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Plot size is of practical importance in any integrated pest management (IPM) study that has a field component. Such studies need to be conducted at a scale relevant to species dynamics because their abundance and distribution in plots might vary according to plot size. An adequate plot size is especially important for researchers, technology providers and regulatory agencies in understanding effects of various insect control technologies on non-target arthropods. Plots that are too small might fail to detect potential harmful effects of these technologies due to arthropod movement and redistribution among plots, or from untreated areas and outside sources. The Arizona cotton system is heavily dependent on technologies for arthropod control, thus we conducted a 2-year replicated field experiment to estimate the optimal plot size for non-target arthropod studies in our system. Experimental treatments consisted of three square plot sizes and three insecticides in a full factorial. We established three plot sizes that measured 144 m2, 324 m2 and 576 m2. For insecticide treatments, we established an untreated check, a positive control insecticide with known negative effects on the arthropod community and a selective insecticide. We investigated how plot size impacts the estimation of treatment effects relative to community structure (27 taxa), community diversity, individual abundance, effect sizes, biological control function of arthropod taxa with a wide range of mobility, including Collops spp., Orius tristicolor, Geocoris spp., Misumenops celer, Drapetis nr. divergens and Chrysoperla carnea s.l.. Square 144 m2 plots supported similar results for all parameters compared with larger plots, and are thus sufficiently large to measure insecticidal effects on non-target arthropods in cotton. Our results are applicable to cotton systems with related pests, predators or other fauna with similar dispersal characteristics. Moreover, these results also might be generalizable to other crop systems with similar fauna.

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BACKGROUND: The population dynamics of polyphagous pests such as Bemisia argentifolii (B. tabaci MEAM1) are governed by complex, interacting factors involving its cultivated and wild host plants, seasonality, movement and demography. To understand mechanisms contributing to population development and pest success within the agroecosystem, contiguous multi-host field sites were established in three environmentally distinct areas in Arizona. Life tables quantified and partition models described mortality sources and rates for immature insect stages on each host plant. RESULTS: Predation and dislodgement were the largest sources of marginal mortality, supplied the highest irreplaceable mortality and predation was the key factor. Rates of mortality were best predicted, in order, by source, temperature, host plant and season. Marginal mortality was highest for fourth-stage nymphs followed by eggs. Mortality rates were predicted in descending order by stage, temperature and season. Survivorship patterns varied among host plants, and generational mortality averaged 70% on spring cantaloupes but nearly 95% on all other hosts. Population density varied seasonally, persisting at low levels on winter hosts and expanding beginning in the spring; perennial hosts and weeds bridge populations year-round. CONCLUSION: Survival on winter hosts such as broccoli, albeit low, enables population continuity, whereas unusually high survivorship on spring crops like cantaloupe is an ecological release propelling population growth and driving regional dynamics in the summer and fall. This detailed understanding of mortality dynamics provides clues to the success of this invasive pest in our agroecosystems and facilitates opportunities for improved pest management at a broader landscape scale. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.

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OBJECTIVES: To assess potential non-target effects of genetically engineered/modified (GM) maize that produces insecticidal proteins from Bacillus thuringiensis (Bt), numerous field experiments have been conducted worldwide. Field data are often variable and influenced by uncontrolled factors and meta-analyses can recognize general effects with increased statistical power compared to individual studies. This database represents a comprehensive collection of experimental field data on non-target invertebrates in Bt and non-Bt maize. It was created for a systematic review with the question if growing Bt maize changes abundance or ecological function of non-target animals compared to growing of non-GM maize. Systematic literature searches identified relevant data. Authors were contacted for additional information or raw data if needed and a critical appraisal scheme was developed and applied to each data record. DATA DESCRIPTION: The database contains 7279 records of non-target invertebrate abundance, activity density, or predation or parasitism extracted from 120 articles. Records for individual species and life stages, but also aggregated data are available. Each record represents a comparison of invertebrates in Bt and non-Bt maize and includes means, standard deviations and sample sizes. Additional variables characterize publication details, experimental setup, cultivars, Bt proteins, geographic location, field management, insecticide treatments, sampling details, and taxonomy.

