The impact of secondary pests on Bacillus thuringiensis (Bt) crops

The intensification of agriculture and the development of synthetic insecticides enabled worldwide grain production to more than double in the last third of the 20th century. However, the heavy dependence and, in some cases, overuse of insecticides has been responsible for negative environmental and ecological impacts across the globe, such as a reduction in biodiversity, insect resistance to insecticides, negative effects on nontarget species (e.g. natural enemies) and the development of secondary pests. The use of recombinant DNA technology to develop genetically engineered insect‐resistant crops could mitigate many of the negative side effects of insecticides. One such genetic alteration enables crops to express toxic crystalline (Cry) proteins from the soil bacteria Bacillus thuringiensis (Bt). Despite the widespread adoption of Bt crops, there are still a range of unanswered questions concerning longer term agro‐ecosystem interactions. For instance, insect species that are not susceptible to the expressed toxin can develop into secondary pests and cause significant damage to the crop. Here, we review the main causes surrounding secondary pest dynamics in Bt crops and the impact of such outbreaks. Regardless of the causes, if nonsusceptible secondary pest populations exceed economic thresholds, insecticide spraying could become the immediate solution at farmers’ disposal, and the sustainable use of this genetic modification technology may be in jeopardy. Based on the literature, recommendations for future research are outlined that will help to improve the knowledge of the possible long‐term ecological trophic interactions of employing this technology.     

Temporal variation of the effects of landscape composition on lacewings (Chrysopidae: Neuroptera) in vineyards

1. Increasing landscape heterogeneity, both in terms of composition and configuration, can promote natural enemies and biological control in agricultural landscapes. However, relatively poor information exists about the effects of landscape heterogeneity on lacewings, which are a major group of predators. Furthermore, temporal changes of landscape effects on natural enemy dynamics remain largely unexplored.
2. Here, we investigated how landscape composition and configuration affect lacewings and their biological control potential on leafhoppers. Lacewings and leafhoppers were sampled from April to July in 10 vineyards located in southwestern France. The vineyards were selected along a gradient of a proportion of semi‐natural habitats in the landscape.
3. The proportion of semi‐natural habitats positively affected the abundance of adults and eggs, as well as species richness, of lacewings, alone or in interaction with the sampling month. Landscape configuration was never found to enhance abundance or species richness of lacewings. Finally, the predator–prey ratio increased through time but did not respond to landscape composition or configuration.
4. Our study highlights that the proportion of semi‐natural habitats increases both abundance and diversity of lacewings in vineyard landscapes but that this effect varies over time. This result indicates the need to assess the variability of landscape effects over time to maximize biological pest control services in agricultural landscapes.

     

Reduced-risk insecticides for control of grape berry moth (Lepidoptera: Tortricidae) and conservation of natural enemies

A 3-yr field study was conducted at commercial grape (Vitis spp.) farms to evaluate insect management programs for control of the grape berry moth, Paralobesia viteana Clemens (Lepidoptera: Tortricidae) and conservation of natural enemies. At each farm, one vineyard received only reduced-risk insecticides for control of second and third generation P. viteana, whereas the comparison vineyard received conventional insecticides. Both vineyards received a conventional insecticide application for control of first generation P. viteana and other insect pests. Monitoring with pheromone traps showed no differences between programs in the total number of adult male moths trapped in vineyards, and oviposition by P. viteana was similar between the two programs in all 3 yr. During weekly samples of crop infestation, both programs had a similar percentage of clusters infested by P. viteana larvae. Berries infested by P. viteana were collected from vineyard borders during the second and third P. viteana generations and held under controlled conditions. In eight of the nine berry samples, survival of larvae was significantly lower in berries collected from vineyards managed under the reduced-risk insecticide program compared with the conventional program. Parasitism of P. citeana larvae in these samples was not consistently different between the two insecticide programs over 3 yr, and similar captures of natural enemies were found on yellow sticky traps in the two programs throughout the study. Our results indicate that integrated pest management programs incorporating reduced-risk insecticides for control of P. viteana can obtain similar or greater control of P. viteana compared with programs based solely on conventional insecticides, but they may not lead to measurable long-term increases in parasitism of P. viteana.     

