More intraguild prey than pest species in arachnid diets may compromise biological control in apple orchards

Understanding the full diet of natural enemies is necessary for evaluating their role as biocontrol agents, because many enemy species do not only feed on pests but also on other natural enemies. Such intraguild predation can compromise pest control if the consumed enemies are actually better for pest control than their predators. In this study, we used gut metabarcoding to quantify diets of all common arachnid species in Swedish and Spanish apple orchards. For this purpose, we designed new primers that reduce amplification of arachnid predators while retaining high amplification of all prey groups. Results suggest that most arachnids consume a large range of putative pest species on apple but also a high proportion of other natural enemies, where the latter constitute almost a third of all prey sequences. Intraguild predation also varied between regions, with a larger content of heteropteran bugs in arachnid guts from Spanish orchards, but not between orchard types. There was also a tendency for cursorial spiders to have more intraguild prey in the gut than web spiders. Two groups that may be overlooked as important biocontrol agents in apple orchards seem to be theridiid web spiders and opilionids, where the latter had several small-bodied pest species in the gut. These results thus provide important guidance for what arachnid groups should be targets of management actions, even though additional information is needed to quantify all direct and indirect interactions occurring in the complex arthropod food webs in fruit orchards.     

Intraguild predation in biological control: consideration of multiple resource species

Theoretical models of intraguild predation (IGP) predict that IGP decreases the effectiveness of biological control while many empirical studies do not agree with this prediction. In this study, I discuss the importance of explicit consideration of multiple resource species that has been neglected in most theoretical IGP models. In the previous models of IGP, a single resource species represented the pest species. However, there are multiple resource species (e.g., multiple pest species or aggregates of pest and non-pest species) in real systems. This study shows that models with multiple resource species can predict a variety of outcomes including those consistent with the empirical observations. The explicit consideration of resource species is useful for the future development of theories in biological control. Handling Editor: Helen Roy     

Intraguild predation of the aphid parasitoid Aphelinus certus by Coccinella septempunctata and Harmonia axyridis

Coincidental intraguild predation is expected to be less disruptive to biological control than omnivorous intraguild predation, and strong intraguild predation is not expected to occur in natural systems. Coincidental intraguild predation in a foodweb involving introduced pest and natural enemy species was examined to determine whether intraguild predation would be disruptive of biological control services in soybean agroecosystems. Introduced natural enemies are important regulators of soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), populations in North America. Seven-spotted lady beetles, Coccinella septempunctata L., and multicolored Asian lady beetles, Harmonia axyridis Pallas (Coleoptera: Coccinellidae), are key predators of soybean aphid in North America while the chalcidoid wasp, Aphelinus certus Yasnosh (Hymenoptera: Aphelinidae), is the most common parasitoid of soybean aphid in Ontario, Canada. Predation of parasitized soybean aphids at two stages (newly parasitized aphids and mummified aphids) by adults and third instar larvae of both C. septempunctata and H. axyridis was examined in laboratory experiments. In choice experiments, all stages of lady beetles preferred non-parasitized aphids over mummified aphids. In cage experiments, third instar larvae and male and female adults of both lady beetles did not discriminate between newly parasitized and non-parasitized aphids. The influence of coincidental intraguild predation on the efficacy of parasitoids as biological control agents, and implications for soybean aphid management decisions based on natural enemies, are discussed.     

Weeds mediate the level of intraguild predation in arthropod food webs

Intraguild predation, which is common for generalist predators, is a specific form of omnivory that may suppress the biological control of a pest. The dietary flexibility of a given organism depends on the choice of the C3 (banana crop) and the C4 (weeds) pathways they use and on the trophic level on which they feed. Understanding the conditions in which intraguild predation decreases biological control is a major issue in agroecosystems. We tested whether the contribution of different primary producer pathways in diets of generalist predators mediates the level of intraguild predation. We studied 10 agroecosystems in which banana plants (C3 metabolism) were diversely associated with weeds (C4 metabolism). Diversity in litter macrofauna was relatively low, with a mean between three and eight species per trap. Measurement of stable isotopes showed a significant decrease in the δ¹⁵N values of generalist predators when the C4 pathway contributed more than the C3 pathway to their diet. We rejected hypotheses that an increase in the abundance of prey and that a decrease in prey's δ¹⁵N values occur when the C4 pathway contributes more than the C3 pathway to their diet. The results are consistent with the diet modification hypothesis, that is, intraguild predation is lower when the C4 (weeds) pathway is preferred to the C3 pathway. Our results suggest that when the C4 pathway of weeds is more exploited by herbivores (or detritivores), generalist predators tend to consume these herbivores and thus neglect the intraguild prey. The diverse C4 plant community probably supports a diverse herbivore community that provides alternative prey. Our results provide evidence that increasing plant diversity in agroecosystems should decrease intraguild predation of generalist predators and should therefore improve pest regulation. In an applied perspective, plant diversity could be increased by establishing a more diverse cover‐crop community.     