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BACKGROUND: Hundreds of studies on environmental effects of genetically modified (GM) crops became available over the past 25 years. For maize producing insecticidal proteins from Bacillus thuringiensis (Bt), potential adverse effects on non-target organisms are a major area of concern and addressed in risk assessments. Reviews and meta-analyses have helped various stakeholders to address uncertainties regarding environmental impacts of the technology. Many field studies from Europe and other parts of the world have been published in the last decade, and those data are often not covered by previous meta-analyses. Therefore, we conducted a systematic review to answer the question: "Does the growing of Bt maize change abundance or ecological function of non-target animals compared to the growing of non-GM maize?" METHODS: Literature published until August 2019 was searched systematically in 12 bibliographic databases, 17 specialized webpages, and reference sections of 78 review articles. Defined eligibility criteria were applied to screen titles, abstracts, and full texts of the retrieved references. A custom-made database was developed with quantitative data on invertebrate abundance, activity density, or predation/parasitism rates. Eligible data that did not fit the quantitative database were captured in detailed tables and summarized narratively. For the first time, a critical appraisal scheme for field studies on non-targets in GM crops was developed to estimate the risk of bias (internal validity) and the suitability to answer the review question (external validity) of all primary data. Meta-analyses on different taxonomic levels, functional groups, and types of Bt maize were conducted. Untreated Bt maize was either compared with untreated non-Bt maize, or with insecticide-treated non-Bt maize. The influence of contributions by private sector product developers on reported effects was investigated. REVIEW FINDINGS: The database on non-target effects of Bt maize field trials contains more than 7200 records from 233 experiments and 120 articles. Meta-analyses on different taxonomic levels revealed only few and often non-robust significant effect sizes when both Bt maize and non-Bt maize were untreated. Bt maize harboured fewer parasitoids (Braconidae, Tachinidae) of the European corn borer, the main target pest of Lepidoptera-active Bt maize, compared with non-Bt maize. Similarly, sap beetles (Nitidulidae), that are associated with Lepidoptera damage, were recorded less in Bt maize. In some analyses, a negative effect of Bt maize was observed for rove beetles (Staphylinidae) and hoverflies (Syrphidae) and a positive effect for ladybeetles (Coccinellidae), flower bugs (Anthocoridae), and lacewings (Neuroptera). However, those effects were not consistent for different analyses and often related to individual articles. When untreated Bt maize was compared with pyrethroid-treated non-Bt maize, more effect sizes were significant. In particular, populations of predators were reduced after pyrethroid treatment, while few data were available for other insecticides. Funnel plots showed no evidence for publication bias and the analyses of private sector contribution revealed no evidence for influence of vested interests. Conclusions about potential effects of Bt maize on vertebrates or on animals inhabiting off-crop habitats were not possible, because only few such studies fitting the format of direct Bt/non-Bt comparisons on plot or field level were identified. CONCLUSIONS: The current work largely confirmed previously published results. The effects of Bt maize on the community of non-target invertebrates inhabiting maize fields were small and mostly neutral, especially when compared with the effects of broad-spectrum pyrethroid insecticide treatments.