Response of wild bee diversity,abundance, and functional traits to vineyard inter‐row management intensity and landscape diversity across Europe

Agricultural intensification is a major driver of wild bee decline. Vineyards may be inhabited by plant and animal species, especially when the inter‐row space is vegetated with spontaneous vegetation or cover crops. Wild bees depend on floral resources and suitable nesting sites which may be found in vineyard inter‐rows or in viticultural landscapes. Inter‐row vegetation is managed by mulching, tillage, and/or herbicide application and results in habitat degradation when applied intensively. Here, we hypothesize that lower vegetation management intensities, higher floral resources, and landscape diversity affect wild bee diversity and abundance dependent on their functional traits. We sampled wild bees semi‐quantitatively in 63 vineyards representing different vegetation management intensities across Europe in 2016. A proxy for floral resource availability was based on visual flower cover estimations. Management intensity was assessed by vegetation cover (%) twice a year per vineyard. The Shannon Landscape Diversity Index was used as a proxy for landscape diversity within a 750 m radius around each vineyard center point. Wild bee communities were clustered by country. At the country level, between 20 and 64 wild bee species were identified. Increased floral resource availability and extensive vegetation management both affected wild bee diversity and abundance in vineyards strongly positively. Increased landscape diversity had a small positive effect on wild bee diversity but compensated for the negative effect of low floral resource availability by increasing eusocial bee abundance. We conclude that wild bee diversity and abundance in vineyards is efficiently promoted by increasing floral resources and reducing vegetation management frequency. High landscape diversity further compensates for low floral resources in vineyards and increases pollinating insect abundance in viticulture landscapes.     

Insecticide‐induced hormesis in an insecticide‐resistant strain of the maize weevil,Sitophilus zeamais

Sublethal responses to insecticides are frequently neglected in studies of insecticide resistance, although stimulatory effects associated with low doses of compounds toxic at higher doses, such as insecticides, have been recognized as a general toxicological phenomenon. Evidence for this biphasic dose–response relationship, or hormesis, was recognized as one of the potential causes underlying pest resurgence and secondary pest outbreaks. Hormesis has also potentially important implications for managing insecticide‐resistant populations of insect‐pest species, but evidence of its occurrence in such context is lacking and fitness parameters are seldom considered in these studies. Here, we reported the stimulatory effect of sublethal doses of the pyrethroid insecticide deltamethrin sprayed on maize grains infested with a pyrethroid‐resistant strain of the maize weevil (Sitophilus zeamais) (Coleoptera: Curculionidae). The parameters estimated from the fertility tables of resistant insects exposed to deltamethrin indicated a peak in the net reproductive rate at 0.05 ppm consequently leading to a peak in the intrinsic rate of population growth at this dose. The phenomenon is consistent with insecticide‐induced hormesis and its potential management implications are discussed.     

Landscape genomics of Colorado potato beetle provides evidence of polygenic adaptation to insecticides

The ability of insect pests to rapidly and repeatedly adapt to insecticides has long challenged entomologists and evolutionary biologists. Since Crow's seminal paper on insecticide resistance in 1957, new data and insights continue to emerge that are relevant to the old questions about how insecticide resistance evolves: such as whether it is predominantly mono‐ or polygenic, and evolving from standing vs. de novo genetic variation. Many studies support the monogenic hypothesis, and current management recommendations assume single‐ or two‐locus models. But inferences could be improved by integrating data from a broader sample of pest populations and genomes. Here, we generate evidence relevant to these questions by applying a landscape genomics framework to the study of insecticide resistance in a major agricultural pest, Colorado potato beetle, Leptinotarsa decemlineata (Say). Genome–environment association tests using genomic variation from 16 populations spanning gradients of landscape variables associated with insecticide exposure over time revealed 42 strong candidate insecticide resistance genes, with potentially overlapping roles in multiple resistance mechanisms. Measurements of resistance to a widely used insecticide, imidacloprid, among 47 L. decemlineata populations revealed heterogeneity at a small (2 km) scale and no spatial signature of origin or spread throughout the landscape. Analysis of nucleotide diversity suggested candidate resistance loci have undergone varying degrees of selective sweeps, often maintaining similar levels of nucleotide diversity to neutral loci. This study suggests that many genes are involved in insecticide resistance in L. decemlineata and that resistance likely evolves from both de novo and standing genetic variation.     

