Colonization of apple orchards by predators of <Emphasis Type="Italic">Dysaphis plantaginea</Emphasis>: sequential arrival,response to prey abundance and consequences for biological control

Episyrphus balteatus (DeGeer) (Diptera: Syrphidae), Adalia bipunctata (L.) (Coleoptera: Coccinellidae) and Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae) are the three most abundant natural enemies of Dysaphis plantaginea Passerini (Homoptera: Aphididae) in Asturian (NW Spain) apple orchards. They attack this aphid in sequence: E. balteatus arrived first, followed by A. bipunctata and then by A. aphidimyza. The cecidomyiids arrived too late to have a regulating effect. The syrphids laid an average of 2.3 ± 1.7 eggs per aphid colony and the coccinellids 18.4 ± 9.9 regardless of the degree of the infestation rates of the apple shoots. This value corresponds to the size of an egg batch laid by one female. Therefore, these aphid predators did not respond numerically to the abundance of the pest. The results of this study indicate that natural populations of syrphids and ladybird beetles are unable to control D. plantaginea, and therefore a more complex strategy than waiting for natural enemies is required.     

Temporal and spatial variability of rosy apple aphid Dysaphis plantaginea populations: is there a role of the alternate host plant Plantago major?

• 1 The rosy apple aphid Dysaphis plantaginea (Passerini) (Homoptera: Aphididae) is a pest of economic importance to the apple industry worldwide, particularly in organic apple orchards where no acceptable controls are available. In the Similkameen Valley of British Columbia, Canada, the rosy apple aphid population size varies widely between orchards and between years. To explain this variation, potential environmental correlates of aphid density were evaluated. The architecture of the alternate host was also evaluated for its effect on rosy apple aphid summer survival and reproduction.
• 2 The percentage of trees infested by rosy apple aphids among orchards was in the range 8–94% for trees having at least one cluster with more than ten aphids in 2007 and in the range 0–39% in 2008.
• 3 A general linear model correlating aphid densities to the environmental variables of abundance of the alternate host (plantain Plantago spp.), foliar nitrogen, tree age and planting density, and reduced by stepwise regression, indicated that foliar nitrogen and tree age explained 33% of the variation. Abundance of the summer, alternate food plant, plantain, was not related to later aphid densities on apple trees.
• 4 Plantain architecture, however, influenced aphid numbers and 25‐fold more aphids were found on low‐lying plantain leaves than on more upright leaves. Experimental manipulation of leaf angle and leaf size showed that significantly more aphids occurred on low angle, large leaves. Finally, mowing that encouraged low lying plants prior to spring aphid migration was associated with a four‐fold greater number of both winged and wingless aphids on the plantain.

     

Effects of augmentative releases of eggs and larvae of the ladybird beetle, Adalia bipunctata, on the abundance of the rosy apple aphid, Dysaphis plantaginea, in organic apple orchards

The impact of augmentative releases of larvae and eggs of the indigenous ladybird beetle Adalia bipunctata (L.) (Coleoptera: Coccinellidae) against the rosy apple aphid Dysaphis plantaginea Pass. (Homoptera: Aphididae), a major pest insect on apple trees, was assessed in field experiments in Switzerland, during 1997. In a first experiment, eggs and larvae were released on 3-year old apple trees infested with five aphids at four different predator-prey ratios (0:5, 1:5, 1:1, 5:1). In a second experiment, eggs and larvae were released at a predator-prey ratio of 5:1 on branches of apple trees naturally infested with aphids. In both experiments, the interaction with ants was taken into account and the releases were done at two different times in spring. The results showed that an augmentative release of larvae significantly prevented the build-up of colonies of D. plantaginea. Significant reductions in aphid numbers were recorded at the two highest predator-prey ratios, 1:1 and 5:1. Larvae were efficient just before flowering of apple trees at a time when growers normally have to spray their trees. On trees where ants were present the larvae of A. bipunctata were significantly less efficient. Effects of eggs of A. bipunctata, however, were less reliable. At the first date of release (5 April), they did not hatch, probably as a consequence of bad weather conditions.     