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Conservation biological control is a fundamental tactic in integrated pest management (IPM). Greater biological control services can be achieved by enhancing agroecosystems to be more favorable to the presence, survival, and growth of natural enemy populations. One approach that has been tested in numerous agricultural systems is the deployment of synthetic chemicals that mimic those produced by the plant when under attack by pests. These signals may attract arthropod natural enemies to crop habitats and thus potentially improve biological control activity locally. A 2-yr field study was conducted in the cotton agroecosystem to evaluate the potential of synthetic methyl salicylate (MeSA) to attract native arthropod natural enemies and to enhance biological control services on two key pests. Slow-release packets of MeSA were deployed in replicated cotton plots season long. The abundance of multiple taxa of natural enemies and two major pests were monitored weekly by several sampling methods. The deployment of MeSA failed to increase natural enemy abundance and pest densities did not decline. Predator to prey ratios, used as a proxy to estimate biological control function, also largely failed to increase with MeSA deployment. One exception was a season-long increase in the ratio of Orius tristicolor (White) (Hemiptera: Anthocoridae) to Bemisia argentifolii Bellows and Perring (= Bemisia tabaci MEAM1) (Hemiptera: Aleyrodidae) adults within the context of biological control informed action thresholds. Overall results suggest that MeSA would not likely enhance conservation biological control by the natural enemy community typical of U.S. western cotton production systems.

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Invasive organisms pose a global threat and are exceptionally difficult to eradicate after they become abundant in their new habitats. We report a successful multitactic strategy for combating the pink bollworm (Pectinophora gossypiella), one of the world's most invasive pests. A coordinated program in the southwestern United States and northern Mexico included releases of billions of sterile pink bollworm moths from airplanes and planting of cotton engineered to produce insecticidal proteins from the bacterium Bacillus thuringiensis (Bt). An analysis of computer simulations and 21 y of field data from Arizona demonstrate that the transgenic Bt cotton and sterile insect releases interacted synergistically to reduce the pest's population size. In Arizona, the program started in 2006 and decreased the pest's estimated statewide population size from over 2 billion in 2005 to zero in 2013. Complementary regional efforts eradicated this pest throughout the cotton-growing areas of the continental United States and northern Mexico a century after it had invaded both countries. The removal of this pest saved farmers in the United States $192 million from 2014 to 2019. It also eliminated the environmental and safety hazards associated with insecticide sprays that had previously targeted the pink bollworm and facilitated an 82% reduction in insecticides used against all cotton pests in Arizona. The economic and social benefits achieved demonstrate the advantages of using agricultural biotechnology in concert with classical pest control tactics.

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An amendment to this paper has been published and can be accessed via a link at the top of the paper.

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The Green Revolution is credited with alleviating famine, mitigating poverty and driving aggregate economic growth since the 1960s. In Asia, high-input technology packages secured a tripling of rice output, with germplasm improvements providing benefits beyond US$4.3 billion yr-1. Here, we unveil the magnitude and macro-economic relevance of parallel nature-based contributions to productivity growth in non-rice crops over the period 1918-2018 (across 23 different Asia-Pacific geopolitical entities). We empirically demonstrate how biological control resolved invasive pest threats in multiple agricultural commodities, ensuring annually accruing (on-farm) benefits of US$14.6-19.5 billion yr-1. Scientifically guided biological control of 43 exotic invertebrate pests permitted 73-100% yield-loss recovery in critical food, feed and fibre crops including banana, breadfruit, cassava and coconut. Biological control thereby promoted rural growth and prosperity even in marginal, poorly endowed, non-rice environments. By placing agro-ecological innovations on equal footing with input-intensive measures, our work provides lessons for future efforts to mitigate invasive species, restore ecological resilience and sustainably raise output of global agrifood systems.

