Some factors influencing the occurrence and site of oviposition by aphidophagous Syrphidae (Diptera)

Syrphids were not deterred from ovipositing on suitable plants by the presence of other syrphid eggs, larvae or larval gut contents. Heavy infestations of Aphis fabae Scop, on bean plants (Viciafaba L.) induced Platycheirus manicatus (Meig.) to lay preferentially on neighbouring uninfested plants. Such behaviour was not found for Syrphus species. Nor did flowers, a source of adult food, stimulate oviposition nearby. The site of oviposition in relation to Brevicoryne brassicae L. on brussels sprouts (Brassicae oleracea gemnifera L.) varied markedly according to species: in similar conditions Syrphus luniger Meig. laid over 50% of its eggs touching aphids and less than 1% on uninfested plants, whereas P. manicatus laid less than 5% touching aphids and over 50% on uninfested plants. Most eggs were laid close to the edge of the leaf, especially by Platycheirus species, e.g. P. scutatus Meig. which laid over 90% within 5 mm of the edge of the leaf.     

Some host-plant factors affecting oviposition by aphidophagous Syrphidae (Diptera)

Although most aphidophagous syrphid species lay their eggs close to aphids, Platycheirus peltatus (Meig.), Melanostoma scalare (F.) and M. mellinum (L.) laid freely on uninfested brussels sprout plants. The existence of species that will lay in the absence of aphids may be valuable in biological control and also provides a useful tool for the investigation of non-aphid oviposition stimuli. Plant species and plant appearance were both important factors in P. peltatus oviposition. Particular sites on the plant were chosen for oviposition by Syrphus balteatus (Deg.). The nature of the substrate at the oviposition site affected the size of the egg batches laid there by M. scalare, M. mellinum, P. peltatus and P. clypeatus (Meig.). The possibility is discussed that plant-location is the older-established system of host-finding and that an aphid-location system has gradually superseded it during the evolution of the obligatorily aphidophagous Syrphinae.     

Hoverfly oviposition response to aphids in broccoli fields

The oviposition response of predacious hoverflies (Diptera: Syrphidae) to Brevicoryne brassicae L. and Myzus persicae (Sulzer) (Homoptera: Aphididae) in commercial broccoli, Brassica oleracea var. italica L., Plenck (Brassicaceae), fields was investigated at two sites over the course of a growing season. The hoverfly oviposition responses to these aphid species on different parts of the broccoli plant canopy were also examined. There were no hoverfly eggs on broccoli plants without aphids, egg numbers were very low on plants with fewer than 50 aphids, and no peak in oviposition relative to aphid numbers was observed. Within individual plants that were colonized by aphids, there was some oviposition on individual leaves without aphids, and no hoverfly eggs were seen on leaves that had more than 400 aphids. Leaves in the broccoli plant canopy, and the datasets associated with them, were divided into three sections vertically, ‘upper’, ‘middle’, and ‘lower’. Brevicoryne brassicae was more abundant in the upper and middle canopy sections, while M. persicae was found mostly in the lower section. The rate of hoverfly oviposition per aphid was higher in the upper section than in the two other sections. Modeling of the oviposition response using logistic regression showed that the presence of hoverfly eggs was positively correlated with numbers of each aphid species and sampling date.     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Infection with an insect virus affects olfactory behaviour and interactions with host plant and natural enemies in an aphid

Aphid ecology and population dynamics are affected by a series of factors including behavioural responses to ecologically relevant chemical cues, capacity for population growth, and interactions with host plants and natural enemies. Using the aphid Rhopalosiphum padi (L.) (Homoptera: Aphididae), we showed that these factors were affected by infection with Rhopalosiphum padi virus (RhPV). Uninfected aphids were attracted to odour of uninfected aphids on the host plant, an aggregation mechanism. However, infected aphids were not attracted, and neither infected nor uninfected aphids were attracted to infected aphids on the plant. Infected aphids did not respond to methyl salicylate, a cue denoting host suitability. Infected aphids were more behaviourally sensitive to aphid alarm pheromone, and left the host plant more readily in response to it. RhPV reduced the lifespan and population growth rate of the aphid. The predacious ladybird, Coccinella septempunctata (L.) (Coleoptera: Coccinellidae), consumed more infected aphids than uninfected aphids in a 24‐h period, and the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) attacked more infected than uninfected aphids. However, the proportion of mummies formed was lower with infected aphids. The results represent further evidence that associated organisms can affect the behaviour and ecology of their aphid hosts.     