Molecular interactions between rosy apple aphids, Dysaphis plantaginea, and resistant and susceptible cultivars of its primary host Malus domestica

The use of crop varieties resistant or tolerant to insect pests or other stress factors is one approach in non‐chemical crop‐protection. Knowledge of the biochemical and molecular background of insect–plant interactions is a prerequisite for optimizing breeding for resistance. However, the resistance genes involved in plant–aphid interactions have so far only been identified and characterized in very few plant species. Our work aims to elucidate the molecular and biochemical mechanisms involved in resistance of apple trees, Malus domestica L. (Rosaceae), against its primary aphid pest, the rosy apple aphid, Dysaphis plantaginea (Passerini) (Homoptera: Aphididae), which is considered a serious economic pest of apple. Gene expression in both resistant and susceptible apple cultivars after infestation with rosy apple aphids was investigated by employing the cDNA‐AFLP method (cDNA–Amplified Fragment Length Polymorphism). From approximately 12 500 cDNA fragments detected on polyacrylamide gels, 21 bands were apparently up‐ or down‐regulated only in the resistant cultivar ‘Florina’ after aphid infestation compared to the susceptible cultivar ‘Topaz’ and/or mechanically wounded or non‐infested leaves. These fragments were cloned, sequenced, and the pattern of gene expression for six fragments was subsequently verified by virtual Northern blots. Sequence comparisons of these fragments to GenBank accessions revealed homologies to already known genes, most of them isolated from Arabidopsis thaliana L. Among them, a putative RNase‐L‐inhibitor‐like protein, a pectinacetylesterase, an inositol‐phosphatase‐like protein, a precursor of the large chain of the ribulose‐1,5‐biphosphate‐carboxylase, and defence‐related genes such as a vacuolar H(+)‐ATPase subunit‐like protein and an ADP‐ribosylating enzyme were identified. The results are discussed in relation to a putative role of these genes in conferring aphid resistance in apple trees.     

DL‐β‐Aminobutyric acid application negatively affects reproduction and larval development of the rosy apple aphid,Dysaphis plantaginea,on apple

The rosy apple aphid, Dysaphis plantaginea (Passerini) (Hemiptera: Aphididae), is one of the major pests of European apple orchards, commonly controlled by the use of synthetic insecticides. In the present work, the non‐protein amino acid DL‐β‐aminobutyric acid (BABA), known to induce plant resistance against a wide range of abiotic and biotic stresses, has been tested for its protective effect against this pest on apple. We first verified the lack of any contact effect of BABA on the insect itself. Next we applied BABA as a soil drench to apple and monitored its effect on the population development of aphids after artificial infestation. We demonstrated that BABA strongly reduced the population growth and that this compound severely affected various life‐history characteristics of the aphid such as female longevity and fecundity, nymph mortality, and larval development.     

Temperature effects on egg development of the rosy apple aphid and forecasting of egg hatch

The development of Dysaphis plantaginea (Pass.) (Homoptera: Aphididae) winter eggs was studied at six different constant temperatures ranging from 7.5 to 16.5 °C in order to improve the basis for phenological forecasts in early spring. The mortality was generally low at temperatures below 13.5 °C but increased considerably at 16.5 °C. The effect of temperature on development rates could be described with linear regression within the temperature range under study. The lower temperature threshold for development was estimated to be 4.0 °C and the thermal constant 140 day‐degrees. A time‐varying distributed delay approach was used to establish a temperature driven phenology model for winter egg hatch of D. plantaginea considering the intrinsic variability in development time. The model parameters such as temperature‐dependent development times and corresponding variances were quantified based on the experimental data. When compared with independent observations on egg hatch under semifield conditions, the model gave satisfactory validation results. It can be used as forecasting tool for the optimal timing of monitoring and control measures for D. plantaginea in early spring.     

