The impact of an ant–aphid mutualism on the functional composition of the secondary parasitoid community

1. Mutualistic and antagonistic interactions, although often studied independently, may affect each other, and food web dynamics are likely to be determined by the two processes working in concert. 2. The structure, and hence dynamics, of food webs depends on the relative abundances of generalist and specialist feeding guilds. Secondary parasitoids of aphids can be divided into two feeding guilds: (i) the more specialised endoparasitoids, which attack the primary parasitoid larvae in the still living aphid, and (ii) the generalist ectoparasitoids, which attack the pre‐pupa of the primary or secondary parasitoid in the mummified aphid. 3. We studied the effect of an ant–aphid mutualism on the relative abundance of these two functional groups of secondary parasitoids. We hypothesised that generalists will be negatively affected by the presence of ants, thus leading to a greater dominance of specialists. 4. We manipulated the access of ants (Lasius niger) to aphid colonies in which we placed parasitised aphids. Aphid mummies were collected and reared to determine the levels of endo‐ and ecto‐secondary parasitism. 5. When aphids were attended by L. niger the proportion of secondary parasitism by ectoparasitoids dropped from 26 to 8% of the total number of parasitised aphids, with Pachyneuron aphidis most strongly affected, while endoparasitoids as a group did not respond. However, among these Syrphophagus mamitus profited from ant attendance becoming the dominant secondary parasitoid, while parasitisation rates of Alloxysta and Phaenoglyphis declined. 6. The shift to S. mamitus as dominant secondary parasitoid in ant‐attended aphid colonies is likely due to the behavioural plasticity of this species in response to ant aggression, and a release from tertiary parasitism by generalist ectoparasitoids. 7. The reduction of secondary parasitism by generalist ectoparasitoids reduces the potential for apparent competition among primary parasitoids with consequences for the dynamics of the wider food web.     

Host specialization in habitat specialists and generalists

Generalists and specialists use different cues to find their habitat and essential resources. While generalists have the advantage of exploiting a wider range of resources, they are predicted to be less efficient in using one particular resource compared to specialists. The level of specialization of parasitoids can be either at the habitat or at the host level; strategies used by either type are expected to differ. We examined interactions between three aphid parasitoid species that are a habitat specialist Aphidius rhopalosiphi, a habitat generalist Aphidius ervi, and a host generalist Praon volucre on three cereal aphids, Sitobion avenae, Metopolophium dirhodum and Rhopalosiphum padi. We compared total parasitism rate across behavioral and physiological variation in a non-choice test. Next, we addressed total parasitism in two phases to examine: (1) the response of parasitoids to different hosts through the behavioral sequence from antennation through oviposition, and (2) the physiological suitability of different hosts for oviposition and larval development. Parasitization typically involved the following behavioral steps: (1) antennal contact, (2) abdominal bending, and (3) ovipositor insertion (acceptance). A. rhopalosiphi had the same number of antennal contacts with the three aphids but showed fewer instances of abdominal bending towards R. padi. Pre-contact host preference was found for A. ervi but it did not correspond to the level of acceptance. The number of antennal contacts by P. volucre corresponded to the parasitization level of the aphid species but more mummies were produced on M. dirhodum than on R. padi. These results suggest that parasitoid species that are habitat specialists react similarly to the different host species present in the same habitat, whereas generalist species exhibit clear preferences during host selection. Preferences were, however, not always related to host suitability.     

