Ontogenetic change of prey preference in the generalist predator Zelus renardii and its influence on predator–predator interactions

1. Two field experiments were conducted to test the hypothesis that the intensity of predation by a generalist predator on two species of prey changes with the developmental stage of the predator. The generalist predator studied was Zelus renardii Kolenati (Hemiptera: Reduviidae) and the prey were the lacewing larva, Chrysoperla carnea Stephens, and the cotton aphid, Aphis gossypii Glover.
2. Zelus renardii and lacewings feed on aphids, thereby acting as potential competitors. In addition, Z. renardii feeds on lacewings. Thus, Z. renardii is an intraguild predator of lacewings.
3. Zelus renardii exhibited changes in prey preferences across developmental stages. The older stages of Z. renardii exerted greater mortality on lacewings and fed on larger lacewing larvae than did the younger stages.
4. Lacewings suppressed aphid population growth strongly. In contrast, none of the stages of Z. renardii was an effective control agent of the cotton aphid.
5. The addition of Z. renardii frequently disrupted the effective control of aphids generated by lacewings. In one of the two replicates of the experiment, the disruption increased with the developmental stage of Z. renardii , paralleling the increase in lacewing mortality.
6. Although the developmental stage of Z. renardii can influence the prevalence of intraguild predation and the intensity of the disruption of the aphid biological control, these experiments have demonstrated that even the youngest instars of Z. renardii can cause substantial lacewing mortality and release aphid populations from regulation.     

Influence of the margin vegetation on the conservation of aphid biological control in apple orchards

The influence of three margin strip treatments (wildflower strips, grass strips and spontaneous vegetation) adjacent to apple orchards on the biological control of Dysaphis plantaginea Passerini (Hemiptera: Aphididae) was compared during two consecutive years. The wildflower strips provided the highest amount of floral resources. Within the margin strips, hoverflies responded positively to higher resource provisioning whereas ladybird abundance did not differ between strip treatments. Within the orchards, the presence of parasitoids, hoverflies, and ladybirds in aphid colonies and the predation of sentinel aphids were not significantly affected by the adjacent strip treatments. The number of natural enemies observed in aphid colonies was mainly driven by aphid number. Aphid numbers were higher close to the margin strips suggesting that aphid colonization from orchard edges may counteract the positive effect of wildflower strips on natural enemy abundance and on a reduction of aphid infestation. The results confirm the positive influence of floral resource provisioning by wildflower strips on the conservation of aphid natural enemies, but also suggest that effects of wildflower strips on aphid regulation inside orchards are not very strong compared with spontaneous vegetation naturally occurring in the margins.     

Implications of floral resources for predation by an omnivorous lacewing

Understanding the relationship between a predator and its prey requires consideration of the other food resources used by the predator. In the case of true omnivores, these include plant-provided foods such as leaf tissue and nectar. The presence of plant resources can increase or decrease predation depending on the degree to which they are complementary to, or substitutable for, the prey. This has implications for the role of omnivores in biological control and some groups, notably heteropteran bugs and phytoseiid mites, have been studied in this context. However, few experiments have considered the effects of plant resources both on prey consumption by individual omnivores (which have an immediate effect on the prey population) and on attributes such as longevity and fecundity which act over the longer term to affect predation at the population level. In this study, a model system comprising an omnivorous adult lacewing (Micromus tasmaniae), buckwheat flowers and aphid prey was used to investigate how floral resources affected per capita predation rate, longevity and fecundity of the lacewing. Flowers reduced prey consumption. In the absence of prey, longevity was higher for lacewings with flowers than those without. In an experiment where aphids were provided in abundance, lacewing fecundity was unaffected by flowers. However, when aphids were less abundant, providing flowers decreased the pre-oviposition period and increased the daily oviposition rate. The results demonstrate that floral resources can mediate omnivore–prey relationships and that, in the context of biological control, their effects may be either positive or negative.     