The induction of leaf-roll galls by the apple aphids Dysaphis devecta and D. plantaginea

Dysaphis devecta causes the leaves of its host plant to roll laterally whereas D. plantaginea causes them to roll longitudinally. Both species of aphid are phloem feeders. D. devecta prefers to feed on the smaller veins in the lamina whereas D.plantaginea chooses the midrib. However, groups of D. devecta or D. plantaginea confined to the stem of an apple seedling induced young leaves several centimetres away to develop leaf rolls characteristic of each species. A single larva of D. devecta or D. plantaginea can induce a leaf-roll or a stem-bend on an apple seedling within 24 h. It is suggested that characteristic abnormalities in leaves attacked by D. devecta and D. plantaginea are caused by specific substances in the saliva of each aphid.     

Effects of autumn kaolin and pyrethrin treatments on the spring population of Dysaphis plantaginea in apple orchards

The impact of treatments with the pyrethrin insecticide Pyrethrum FS and the repellent kaolin product Surround^® WP to control the autumn forms of Dysaphis plantaginea Pass. (Hom., Aphididae), a major insect pest of apple, was assessed in 1‐year field experiments in Switzerland. Single and multiple applications of pyrethrin and kaolin were tested at different dates after apple harvesting in autumn 2002 when sexuales of the rosy apple aphid were present. Repeated applications of Surround^® WP in autumn significantly reduced the number of females in autumn and, consequently, the number of hatched fundatrices in spring. Single kaolin treatments were less effective. Unexpectedly, neither single nor multiple applications of the contact insecticide Pyrethrum FS had a knock‐down effect on females in autumn. However, pyrethrin significantly reduced the number of hatched fundatrices in spring. Neither pesticide completely controlled the rosy apple aphid but with a more detailed analysis of factors influencing the efficacy of autumn treatments a new approach to control this serious pest could be achieved.     

The effect of ant attendance on the success of rosy apple aphid populations, natural enemy abundance and apple damage in orchards

1 The rosy apple aphid, Dysaphis plantaginea, is the most serious pest of apple in Europe and, although conventionally controlled by insecticides, alternative management measures are being sought. Colonies of D. plantaginea are commonly attended by ants, yet the effects of this relationship have received little attention. 2 An ant exclusion study was conducted in two distant orchards within the U.K. At both sites, ants were excluded from a subset of D. plantaginea infested trees at the beginning of the season and populations were monitored. The number of natural enemies observed on trees was also recorded and, before harvest, the percentage of apples damaged by D. plantaginea calculated. 3 Overall, the exclusion of ants reduced the growth and eventual size of D. plantaginea populations. On trees accessed by ants, greater numbers of natural enemies were recorded, presumably because aphid populations were often greater on such trees. However, this increased natural enemy presence was diluted by the larger aphid populations such that individual aphids on ant‐attended trees were subjected to a lower natural enemy pressure compared with those on ant‐excluded trees. 4 At harvest, apple trees that had been accessed by ants bore a greater proportion of apples damaged by D. plantaginea. There were also differences in cultivar susceptibility to D. plantaginea damage. 5 The present study highlights the importance of the ant–D. plantaginea relationship and it ia suggested that ant manipulation, whether physically or by semiochemicals that disrupt the relationship, should be considered as a more prominent component in the development of future integrated pest management strategies.     

