Effects of a simple plant morphological mutation on the arthropod community and the impacts of predators on a principal insect herbivore

Plant features that enhance predator effectiveness can be considered extrinsic-resistance factors because they result in reduced insect herbivory. In this paper we test the hypothesis that reduced epicuticular wax (EW) in Pisum sativum L. is an extrinsic-resistance factor contributing to field resistance to Acyrthosiphon pisum (Harris). We monitored pea aphid populations in the field on reduced EW and normal EW near isolines of peas for two seasons and confirmed that aphid populations are lower on reduced EW peas than on normal EW peas. We also monitored predators within the canopies of the two pea lines to discover community level patterns in response to differences in EW. We found that while predator numbers were similar between the two lines, there were more syrphids on the normal EW peas, and a trend towards more coccinellids on reduced EW peas. We tested the impact of predators on pea aphids on the two EW lines by monitoring their population levels in cages that excluded predators, and in cages that allowed predators to enter. We found that pea aphid populations were similar on the two EW lines when predators were excluded. When predators were allowed access to the plants, pea aphid populations were reduced more on reduced EW peas than on normal EW peas. We also examined the intrinsic resistance to aphids in reduced EW peas with laboratory dual-choice tests comparing aphid response to reduced EW and normal EW peas, and found that walking, apterous aphids displayed no preference for one pea line over the other. Bioassays to measure growth and fecundity of the pea aphid on the two EW types in the greenhouse and in the field showed that intrinsic rate of increase, and other life table parameters, were not different for aphids on the two lines. Together these results support the hypothesis that reduced EW in peas is a predator-dependent extrinsic resistance factor. Genetically reducing EW bloom in peas and other waxy crop plants might improve the effectiveness of arthropod natural enemies of insect pests. More generally, the results show that a subtle change in plant morphology can substantially influence the impact of predators on insect herbivore populations. The benefit of extrinsic resistance to herbivory conferred by reduced EW may balance any benefits of a prominent EW bloom, thereby sustaining EW polymorphisms in some natural plant populations.     

Plant waxy bloom on peas affects infection of pea aphids by Pandora neoaphidis

This study examined the effects of the surface wax bloom of pea plants, Pisum sativum, on infection of pea aphids, Acyrthosiphon pisum, by the fungal pathogen Pandora neoaphidis. In prior field surveys, a higher proportion of P. neoaphidis-killed pea aphids (cadavers) had been observed on a pea line with reduced wax bloom, as compared with a sister line with normal surface wax bloom. Laboratory bioassays were conducted in order to examine the mechanisms. After plants of each line infested with aphids were exposed to similar densities of conidia, the rate of accumulation of cadavers on the reduced wax line was significantly greater than on the normal wax bloom line; at the end of the experiment (13d), the proportion of aphid cadavers on the reduced wax line was approximately four times that on the normal wax bloom line. When plants were exposed to conidia first and then infested with aphids, the rate of accumulation of cadavers was slightly but significantly greater on the reduced wax line, and infection at the end of the experiment (16d) did not differ between the lines. When aphids were exposed first and then released onto the plants, no differences in the proportion of aphid cadavers were observed between the pea lines. Greater infection of pea aphid on reduced wax peas appears to depend upon plants being exposed to inoculum while aphids are settled in typical feeding positions on the plant. Additional experiments demonstrated increased adhesion and germination by P. neoaphidis conidia to leaf surfaces of the reduced wax line as compared with normal wax line, and this could help explain the higher infection rate by P. neoaphidis on the reduced wax line. In bioassays using surface waxes extracted from the two lines, there was no effect of wax source on germination of P. neoaphidis conidia.     

Delineating the effects of a plant trait on interactions among associated insects

Plant traits can affect ecological interactions between plants, herbivores, and predators. Our study tests whether reduced leaf wax in peas alters the interaction between the pea aphid ( Acyrthosiphon pisum), a foliar-foraging predator (a lady beetle, Hippodamia convergens) and a ground-foraging predator (a ground beetle, Poecilus scitulus). We performed a 2×2×2 factorial experiment in which wax level, presence of H. convergens, and presence of P. scitulus were manipulated. Experimental arenas consisted of a cage surrounding three pea plants. One plant in each cage was stocked with 15 pea aphids. In greenhouse and field cage experiments, we assessed the effect of each factor and their interactions on aphid density. As in previous studies, H. convergens foraged for aphids more effectively on reduced wax peas than on normal peas. Other interactions among H. convergens, P. scitulus , and A. pisum were the same on both types of peas. We consider how aphid movement, plant growth, and a high frequency of predation by P. scitulus on H. convergens influenced pea aphid density.     