The oviposition behavior ofAphidius ervi andMonoctonus paulensis (Hymenoptera: Aphididae) encountering different host species (Homoptera: Aphididae) in sequential patches

Two-day-old mated females ofAphidius ervi Haliday andMonoctonus paulensis (Ashmead) were each provided with two sequential host patches. Patches were comprised of plastic petri dishes containing either 15 pea aphids,Acyrthosiphum pisum (Harris), or 15 alfalfa aphids,Macrosiphum creelii Davis. Both wasp species parasitized more hosts in patches containing pea aphids than in those containing alfalfa aphids, regardless of sequence. Females ofA. ervi also laid more eggs per aphid in patches containing pea aphids than in patches containing alfalfa aphids. When both patches contained alfalfa aphids,M. paulensis females parsitized more aphids in the second patch than in the first. Fewer alfalfa aphids were parasitized in the second patch when the first patch contained pea aphids, and fewer eggs were laid per alfalfa aphid. Parasitoid females of both species exhibited consistently higher rates of oviposition into their preferred host species and adjusted their reproductive allocation to hosts and host patches as a function of their experience in previous patches.     

Effects of integrating companion cropping and nitrogen application on the performance and infestation of collards by Brevicoryne brassicae

Sustainable management of cabbage aphids, Brevicoryne brassicae (L.) (Hemiptera: Aphididae), is a major goal for collard, Brassica oleracea (L.) var. acephala (Brassicaceae), growers globally. Host finding ability of insect pests is significantly affected by diversified cropping systems, and this approach is being utilized currently as a pest management tool. Soil nutrition and its interaction with the cropping systems could have a significant effect on the general performance of collards and the infestation by cabbage aphids. In a search for a sustainable cabbage aphid control, a two‐season field experiment was carried out with two intercrops, collards and chilli, Capsicum frutescens (L.) (Solanaceae), and collards and spring onions, Allium cepa (L.) (Alliaceae), and a collard monoculture. For each of the cropping systems, nitrogen (N) was applied to the soil as a top‐dress at 20, 25, 30, and 35 g per collard plant. The response factors monitored were collard yield (fresh weight) and aphid infestation on collards. Spring onion‐collard intercrop had the lowest aphid density and the highest yield. Collard monoculture had the highest aphid infestation and the lowest yield. High levels of N led to increased infestation of collards by aphids, but also led to a significant increase in the yield of collards. Significant interactions between the N rates and the cropping systems were observed on some sampling dates, with the highest yield being realized under a combination of spring onion‐collard intercrop at a N rate of 30 g per plant. High aphid density led to a decrease in the yield of collards. It was concluded that with a spring onion‐collard intercrop, the soil N level could be raised from the blanket rate of 20–30 g per plant and this would lead to an increase in yield.     

Mechanisms of species‐sorting: effect of habitat occupancy on aphids' host plant selection

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Impact of wheat-mung bean intercropping on English grain aphid (Hemiptera: Aphididae) populations and its natural enemy