Problems Inherent to Augmentation of Natural Enemies in Open Agriculture

Augmentation biological control has successfully replaced a lot of insecticide use in ‘closed system’ agriculture (e.g., greenhouses). The profitable commercialization of biocontrol agents in greenhouses has created an incentive to expand markets for mass-reared beneficial insects into open agricultural systems, often without sufficient scientific justification. However, the semi-contained nature of greenhouse culture is often critical to the success of augmentation and can serve to mask potential pitfalls and intrinsic limitations of this approach in open systems. Factors contributing to greenhouse successes include the reduced biological diversity of contained agroecosystems, the prevention of agent dispersal, the ability to maintain environmental conditions within a range favorable for the agent, the exclusion of competitors and natural enemies of the agent that might otherwise diminish its efficacy, and the absence of alternative prey/hosts that could divert predation/parasitism from the target pest. There are also problems arising from collection of source material from locally adapted populations, and the inadvertent imposition of artificial selection in the course of laboratory rearing. Besides highlighting these pitfalls, this paper aims to encourage more consideration of conservation approaches prior to investment in augmentation programs which entice farmers into perpetual cycles of ‘rear and release.’ I argue that although augmentation can benefit agriculture whenever it replaces pesticide applications, it does not constitute an ecologically sustainable solution because it requires continued inputs, and it can distract research attention away from more sustainable objectives. Sustainable biological control is best achieved through modifications to cultural practices that increasingly ‘naturalize’ agroecosystems, thus facilitating the natural recruitment and persistence of beneficial arthropod fauna, combined with habitat management geared to increasing overall plant and arthropod diversity in the agroecosystem.     

Trophic supplements to intraguild predation

Intraguild predation (IGP) is a dominant community module in terrestrial food webs that occurs when multiple consumers feed both on each other and on a shared prey. This specific form of omnivory is common in terrestrial communities and is of particular interest for conservation biology and biological control given its potential to disrupt management of threatened or pest species. Extensive theory exists to describe the dynamics of three-species IGP, but these models have largely overlooked the potential for other, exterior interactions, to alter the dynamics within the IGP module. We investigated how three forms of feeding outside of the IGP module by intraguild predators (i.e. trophic supplementation) affect the dynamics of the predators (both IG predator and IG prey) and their shared resource. Specifically, we examined how the provision of a constant donor-controlled resource, the availability of an alternative prey species, and predator plant-feeding affect the dynamics of IGP models. All three forms of trophic supplements modified the basic expectations of IGP theory in two important ways, and their effects were similar. First, coexistence was possible without the IG prey being a superior competitor for the original shared resource if the IG prey could effectively exploit one of the types of trophic supplements. However, supplements to the IG predator restricted the potential for coexistence. Second, supplements to the IG prey ameliorated the disruptive effects of the IG predator on the suppression of the shared resource, promoting effective control of the resource in the presence of both predators. Consideration of these three forms of trophic supplementation, all well documented in natural communities, adds substantial realism and predictive power to intraguild predation theory.     

Long handling times of Praon unicum by Harmonia axyridis result in protection from predation

The intraguild predator Harmonia axyridis has a longer handling time of Praon unicum mummies in contrast to Aphidius matricariae mummies and the pest aphid Myzus persicae. In addition, the rejection rate of P. unicum is higher as compared to the A. matricariae and M. persicae. Harmonia Axyridis also has a shorter residence times when foraging in P. unicum patches. The longer handling time may provide P. unicum with a refuge from intraguild predation by H. axyridis. Thus, Praon unicum could be a better biocontrol agent than A. matricariae in the presence of intraguild predation, as it will face lower predation rates.     