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Bt cotton was genetically modified to produce insecticidal proteins targeting Lepidopteran pests and is therefore only minimally affected by caterpillar damage. This could lead to reduced levels of inherent, systemically inducible defensive compounds in Bt cotton which might benefit other important cotton herbivores such as plant bugs. We studied the effects of plant defense induction on the performance of the plant bug Lygus hesperus by caging nymphs on different food sources (bolls/squares) of Bt and non-Bt cotton which were either undamaged, damaged by Bt tolerant caterpillars, or treated with jasmonic acid (JA). Terpenoid induction patterns of JA-treated and L. hesperus-damaged plants were characterized for different plant structures and artificial diet assays using purified terpenoids (gossypol/heliocide H1/4) were conducted. Nymphs were negatively affected if kept on plants damaged by caterpillars or sprayed with JA. Performance of nymphs was increased if they fed on squares and by the Bt-trait which had a positive effect on boll quality as food. In general, JA-sprayed plants (but not L. hesperus infested plants) showed increased levels of terpenoids in the plant structures analyzed, which was especially pronounced in Bt cotton. Nymphs were not negatively affected by terpenoids in artificial diet assays indicating that other inducible cotton responses are responsible for the found negative effects on L. hesperus. Overall, genetically engineered plant defenses can benefit plant bugs by releasing them from plant-mediated indirect competition with lepidopterans which might contribute to increasing numbers of hemipterans in Bt cotton.

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Eggplant or brinjal (Solanum melongena) is a popular vegetable grown throughout Asia where it is attacked by brinjal fruit and shoot borer (BFSB) (Leucinodes orbonalis). Yield losses in Bangladesh have been reported up to 86% and farmers rely primarily on frequent insecticide applications to reduce injury. Bangladesh has developed and released four brinjal varieties producing Cry1Ac (Bt brinjal) and is the first country to do so. We report on the first replicated field trials comparing four Bt brinjal varieties to their non-Bt isolines, with and without standard insecticide spray regimes. Results of the two-year study (2016-17) indicated Bt varieties had increased fruit production and minimal BFSB fruit infestation compared with their respective non-Bt isolines. Fruit infestation for Bt varieties varied from 0-2.27% in 2016, 0% in 2017, and was not significantly affected by the spray regime in either year. In contrast, fruit infestation in non-Bt lines reached 36.70% in 2016 and 45.51% in 2017, even with weekly spraying. An economic analysis revealed that all Bt lines had higher gross returns than their non-Bt isolines. The non-sprayed non-Bt isolines resulted in negative returns in most cases. Maximum fruit yield was obtained from sprayed plots compared to non-sprayed plots, indicating that other insects such as whiteflies, thrips and mites can reduce plant vigor and subsequent fruit weight. Statistically similar densities of non-target arthropods, including beneficial arthropods, were generally observed in both Bt and non-Bt varieties. An additional trial that focused on a single Bt variety and its isoline provided similar results on infestation levels, with and without sprays, and similarly demonstrated higher gross returns and no significant effects on non-target arthropods. Together, these studies indicate that the four Bt brinjal lines are extremely effective at controlling BFSB in Bangladesh without affecting other arthropods, and provide greater economic returns than their non-Bt isolines.

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Conservation biological control can be an effective tactic for minimizing insect-induced damage to agricultural production. In the Arizona cotton system, a suite of generalist arthropod predators provides critical regulation of Bemisia tabaci Gennadius (MEAM1) (Hemiptera: Aleyrodidae) and other pests. Arthropod predator and B. tabaci populations were manipulated with a range of broad-spectrum and selective insecticide exclusions to vary predator to prey interactions in a 2-yr field study. Predator to prey ratios associated with B. tabaci densities near the existing action threshold were estimated for six predator species found to be negatively associated with either adult and/or large nymphs of B. tabaci [Misumenops celer (Hentz) (Araneae: Thomisidae), Drapetis nr divergens (Diptera: Empididae), Geocoris pallens Stäl (Hemiptera: Geocoridae), Orius tristicolor (White) (Hemiptera: Anthocoridae), Chrysoperla carnea s.l. (Neuroptera: Chrysopidae), and Collops spp. (Coleoptera: Melyridae)] with the first three most consistently associated with declining B. tabaci densities. Ratios ranged from 1 M. celer per 100 sweeps to 1 B. tabaci adult per leaf to 44 D. nr. divergens per 100 sweeps to 1 large nymph per leaf disk. These ratios represent biological control informed thresholds that might serve as simple-to-use decision tool for reducing risk in the current B. tabaci integrated pest management strategy. The identification of key predators within the large, flexible food web of the cotton agro-ecosystem and estimation of predator to B. tabaci ratios clarifies the role of key predators in B. tabaci suppression, yielding potential decision-making advantages that could contribute to further improving economic and environmental sustainability of insect management in the cotton system.

