across southern Wisconsin and Michigan, USA. We used structural equation modeling to ask (1) whether natural enemies influence biocontrol of soybean aphids (SBA) and soybean yield and (2) whether landscape effects on natural enemies influence the strength of the trophic cascades. We found that natural enemies (NE) suppressed aphid populations in both years of our study, and, in 2011, the yield of soybean plants exposed to natural enemies was 37% higher than the yield of plants with aphid populations protected from natural enemies. The strength of the :rophic cascade was also influenced by landscape context. We found that landscapes with a higher proportion of soybean and higher diversity habitats resulted in more NE, fewer aphids, and, in some cases, a trend toward greater soybean yield. These results indicate that landscape context is important for understanding spatial variability in biocontrol and yield, but other factors, such as environmental variability and compensatory growth, might overwhelm the beneficial effects of biocontrol on crop yield.

Agriculture is being challenged to provide food, and increasingly fuel, for an expanding global population. Producing bioenergy crops on marginal lands--farmland suboptimal for food crops--could help meet energy goals while minimizing competition with food production. However, the ecological costs and benefits of growing bioenergy feedstocks--primarily annual grain crops--on marginal lands have been questioned. Here we show that perennial bioenergy crops provide an alternative to annual grains that increases biodiversity of multiple taxa and sustain a variety of ecosystem functions, promoting the creation of multifunctional agricultural landscapes. We found that switchgrass and prairie plantings harbored significantly greater plant, methanotrophic bacteria, arthropod, and bird diversity than maize. Although biomass production was greater in maize, all other ecosystem services, including methane consumption, pest suppression, pollination, and conservation of grassland birds, were higher in perennial grasslands. Moreover, we found that the linkage between biodiversity and ecosystem services is dependent not only on the choice of bioenergy crop but also on its location relative to other habitats, with local landscape context as important as crop choice in determining provision of some services. Our study suggests that bioenergy policy that supports coordinated land use can diversify agricultural landscapes and sustain multiple critical ecosystem services., Competing Interests: The authors declare no conflict of interest.

Biomass crops grown on marginal soils are expected to fuel an emerging bioenergy industry in the United States. Bioenergy crop choice and position in the landscape could have important impacts on a range of ecosystem services, including natural pest-suppression (biocontrol services) provided by predatory arthropods. In this study we use predation rates of three sentinel crop pests to develop a biocontrol index (BCI) summarizing pest-suppression potential in corn and perennial grass-based bioenergy crops in southern Wisconsin, lower Michigan, and northern Illinois. We show that BCI is higher in perennial grasslands than in corn, and increases with the amount of perennial grassland in the surrounding landscape. We develop an empirical model for predicting BCI from information on energy crop and landscape characteristics, and use the model in a qualitative assessment of changes in biocontrol services for annual croplands on prime agricultural soils under two contrasting bioenergy scenarios. Our analysis suggests that the expansion of annual energy crops onto 1.2 million ha of existing perennial grasslands on marginal soils could reduce BCI between -10 and -64% for nearly half of the annual cropland in the region. In contrast, replacement of the 1.1 million ha of existing annual crops on marginal land with perennial energy crops could increase BCI by 13 to 205% on over half of the annual cropland in the region. Through comparisons with other independent studies, we find that our biocontrol index is negatively related to insecticide use across the Midwest, suggesting that strategically positioned, perennial bioenergy crops could reduce insect damage and insecticide use on neighboring food and forage crops. We suggest that properly validated environmental indices can be used in decision support systems to facilitate integrated assessments of the environmental and economic impacts of different bioenergy policies.

Determining when multiple predator species provide better pest suppression than single species is a key step towards developing ecologically-informed biological control strategies. Theory and experiments predict that resource partitioning among functionally different predator species can strengthen prey suppression, because as a group they can access more prey types than functionally redundant predators. However, this prediction assumes that competition limits predation by functionally similar predators. Differences in prey density can alter the strength of competition, suggesting that prey abundance may modulate the effect of combining functionally diverse species. The experiment documented here examined the potential for functional differences among predator species to promote suppression of an insect pest, the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), at different prey densities. Predation was compared at two prey densities between microcosms that contained one predator species or two functionally distinct species: the lady beetle, Coleomegilla maculata De Geer (Coleoptera: Coccinellidae) that kills early L. decemlineata instars, and the soldier bug, Podisus maculiventris Say (Hemiptera: Pentatomidae) that kills late instars. The data show that combining these predators increased predation only when prey densities were low. This suggests that multiple predator species may only provide greater biological control than single species in systems where prey is limiting.