(E)-β-Farnesene synthase genes affect aphid (Myzus persicae) infestation in tobacco (Nicotiana tabacum)

Aphids are major agricultural pests which cause significant yield losses of the crop plants each year. (E)-β-farnesene (EβF) is the alarm pheromone involved in the chemical communication between aphids and particularly in the avoidance of predation. In the present study, two EβF synthase genes were isolated from sweet wormwood and designated as AaβFS1 and AaβFS2, respectively. Overexpression of AaβFS1 or AaβFS2 in tobacco plants resulted in the emission of EβF ranging from 1.55 to 4.65 ng/day/g fresh tissues. Tritrophic interactions involving the peach aphids (Myzus persicae), predatory lacewings (Chrysopa septempunctata) demonstrated that the transgenic tobacco expressing AaβFS1 and AaβFS2 could repel peach aphids, but not as strongly as expected. However, AaβFS1 and AaβFS2 lines exhibited strong and statistically significant attraction to lacewings. Further experiments combining aphids and lacewing larvae in an octagon arrangement showed transgenic tobacco plants could repel aphids and attract lacewing larvae, thus minimizing aphid infestation. Therefore, we demonstrated a potentially valuable strategy of using EβF synthase genes from sweet wormwood for aphid control in tobacco or other economic important crops in an environmentally benign way.     

Combining lacewings and parasitoids for biological control of foxglove aphids in sweet pepper

The role of natural enemy diversity in biological pest control has been debated in many studies, and understanding how interactions amongst predators and parasitoids affect herbivore populations is crucial for pest management. In this study, we assessed the individual and combined use of two species of natural enemies, the parasitoid Aphidius ervi Haliday, and the predatory brown lacewing Micromus variegatus (Fabricius), on their shared prey, the foxglove aphid, Aulacorthum solani (Kaltenbach), on sweet pepper. We hypothesized that the presence of intraguild predation (IGP) and predator facilitation (through induced aphid dropping behaviour) might have both negative and positive effects on aphid control, respectively. Our greenhouse trial showed that overall, the greatest suppression of aphids occurred in the treatment with both the parasitoid and the lacewing. While the combination of lacewings and parasitoids significantly increased aphid control compared to the use of parasitoids alone, the effect was not significantly different to the treatment with only predators, although there was a clear trend of enhanced suppression. Thus, the combined effects of both species of natural enemies were between additive and non‐additive, suggesting that the combination is neither positive nor negative for aphid control. High levels of IGP, as proven in the laboratory, were probably compensated for by the strong aphid suppression provided by the lacewings, whether or not supplemented with some level of predator facilitation. For aphid management over a longer time scale, it might still be useful to combine lacewings and parasitoids to ensure stable and resilient aphid control.     

Manipulation of oviposition and overwintering site choice of common green lacewings with synthetic lure (Neuroptera: Chrysopidae)

Common green lacewings (Chrysoperla carnea species complex) are important predators of several insect pests, particularly aphids. Due to their potential in biological control, several efforts have been made to find means to retain common green lacewing adults or even to increase their numbers at a given site. In the course of field experiments conducted in Hungary and Norway, we tested the effect of a known ternary bait (phenylacetaldehyde, methyl salicylate, acetic acid) on oviposition and overwintering site choice of common green lacewings. In an orchard in Hungary, overwintering chambers were set up with and without baits. For the experiment on oviposition in Norway, a number of lacewing eggs were compared on trees with and without baits in selected sectors of the canopy. With the ternary bait, more common green lacewings could be attracted to overwintering chambers, and also significantly more eggs were found in the vicinity of baits than in other selected parts of the canopy, suggesting that this bait has the potential to manipulate both overwintering site choice and oviposition site choice of these important beneficial insects.     

Control of potato aphids by the addition of barley strips in potato fields: a successful example of vegetation management

The densities of barley and potato aphids, their natural enemies and hyperparasitoids were assessed in three experimental potato fields as a case study to investigate the effectiveness of the addition of barley strips in potato fields for conservation biological control. These fields were located in a low plant-diversity landscape, but common aphid species and their natural enemies were present. The barley strips in the potato fields were found to support different species of aphids of potato, but these different sets of aphids shared a common set of natural enemies. The amount of time between peak aphid densities and peaks of their natural enemies' populations was shorter in the potato fields than in the barley strips. The levels of winged aphids in a potato monoculture field were significantly higher than those in a field with barley strips. The wingless and winged aphid populations in the field without barley strips was almost three times higher than in the fields with the barley strips, as measured at the peak aphid density. This result is one of few examples of the application of the conservation effect of greenhouse banker plants on outdoor crops.     

Interaction of ants with aphid enemies: Do inexperienced ants specializing in honeydew collection recognize aphidophages at their first contact?