Comparison of predator–prey interactions with and without intraguild predation by manipulation of the nitrogen source

Theory predicts that intraguild predation leads to different community dynamics than the trophic cascades of a linear food chain. However, experimental comparisons of these two food‐web modules are rare. Mixotrophic plankton species combine photoautotrophic and heterotrophic nutrition by grazing upon other phytoplankton species. We found that the mixotrophic chrysophyte Ochromonas can grow autotrophically on ammonium, but not on nitrate. This offered a unique opportunity to compare predator–prey interactions in the presence and absence of intraguild predation, without changing the species composition of the community. With ammonium as nitrogen source, Ochromonas can compete with its autotrophic prey for nitrogen and therefore acts as intraguild predator. With nitrate, Ochromonas acts solely as predator, and is not in competition with its prey for nitrogen. We parameterized a simple intraguild predation model based on chemostat experiments with monocultures of Ochromonas and the toxic cyanobacterium Microcystis. Subsequently, we tested the model predictions by inoculating Ochromonas into the Microcystis monocultures, and vice versa. The results showed that Microcystis was a better competitor for ammonium than Ochromonas. In agreement with theoretical predictions, Microcystis was much more strongly suppressed by intraguild predation on ammonium than by top–down predation on nitrate. Yet, Microcystis persisted at very low population densities, because the type III functional response of Ochromonas implied that the grazing pressure upon Microcystis became low when Microcystis was rare. Our results provide experimental support for intraguild predation theory, and indicate that intraguild predation may enable biological control of microbial pest species.     

有益真绥螨与巴氏新小绥螨的集团内捕食和同类相残作用

【目的】有益真绥螨Euseius utilis是北方地区广泛分布的一种多食性植绥螨,而巴氏新小绥螨Neoseiulus barkeri目前在我国广泛应用于农业害虫的生物防治中。本研究旨在对巴氏新小绥螨在本地的应用进行风险评估及为与有益真绥螨的联合释放提供依据。【方法】本研究在室内通过一系列实验,比较了实验室饲养的巴氏新小绥螨和采自内蒙古农业大学校园的有益真绥螨的雌成螨对同种或异种未成熟螨的捕食量、存活时间及产卵量的差异,检测了有益真绥螨与巴氏新小绥螨两种植绥螨相互之间的攻击强度以及种内和种间的相互作用。【结果】两种植绥螨都难以刺吸同种或异种植绥螨的卵,而对同种或异种植绥螨幼螨的捕食量最大,其次是对若螨。在无共同食物的情况下,有益真绥螨雌成螨对同种植绥螨幼螨的攻击性比对异种植绥螨幼螨的攻击性强(BreslowDay检验:χ~2=13.84,df=1,P0.001),且有益真绥螨对同种植绥螨幼螨的捕食量(9.10±1.65头)高于对异种植绥螨幼螨的捕食量(5.31±1.43头)(T检验:t=5.487,P0.001),巴氏新小绥螨对异种植绥螨幼螨的捕食量(7.48±0.75头)高于对同种植绥螨幼螨的捕食量(4.75±0.58头)(T检验:t=9.110,P0.05)。【结论】有益真绥螨更倾向于捕食同种幼螨而发生同类相残;巴氏新小绥螨更倾向于捕食异种幼螨而发生集团内捕食。     

Optimal intraguild foraging and population stability

This article explores effects of adaptive intraguild predation on species coexistence and community structure in three species' food webs. Two Lotka-Volterra models that assume a trade-off between competition and predation strength are considered in detail. The first model does not explicitly model resource dynamics and is considered with both nonadaptive and adaptive intraguild predation; in the latter case predators choose their diet in order to maximize their instantaneous population growth rate. The second model includes resource population dynamics. Effects of adaptive intraguild predation on the community structure along a gradient in environment productivity are analyzed and compared with some experimental results of protist food webs. Conditions under which intraguild predation is adaptive are discussed for both models. It is proved that if intraguild predators are perfect optimizers then intraguild predation should decrease with increasing environmental productivity and adaptive intraguild predation is a stabilizing factor provided environmental productivity is high enough.     

Ecology and biological control application of multicoloured Asian ladybird, Harmonia axyridis: A review

The ecology of and biological control by multicoloured Asian ladybird beetle, Harmonia axyridis (Pallas) are reviewed. Our emphasis is on assembling and interpreting information on general characteristics, invasion and establishment, sexual activity, foraging and predation, development, survival and reproduction, predator-predator interactions, natural enemies, biocontrol, non-target effects and status of H. axyridis as a pest of fruits. Colonization of H. axyridis for aphid biocontrol in the USA have been successful in terms of its establishment, but its abundance is turning out to be a nuisance to humans. Its harmful non-target impact on beneficial organisms, humans and native species is becoming a debatable issue. The question on its present position, whether it is a biocontrol agent or pest, is a critical issue and discussed. Inferences from the empirical data are made and new avenues for future research are suggested.     