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Cabbage looper, Trichoplusia ni (Hübner) is an important lepidopteran pest on many vegetable and greenhouse crops, and some field crops. Although there are no commercial transgenic Bt vegetable or greenhouse crops, T. ni is a target of Bollgard II cotton, which produces Cry1Ac and Cry2Ab. We expand on previous work that examined the effect of Bt crops on parasitoids using Bt-resistant lepidopteran populations as hosts. Cry1Ac/Cry2Ab-resistant T. ni larvae were used to eliminate host quality effects and to evaluate the direct effects of Bt cotton on the parasitoids Copidosoma floridanum (Ashmead) and Cotesia marginiventris (Cresson). These tri-trophic studies confirm that Bt cotton had no significant impact on development, success of parasitism, survival and adult longevity of C. marginiventris when using Bt-resistant T. ni fed on Bt cotton. Similarly, this Bt cotton had no significant impact on the development, mummy weight and the number of progeny produced by C. floridanum. Our studies verified that lyophilized Bt crop tissue maintained its insecticidal bioactivity when incorporated into an artificial diet, demonstrating that hosts and parasitoids were exposed to active Cry proteins. The egg-larval parasitoid C. floridanum, or similar species that consume their entire host, should be considered useful surrogates in risk assessment of Bt crops to non-target arthropods.

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Life tables provide a means of measuring the schedules of birth and death from populations over time. They also can be used to quantify the sources and rates of mortality in populations, which has a variety of applications in ecology, including agricultural ecosystems. Horizontal, or cohort-based, life tables provide for the most direct and accurate method of quantifying vital population rates because they follow a group of individuals in a population from birth to death. Here, protocols are presented for conducting and analyzing cohort-based life tables in the field that takes advantage of the sessile nature of the immature life stages of a global insect pest, Bemisia tabaci. Individual insects are located on the underside of cotton leaves and are marked by drawing a small circle around the insect with a non-toxic pen. This insect can then be observed repeatedly over time with the aid of hand lenses to measure development from one stage to the next and to identify stage-specific causes of death associated with natural and introduced mortality forces. Analyses explain how to correctly measure multiple mortality forces that act contemporaneously within each stage and how to use such data to provide meaningful population dynamic metrics. The method does not directly account for adult survival and reproduction, which limits inference to dynamics of immature stages. An example is presented that focused on measuring the impact of bottom-up (plant quality) and top-down (natural enemies) effects on the mortality dynamics of B. tabaci in the cotton system.

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In its defense against herbivores, cotton (Gossypium sp.) relies in part on the production of a set of inducible, non-volatile terpenoids. Under uniform damage levels, in planta allocation of induced cotton terpenoids has been found to be highest in youngest leaves, supporting assumptions of the optimal defense theory (ODT) which predicts that plants allocate defense compounds to tissues depending on their value and the likelihood of herbivore attack. However, our knowledge is limited on how varying, and thus more realistic, damage levels might affect cotton defense organization. We hypothesized that the allocation of terpenoids and densities of terpenoid-storing glands in leaves aligns with assumptions of the ODT, even when plants are subjected to temporally, spatially and quantitatively varying caterpillar (Heliothis virescens) damage. As expected, cotton plants allocated most of their defenses to their youngest leaves regardless of damage location. However, defense induction in older leaves varied with damage location. For at least 14 days after damage treatments ended, plants reallocated defense resources from previously young leaves to newly developed leaves. Furthermore, we observed a positive hyperbolic relationship between leaf damage area and both terpenoid concentrations and gland densities, indicating that cotton plants can fine-tune defense allocation. Although it appears that factors like vascular constraints and chemical properties of individual defense compounds can affect defense levels, our results overall demonstrate that induced defense organization of cotton subjected to varying damage treatments is in alignment with key assumptions of the ODT.