Conservation biological control of rosy apple aphid, Dysaphis plantaginea (Passerini), in eastern North America

Because of the potentially serious damage rosy apple aphid, Dysaphis plantaginea (Passerini) (Homoptera: Aphididae), can cause to apple fruit and branch development, prophylactic insecticides are often used for control. If biological control could be relied on, the amount of pesticide applied in orchards could be reduced. This study examined biological control of rosy apple aphid in eastern West Virginia and the potential for enhancement through conservation biological control, in particular, the effect of interplanting extrafloral nectar-bearing peach trees. By 20 d after first bloom, only 2% of fundatrices initially present survived to form colonies based on regression of data from 687 colonies. Exclusion studies showed that many of the early colonies were probably destroyed by predation; the major predator responsible seemed to be adult Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Mortality before apple bloom was most important in controlling rosy apple aphid population growth but by itself is not sufficiently reliable to prevent economic injury. Interplanting of extrafloral nectar-bearing trees did not increase biological control, and interplanting with 50% trees with extrafloral nectar glands reduced biological control. The number of leaf curl colonies in the 50% interplanted orchards was lower than in monoculture orchards, suggesting a preference of alate oviparae for more diverse habitats, supporting the resource concentration hypothesis but not at a level sufficient to prevent injury. Predation and parasitism after the formation of leaf curl colonies was not adequate to control rosy apple aphid populations.     

Isolation of microsatellite markers from the rosy apple aphid Dysaphis plantaginea

We report the development of five polymorphic microsatellite loci from the aphid Dysaphis plantaginea, a serious pest on apple. Using a multimotif enrichment protocol, we have isolated loci that are proving to be valuable tools for studying aphid movement at both between‐county and within‐orchard scales.     

Seasonal coincidence between rosy apple aphid fundatrices and their parasitoids: implications for an early biological control of Dysaphis plantaginea

The potential contribution of the aphid parasitoid Ephedrus persicae Froggatt (Hymenoptera: Braconidae, Aphidiinae) in regulating stem mothers of the rosy apple aphid Dysaphis plantaginea (Passerini) (Homoptera: Aphididae) was investigated in culture conditions using both species' phenological emergence data in spring and host stage suitability trials. In 2003, emergence of E. persicae started on April 1st, i.e., 108.2 degree-days (base: 4.5 degrees C; start date: January 1st) later than its host on March 10th. When less than 3% of parasitoids had emerged on April 12th, more than 97% of D. plantaginea stem mothers had moulted beyond their 1st instar. The latest parasitoids left their diapause mummies in late April as the majority of fundatrices had reached the adult stage. Parasitisation trials demonstrated the suitability for E. persicae of all developmental stages of rosy apple aphid fundatrices, including mature individuals. No offspring were deposited by fundatrices parasitised as Ist-instar nymphs but later instars did produce a progeny whose size was substantially reduced compared with unparasitised individuals, and scaled against their age at the time of parasitisation. Although the temperature accumulation required for emergence in spring is reached later for E. persicae than for its host aphids, thus allowing part of the stem mother population to temporarily escape parasitisation and reach maturity, the marked reduction in fecundity of individuals parasitised even as late-instar nymphs does confer to E. persicae a non-negligible role in potentially regulating D. plantaginea.     

Effects of augmentative releases of the coccinellid, Adalia bipunctata, and of insecticide treatments in autumn on the spring population of aphids of the genus Dysaphis in apple orchards

The impact of augmentative releases of indigenous predators and insecticide applications to control the autumn aphid forms of the genus Dysaphis (Homoptera: Aphididae), major pest insects on apple trees, was assessed in one-year field experiments in Switzerland. Eggs and larvae of the two-spot ladybird beetle Adalia bipunctata (L.) (Coleoptera: Coccinellidae) were released on 4-year old apple trees in various numbers at five different dates in autumn 1998 when sexuales of the aphids were present. Additionally, Pyrethrum HP was sprayed at the same five dates to compare the effectiveness of these augmentative releases to a commonly applied insecticide. Augmentative releases of larvae before mid-October significantly prevented the deposition of overwintering eggs by aphids of the genus Dysaphis and consequently reduced the number of hatched fundatrices in spring, 1999. There was a significant negative functional response among the number of released coccinellid larvae in autumn 1998 and the number of observed fundatrices on apple trees in spring 1999. Applications of Pyrethrum HP before mid-October were more effective than augmentative releases of larvae of A. bipunctata. The release of coccinellid eggs had no impact on the number of fundatrices of the genus Dysaphis in the next spring because they did not hatch due to bad weather conditions. The weather conditions in autumn seemed to have an impact on the autumn migration of the winged aphids back to their primary host. The prevention of egg deposition of aphids in autumn is a promising control strategy and deserves further exploration for practical use.     