不同农业景观结构对麦蚜种群动态的影响

研究表明农业景观结构的复杂性与害虫种群发生强度关系密切,然而在不同农业景观结构下研究麦蚜的发生、种群及寄生蜂的变化还不多。设计了不同的麦田景观结构,调查研究了不同麦田景观结构对麦蚜种群的影响。在简单与复杂两种农业景观结构下,分析了不同种类麦蚜的入田时间、入田量、种群增长率、种群密度及寄生性天敌的多样性与寄生率。结果表明:景观结构对不同种类麦蚜影响不同,但复杂农业景观下麦蚜迁飞入田时间都要晚于简单农业景观(连片种植)下的入田时间,复杂农业景观下有翅蚜的迁入量显著低于简单景观下有翅蚜的迁入量,并且复杂农业景观下麦蚜种群增长速率高于简单农业景观下的增长速率。不同种类麦蚜对景观结构的不同反应可能与形态学与生活史特征有关,两种不同农业景观结构下寄生性天敌的多样性与寄生率无显著差异。复杂景观结构下的麦蚜有翅蚜低的迁入量、高的增长速率可能与生境高度破碎化有关,其中与温室大棚塑料白色反光有的很大的影响。生境破碎化影响了麦蚜对寄主植物寻找以及天敌对猎物的寻找效应。     

Drought and Root Herbivory Interact to Alter the Response of Above-Ground Parasitoids to Aphid Infested Plants and Associated Plant Volatile Signals

Multitrophic interactions are likely to be altered by climate change but there is little empirical evidence relating the responses of herbivores and parasitoids to abiotic factors. Here we investigated the effects of drought on an above/below-ground system comprising a generalist and a specialist aphid species (foliar herbivores), their parasitoids, and a dipteran species (root herbivore).We tested the hypotheses that: (1) high levels of drought stress and below-ground herbivory interact to reduce the performance of parasitoids developing in aphids; (2) drought stress and root herbivory change the profile of volatile organic chemicals (VOCs) emitted by the host plant; (3) parasitoids avoid ovipositing in aphids feeding on plants under drought stress and root herbivory. We examined the effect of drought, with and without root herbivory, on the olfactory response of parasitoids (preference), plant volatile emissions, parasitism success (performance), and the effect of drought on root herbivory. Under drought, percentage parasitism of aphids was reduced by about 40–55% compared with well watered plants. There was a significant interaction between drought and root herbivory on the efficacy of the two parasitoid species, drought stress partially reversing the negative effect of root herbivory on percent parasitism. In the absence of drought, root herbivory significantly reduced the performance (e.g. fecundity) of both parasitoid species developing in foliar herbivores. Plant emissions of VOCs were reduced by drought and root herbivores, and in olfactometer experiments parasitoids preferred the odour from well-watered plants compared with other treatments. The present work demonstrates that drought stress can change the outcome of interactions between herbivores feeding above- and below-ground and their parasitoids, mediated by changes in the chemical signals from plants to parasitoids. This provides a new insight into how the structure of terrestrial communities may be affected by drought.     

The tri-trophic interactions hypothesis: interactive effects of host plant quality, diet breadth and natural enemies on herbivores

Several influential hypotheses in plant-herbivore and herbivore-predator interactions consider the interactive effects of plant quality, herbivore diet breadth, and predation on herbivore performance. Yet individually and collectively, these hypotheses fail to address the simultaneous influence of all three factors. Here we review existing hypotheses, and propose the tri-trophic interactions (TTI) hypothesis to consolidate and integrate their predictions. The TTI hypothesis predicts that dietary specialist herbivores (as compared to generalists) should escape predators and be competitively dominant due to faster growth rates, and that such differences should be greater on low quality (as compared to high quality) host plants. To provide a preliminary test of these predictions, we conducted an empirical study comparing the effects of plant (Baccharis salicifolia) quality and predators between a specialist (Uroleucon macolai) and a generalist (Aphis gossypii) aphid herbivore. Consistent with predictions, these three factors interactively determine herbivore performance in ways not addressed by existing hypotheses. Compared to the specialist, the generalist was less fecund, competitively inferior, and more sensitive to low plant quality. Correspondingly, predator effects were contingent upon plant quality only for the generalist. Contrary to predictions, predator effects were weaker for the generalist and on low-quality plants, likely due to density-dependent benefits provided to the generalist by mutualist ants. Because the TTI hypothesis predicts the superior performance of specialists, mutualist ants may be critical to A. gossypii persistence under competition from U. macolai. In summary, the integrative nature of the TTI hypothesis offers novel insight into the determinants of plant-herbivore and herbivore-predator interactions and the coexistence of specialist and generalist herbivores.     