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Crops producing insecticidal crystal (Cry) proteins from the bacterium, Bacillus thuringiensis (Bt), are an important tool for managing lepidopteran pests on cotton and maize. However, the effects of these Bt crops on non-target organisms, especially natural enemies that provide biological control services, are required to be addressed in an environmental risk assessment. Amblyseius andersoni (Acari: Phytoseiidae) is a cosmopolitan predator of the two-spotted spider mite, Tetranychus urticae (Acari: Tetranychidae), a significant pest of cotton and maize. Tri-trophic studies were conducted to assess the potential effects of Cry1Ac/Cry2Ab cotton and Cry1F maize on life history parameters (survival rate, development time, fecundity and egg hatching rate) of A. andersoni. We confirmed that these Bt crops have no effects on the biology of T. urticae and, in turn, that there were no differences in any of the life history parameters of A. andersoni when it fed on T. urticae feeding on Cry1Ac/Cry2Ab or non-Bt cotton and Cry1F or non-Bt maize. Use of a susceptible insect assay demonstrated that T. urticae contained biologically active Cry proteins. Cry proteins concentrations declined greatly as they moved from plants to herbivores to predators and protein concentration did not appear to be related to mite density. Free-choice experiments revealed that A. andersoni had no preference for Cry1Ac/Cry2Ab cotton or Cry1F maize-reared T. urticae compared with those reared on non-Bt cotton or maize. Collectively these results provide strong evidence that these crops can complement other integrated pest management tactics including biological control.

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Biological control is an underlying pillar of integrated pest management, yet little focus has been placed on assigning economic value to this key ecosystem service. Setting biological control on a firm economic foundation would help to broaden its utility and adoption for sustainable crop protection. Here we discuss approaches and methods available for valuation of biological control of arthropod pests by arthropod natural enemies and summarize economic evaluations in classical, augmentative, and conservation biological control. Emphasis is placed on valuation of conservation biological control, which has received little attention. We identify some of the challenges of and opportunities for applying economics to biological control to advance integrated pest management. Interaction among diverse scientists and stakeholders will be required to measure the direct and indirect costs and benefits of biological control that will allow farmers and others to internalize the benefits that incentivize and accelerate adoption for private and public good.

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A concern with Bt-transgenic insect-resistant plants is their potential to harm non-target organisms. Early studies reported that Cry1Ab-producing Bt maize and purified Cry1Ab harmed larvae of the green lacewing, Chrysoperla carnea. Although these effects could not be confirmed in subsequent studies, some authors still refer to them as evidence that Bt maize harms beneficial species. We provide a comprehensive review of the studies evaluating the effects of Bt (Cry1Ab) maize on C. carnea. The evidence indicates that this important predator is not affected by Bt maize or by the produced Cry1Ab protein. We discuss how conceptual models can assist environmental risk assessments, and we emphasize the importance of robust and reproducible studies.