Onion maggot, Delia antiqua (Meigen) (Diptera: Anthomyiidae), is an important pest of onion, Allium cepa L., in northern temperate areas, especially in the Great Lakes region of North America Management of D. antiqua relies on insecticide use at planting, but insecticide resistance can cause control failures that threaten the long-term viability of this strategy. Delaying the time onions are planted was investigated as an alternative management approach for D. antiqua and the ecological and behavioral mechanisms underlying host age and insect relationships were examined in laboratory and field experiments. Delaying onion planting by two to four weeks reduced damage to onions by 35 and 90%, respectively. Onions planted later emerged later and this reduced the period overwintered flies had to oviposit on the plants. Moreover, flies tended to lay few to no eggs on these young, late-planted onions. As anticipated, D. antiqua laid 4-8 times more eggs on older onions than on young onions, and older onions were more resilient to injury caused by D. antiqua neonates compared with younger onions. However, the resiliency to maggot attack lessened as the density of D. antiqua increased from 2 to 10 eggs per plant, which probably explains why greater levels of maggot damage are typically observed in early onion plantings compared with later plantings. Delaying onion planting until mid-May reduced D. antiqua damage without jeopardizing the period required to produce marketable yield, but this cultural tactic must be combined with other management strategies to prevent economic loss.

Agronomic intensification has transformed many agricultural landscapes into expansive monocultures with little natural habitat. A pervasive concern is that such landscape simplification results in an increase in insect pest pressure, and thus an increased need for insecticides. We tested this hypothesis across a range of cropping systems in the Midwestern United States, using remotely sensed land cover data, data from a national census of farm management practices, and data from a regional crop pest monitoring network. We found that, independent of several other factors, the proportion of harvested cropland treated with insecticides increased with the proportion and patch size of cropland and decreased with the proportion of seminatural habitat in a county. We also found a positive relationship between the proportion of harvested cropland treated with insecticides and crop pest abundance, and a positive relationship between crop pest abundance and the proportion cropland in a county. These results provide broad correlative support for the hypothesized link between landscape simplification, pest pressure, and insecticide use. Using regression coefficients from our analysis, we estimate that, across the seven-state region in 2007, landscape simplification was associated with insecticide application to 1.4 million hectares and an increase in direct costs totaling between $34 and $103 million. Both the direct and indirect environmental costs of landscape simplification should be considered in design of land use policy that balances multiple ecosystem goods and services.

Agricultural habitats are the recipients of ecosystem services provided by mobile arthropods like pollinators and natural enemies. These organisms can disperse between non-crop habitats and crops and can benefit from resources in non-crop areas. Thus, it may be important to conserve non-crop habitats to maintain ecosystem services within crops. However, even single ecosystem services, such as pollination or predation of pests, may be provided by a complex of species. Different species may respond to landscape structure at differing spatial scales. This raises the question: What scales should conservation take place at to sustain an ecosystem service? Here, we examine variation in predation of two potato pests in response to landscape structure. Specifically, predation of Colorado potato beetle, Leptinotarsa decemlineata, and green peach aphid, Myzus persicae, were quantified in potato and field-margin habitats set in landscapes that varied at different scales. At a local scale (meters), potato fields of different sizes were bordered by different areas of grassy field margin. At a broad scale (kilometers), these habitats were set in landscapes composed of varying percentages of non-crop habitat. Predation of both pests was associated with landscape structure, but this relationship occurred at different scales for each pest and interacted differently with habitat type. Predation of L. decemlineata eggs was greater in field margins than in the potato crop and increased in both habitats when field margins were large relative to potatoes. In contrast, aphid predation in field margins increased with the area of non-crop habitat within 1.5 km, but did not change in adjacent potato. These data emphasize that, even a single ecosystem service, such as predation of pests, can be influenced by landscape structure at multiple scales. More generally, it may be necessary to conserve heterogeneity both at local scales (individual farm fields) and broad scales (entire mosaic landscapes) to maintain ecosystem services. This may be especially true for ecosystem services that are the aggregate result of the actions of many species or ecological processes.