Intraguild predation promotes complex alternative states along a productivity gradient

Intraguild predation is the simplest, ubiquitous form of trophic omnivory, known to greatly influence the structure and functioning of natural and managed food webs. Although alternative states are fundamental to intraguild predation dynamics, only necessary conditions for alternative states have been previously reported. Using simple models, we found complex but systematic patterns in which different alternative states occur along a productivity gradient, and clarified the sufficient conditions to separate these patterns. We found that two quantities known to control the necessary conditions also determine the sufficient conditions: (1) relative energy transfer efficiency through alternative trophic pathways to an intraguild predator, and (2) relative resource exploitation ability between intraguild prey and predator. These governing quantities suggest how body size and stoichiometric relations between intraguild prey and predators can influence the possibility of alternative states. Our results indicate that food webs involving intraguild predation have a high potential of complex alternative states, and their management can be highly precarious.     

Species complementarity in two myrmecophilous lady beetle species in a coffee agroecosystem: implications for biological control

Natural enemy diversity may be beneficial, through species complementarity, or detrimental, through antagonistic interactions, such as competition or intraguild predation, for the biological control of agricultural pests. We studied two coexisting myrmecophilous coccinellid beetles, Azya orbigera (Mulsant) (Coleoptera: Coccinellidae) and an undescribed species in the genus Diomus (Coleoptera: Coccinellidae), in a coffee agroecosystem in Chiapas, Mexico. As both beetles specialize on the same prey, the green coffee scale pest, Coccus viridis (Green) (Hemiptera: Coccidae), we studied the beetles’ behavior and distribution to determine if they niche partition in order to avoid extreme competition. Through field surveys and lab experiments we detected spatial segregation but not resource partitioning among A. orbigera and Diomus sp. We posit that the presence of both species can lead to improved biocontrol of C. viridis populations through species complementarity. Our work supports the growing evidence that natural enemy diversity can provide enhanced conservation biological control.     

Tracking the role of alternative prey in soybean aphid predation by Orius insidiosus: a molecular approach

The soybean aphid, Aphis glycines (Hemiptera: Aphididae), is a pest of soybeans in Asia, and in recent years has caused extensive damage to soybeans in North America. Within these agroecosystems, generalist predators form an important component of the assemblage of natural enemies, and can exert significant pressure on prey populations. These food webs are complex and molecular gut-content analyses offer nondisruptive approaches for examining trophic linkages in the field. We describe the development of a molecular detection system to examine the feeding behaviour of Orius insidiosus (Hemiptera: Anthocoridae) upon soybean aphids, an alternative prey item, Neohydatothrips variabilis (Thysanoptera: Thripidae), and an intraguild prey species, Harmonia axyridis (Coleoptera: Coccinellidae). Specific primer pairs were designed to target prey and were used to examine key trophic connections within this soybean food web. In total, 32% of O. insidiosus were found to have preyed upon A. glycines, but disproportionately high consumption occurred early in the season, when aphid densities were low. The intensity of early season predation indicates that O. insidiosus are important biological control agents of A. glycines, although data suggest that N. variabilis constitute a significant proportion of the diet of these generalist predators. No Orius were found to contain DNA of H. axyridis, suggesting intraguild predation upon these important late-season predators during 2005 was low. In their entirety, these results implicate O. insidiosus as a valuable natural enemy of A. glycines in this soybean agroecosystem.     