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Laboratory studies were conducted to investigate tritrophic transfer of insecticidal Cry proteins from transgenic cotton to an herbivore and its predator, and to examine effects of these proteins on the predator's development, survival, and reproduction. Cry1Ac and Cry2Ab proteins from the bacterium Bacillus thuringiensis (Bt) produced in Bollgard-II (BG-II, Event 15985) cotton plants were acquired by Thrips tabaci Lindeman (Thysanoptera: Thripidae), an important sucking pest of cotton, and its generalist predator, Orius insidiosus (Say) (Hemiptera: Anthocoridae). The average protein titers in BG-II cotton leaves were 1,256 and 43,637 ng Cry1Ac and Cry2Ab per gram fresh leaf tissue, respectively. At the second trophic level, larvae of T. tabaci reared on BG-II cotton for 48-96 h had 22.1 and 2.1% of the Cry1Ac and Cry2Ab levels expressed in leaves, respectively. At the third trophic level, O. insidiosus that fed on T. tabaci larvae had 4.4 and 0.3% of the Cry1Ac and Cry2Ab protein levels, respectively, expressed in BG-II plants. O. insidiosus survivorship, time of nymphal development, adult weight, preoviposition and postoviposition periods, fecundity, and adult longevity were not adversely affected owing to consumption of T. tabaci larvae that had fed on BG-II cotton compared with non-Bt cotton. Our results indicate that O. insidiosus, a common predator of T. tabaci, is not harmed by BG-II cotton when exposed to Bt proteins through its prey. Thus, O. insidiosus can continue to provide important biological control services in the cotton ecosystem when BG-II cotton is used to control primary lepidopteran pests.

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Variation in plant quality and natural enemy abundance plays an important role in insect population dynamics. In manipulative field studies, we evaluated the impact of varying irrigation levels and insecticide type on densities of Lygus hesperus Knight and the arthropod predator community in cotton. Three watering levels were established via irrigations timed according to three levels of percent soil water depletion (SWD): 20, 40, or 60, where 40% SWD is considered standard grower practice, 60% represents a deficit condition likely to impose plant productivity losses, and 20% represents surplus conditions with likely consequences on excessive vegetative plant production. The two key L. hesperus insecticides used were the broad-spectrum insecticide acephate and the selective insecticide flonicamid, along with an untreated check. We hypothesized that densities of L. hesperus and its associated predators would be elevated at higher irrigation levels and that insecticides would differentially impact L. hesperus and predator dynamics depending on their selectivity. L. hesperus were more abundant at the higher irrigation level (20% SWD) but the predator densities were unaffected by irrigation levels. Both L. hesperus and its predators were affected by the selectivity of the insecticide with highest L. hesperus densities and lowest predator abundance where the broad spectrum insecticide (acephate) was used. There were no direct interactions between irrigation level and insecticides, indicating that insecticide effects on L. hesperus and its predators were not influenced by the irrigation levels used here. The implications of these findings on the overall ecology of insect-plant dynamics and yield in cotton are discussed.

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Geocoris punctipes (Say) and Orius insidiosus (Say) are generalist predators found in a wide range of crops, including cotton (Gossypium hirsutum L.) and maize (Zea mays L.), where they provide important biological control services by feeding on an array of pests, including eggs and small larvae of caterpillars. A high percentage of cotton and maize in the United States and several other countries are transgenic cultivars that produce one or more of the insecticidal Cry proteins of Bacillus thuringiensis Berliner (Bt). Here we quantify effects of three Cry proteins on the life history of these predators over two generations when they are exposed to these Cry proteins indirectly through their prey. To eliminate the confounding prey quality effects that can be introduced by Bt-susceptible prey, we used Cry1Ac/Cry2Ab-resistant Trichoplusia ni (Hübner) and Cry1 F-resistant Spodoptera frugiperda (J.E. Smith) in a series of tri-trophic studies. Survival, development, adult mass, fecundity, and fertility were similar when predators consumed larvae feeding on Cry1Ac/Cry2Ab cotton or Cry1 F maize compared with prey feeding on isogenic or near-isogenic cotton or maize. Repeated exposure of the same initial cohort over a second generation also resulted in no differences in life-history traits when feeding on non-Bt- or Bt-fed prey. Enzyme-linked immunosorbent assay showed that predators were exposed to Bt Cry proteins from their prey and that these proteins became increasingly diluted as they moved up the food chain. Results show a clear lack of effect of three common and widespread Cry proteins on these two important predator species. The use of resistant insects to eliminate prey quality effects provides a robust and meaningful assessment of exposure and hazard.

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