The effects of intercropping wheat, Triticum aestivum L., with mung bean, Vigna radiate L., on the populations of English grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae), and its natural enemies were evaluated by field and laboratory experiments. The population densities of aphids and their natural enemies were evaluated in the intercropped field against different row ratio combinations of wheat-mung bean. Results showed that wheat-mung bean intercropping caused a drop in aphid densities, and the ratio 12 wheat: 4 mung bean brought about the largest drop (> 8%). In addition, the population densities of coccinellids (ladybirds) and parasitoids and the species diversity of all the natural enemies of aphid were higher in the intercropped field than in the field planted only with wheat. However, intercropping did not influence the community indices (evenness and index of dominance concentration) of the natural enemies. Y-tube olfactometer bioassays were carried out in the laboratory to test whether odor blends of host and nonhost plants affect the host selection of S. avenae. Bioassays indicated that both apterous and alate aphids significantly preferred host plant odor over odor blends of host and intercropped species. Hence, the olfactory-based host location of aphids in the field might be affected by intercropping. The intercropping experiment clearly showed that increased crop species diversity suppresses aphid population growth and preserves the population of natural enemies of aphids. Our results also provide support for the "resource concentration hypothesis" and the "enemies hypothesis".     

Glucosinolate profile and oviposition behavior in relation to the susceptibilities of Brassicaceae to the cabbage seedpod weevil

Understanding how host‐plant characteristics affect behavioral and physiological responses of insect herbivores is of considerable importance in the development of resistant crop germplasm. Feeding, oviposition preference, larval development, and oviposition behavior of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (= Ceutorhynchus assimilis Payk.) (Coleoptera: Curculionidae), were investigated on eight Brassicaceae species that differed in their glucosinolate profiles. The least preferred host plants for feeding and oviposition were the Sinapis alba L. lines while the Brassica carinata L. line was most preferred. Larval development occurred most rapidly on Brassica rapa L. and slowest on S. alba. Larval weight was highest on B. napus L. and lowest on S. alba. Total glucosinolate levels did not influence C. obstrictus larval growth or development; however high levels of specific glucosinolates such as p‐hydroxybenzyl and 3‐butenyl glucosinolate were associated with increased developmental time or reduced weight. The time required for oviposition behavioral events was measured on different host‐plant species: B. rapa, B. napus, B. napus×S. alba, B. tournefortii Gouan., B. juncea (L.) Czern, B. carinata, B. nigra (L.) Koch., and S. alba. The early steps in the sequence were completed faster on more susceptible host plants (B. carinata, B. napus, and B. rapa) than on relatively resistant ones (B. tournefortii and B. juncea). Females explored pods of B. nigra and S. alba, but oviposition occurred only rarely on these species. There was no significant difference in the location on the pod on which oviposition occurred among the different plant species. Mean eggs laid per female weevil were highest on the B. napus×S. alba hybrid and lowest on B. nigra and S. alba.     

Predatory hoverflies select their oviposition site according to aphid host plant and aphid species

The hoverfly Episyrphus balteatus De Geer (Diptera: Syrphidae) is an abundant and efficient aphid‐specific predator. Several aphidophagous parasitoids and predators are known to respond positively to aphid‐infested plants. Semiochemicals from the latter association usually mediate predator/parasitoid foraging behavior toward sites appropriate for offspring fitness. In this study, we investigated the effect of aphid host plant and aphid species on foraging and oviposition behavior of E. balteatus. Behavioral observations were conducted using the Noldus Observer v. 5.0, which allows observed insect behavior to be subdivided into different stages. Additionally, the influence of aphid species and aphid host plant on offspring fitness was tested in a second set of experiments. Acyrthosiphon pisum Harris and Megoura viciae Buckton were equally attractive for E. balteatus whereas Aphis fabae Scopoli (all Homoptera: Aphididae) were less attractive. These results were correlated with (i) the number of eggs laid, which was significantly higher for the two first aphid species, and (ii) the fitness of hoverfly larvae, pupae, and adults. Two solanaceous plant species, Solanum nigrum L. and Solanum tuberosum L. (Solanaceae), which were infested with Myzus persicae Sulzer (Homoptera: Aphididae), were also compared using the same approach. Discrimination between these two M. persicae host plants was observed, with S. tuberosum being preferred as an oviposition site by the predatory hoverfly. Larval and adult fitness was correlated with the behavioral observations. Our results demonstrated the importance of the prey–host plant association on the choice of the oviposition site by an aphid predator, which is here shown to be related to offspring fitness.     