Virus mediated trophic interactions between aphids and their natural enemies

Microbial endosymbionts alter the phenotype of their host which may have cascading effects at both population and community levels. However, we currently lack information on whether the effects of viruses on both host phenotypic traits and host population demography can modify interactions with upper trophic levels. To fill this gap, we investigated whether a prevalent densovirus infecting the aphid Myzus persicae (i.e. MpDNV) can modify trophic interactions between host aphids and their natural enemies (i.e. predators and parasitoids) by influencing aphid phenotypic traits (i.e. body mass and defensive behaviours), population demography (i.e. density and age-structure) and susceptibility towards both predation and parasitism. We found that the virus decreased aphid body mass but did not influence their behavioural defences. At the population level, the virus had a minor effect on aphid adult mortality whereas it strongly reduced the density of nymphs and influenced the stage structure of aphid populations. In addition, the virus enhanced the susceptibility of aphids to parasitism regardless of the parasitoid species. Predation rate on adult aphids was not influenced by the virus but ladybeetle predators strongly decreased the number of aphid nymphs, especially for uninfected ones compared to infected ones. As a result, the virus decreased predator effect on aphid populations. By reducing both aphid quality and availability, increasing their susceptibility to parasitism, and modulating predator effect on aphid populations, we highlighted that viral endosymbionts can be prevalent drivers of their host ecology as they modify their phenotypes and interspecific interactions. These virus-mediated ecological effects may have consequences on enemies foraging strategies as well as trophic webs dynamics and structure.     

Effects of inter-annual landscape change on interactions between cereal aphids and their natural enemies

The agricultural intensification and the subsequent habitat changes in agroecosystem can strongly affect biological control services. We here examine the influence of inter-annual landscape change in wheat field area on interactions of cereal aphids and their natural enemies, as well as the efficacy of biological control using data collected from a 4-year experiment in Northwest China. Two hypotheses were tested. (i) Population densities decline following an inter-annual expansion of wheat crop proportion cover due to dilution and crowding effects. (ii) Species that are specialists or at higher trophic levels are more sensitive to bottom-up disturbance by inter-annual change in percent cover of wheat crop. Results showed the population densities of one cereal aphid (Macrosiphum avenae), one parasitic wasp (Aphidius avenae), two specialist predators (ladybirds: Hippodamia variegata and H. tredecimpunctata) and one hyperparasitic wasp (Pachyneuron aphidis) declined following the expansion of wheat crop areas, supporting the predictions of inter-annual dilution and crowding effects. In contrast, the populations of one cereal aphid (Schizaphis graminum), one parasitic wasp (A. gifuensis), two generalist predators (spiders: Pardosa astrigera; carabid beetles: Chlaenius pallipes), and two hyperparasitic wasps (Asaphes suspensus, and Alloxysta sp.) did not respond to inter-annual landscape change. The two hypotheses were partially supported but with noticeable exceptions, and the bio-control efficiency declined with the increase of the proportion cover of wheat field in agricultural landscape. Overall, different responses of cereal aphids and their natural enemies make it difficult but still possible to optimize inter-annual landscape change for enhancing the parasitism rate and predator-prey ratio.     