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.     

Hypersensitivity in apple to feeding by Dysaphis plantaginea: effects on aphid biology

Effects on rosy apple aphid (Dysaphis plantaginea) of hypersensitivity to aphid feeding in apple were assessed in a series of insectary experiments in 1980–82. Hypersensitive plants rapidly produced tissue necroses at aphid feeding sites. Inoculation with apterae demonstrated that, in comparison with aphids on susceptible plants, aphids on hypersensitive plants showed symptoms characteristic of host antibiosis, including increased mortality, failure to reproduce and slower development. Embryo number indicated that fecundity of surviving adults was lower on hypersensitive plants and mean relative growth rate of aphids was reduced. Hypersensitivity did not prevent colonisation by gynoparae returning from the summer host, nor subsequent egg-laying. It conferred, however, a strong resistance against colonisation after egg-hatch the following spring. The potential of hypersensitivity as a field-resistance is discussed.     

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.     

The potential of three native insect predators to control the rosy apple aphid, Dysaphis plantaginea

The potential of three aphidophagous predators, Adalia bipunctata, Aphidoletes aphidimyza, and Episyrphus balteatus to control the rosy apple aphid, Dysaphis plantaginea Pass., a major pest on apple in Europe, was assessed by means of laboratory and field cage experiments in Northern Switzerland. Under laboratory conditions, all three predators efficiently preyed upon D. plantaginea on apple seedlings. The searching success of larvae of A. bipunctata for individual aphids was not dependent on the size of branches of apple trees varying in leaf surface area from 150 cm² to 960 cm². Fifty and 70% of individual aphids were found and killed 6 hours and 48 hours, respectively, after release of single second instar larva of A. bipunctata. In a first field cage experiment in 1996, A. bipunctata, and to a lesser extent E. balteatus, proved to be effective and consistent predators of D. plantaginea during spring conditions, being little affected by cool temperatures and wet weather. In a subsequent field cage experiment in 1997, larvae of A. bipunctata and E. balteatus were released singly and in combination on aphid infested apple seedlings to study interactions between these two promising control agents. Both species had a significant negative effect on aphid population increase. The two species did not significantly interact and thus, their joint effect is best explained by an additive model. Combined releases of the two predator species reduced aphid densities to 5% of the control. This indicates the potential for augmentative releases of these native aphid predators to control D. plantaginea.     

Mortality factors affecting immature stages of codling moth,Cydia pomonella (Lepidoptera: Tortricidae), and the impact of parasitoid complex

Codling moth, Cydia pomonella Linnaeus (Lepidoptera: Tortricidae), is a serious pest of apples worldwide. This study aimed to evaluate the mortality rate of codling moth eggs, larvae and pupae in the field in commercial and neglected apple and walnut orchards over two years, and to investigate the biodiversity and intensity of parasitoids associated with codling moth in the orchards. Five patches of wax paper containing 1-day-old codling moth eggs were placed in a neglected orchard in order to evaluate parasitism rates. Corrugated cardboard bands were placed around the trunk of 15 trees during late spring and the beginning of summer through to fruiting season to capture and measure parasitism of codling moth larvae. 5285 larvae in total were collected during this study. Mortality rate (egg?+?larvae?+?pupae) varied between the commercial and neglected orchards, reaching a maximum of (42.89% and 66.67%) in neglected apple orchards and (61.03% and 74.76%) in the neglected walnut orchard in 2003 and 2004, respectively. Trichogramma cacoeciae (Hymenoptera: Tichogrammatidae) was the only egg parasitoid recorded. Eight hymenopteran larval and pupal parasitoids belonging to several subfamilies were recorded: Cheloninae, Agathidinae, Cremastinae, Haltichellinae, Chalcidinae, Anomalinae, and Pteromalinae and one dipteran belonging to Tachininae. In conclusion, mortality factors, mainly by parasitoids, are contributing to a general reduction in codling moth larvae populations particularly in neglected orchards. The hymenopteran Ascogaster quadridentata and the dipteran Neoplectops pomonellae can contribute to biological control programmes against codling moth in the coastal region and other regions.     