Susceptibility of Chrysochus auratus,a natural enemy of spreading dogbane,to insecticides used in wild blueberry production

Insect pest management in wild blueberry (Vaccinium angustifolium Aiton) usually involves insecticidal sprays, which may have detrimental effects on non‐target beneficial insects. Dogbane beetle (Chrysochus auratus Fabricius) (Coleoptera: Chrysomelidae) feeds almost exclusively on spreading dogbane (Apocynum androsaemifolium L.), an increasing weed problem in wild blueberry production. Because C. auratus is an important natural enemy of spreading dogbane, we assessed its susceptibility to several insecticides it may be exposed to during insect pest management. In laboratory bioassays, we found adult dogbane beetles were highly susceptible to field rates of phosmet (Imidan) and acetamiprid (Assail) by direct topical contact and ingestion of treated foliage, whereas no mortality was seen with spirotetramat (Movento) and chlorantraniliprole (Altacor). Topical applications of spinetoram (Delegate) did not cause significant mortality of beetles, but high mortality to beetles was found when they ingested spinetoram‐treated foliage. The results suggest that while some insecticides used in blueberry management will be hazardous to C. auratus, options are available that will cause little harm to this natural enemy.     

A global invasion by the thrip,Frankliniella occidentalis: Current virus vector status and its management

Western flower thrip, Frankliniella occidentalis (Pergande), is among the most economically important agricultural pests globally, attacking a wide range of vegetable and horticultural crops. In addition to causing extensive crop damage, the species is notorious for vectoring destructive plant viruses, mainly belonging to the genera Orthotospovirus, Ilarvirus, Alphacarmovirus and Machlomovirus. Once infected by orthotospoviruses, thrips can remain virulent throughout their lifespan and continue transmitting viruses to host plants when and wherever they feed. These irruptive viral outbreaks in crops will permanently disrupt functional integrated pest management systems, and typically require a remedial treatment involving insecticides, contributing to further development of insecticide resistance. To mitigate against this continuing cycle, the most effective management is early and comprehensive surveillance of the pest species and recognition of plant viruses in the field. This review provides information on the pest status of F. occidentalis, discusses the current global status of the viruses vectored by this thrip species, examines the mechanisms involved in transmitting virus‐induced diseases by thrips, and reviews different management strategies, highlighting the potential management tactics developed for various cropping systems. The early surveillance and the utilization of potential methods for control of both F. occidentalis and viruses are proposed.     

Tackling diamondback moth Plutella xylostella (L.) resistance: a review on the current research on vegetable integrated pest management in Zimbabwe

The diamondback moth (DBM), Plutella xylostella (L.) is a destructive pest of brassicas globally. Control of the pest is dominated by insecticides. Studies have shown that in some African countries, there is a great reliance on broad spectrum insecticides such as pyrethroids, organophosphates and carbamates, that are applied weekly or twice per week. Use of unregistered insecticides has also been reported. The quality of insecticide application has also been reported to be poor or ineffective. It is therefore not surprising that DBM is fast developing resistance to the major insecticides used against it. Adopting an integrated pest management strategy may be a good arsenal to use against the pest.     

Coccinellid response to landscape composition and configuration

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Mitigating the effects of insecticides on arthropod biological control at field and landscape scales

Integrated pest management (IPM) programs emphasize the combination of tactics, such as chemical and biological control, to maintain pest populations below economic thresholds. Although combining tactics may provide better long-term sustainable pest suppression than one tactic alone, in many cases, insecticides and natural enemies are incompatible. Insecticides can disrupt natural enemies through lethal and sub-lethal means causing pest resurgence or secondary pest outbreaks. Legislative actions such as the Food Quality Protection Act (US) and the Directive on Sustainable Use of Pesticides (EU) have placed greater restrictions on insecticides used in agriculture, potentially enhancing biological control. Here we focus on the effects of insecticides on biological control, and potential mitigation measures that can operate at different scales. At the farm scale, natural enemies can be conserved through the use of selective insecticides, low doses, special formulations, creation of refugia, special application methods, and targeted applications (temporal or spatial). At the landscape scale, habitat quality and composition affect the magnitude of biological control services, and the degree of mitigation against the effects of pesticides on natural enemies. Current research is teasing apart the relative importance of local and landscape effects of pesticides on natural enemies and the ecosystem services they provide, and the further development of this area will ultimately inform the decisions of policy makers and land managers in terms of how to mitigate pesticide effects through habitat manipulation.     