麦蚜和寄生蜂对农业景观格局的响应及其关键景观因子分析

大量研究表明多样性的农业景观格局能够影响蚜虫及寄生蜂的分布。本文利用并设计了两种尺度的麦田农业景观格局,依据麦蚜种群发生特点,分为迁入期、增长期与高峰期三个时期,论述了不同尺度下农业景观元素对麦蚜及寄生蜂系统的影响,通过逐步回归筛选了不同时期麦蚜及寄生蜂分布的关键景观元素,最后通过CANOCO软件模拟了麦蚜及寄生蜂的分布排序格局。结果表明不同景观因子对麦蚜及寄生蜂种群影响不同,迁入期两种尺度下裸地最有利于两种有翅蚜的迁飞入田,塑料大棚对有翅蚜种群的入田有抑制作用。增长期草地与林地生境对麦蚜种群增长率促进最大,塑料大棚同样抑制了麦蚜的种群增长率;但塑料大棚、草地与林地对蚜茧蜂的种群增长率有促进作用,非麦类作物生境对蚜茧蜂种群增长率抑制作用最大;草地与林地有利于重寄生蜂的种群增长率;高峰期裸地比例大的农业景观下麦蚜的最大种群密度较大,草地与林地对蚜茧蜂与重寄生蜂的最大种群密度均有促进作用。两种尺度下的研究结果一致。不同麦物种的对不同景观元素反应与形态学与生活史特征有关,而且景观结构中特定的植物种类、非作物植物的密度与物候期都可能影响寄生蜂群落的多样性与功能。     

Food web structure and biocontrol in a four-trophic level system across a landscape complexity gradient

Decline in landscape complexity owing to agricultural intensification may affect biodiversity, food web complexity and associated ecological processes such as biological control, but such relationships are poorly understood. Here, we analysed food webs of cereal aphids, their primary parasitoids and hyperparasitoids in 18 agricultural landscapes differing in structural complexity (42-93% arable land). Despite little variation in the richness of each trophic group, we found considerable changes in trophic link properties across the landscape complexity gradient. Unexpectedly, aphid-parasitoid food webs exhibited a lower complexity (lower linkage density, interaction diversity and generality) in structurally complex landscapes, which was related to the dominance of one aphid species in complex landscapes. Nevertheless, primary parasitism, as well as hyperparasitism, was higher in complex landscapes, with primary parasitism reaching levels for potentially successful biological control. In conclusion, landscape complexity appeared to foster higher parasitism rates, but simpler food webs, thereby casting doubt on the general importance of food web complexity for ecosystem functioning.     

Top-down and bottom-up effects on the spatiotemporal dynamics of cereal aphids: testing scaling theory for local density

The relationship between density and area depends on local growth rates and the area-dependence of migration rates. These rates vary among taxa due to dispersal behaviour, plot productivity and natural enemy impact. Previous studies in aphids suggest that aphid densities are highest in patches of intermediate sizes, and lower in small and large patches. The suggested mechanism causing these patterns is that the dispersal behaviour in aphids creates a mixture of area- and perimeter-dependent migration rates. In this paper, we used these predictions to examine the additional consequences of nutrient availability and natural enemies on the density-area relationship. The derived predictions were compared to data from a system with three aphid species, a set of aphid parasitoids and generalist natural enemies, and at two levels of plant nutrient availability. We find that predictions from the model based only on dispersal and local growth agree with the temporal dynamics of density-area relationships for aphids in high nutrient patches. In patches with low nutrients, high parasitism rates appeared to cause a negative density-area relationship for aphids, thereby deviating from predictions driven by the aphids' dispersal behavior. Hence, the dispersal model with scale-dependent migration rates can provide a useful tool for understanding insect distribution in patch size gradients, but the relative importance of top-down effects can completely change with plot productivity.     

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.     

Increased nitrogen fertilization inhibits the biocontrol activity promoted by the intercropping partner plant

The examination of the compatibility between agricultural practices and biocontrol activities is crucial for establishing an efficient, eco-friendly, and sustainable pest management program. In this study, we examined the population dynamics of two specialist aphids, the English grain aphid (Sitobion avenae) on potted wheat and the pea aphid (Acyrthosiphon pisum) on potted alfalfa, as well as the biocontrol activity of a generalist predator, the harlequin ladybird beetle (Harmonia axyridis). We investigated their responses to the presence of the intercropping partner plant species (alfalfa and wheat, respectively) through plant volatiles or visual cues at three nitrogen fertilizer levels in a greenhouse. In the absence of the predator, the English grain aphid population growth rate increased significantly with increasing nitrogen levels, whereas the pea aphid population increased significantly more slowly in response to high nitrogen levels. The English grain aphid and pea aphid population dynamics were unaffected by the presence of the intercropping partner. However, the presence of the intercropping partner enhanced the control of both aphid populations by the harlequin ladybird beetle. Increasing nitrogen fertilizer levels decreased the predation rates, which were otherwise increased by the intercropping partner. The beneficial effects of the intercropping partner were eventually non-existent at the highest nitrogen level tested. These results imply that the interaction between the presence of intercropping partner and the nitrogen fertilizer application affects the biocontrol activity of the natural enemies of insect pests. Thus, the compatibility between agricultural intensification and biocontrol strategies in integrated pest management programs need to be investigated.     