Preference of cereal aphids for different varieties of winter wheat

This paper reports on the development of a simple and robust preference meter (developed in-house) to score the host choice behavior of apterous aphids. With this tool, the preferences of two important cereal aphids Sitobion avenae (Fab.) and Metopolophium dirhodum (Walker) were investigated against four different varieties of winter wheat (Triticum aestivum L.) with a different susceptibility for Fusarium head blight (FHB). Differences in the choice behavior of both aphid species were observed for different wheat varieties. The preferred wheat variety of S. avenae and M. dirhodum was not the same. Also, both aphid species clearly had a differential preference for seedlings and ears. Using seedlings, M. dirhodum was about 1.8 times more rapid in making its choice than S. avenae. In separate experiments with ears, S. avenae was 4.5 times faster than in the experiments with seedlings. In the present study, we aim to highlight differences in preference behavior in relation to potential mechanisms for host selection.     

Field manipulation of populations of individual staphylinid species in cereals and their impact on aphid populations

Abstract.

• 1 Four species of staphylinids were caged separately on wheat and their effect on the population development of the grain aphid, Sitobion avenae (F.), was investigated over two seasons.
• 2 A novel field cage design was used to exclude all but the staphylinid species under investigation.
• 3 Predator effects were compared with total-exclusion cages in which all predators and parasitoids were kept out.
• 4 During a high-density phase of aphid population development only Philonthus cognutus (Stephens) significantly influenced aphid population numbers.
• 5 At low aphid densities, Tachyporus obtusus (L.), T.chrysomelinus (L.) and P.cognatus reduced aphid population levels.
• 6 The experiments showed that Tachyporus spp. can reduce the numbers of cereal aphids prior to the exponential phase of aphid population increase and that P.cognatus causes some reduction at both the stages of aphid population growth investigated.

     

The effect of rearing history and aphid density on volatile‐mediated foraging behaviour of Diaeretiella rapae

1. Parasitoid females foraging for hosts rely on cues derived from the insect host, the host plant and/or their interaction, and all of these can be learned during the immature and adult stages. 2. The present study investigated the importance of rearing history on foraging behaviour of Diaeretiella rapae, an endoparasitoid often associated with aphids feeding on brassicaceous plant species. 3. Parasitoids were reared on one of the four possible combinations, comprising two brassicaceous host plant species, Brassica nigra or Raphanus sativus, and two aphid species Brevicoryne brassicae or Myzus persicae. These parasitoids were tested in a Y‐tube olfactometer and given the choice between volatiles emitted by an aphid‐infested plant (25 or 100 aphids per plant) and an uninfested control plant. The parasitoid's responses were compared when offered: (i) the same plant–aphid combination as the one on which it had been reared; (ii) the same host plant infested with the alternative aphid species; or (iii) an alternative plant with the alternative aphid species. 4. Aphid density did affect the behavioural responses to the various odour sources, but rearing history did not. Diaeretiella rapae only preferred aphid‐induced to non‐induced plant volatiles at low aphid infestation level, whereas they did not discriminate between volatiles at high aphid infestation level. 5. It is concluded that aphid‐induced volatiles of brassicaceous plants play an important role during host habitat location, but seem less important for parasitoids to locate the aphid host itself. The data are discussed in the light of manipulation of host plant defences by aphids.     