Behavioural responses of the aphid parasitoid Diaeretiella rapae to volatiles from Arabidopsis thaliana induced by Myzus persicae

In response to herbivory by insects, several plant species have been shown to produce volatiles that attract the natural enemies of those herbivores. Using a Y‐tube olfactometer, we investigated responses of the aphid parasitoid Diaeretiella rapae MacIntosh (Hymenoptera: Aphidiidae) to volatiles from Arabidopsis thaliana Columbia (Brassicaceae) plants that were either undamaged, infested by the peach‐potato aphid, Myzus persicae Sulzer (Homoptera: Aphididae), or mechanically damaged, as well as to volatiles from just the aphid or its honeydew. In dual‐choice experiments, female D. rapae given oviposition experience on A. thaliana infested with M. persicae were significantly attracted to volatiles from A. thaliana infested with M. persicae over volatiles from undamaged A. thaliana and similarly were significantly attracted to plants that had been previously infested by M. persicae, but from which the aphids were removed, over undamaged plants. Diaeretiella rapae did not respond to volatiles from M. persicae alone, their honeydew, or plants mechanically damaged with either a pin or scissors. We conclude that an interaction between the plant and the aphid induces A. thaliana to produce volatiles, which D. rapae can learn and respond to. Poor responses of D. rapae to volatiles from an A. thaliana plant that had two leaves infested with M. persicae, with the two infested leaves being removed before testing, suggested the possibility that, at this stage of infestation, the majority of volatile production induced by M. persicae may be localized to the infested tissues of the plant. We conclude that this tritrophic interaction is a suitable model system for future investigations of the biochemical pathways involved in the production of aphid‐induced volatiles attractive to natural enemies.     

Role of ants in structuring the aphid community on apple

1. The aphids Dysaphis plantaginea Passerini, Aphis spp. (Aphis pomi De Geer and Aphis spiraecola Patch), and Eriosoma lanigerum Hausmann are commonly found together in apple orchards. Ants establish a mutualistic relationship with the myrmecophilous aphids D. plantaginea and Aphis spp. but not with E. lanigerum. 2. Field surveys and one experiment manipulating the presence of ants and the aphid species were conducted to test the hypothesis that ants play a role in structuring the community of these aphids on apple. 3. Ants tended D. plantaginea and Aphis spp. but not E. lanigerum colonies. In the field, D. plantaginea performed better in the presence of ants while no effect was observed in Aphis spp. Contrarily, populations of Aphis spp. in the manipulative experiment performed better in the presence of ants while no differences were observed for D. plantaginea. Such differences between field and manipulative conditions could be related to thermal tolerance, phenology, and life cycles. In contrast, populations of E. lanigerum were reduced in the presence of ants. 4. Ants also had a significant negative effect on the abundance of natural enemies, which could partially explain the benefits to the tended aphids. However, while ants did not provide a benefit to Aphis spp. when it was reared alone, in the presence of other species ant attendance increased Aphis abundance by 256% and simultaneously reduced E. lanigerum abundance by 63%. Therefore, ants benefited Aphis by reducing competition with other aphid species, which involves a different mechanism, explaining the benefit of ant attendance. Considering all the aphid species together, ants had a net positive effect on aphid abundance, which was consequently considered harmful for the plant. 5. Our results highlighted the role that ants play in structuring apple aphid communities and give support to the observed pattern that ants can benefit tended aphids while simultaneously reducing the abundance of untended herbivores.