Monoclonal antibodies reveal the potential of the tetragnathid spider Pachygnatha degeeri (Araneae: Tetragnathidae) as an aphid predator

The drive towards a more sustainable and integrated approach to pest management has engendered a renewed interest in conservation biological control, the role of natural enemy communities and their interactions with prey. Monoclonal antibodies have provided significant advances in enhancing our knowledge of trophic interactions and can be employed to help quantify predation on target species. The tetragnathid spider Pachygnatha degeeri Sundevall was collected from fields of winter wheat in the UK and assayed by ELISA for aphid proteins. It was demonstrated that this spider did not simply consume greater quantities of aphids because it was bigger. In addition, P. degeeri contained significantly greater concentrations of aphid in their guts than other spiders, showing that aphids comprised a greater proportion of their diet. Although P. degeeri constituted only 6% of the spider population numerically, females and males respectively contained 16% and 37% of total aphid proteins within all spiders screened, significantly more than their density would predict. These spiders also preyed upon aphids at a disproportionately high rate in June, during the aphid establishment phase, theoretically the best time for limiting growth in the aphid population. Although less abundant than other generalist predators, the capability of these hunting spiders to consume large numbers of aphids highlights them as a more significant component of the predator complex than had previously been realized. Limitation of aphid numbers early in the year by generalist predators provides more time for the specialist aphid predators and parasitoids to move in.     

Aphid population dynamics: does resistance to parasitism influence population size?

1. Insect population size is regulated by both intrinsic traits of organisms and extrinsic factors. The impacts of natural enemies are typically considered to be extrinsic factors, however insects have traits that affect their vulnerability to attack by natural enemies, and thus intrinsic and extrinsic factors can interact in their effects on population size. 2. Pea aphids Acyrthosiphon pisum Harris (Hemiptera: Aphididae) in New York and Maryland that are specialised on alfalfa are approximately two times more physiologically resistant to parasitism by Aphidius ervi Haliday (Hymenoptera: Braconidae) than pea aphids specialised on clover. To assess the potential influence of this genetically based difference in resistance to parasitism on pea aphid population dynamics, pea aphids, A. ervi, and other natural enemies of aphids in clover and alfalfa fields were sampled. 3. Rates of successful parasitism by A. ervi were higher and pea aphid population sizes were lower in clover, where the aphids are less resistant to parasitism. In contrast, mortality due to a fungal pathogen of pea aphids was higher in alfalfa. Generalist aphid predators did not differ significantly in density between the crops. 4. To explore whether intrinsic resistance to parasitism influences field dynamics, the relationship between resistance and successful field parasitism in 12 populations was analysed. The average level of resistance of a population strongly predicts rates of successful parasitism in the field. The ability of the parasitoid to regulate the aphid may vary among pea aphid populations of different levels of resistance.     