Intraguild predation of parasitoids by Solenopsis invicta: a non-disruptive interaction

The outcome of intraguild predation among natural enemies can have significant ramifications for herbivore suppression and biological control. Manipulating habitat complexity may alter the strength of intraguild predation, since changes in habitat complexity are often associated with concomitant changes in natural enemy abundance. Using a combination of greenhouse and field experiments, we determined if asymmetric intraguild predation by a pervasive generalist predator, the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), disrupts important parasitoids in a collard, Brassica oleracea L. (Brassicaceae), agroecosystem. The effect of habitat complexity on this interaction was assessed by conducting field experiments in a simple, collard monocrop and a more complex, collard‐white clover intercrop. Neither the density of adult parasitoids nor the percentage parasitism of caterpillars was affected when S. invicta abundance was manipulated. Solenopsis invicta reduced the survival of the diamondback moth larvae, Plutella xylostella (L.) (Lepidoptera: Plutellidae) by 26% and 42% in greenhouse and field experiments, respectively, but there was no preference of S. invicta for parasitized or unparasitized caterpillars. An increase in habitat complexity significantly affected the abundance of both S. invicta and parasitoids, but had no overall effect on their interaction. The results of this study suggest that although S. invicta is an intraguild predator of parasitoids because it preys upon parasitized caterpillars, the action of S. invicta may not compromise overall biological control. This study's findings are important, because they suggest that the presence and conservation of multiple natural enemies may result in sustained pest suppression in agroecosystems, even complex systems containing many species of natural enemies including strong intraguild predators such as S. invicta.     

Invasion theory and biological control

Recent advances in the mathematical theory of invasion dynamics have much to offer to biological control. Here we synthesize several results concerning the spatiotemporal dynamics that occur when a biocontrol agent spreads into a population of an invading pest species. We outline conditions under which specialist and generalist predators can influence the density and rate of spatial spread of the pest, including the rather stringent conditions under which a specialist predator can successfully reverse a pest invasion. We next discuss the connections between long distance dispersal and invasive spread, emphasizing the different consequences of fast spreading pests and predators. Recent theory has considered the effects of population stage-structure on invasion dynamics, and we discuss how population demography affects the biological control of invading pests. Because low population densities generally characterize early stages of an invasion, we discuss the lessons invasion theory teaches concerning the detectability of invasions. Stochasticity and density-dependent dynamics are common features of many real invasions, influencing both the spatial character (e.g. patchiness) of pest invasions and the success of biocontrol agents. We conclude by outlining theoretical results delineating how stochastic effects and complex dynamics generated by density dependence can facilitate or impede biological pest control.     

Mathematical modeling of the population dynamics of a distinct interactions type system with local dispersal

Distinct biotic interactions in multi-species communities are a ubiquitous force in the natural ecosystem, and this force is an essential determinant of community stability and species coexistence outcomes. We conduct numerical simulations and bifurcation analysis of partial differential equations to gain better understanding and ecological insights into how predation (a), predator handling time (h), and local dispersal affect multi-species community dynamics. This system consists of resource-mutualist-exploiter-competitor interactions and local dispersal. From the inspection of our numerical simulations and co-dimension one bifurcation analysis findings, we discover several critical values that correspond to transcritical bifurcation, subcritical and supercritical Hopf bifurcations. This occurs as we vary the bifurcation parameters a and h in this complex ecological system under symmetric and asymmetric dispersal scenarios. Furthermore, the interplay between these local bifurcation points results in an exciting co-dimension two bifurcations, i.e., Bogdanov-Takens and cusp bifurcation points, respectively, which act as the synchronization points in this complex ecological system. From an ecological viewpoint, we find that (i) the effect of the no-dispersal scenario supports the maintenance of species biodiversity when the predation strength is moderate; (ii) symmetric dispersal induces both subcritical and supercritical Hopf bifurcation and support species diversity for moderate predation strength; and (iii) asymmetric dispersal promotes species diversity as it simplifies the bifurcation changes in dynamics by eliminating the subcritical bifurcations that trigger uncertainty, and this dispersal mechanism mediates species coexistence outcomes. Fundamentally, stable limit cycles have been reported as predator handling time varies in some ecological models; however, we observed in our bifurcation analysis the emergence of the unstable limit cycle as predator handling time changes. We discover that intense predator handling time destabilizes this complex ecological community. In general, our results demonstrate the influential roles of predation, predator handling time, and local dispersal in determining this system’s coexistence dynamics. This knowledge provides a better understanding of species conservation and biological control management.     