Honeydew collectors of Formica pratensis taken from the nature (control) and laboratory-reared “naïve” ants, which had never met either “mature” workers or aphids and aphidophages, were observed during their interactions with various aphid enemies: adults and larvae of ladybirds and lacewings, and larvae of syrphid flies. The naïve ants were significantly more aggressive towards adults than towards larvae of aphidophages. More than 70% of the naïve ants treated ladybirds and lacewings as enemies at their first encounter and attacked them immediately without any prior antennation. The frequency of aggressive reactions (body jerking and bites) towards larvae was significantly higher in the control group, whereas the percentage of ants showing explorative behavior was significantly higher in the naïve ants. Overall, experience proved to be not important for displaying the key behavioral reactions towards adult ladybirds and lacewings underlying the protection of trophobionts from these natural enemies. However, accumulation of experience is assumed to play an important role in the recognition of aphidophage larvae and formation of aggressive behavior towards them.     

Energy budgets of the Chinese green lacewing （Neuroptera： Chrysopidae） and its potential for biological control of the cotton aphid （Homoptera： Aphididae）

Energy budgets of larval stages of the Chinese green lacewing, Chrysopa sinica (Tjeder) (Neuroptera: Chrysopidae) were determined under laboratory conditions at photo‐period of 14:10 L:D, 27 ± 1°C and 75%± 2% RH. The energy used as ingestion, assimilation, respiration, productivity and feces was constructed for each developmental stage. In addition, under these experimental conditions, the potential of C. sinica as a biological control agent was evaluated according to the ingestion by this predator and the energy content of cotton aphid, Aphis gossypii (Glover) (Homoptera: Aphididae). The larval stage of C. sinica was able to consume 1281.4 1‐day‐old aphids, 1018.7 2‐day‐old aphids, 626.9 3‐day‐old aphids, 393.5 4‐day‐old aphids, 312.1 5‐day‐old aphids or 203.5 9‐day‐old aphids, respectively. No significant difference was detected between the estimated number of aphids consumed by the lacewings using energetic methods and the actual number of aphids consumed by the lacewings in this experiment. Our results showed that C. sinica is an important natural enemy of the cotton aphid, and energetic methods are very useful to quantify biological control efficacy of natural enemies.     

Assessment of fennel aphids (Hemiptera: Aphididae) and their predators in fennel intercropped with cotton with colored fibers

The fennel aphid, Hyadaphis foeniculi (Passerini) (Hemiptera: Aphididae) is a major pest of fennel, Foeniculum vulgare Miller in northeast region of Brazil. We hypothesize that intercropping can be used as an alternative pest management strategy to reduce aphid yield loss in fennel. Thus, we investigated the severity of fennel plant damage in relation to infestation by the fennel aphid and predation by Cycloneda sanguinea (L.) (Coleoptera: Coccinellidae) (spotless lady beetle), green lacewing, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae), and Scymnus spp. (Coleoptera: Coccinellidae) in sole fennel plots and plots of fennel intercropped with cotton with colored fibers. The fennel aphid populations in nontreated plots were significantly larger in sole fennel plots than in intercropped plots. The highest densities of C. sanguinea, green lacewings and Scymnus spp., associated with the suppression of fennel aphid populations was found in fennel in the intercropping systems. Fennel aphids reduced the fennel seed yield by 80% in the sole fennel plots compared with approximately 30% for all intercropping systems. The results obtained in this research are of practical significance for designing appropriate strategies for fennel aphid control in fennel-cotton intercropping systems. In summary, intercropping fennel with cotton with colored fibers apparently promoted biocontrol of fennel aphid in fennel.     

Potential for augmentative biological control of black bean aphid in California sugarbeet

In northern California, black bean aphid (Aphis fabae Scopoli complex) can be a major pest of sugarbeet, particularly in spring-planted fields. The major natural enemies of the aphid are predators, including the coccinellidsHippodamia convergens Guerin,Coccinella novemnotata Herbst &C. septempunctata L., and the chrysopidsChrysopa oculata Say &C. nigricornis Burmeister. Augmentative releases of eggs of eitherChrysoperla carnea (Stephens) orC. rufilabris (Burmeister) failed to significantly reduce aphid populations under field conditions. This was attributed in part to incompatibility between black bean aphids and lacewing larvae from commercial sources. Application of food sprays containing yeast hydrolysate, sucrose and molasses resulted in increased densities of eggs and adults ofC. carnea in treated plots. These results suggest that the potential for augmentative biological control of black bean aphid through the application of food sprays is greater than that for release of commercially available lacewings. Regardless of method, an holistic approach to augmentation that takes into account the ecological structure of the target agroecosystem will be required. Some aspects of “pre-emptive” biological control are discussed.     