Diet breadth and its relationship with genetic diversity and differentiation: the case of southern beech aphids (Hemiptera: Aphididae)

Herbivorous insect species with narrow diet breadth are expected to be more prone to genetic differentiation than insect species with a wider diet breadth. However, a generalist can behave as a local specialist if a single host-plant species is locally available, while a specialist can eventually behave as a generalist if its preferred host is not available. These problems can be addressed by comparing closely related species differing in diet breadth with overlapping distributions of insect and host populations. In this work, diet breadth, genetic diversity and population differentiation of congeneric aphid species from southern beech forests in Chile were compared. While at the species level no major differences in genetic diversity were found, a general trend towards higher genetic diversity as diet breadth increased was apparent. The aphid species with wider diet breadth, Neuquenaphis edwardsi (Laing), showed the highest genetic diversity, while the specialist Neuquenaphis staryi Quednau & Remaudière showed the lowest. These differences were less distinct when the comparisons were made in the same locality and over the same host. Comparison of allopatric populations indicates that genetic differentiation was higher for the specialists, Neuquenaphis similis Hille Ris Lambers and N. staryi, than for the generalist N. edwardsi. Over the same host at different locations, genetic differentiation among populations of N. edwardsi was higher than among populations of N. similis. The results support the assumption that specialists should show more pronounced genetic structuring than generalists, although the geographical distribution of host plants may be playing an important role.     

Biodiversity and biocontrol: emergent impacts of a multi-enemy assemblage on pest suppression and crop yield in an agroecosystem

The suppression of agricultural pests has often been proposed as an important service of natural enemy diversity, but few experiments have tested this assertion. In this study we present empirical evidence that increasing the richness of a particular guild of natural enemies can reduce the density of a widespread group of herbivorous pests and, in turn, increase the yield of an economically important crop. We performed an experiment in large field enclosures where we manipulated the presence/absence of three of the most important natural enemies (the coccinellid beetle Harmonia axyridis, the damsel bug Nabis sp., and the parasitic wasp Aphidius ervi) of pea aphids (Acyrthosiphon pisum) that feed on alfalfa (Medicago sativa). When all three enemy species were together, the population density of the pea aphid was suppressed more than could be predicted from the summed impact of each enemy species alone. As crop yield was negatively related to pea aphid density, there was a concomitant non‐additive increase in the production of alfalfa in enclosures containing the more diverse enemy guild. This trophic cascade appeared to be influenced by an indirect interaction involving a second herbivore inhabiting the system – the cowpea aphid, Aphis craccivora. Data suggest that high relative densities of cowpea aphids inhibited parasitism of pea aphids by the specialist parasitoid, A. ervi. Therefore, when natural enemies were together and densities of cowpea aphids were reduced by generalist predators, parasitism of pea aphids increased. This interaction modification is similar to other types of indirect interactions among enemy species (e.g. predator–predator facilitation) that can enhance the suppression of agricultural pests. Results of our study, and those of others performed in agroecosystems, complement the broader debate over how biodiversity influences ecosystem functioning by specifically focusing on systems that produce goods of immediate relevance to human society.     

Entomopathogenic fungi stimulate transgenerational wing induction in pea aphids,Acyrthosiphon pisum (Hemiptera: Aphididae)

1. Aphid natural enemies include not only predators and parasitoids but also pathogens, of which fungi are the most studied for biological control. While wing formation in aphids is induced by abiotic conditions, it is also affected by biotic interactions with their arthropod natural enemies. Wing induction via interactions with arthropod natural enemies is mediated by the increase in their physical contact when alarmed (pseudo‐crowding). Pathogenic fungi do not trigger this alarm behaviour in aphids and, therefore, no pseudo‐crowding occurs. 2. We hypothesise that, while pathogenic fungi will stimulate maternally induced wing formation, the mechanism is different and is influenced by pathogen specificity. We tested this hypothesis using two entomopathogenic fungi, Pandora neoaphidis and Beauveria bassiana, an aphid specialist and a generalist respectively, on the pea aphid, Acyrthosiphon pisum Harris. 3. We first demonstrate that pea aphids infected with either pathogen and maintained in groups on broad bean plants produced a higher proportion of winged morphs than uninfected control aphids. We then show that, when maintained in isolation, aphids infected with either pathogen also produced higher proportions of winged offspring than control aphids. There was no difference between P. neoaphidis and B. bassiana in their effects on wing induction in either experiment. 4. Unlike the effect of predators and parasitoids on pea aphid wing induction, the effect of pathogens is independent of physical contact with other aphids, suggesting that physiological cues induce wing formation in infected aphids. It is possible that aphids benefit from wing induction by escaping infected patches whilst pathogens may benefit through dispersion. Possible mechanisms of wing induction are discussed.     