Food supplementation to optimize inoculative release of the predatory bug Macrolophus pygmaeus in sweet pepper

Biological control is widespread in management of greenhouse sweet pepper crops. Several species of predatory mites, bugs, and parasitoids are used against a wide range of pest species. However, biological control of particular pests like aphids, caterpillars, and the tobacco whitefly, Bemisia tabaci Gennadius, remains problematic. Macrolophus pygmaeus Rambur (Hemiptera: Miridae) is a generalist predatory bug which is used on a large scale in Western European tomato greenhouses. It has already been demonstrated that M. pygmaeus is a valuable biocontrol option in sweet pepper crops, but it has yet to find its way into common practice. Macrolophus pygmaeus should be introduced at the start of the growing season and determining an optimal release strategy is a key step in this process. In tomato crops, M. pygmaeus requires supplemental food releases to reach sufficient population numbers and dispersal levels. In this study, the need for food supplementation in sweet pepper is investigated. Three strategies were tested: (1) no food supplementation, (2) local food supplementation, and (3) full field food supplementation. Both population numbers and dispersal rates of the second generation were higher under the third strategy. Macrolophus pygmaeus oviposits near food sources, therefore dispersal rates are higher when food is more spread out. Pest control was achieved in all treatments, but faster and at lower pest levels under the full field strategy.     

The fear diet: Risk,refuge, and biological control by omnivorous weed seed predators

Weed seed biocontrol by omnivorous mice and insects can limit weed seedbanks, but this ecosystem service can be difficult to predict given the broad diet breadth of seed predators and their potential for intraguild predation. Seed foraging behavior is further modified by fluctuating cues of predation risk from higher trophic levels and the availability of refuge habitat. Uncertainty about whether co-occurring insects and mice additively contribute to weed biocontrol or interfere with each other via intraguild predation limits our ability to recommend habitat management strategies that reliably promote seed destruction. Using seed removal assays, fluorescent powder tracking, and stable isotope analyses, we assessed effects of a predation risk cue (moonlight) on mouse foraging patterns in a patchwork of vegetated and exposed plots in a cultivated field. Mouse foraging activity decreased on exposed ground during the full moon, compared to dark nights, yet foraging movements were unaffected by moon cycle within refuge patches. Weed seed consumption was more than three times higher in cover than exposed soil, and 78% of that difference was attributable to invertebrate granivores. Mice and invertebrate granivores both exhibited higher foraging activity in cover, indicating co-occurrence of intraguild predators and prey. However, stable isotope analyses of fecal samples revealed that mice captured in refuge habitats fed at slightly lower trophic levels than those in exposed habitats (suggesting minimal intraguild predation in refuge habitat), and mouse diet was unaffected by moonlight. Despite increased availability of invertebrate prey in cover patches, mice do not appear to preferentially exploit prey when avoiding their own predators or interfere with weed seed predation. Therefore, functional redundancy of mice and invertebrate seed predators in cover crops and other refuge habitats may strengthen and stabilize weed seed biocontrol.     

Biological control of aphids in the presence of thrips and their enemies

Generalist predators are often used in biological control programs, although they can be detrimental for pest control through interference with other natural enemies. Here, we assess the effects of generalist natural enemies on the control of two major pest species in sweet pepper: the green peach aphid Myzus persicae (Sulzer) and the western flower thrips Frankliniella occidentalis (Pergande). In greenhouses, two commonly used specialist natural enemies of aphids, the parasitoid Aphidius colemani Viereck and the predatory midge Aphidoletes aphidimyza (Rondani), were released together with either Neoseiulus cucumeris Oudemans, a predator of thrips and a hyperpredator of A. aphidimyza, or Orius majusculus (Reuter), a predator of thrips and aphids and intraguild predator of both specialist natural enemies. The combined use of O. majusculus, predatory midges and parasitoids clearly enhanced the suppression of aphids and consequently decreased the number of honeydew-contaminated fruits. Although intraguild predation by O. majusculus on predatory midges and parasitoids will have affected control of aphids negatively, this was apparently offset by the consumption of aphids by O. majusculus. In contrast, the hyperpredator N. cucumeris does not prey upon aphids, but seemed to release aphids from control by consuming eggs of the midge. Both N. cucumeris and O. majusculus did not affect rates of aphid parasitism by A. colemani. Thrips were also controlled effectively by O. majusculus. A laboratory experiment showed that adult predatory bugs feed on thrips as well as aphids and have no clear preference. Thus, the presence of thrips probably promoted the establishment of the predatory bugs and thereby the control of aphids. Our study shows that intraguild predation, which is potentially negative for biological control, may be more than compensated by positive effects of generalist predators, such as the control of multiple pests, and the establishment of natural enemies prior to pest invasions. Future work on biological control should focus on the impact of species interactions in communities of herbivorous arthropods and their enemies.