Waxy bloom in peas influences the performance and behavior of Aphidius ervi, a parasitoid of the pea aphid

Leaf surface waxy bloom can influence the predator–prey interactions that take place on peas. We tested whether the interaction between the pea aphid Acyrthosiphon pisum Harris (Homoptera: Aphidae) and a parasitoid, Aphidius ervi Haliday (Hymenoptera: Aphidiidae), is affected by reduced wax. We performed greenhouse experiments comparing aphid parasitism by individual A. ervi on two varieties of reduced wax peas to two normal wax sister varieties. We also observed the behavior of individual A. ervi in the greenhouse and measured field parasitism in small plots of reduced wax and normal wax peas. In the greenhouse, individual A. ervi parasitized more aphids on the reduced wax varieties than on their normal wax counterparts. Wasps spent more time actively foraging on reduced wax pea plants, which may contribute to the higher parasitism observed on those varieties. The greenhouse results suggested that the improved performance of individual A. ervi on reduced wax peas might contribute to a higher parasitism on reduced wax peas in the field. Field parasitism was significantly higher in reduced wax pea plots during 2000. Overall parasitism was higher in 2000 than in 2001 and 2002. In the latter years, parasitism was higher on reduced wax plants, but not significantly different from normal wax plants. Improved foraging by individual A. ervi resembles improved foraging by other carnivorous insects on reduced wax peas. The advantages of reduced wax for biological control of the pea aphid may hold when any of several different natural enemies is abundant.     

A plant surface mutation mediates predator interference among ladybird larvae

Abstract.  1. We investigated mechanisms causing predator–predator interference between fourth instar Hippodamia convergens larvae foraging for pea aphids on pea plants, Pisum sativum , with a wild-type wax bloom, and the lack of such interference between larvae foraging on pea plants with a reduced-wax bloom caused by the single gene mutation wel .
2. Observations showed that behavioural interactions between larvae were not affected by wax phenotype. Specifically, larvae did not encounter one another more frequently on normal-wax peas as may have been predicted because reduced ability by coccinellids to attach to normal-wax plant surfaces could restrict them to foraging on only some parts of these plants.
3. In a controlled bioassay on normal-wax peas, H. convergens larvae avoided leaflets previously exposed to another larva. On reduced-wax peas, this effect was not detected.
4. In microcosm experiments, inter-predator interference in terms of prey consumption occurred on normal-wax peas, but not on reduced-wax peas. The interference on normal-wax peas occurred whether two H. convergens larvae were placed on a pea aphid-infested, normal-wax plant simultaneously or sequentially.
5. We conclude that the observed inter-predator interference is not as a result of direct physical contact, but rather arises because of (i) inhibition of foraging by chemical trails left by other larvae, (ii) the inability of larvae to access portions of the normal-wax plants creating aphid refugia, or (iii) a combination of these factors.     

Non‐consumptive effects of a natural enemy on a non‐prey herbivore population

1. Predator–prey interactions have traditionally focused on the consumptive effects that predators have on prey. However, predators can also reduce the abundance of prey through behaviourally‐mediated non‐consumptive effects. For example, pea aphids (Acyrthosiphon pisum Harris) drop from their host plants in response to the risk of attack, reducing population sizes as a consequence of lost feeding opportunities. 2. The objective of the present study was to determine whether the non‐consumptive effects of predators could extend to non‐prey herbivore populations as a result of non‐lethal incidental interactions between herbivores and foraging natural enemies. 3. Polyculture habitats consisting of green peach aphids (Myzus persicae Sulzer) feeding on collards and pea aphids feeding on fava beans were established in greenhouse cages. Aphidius colemani Viereck, a generalist parasitoid that attacks green peach aphids but not pea aphids, was released into half of the cages and the abundance of the non‐host pea aphid was assessed. 4. Parasitoids reduced the population growth of the non‐host pea aphid by increasing the frequency of defensive drops; but this effect was dependent on the presence of green peach aphids. 5. Parasitoids probably elicited the pea aphid dropping behaviour through physical contact with pea aphids while foraging for green peach aphids. It is unlikely that pea aphids were responding to volatile alarm chemicals emitted by green peach aphids in the presence of the parasitoid. 6. In conclusion, the escape response of the pea aphid provided the opportunity for a parasitoid to have non‐target effects on an herbivore with which it did not engage in a trophic interaction. The implication is that natural enemies with narrow diet breadths have the potential to influence the abundance of a broad range of prey and non‐prey species via non‐consumptive effects.     

Attachment forces of pea aphids (Acyrthosiphon pisum) on different legume species

1. Sympatric populations of insects adapted to different host plants are good model systems not only to study how they adapt to the chemistry of their food plant, but also to investigate whether morphological modifications evolved enabling them to live successfully on a certain plant species. 2. The pea aphid, Acyrthosiphon pisum (Harris) encompasses at least 11 genetically distinct sympatric host races, each showing a preference for a certain legume species. The leaflet surfaces of these legumes differ considerably in their wax coverage. 3. It was investigated whether the attachment structures of three pea aphid genotypes from different host races are adapted to the different surface properties of their host plants and whether they show differences in their attachment ability on the respective host and non‐host plants. 4. The surface morphology of plants and aphid tarsi was examined using SEM (scanning electron microscopy). The ability of the aphids to walk on specific surfaces was tested using traction force measurements. 5. The presence of wax blooms on the leaflets lowers the aphids' attachment ability considerably and diminishes their subsequent attachment on ‘neutral’ surfaces like glass. The pea aphid host races differ in their ability to walk on certain surfaces. However, the genotype from the adapted aphid host race was not necessarily the one with the best walking performance on their host plant. All aphids, regardless of the original host plant, were most efficient on the neutral control surface glass. The general host plant Vicia faba was the plant with the most favourable surface for all aphid host races.     