Density-dependent parasitism of cowpea aphid,Aphis craccivora by Lysiphlebus fabarum,Binodoxys acalephae,and Aphidius matricariae (Hymenoptera: Braconidae: Aphidiinae)

Biological control, as a major component of pest management strategies, uses natural biological agents to reduce pest populations. Studying the interaction among Aphis craccivora and its parasitoids including, Lysiphlebus fabarum, Binodoxys acalephae, and Aphidius matricariae in 2016 and 2017 in Tehran Parke-Shahr, showed positive, significant correlations in all cases between the densities of three parasitoid species and that of aphid nymphs and adults. The density of the parasitoids increased by increasing the density of the aphids. The parasitoids showed aggregative behavior in response to different densities of the host. There was a positive density-dependent correlation between the density of A. craccivora and rate of parasitism. Parasitism rates of nymphs and adult aphids by L. fabarum, B. acalephae, and A. matricariae increased or decreased along with decline or increase in the population of the aphid host. In 2016 spring, the highest rates of parasitism on aphid nymphs by L. fabarum, B. acalephae, and A. matricariae were 46.82, 23.09, and 17.16%, respectively. In 2017 spring, the highest rates of parasitism on aphid nymphs by L. fabarum, B. acalephae, and A. matricariae were 48.97, 21.77, and 15.06%, respectively. So, given the accordance between changes in aphid population and that of parasitoids, and parasitoids’ efficacy in Tehran’s polluted air, they can be used as biological agents in the management of A. craccivora population.     

On the role of sinigrin (mustard oil) in a tritrophic context: plant–aphid–aphidophagous hoverfly

1. Plant secondary metabolites can govern prey–predator interactions by altering the diet breadth of predators and sometimes provide an ecological refuge to prey. Brassicaceae plants and their specialist pests can be used as a model system for understanding the role of chemically mediated effects restricting the diet breadth of natural enemies, and consequently the occurrence of enemy‐free space for the specialist pest. 2. The objective of the present study was to test the performance of the generalist predator Episyrphus balteatus De Geer (Diptera: Syrphidae) fed on the specialist herbivore Brevicoryne brassicae L.(Homoptera: Aphididae), reared on two different brassica species: black mustard (Brassica nigra), a wild species with high levels of sinigrin; and canola (Brassica napus), a cultivated species without sinigrin. 3. The preference and performance of the predator and the performance of the prey were measured. Sinigrin was quantified by high‐performance liquid chromatography in both leaf samples and aphids reared on the two host plants. 4. The cabbage aphid performed better on canola than on black mustard. The performance of the predator on this aphid when reared on canola was clearly better than when reared on black mustard. Females had a higher overall preference for cabbage aphids reared on canola than on black mustard. 5. The ability of aphids reared on plants with high glucosinolate content to reduce the performance of their generalist predators indicates that the presence of B. nigra may provide enemy‐free space for the cabbage aphid from its predator, a concept that has useful application in the context of biological control for agricultural systems.     

Evidence for a trade-off between host-range breadth and host-use efficiency in aphid parasitoids

"The jack-of-all-trades is a master of none" describes the widely held belief that engaging in many tasks comes at the cost of being unable to do those tasks well. However, empirical evidence for generalist fitness costs remains scarce. We used published data from a long-term field survey of aphid parasitoids to determine whether relative specialists are more abundant than generalists on their shared hosts, a pattern that would be expected if generalists suffer a trade-off between host-range breadth and host-use efficiency. Relative specialists were more abundant than generalists on their shared hosts, but only when we used a measure of specialization that accounts for the taxonomic differences among parasitoids' hosts. These results suggest that a generalist-specialist trade-off exists within this group of parasitoids and that the generalist fitness cost depends on the taxonomic breadth, rather than the number, of host species that are used.