Changes in emergence phenology,fatty acid composition,and xenobiotic-metabolizing enzyme expression is associated with increased insecticide resistance in the Colorado potato beetle

The Colorado potato beetle (Leptinotarsa decemlineata) is a major agricultural pest of solanaceous crops. An effective management strategy employed by agricultural producers to control this pest species is the use of systemic insecticides. Recent emphasis has been placed on the use of neonicotinoid insecticides. Despite efforts to curb resistance development through integrated pest management approaches, resistance to neonicotinoids in L. decemlineata populations continues to increase. One contributing factor may be alterations in insect fatty acids, which have multiple metabolic functions and are associated with the synthesis of xenobiotic-metabolizing enzymes to mitigate the effects of insecticide exposure. In this study, we analyzed the fatty acid composition of L. decemlineata populations collected from an organic production field and from a commercially managed field to determine if fatty acid composition varied between the two populations. We demonstrate that a population of L. decemlineata that has a history of systemic neonicotinoid exposure (commercially managed) has a different lipid composition and differential expression of known metabolic detoxification mechanisms relative to a population that has not been exposed to neonicotinoids (organically managed). The fatty acid data indicated an upregulation of Δ⁶ desaturase in the commercially managed L. decemlineata population and suggest a role for eicosanoids and associated metabolic enzymes as potential modulators of insecticide resistance. We further observed a pattern of delayed emergence within the commercially managed population compared with the organically managed population. Variations in emergence timing together with specific fatty acid regulation may significantly influence the capacity of L. decemlineata to develop insecticide resistance.     

Silicon‐mediated multiple interactions: Simultaneous induction of rice defense and inhibition of larval performance and insecticide tolerance of Chilo suppressalis by sodium silicate

The rice striped stem borer (SSB, Chilo suppressalis) is one of the most destructive pests of rice plants. Si‐mediated rice defense against various pests has been widely reported, and sodium silicate (SS) has been used as an effective source of silicon for application to plants. However, there is quite limited information about the direct effects of Si application on herbivorous insects. SSB larval performance and their insecticide tolerance were examined after they had been reared either on rice plants cultivated in nutrient solution containing 0.5 and 2.0 mM SS or on artificial diets with 0.1% and 0.5% SS. SS amendment in either rice culture medium or artificial diets significantly suppressed the enzymatic activities of acetylcholinesterase, glutathione S‐transferases, and levels of cytochrome P450 protein in the midgut of C. suppressalis larvae. Larvae fed on diets containing SS showed lower insecticide tolerance. Additionally, RNA‐seq analysis showed that SS‐mediated larval insecticide tolerance was closely associated with fatty acid biosynthesis and pyruvate metabolism pathways. Our results suggest that Si not only enhances plant resistance against insect herbivore, but also impairs the insect's capacity to detoxify the insecticides. This should be considered as another important aspect in Si‐mediated plant–insect interaction and may provide a novel approach of pest management.     

Cabbage whiteflies colonise Brassica vegetables primarily from distant,upwind source habitats