Effect of plant nutrition on aphid size,prey consumption,and life history characteristics of green lacewing

Plant quality can directly and indirectly affect the third trophic level. The predation by all the instars of green lacewing, Chrysoperla carnea （S.） （Neuroptera： Chrysopidae） on the cereal aphids, Rhopalosiphum padi （L.）, and Sitobion avenae （F.） at varying nitrogen fertilizer levels was calculated under laboratory conditions. Wheat plants were grown on four nitrogen fertilizer levels and aphids were fed on these plants and subsequently offered as food to the C. carnea. Aphid densities of 10, 30, and 90 were offered to first, second, and third instar larvae of green lacewing. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. Aphid consumption by green lacewings was reduced with the increase in nitrogen content in the host plants of aphids. Predation of both aphid species by first, second, and third instars larvae of C. carnea was highest on aphids reared on plants with the lowest rate of fertilization, suggesting a compensatory consumption to overcome reduced biomass （lower aphid size）. Total biomass devoured by C. carnea on all nitrogen fertilizer treatments was not statistically different. Additionally, the heavier host prey influenced by the plant nutrition had an effect on the life history characteristics of green lacewings. The larval duration, pupal weight, pupal duration, fecundity, and male and female longevity were significantly affected by the level of nitrogen fertilization to the aphid＇s host plants, except for pupal duration when fed on S. avenae. This study showed that quantity of prey supplied to the larvae affects the prey consumption and thereafter the life history characteristics of green lacewings.     

Intraguild predation on the aphid parasitoid <Emphasis Type="Italic">Aphelinus asychis</Emphasis> by the ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

We investigated intraguild predation (IGP) on an aphid parasitoid, Aphelinus asychis Walker (Hymenoptera: Aphelinidae), by the multicolored Asian ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), and used the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae) as the prey/host in the laboratory. The ladybirds reared on artificial diet and on aphids consumed more aphids than mummies, while those reared on parasitized aphids consumed similar numbers of aphids and mummies. The ladybirds chose more mummies in treatments when mummies were more abundant, and more aphids when numbers of aphids and mummies were equal, or when aphids were more abundant. However, at all density treatments, rejection rates of mummies (36%) were much greater than of aphids (2%). H. axyridis prey on more aphids than A. asychis mummies, which enhances biological control by the two species. However, prior feeding experience affected subsequent choice, increasing the competition between natural enemies which would reduce their combined effectiveness for biological control.     

A survey of aphids and aphid parasitoids in cereal fields in Denmark, and the parasitoids' role in biological control

In 1996 and 1997 a field survey of the abundance and species composition of cereal aphid primary and secondary parasitoids in spring barley, winter wheat and durum wheat was conducted in Zealand, Denmark. The purpose was to create a better understanding of the mechanisms underlying aphid–parasitoid dynamics in the field. Such an understanding can be used when developing biological control methods in cereals. In both years aphid attacks in cereals began in late June and never exceeded the economic threshold. In 1996 the first aphids were found in wheat on 26 June; in 1997 the first aphids were found on 24 June on both crops. The highest densities reached in 1996 were an average of six aphids per shoot in winter wheat and one aphid per shoot in spring barley. In 1997 the highest densities reached were 11 aphids per shoot in winter wheat and four aphids per shoot in spring barley. The aphid population collapsed by the end of July to early August in 1996, but it collapsed by mid-July in 1997. The onset and peak of parasitization were delayed in comparison to aphid infestation. Parasitism was 20–60% by the end of the cropping season in spring barley, and 30–80% in winter wheat and durum wheat in 1996. In 1997 parasitism did not exceed 3–11% in barley and was less than 2% in one winter wheat field but more than 40% in the other winter wheat field sampled. In both years most parasitism was due to Aphidiidae (Hymenoptera). The two dominant species were Aphidius ervi Haliday and Aphidius rhopalosiphi De Stefani-Perez. Hyperparasitism began after primary parasitism and increased progressively during the cropping season. The two years were similar in many respects, including for species composition of aphids and parasitoids. The late start of the aphid infestation may have contributed to the high level of parasitization found in 1996, but in 1997 the aphid infestation period was so short that a parasitoid population did not have time to build up.     