Root herbivory in vitro: Interactions between roots and aphids grown in aseptic coculture

Summary An in vitro coculture system has been established to study interactions between roots and aphids. Eight aphid species (Aphis spiraecola P., Trama rara M., Macrosiphum euphorbiae S., Rhopalosiphum padi L., Sitobion avenae F., Rhopalosiphum maidis F., Metopolophium dirhodum W., and Pemphigus populivenae F.) were reared on six species of hairy root cultures, Carthamus tinctorius L. cv N10, Tagetes patula L., Trichosanthes cucumerina L. var anguina, Hyoscyamus muticus L., Nicotiana tabacum L., and Beta vulgaris L. subsp. vulgaris. All species of aphids survived on root cultures for at least 2 d. Three cocultures have been maintained aseptically for periods ranging from 2 mo. to over 2 yr. The coculture of R. padi on C. tinctorius cv N10 (N10-Rp) was used to study morphological and biochemical responses of roots under aphid herbivory. Aphid herbivory caused browning of cultures, reduced root vegetative growth, and increased production of polyacetylenes in C. tinctorius cv N10 roots. Our results suggest that this coculture system may improve our understanding of interactions between aphids and plant roots.     

Fungal endosymbionts affect aphid population size by reduction of adult life span and fecundity

The symbiosis between grasses and endophytic fungi is a common phenomenon and can affect herbivore performance through acquired, chemical plant defence by fungal alkaloids. In laboratory experiments, two species of common grass aphids, Rhopalosiphum padi and Metopolophium dirhodum were tested, in a population experiment (on four plant cultivars) and individually (on one plant cultivar) for the effects of the endophyte, Neotyphodium lolii, that forms symbiotic associations with perennial ryegrass Lolium perenne. In the population experiment that lasted for four aphid generations both aphid species showed decreased population sizes when feeding on each of the four endophyte-infected cultivars. Individuals of R. padi tested individually showed reduced adult life span and fecundity when feeding on infected plants. Individuals of M. dirhodum showed no response in any of the traits measured. This suggests that R. padi individuals are more sensitive to endophyte infection than M. dirhodum individuals. However, all infected grass cultivars reduced population sizes of both aphid species over four generations. Therefore, fungal endophytes can reduce populations of aphid herbivores independent of plant cultivars.     

Spatial patterns of goldenrod aphids and the response of enemies to patch density

Summary The two aphid species feeding on goldenrod (Solidago altissima) in northern Florida (U.S.A.) exhibited behavioral differences that resulted in characteristic spatial patterns. Uroleucon nigrotuberculatum alates (winged forms) aggregated when colonizing stems and subsequent non-winged generations were relatively sedentary, resulting in a clumped spatial pattern. U. tissoti colonized stems singly and was more mobile; these behaviors resulted in its more random spatial pattern within fields of goldenrod.Manipulations of aphid density in the field revealed that although patches with high densities of aphids accumulated more predators than patches with few aphids, predation pressure (measured as number of predators per aphid) was lower in dense patches. As a result, aphids in dense patches had a higher per capita change in density than aphids in sparse patches. However, when the fungal pathogen, Neozygites fresenii, became the dominant mortality agent, the influence of aphid density on mortality was reversed; aphids in dense patches were then more vulnerable than aphids in sparse patches. Thus the spatial patterns exhibited by the U. nigrotuberculatum and U. tissoti resulted in differences in their relative vulnerability to different natural enemies.     

Evaluation of induced responses, insect population growth, and host-plant fitness may change the outcome of tests of the preference-performance hypothesis: a case study

The preference‐performance hypothesis predicts that insect preference should correspond to host suitability for offspring development. We studied the pattern of within‐plant preference in the aphid Sipha flava and its consequences for offspring performance on the host‐plant Sorghum halepense, regarding the role of induced responses of plants to aphid feeding. The consequences of within‐plant preference on aphid population growth and host‐plant traits were also evaluated. Our results showed that winged and wingless aphids preferred to settle on mature rather than young leaves. In contrast, aphid individual growth rate was higher on young leaves when compared with mature leaves, suggesting that the outcome of this test rejected the preference‐performance hypothesis. However, the inclusion of the factor ‘previous aphid infestation’ changed the outcome from a maladaptive choice to a neutral one. Thus, individual growth rates of S. flava increased when aphids developed on leaves that had been previously infested. Interestingly, aphid growth rate on previously infested leaves did not differ between young and mature leaves. On the other hand, aphid population reproductive rate was higher and the percentage of winged aphids lower when infestation occurred on mature rather than young leaves. Aphid infestation reduced plant and shoot biomass, and increased leaf mortality. These negative effects on plant traits related to plant fitness were greater when aphid infestation occurred on young leaves. Likewise, whereas infestation on mature leaves did not cause a significant reduction in the number of flowering plants compared with control plants, aphid infestation on young leaves did reduce the number of plants at the flowering stage. Consequently, if both the reproductive rate of aphids in the mid‐term, and host‐plant fitness are taken into account, the results indicate that aphid preference for mature leaves may be an adaptive choice, thus supporting the preference‐performance hypothesis.     