The occurrence of species in rapidly changing environments, such as agricultural landscapes, is affected by their ability to recolonise habitats. Knowledge of the landscape scale affecting colonisation is essential for large‐scale pest management. Colonisation by insects can be affected on multiple landscape scales, as different morphs of a species may have specific dispersal abilities. The cabbage whitefly, Aleyrodes proletella (L.) (Hemiptera: Aleyrodidae), a major pest of Brassica vegetables, is known to colonise Brassica vegetables primarily from fields of oilseed rape, Brassica napus L. (Brassicaceae). We used field mapping and remote sensing to characterise the relevant scales for colonisation of Brussels sprouts by cabbage whiteflies. Surprisingly, oilseed rape fields in wide landscapes (2–8 km around study sites) explained colonisation better than oilseed rape areas in local landscapes (200–1 000 m around study sites). The explained variance increased when additional weight was given to upwind source habitats, indicating wind transport of whitefly colonisers. Low importance of local compared to wide landscape source habitats can be explained by the flight behaviour of whitefly morphs. Migratory morphs show phototactic attraction but are attracted by hosts only during the later phases of flight. Therefore, they ignore host plants close to their origin and disperse several kilometres. Trivial flight morphs rarely move more than a few hundred metres. In conclusion, as most whitefly colonisers reached Brassica vegetables from source habitats at a distance of 2–8 km, predictions on pest pressure and landscape‐scale whitefly management should consider these distances. In contrast, oilseed rape fields in the local landscape, which usually worry farmers, had little effect.     

Oviposition site preference of Barbitistes vicetinus (Orthoptera,Tettigoniidae) during outbreaks

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Colonization of vineyards by Kampimodromus aberrans (Oudemans) (Acari: Phytoseiidae): dispersal from surrounding plants as indicated by random amplified polymorphism DNA typing

Abstract

• 1 Kampimodromus aberrans (Oudemans) is the most important predatory mite found in vineyards of southern France. This mite also occurs in surrounding uncultivated areas from where it disperses to colonize adjacent vineyards
• 2 To determine accurately origins of immigrant mites and to study their establishment in vineyards, a study using RAPD markers (Random Amplified Polymorphism DNA) was performed. Females of K. aberrans were sampled on leaves collected in an experimental vineyard, and from several adjacent areas including neighbouring vine plots and natural plants, both of which harboured high densities of the mite. Samples were taken in May and July before and after major dispersal of K. aberrans into the experimental plot occurred.
• 3 For both dates, genetic distances within population were lower than between populations and three groupings of mites were observed. Strong relationships were observed between (1) females from different parts of the same experimental vineyard (variety Cabernet‐Sauvignon), (2) females from several plants in the woody margin neighbouring this experimental plot, and (3) females from two neighbouring vineyards (variety Carignan). Populations seemed to be structured and no correlation between genetic and geographical distances was observed. Hence, definitive conclusions about origins of migrants were not possible.
• 4 Once in a vineyard, mites are probably exposed to selection pressures (i.e. pesticide applications or vine variety characteristics) that largely determine differentiation of populations. Thus, despite many immigrants moving into vineyards, our study indicated that there was limited survival or reproduction of immigrants. Further studies of within vineyard selection factors and impacts on immigrant mites are needed to determine the influence of natural colonization on grape pest management.

     

Lethal and sublethal effects of insecticides on the survival and reproduction of Brevipalpus yothersi (Acari: Tenuipalpidae)

The overuse of insecticides to control vector insects such as Diaphorina citri Kuwayama in citrus groves has altered the population dynamics of pest mites. Among phytophagous mites, population outbreaks of citrus leprosis mite, Brevipalpus yothersi Baker, have been increasingly intense and frequent in Brazilian citrus groves. Despite the great importance of the B. yothersi mite for citrus production, the lethal and sublethal effects of insecticides on this mite have not yet been studied. Therefore, in this study, the effects of insecticides commonly used for D. citri control on B. yothersi mortality, reproduction, and instantaneous growth rate were assessed. For this, two experiments were carried out, one under controlled conditions and another in a greenhouse. The insecticides tested were beta-cyfluthrin, bifenthrin, buprofezin, chlorpyrifos, dimethoate, pyriproxyfen, and thiamethoxam at 0 (control), 0.0625, 0.125, 0.25, 0.5, 1, and twofold the recommended insecticide concentration for D. citri control. The pyriproxyfen insecticide provided high mortality of B. yothersi even at low concentrations. Furthermore, this insecticide negatively interfered with the reproduction of this mite. Beta-cyfluthrin, bifenthrin, buprofezin, chlorpyrifos, dimethoate, and thiamethoxam, in the tested concentrations, showed low impact on citrus leprosis mite. Regarding the reproduction of the mite, no significant increase in fecundity was observed on B. yothersi females exposed to insecticide residues, regardless of the concentration tested. Therefore, the application of these insecticides in the management of pest insects is unlikely to promote an increase in the citrus leprosis mite population.