Effects of reduced rates of two insecticides on enzyme activity and mortality of an aphid and its lacewing predator

By applying insecticides at lower rates of active ingredients per unit area, survival rates of the pests' natural enemies can be enhanced, whereas pest mortality can remain high. The effects of reduced application rates of the insecticides lambda-cyhalothrin and dimethoate on the mortality of bird cherry-oat aphid, Rhopalosiphon padi (L.), and lacewing Micromus tasmaniae Walker were determined in the laboratory and field. Cholinesterase (ChE) and glutathione S-transferase (GST) activities in survivors provided a measure of sublethal effects and general fitness. In the laboratory, lacewings were less sensitive than aphids to both insecticides, and dimethoate was more toxic than lambda-cyhalothrin. However, these results could not be recreated in the field, in part due to very low recapture rates. In summary, lambda-cyhalothrin seemed to have no effect on aphids, but it was toxic to lacewings. Dimethoate was far less toxic in the field, but aphids were still more sensitive than were lacewings. Cholinesterase activity was reduced by dimethoate exposure in the laboratory in both species, but there were species-specific differences. Dimethoate and lambda-cyhalothrin had no effects on GST activity in either species. The high mortality rate for lacewings and aphids exposed to dimethoate in the field suggests that the application rate could be reduced to as low as 10% of that recommended by manufacturers, and this should still be highly efficacious against aphids, while protecting the predatory lacewing. Measurement of enzyme activity could provide a useful indicator of "fitness" of survivors.     

Flower strips increase the control of rosy apple aphids after parasitoid releases in an apple orchard

Mass releases of two parasitoid species, Aphidius matricariae and Ephedrus cerasicola, may provide an alternative measure to pesticides to control the rosy apple aphid Dysaphis plantaginea in organic apple orchards. As an exploratory study, we tested if the presence of flower strips between apple tree rows could improve the action of three early parasitoid releases––and of other naturally present aphid enemies––on the control of aphid colonies and the number of aphids per tree. Apple trees located at various distances from parasitoid release points were monitored in plots with and without flower strips in an organic apple orchard over two years, along the season of aphid infestation (March to July). Our case study demonstrated that the presence of flowering plant mixes in the alleyways of the apple orchard reduced the presence of D. plantaginea by 33.4%, compared to plots without flower strips, at the infestation peak date. We also showed a negative effect of increasing the distance to parasitoid release points on aphid control. However, our results at the infestation peak date suggest that the presence of flower strips could marginally compensate for the detrimental effect of increasing distance to the release point, probably by improving the persistence and dispersal capacities of natural enemies. Despite high variations in aphid population dynamics between years, we conclude that combining flower strips with early parasitoid releases in apple orchards is promising for biological control of the rosy apple aphid, although the method merits to be further refined.     

Use of ryegrass strips to enhance biological control of aphids by ladybirds in wheat fields

Abstract Non‐crop habitats may play a vital role in conservation biological control. This study tested the effect of ryegrass (Lolium multiflorum L.) strips on aphid and ladybird populations in adjacent winter wheat fields. The field experiment was conducted in three ryegrass‐margin wheat plots and three control plots in 2010 in North China. In spring, the same aphid species, Sitobion miscanthi (Takahashi), was found in both the ryegrass strips and wheat plots. The population density of ladybirds in the ryegrass strips (3.5 ± 0.9/m²) was significantly higher than in the wheat plots (1.5 ± 0.5/m²). We cut the ryegrass, forcing the ladybirds to migrate to the wheat fields. Three and eight days after cutting the ryegrass, the aphid numbers in the ryegrass‐margin wheat plots decreased significantly: they were 19.9% and 53.6%, respectively, lower than in control plots. In the early period of ladybird population development, the percentage of larvae was greater in the ryegrass‐margin wheat plots than in controls, and the peak number of pupae in the ryegrass‐margin wheat plots occurred 5 days earlier than in the control plots. The results suggest that ryegrass strips may promote the development of ladybird populations. Cutting ryegrass can manipulate ladybirds to enhance biological aphid control in wheat fields. The efficiency of this management approach is discussed.     