Leaf surface wax and plant morphology of peas influence insect density

Insect predators and parasitoids adhere better, forage more effectively, and take more aphid prey on pea plants (Pisum sativum L.) (Leguminosae) with mutations that reduce the crystalline wax bloom on the plant surface. To assess the agronomic potential of this trait for pest management, abundance of pea aphids (Acyrthosiphon pisum L.) (Homoptera: Aphididae) and coccinellid predators, and percent parasitism of the aphids were evaluated on pea lines differing in wax bloom and plant architecture over two field seasons. Three pairs of pea lines were evaluated, each pair with a different architecture and differing within the pair in the amount of surface wax bloom (reduced or normal). The trials included plots treated with a narrow spectrum insecticide (pymetrozine) to reduce aphid populations and untreated controls. Reduced wax peas had significantly fewer aphids per plant in 2002 but not in 2003. Total natural enemy abundance was greater on reduced wax than on normal wax pea lines in both years of the study. Pymetrozine reduced aphid densities significantly in both years. Among the four pea lines evaluated for yield, seed yield per plant was affected by plant morphology and insecticide treatment. Yield was greatest on semileafless plants and on pymetrozine sprayed plots in both years. Yield of the reduced wax line in the semileafless background was similar to or exceeded yield in its normal wax sister line, suggesting that this morphological type was best for an agronomically viable reduced wax phenotype. Pea weevil (Bruchus pisorum L.) (Coleoptera: Bruchidae) damage to seed was overall more frequent on seeds from reduced wax varieties than from normal wax varieties. The results illustrate the trade‐offs associated with a reduced wax trait in peas but also show that certain combinations of reduced wax and gross morphology lead to reduced pea aphid populations and yields similar to those of normal wax peas.     

Predator and parasitoid attacking ant-attended aphids: effects of predator presence and attending ant species on emerging parasitoid numbers

Interaction between a predator and a parasitoid attacking ant-attended aphids was examined in a system on photinia plants, consisting of the aphid Aphis spiraecola, the two ants Lasius japonicus and Pristomyrmex pungens, the predatory ladybird beetle Scymnus posticalis, and the parasitoid wasp Lysiphlebus japonicus. The ladybird larvae are densely covered with waxy secretion and are never attacked by attending ants. The parasitoid females are often attacked by ants, but successfully oviposit by avoiding ants. The two ants differ in aggressiveness towards aphid enemies. Impacts of the predator larvae and attending ant species on the number of parasitoid adults emerging from mummies per aphid colony were assessed by manipulating the presence of the predator in introduced aphid colonies attended by either ant. The experiment showed a significant negative impact of the predator on emerging parasitoid numbers. This is due to consumption of healthy aphids by the predator and its predation on parasitized aphids containing the parasitoid larvae (intraguild predation). Additionally, attending ant species significantly affected emerging parasitoid numbers, with more parasitoids in P. pungens-attended colonies. This results from the lower extent of interference with parasitoid oviposition by the less aggressive P. pungens. Furthermore, the predator reduced emerging parasitoid numbers more when P. pungens attended aphids. This may be ascribed to larger numbers of the predator and the resulting higher levels of predation on unparasitized and parasitized aphids in P. pungens-attended colonies. In conclusion, a negative effect of the predator on the parasitoid occurs in ant-attended aphid colonies, and the intensity of the interaction is affected by ant species.