     

Landscape changes have greater effects than climate changes on six insect pests in China

In recent years, global changes are the major causes of frequent, widespread outbreaks of pests in mosaic landscapes, which have received substantial attention worldwide. We collected data on global changes(landscape and climate) and economic damage caused by six main insect pests during 1951–2010 in China. Landscape changes had significant effects on all six insect pests. Pest damage increased significantly with increasing arable land area in agricultural landscapes. However, climate changes had no effect on damage caused by pests, except for the rice leaf roller(Cnaphalocrocis medinalis Guenee) and armyworm(Mythimna separate(Walker)), which caused less damage to crops with increasing mean temperature. Our results indicate that there is slight evidence of possible offset effects of climate changes on the increasing damage from these two agricultural pests. Landscape changes have caused serious outbreaks of several species, which suggests the possibility of the use of landscape design for the control of pest populations through habitat rearrangement. Landscape manipulation may be used as a green method to achieve sustainable pest management with minimal use of insecticides and herbicides.     

Monitoring and mechanisms of organophosphate resistance in San Jose scale,Diaspidiotus perniciosus (Hemiptera: Diaspididae)

San Jose scale, Diaspidiotus perniciosus (Comstock), is a serious pest in Chilean apple tree orchards, and a number of organophosphate insecticides were used to control them for decades. Recently, control failures with these insecticides were reported and linked to insecticide resistance development. In this study, 40 San Jose scale field populations were collected and their susceptibility to two commonly used organophosphate insecticides, that is chlorpyrifos and methidathion, was assessed. The obtained bioassay data suggest moderate levels of resistance to both insecticides when compared to a reference susceptible strain. The highest resistance ratio (RR) detected for chlorpyrifos and methidathion was 31‐fold and 11‐fold, respectively. The bioassay results suggest the occurrence of a significant cross‐resistance between both compounds. Biochemical measurements revealed a role for esterases in conferring resistance to organophosphates, but not modified acetylcholinesterase. The spatial spread and extend of insecticide resistance were also evaluated. Our result shows that no autocorrelation can be assumed, and then, insecticide resistance is caused by random factors.     

Erratum

Abstract: Fertilization levels for ornamental crops may influence pest population dynamics, crop quality, and pest management strategy. We examined the effect of fertilization on population growth and within‐plant distribution of melon or cotton aphid, Aphis gossypii Glover, on potted chrysanthemum, Dendranthema grandiflora (Tzvelev). In terms of pest management implications, we also investigated the effect of fertilization on the number of insecticide applications needed to control A. gossypii on potted chrysanthemum. Population growth rate of A. gossypii increased with fertilization levels from 0 to 38 ppm N and reached a plateau from 38 to 488 ppm N. Increased fertilization beyond 38 ppm N, 10% of the commercial standard, did not result in higher aphid number. Aphids responded to nutrient availability of plants by distributing themselves in areas with higher level of nitrogen. More aphids were found in the apical and middle strata of the plants than the basal stratum, which had the lowest nitrogen content. Leaf nitrogen content increased with increased fertilization level and was consistently higher in the apical and middle strata than the basal stratum. Increased fertilization from 0 to 375 ppm N did not result in higher number of insecticide applications. All three insecticides (bifenthrin, kinoprene or pymetrozine) were effective in keeping the aphid infestation below a pre‐determined level, five aphids per plant, but pymetrozine required the least number of applications. For chrysanthemum, a fast‐growing crop and heavy utilizer of nitrogen, increased fertilization shortened the time to flowering, which would allow growers to harvest their crop sooner and reduce the time for aphid population growth. Reduction in time to harvest could result in significant reduction of insecticide usage by reducing the time for aphid population growth. As a result, high fertilization together with minimal runoff may be a useful tactic to an integrated pest management (IPM) programme for managing A. gossypii on potted chrysanthemums.