Aphidophagous guilds on nettle （Urtica dioica） strips close to fields of green pea, rape and wheat

The common nettle （Urtica dioica L.） is a perennial and cosmopolitan plant species and is known to be the source of food for a great diversity of insects. To understand the importance of the nettle in agro-ecosystems, a field experiment was carried out in an experimental farm at Gembloux （Belgium） to study the effect of nettle margin strips on aphid and aphidophagous populations in close field crops, namely wheat （Triticum aestivum L.）, green pea （Pisum sativum L.） and rape （Brassicae napus L.）. The aphids and related beneficial populations were weekly assessed, from March to August 2005, by visual observations in two plots per field crop. A higher abundance of aphidophagous beneficials was collected in nettle strips when compared to the field crops. Particularly, the presence of predatory anthocorids, mirids and green lacewings was observed on nettle only. Nevertheless, the most abundant aphid predatory family, the Coccinellidae, was distributed in both environments, in nettle strips and in crop fields. The field margin supported a significantly higher density ofHarmonia axyridis than the field crops. In contrast, the field crops, green pea particularly, supported a higher density of Coccinella septempunctata. The distribution of the aphidophagous species, mainly the ladybirds, was discussed in relation to the host plant and related aphid species and their potential effect on integrated pest management.     

The interplay between density- and trait-mediated effects in predator-prey interactions: a case study in aphid wing polymorphism

Natural enemies not only influence prey density but they can also cause the modification of traits in their victims. While such non-lethal effects can be very important for the dynamic and structure of prey populations, little is known about their interaction with the density-mediated effects of natural enemies. We investigated the relationship between predation rate, prey density and trait modification in two aphid-aphid predator interactions. Pea aphids (Acyrthosiphon pisum, Harris) have been shown to produce winged dispersal morphs in response to the presence of ladybirds or parasitoid natural enemies. This trait modification influences the ability of aphids to disperse and to colonise new habitats, and hence has a bearing on the population dynamics of the prey. In two experiments we examined wing induction in pea aphids as a function of the rate of predation when hoverfly larvae (Episyrphus balteatus) and lacewing larvae (Chrysoperla carnea) were allowed to forage in pea aphid colonies. Both hoverfly and lacewing larvae caused a significant increase in the percentage of winged morphs among offspring compared to control treatments, emphasising that wing induction in the presence of natural enemies is a general response in pea aphids. The percentage of winged offspring was, however, dependent on the rate of predation, with a small effect of predation on aphid wing induction at very high and very low predation rates, and a strong response of aphids at medium predation rates. Aphid wing induction was influenced by the interplay between predation rate and the resultant prey density. Our results suggests that density-mediated and trait-mediated effects of natural enemies are closely connected to each other and jointly determine the effect of natural enemies on prey population dynamics.     

Abundance and natural control of the woolly aphid <Emphasis Type="Italic">Eriosoma lanigerum</Emphasis> in an Australian apple orchard IPM program

Woolly aphid (Eriosoma lanigerum Hausmann) (Hemiptera: Aphididae), was monitored over three growing seasons (1995--1998) to assess its abundance and management under apple IPM programs at Bathurst on the Central Tablelands of NSW, Australia. Woolly aphid infestations were found to be extremely low in IPM programs utilising mating disruption and fenoxycarb for codling moth Cydia pomonella L. (Lepidoptera: Tortricidae) control. This was the direct result of increased numbers of natural enemies. No insecticides were applied for woolly aphid control. Under the IPM strategies tested the principal control agent was identified as European earwig (Forficula auricularia L.) (Dermaptera: Forficulidae). Earwigs in combination with Aphelinus mali (Haldeman) (Hymenoptera: Aphelinidae) reduced woolly aphid infestations below the action threshold set by commercial growers. However, A. mali together with other flying natural enemies, e.g., ladybirds, lacewings and hoverflies, did not provide commercially acceptable control of woolly aphid in the absence of earwigs. Under the conventional spray program, using the broad-spectrum insecticide azinphos-methyl for codling moth control, the level of woolly aphid infestation increased with each successive season and biological control was not established. When azinphos-methyl was withdrawn, natural enemies migrated in and provided control of woolly aphid within one season. This is the first study to show that the biological control of woolly aphid can be achieved in a commercially viable IPM program.