Effects of leaf epicuticular wax on the movement, foraging behavior, and attack efficacy of Diaeretiella rapae

Plant morphological characters can affect the foraging abilities of natural enemies. Heavy wax blooms have been shown to impede predators searching for herbivores on various species in the genus Brassica (Brassicaceae). This study determined whether epicuticular wax affected the foraging behavior of the braconid wasp Diaeratiella rapae (M’Intosh) (Hymenoptera: Braconidae) as it searched for its aphid host Brevicoryne brassicae L. (Homoptera: Aphididae) on the leaves of cauliflower varieties with heavy or light wax blooms. Wasps on the variety with a heavier wax bloom foraged more slowly, groomed more often and for longer periods of time, fell from the leaves more often, took longer to find colonies of aphids, and attacked them at a lower rate than wasps foraging on the variety with a lighter wax bloom. When epicuticular wax was removed from the leaves, the wasp's foraging efficiency and efficacy improved significantly on the cauliflower variety with a heavy wax bloom. The amount of epicuticular wax present on a leaf was shown to impede the ability of a parasitoid to forage, locate, and attack its host.     

Influences of fertilization on Aphis gossypii and insecticide usage

Abstract: Fertilization levels for ornamental crops may influence pest population dynamics, crop quality, and pest management strategy. We examined the effect of fertilization on population growth and within‐plant distribution of melon or cotton aphid, Aphis gossypii Glover, on potted chrysanthemum, Dendranthema grandiflora (Tzvelev). In terms of pest management implications, we also investigated the effect of fertilization on the number of insecticide applications needed to control A. gossypii on potted chrysanthemum. Population growth rate of A. gossypii increased with fertilization levels from 0 to 38 ppm N and reached a plateau from 38 to 488 ppm N. Increased fertilization beyond 38 ppm N, 10% of the commercial standard, did not result in higher aphid number. Aphids responded to nutrient availability of plants by distributing themselves in areas with higher level of nitrogen. More aphids were found in the apical and middle strata of the plants than the basal stratum, which had the lowest nitrogen content. Leaf nitrogen content increased with increased fertilization level and was consistently higher in the apical and middle strata than the basal stratum. Increased fertilization from 0 to 375 ppm N did not result in higher number of insecticide applications. All three insecticides (bifenthrin, kinoprene or pymetrozine) were effective in keeping the aphid infestation below a pre‐determined level, five aphids per plant, but pymetrozine required the least number of applications. For chrysanthemum, a fast‐growing crop and heavy utilizer of nitrogen, increased fertilization shortened the time to flowering, which would allow growers to harvest their crop sooner and reduce the time for aphid population growth. Reduction in time to harvest could result in significant reduction of insecticide usage by reducing the time for aphid population growth. As a result, high fertilization together with minimal runoff may be a useful tactic to an integrated pest management (IPM) programme for managing A. gossypii on potted chrysanthemums.     

Effects of Aphis fabae Scop, and of its attendant ants and insect predators on yields of field beans (Vicia faba L.)

Predators (mainly coccinellid adults and larvae and syrphid larvae), although few, were important in decreasing numbers of Aphis fabae on a small plot of field beans during the early stages of infestation in a year favourable to the aphid. At the same time, ants (Lasius niger L.), attending aphids on other plants on the same plot, effectively protected the aphids from predators for about 2 weeks, enabling the attended aphids to multiply faster than the unattended. When all aphid populations started to decline, predators became more numerous and accelerated the decline on both sets of plants. Bean plants without aphids yielded fifty-six seeds per plant; those with aphids but free from ants gave seventeen; and those with ant-attended aphids, eight seeds per plant. The damage and loss of yield was caused by the large aphid populations that developed when the pods were maturing, and not by the fewer aphids present when the plants were in flower. It appears that small, temporary infestations during flowering might increase the yield of field beans.     

Erratum

Abstract: Fertilization levels for ornamental crops may influence pest population dynamics, crop quality, and pest management strategy. We examined the effect of fertilization on population growth and within‐plant distribution of melon or cotton aphid, Aphis gossypii Glover, on potted chrysanthemum, Dendranthema grandiflora (Tzvelev). In terms of pest management implications, we also investigated the effect of fertilization on the number of insecticide applications needed to control A. gossypii on potted chrysanthemum. Population growth rate of A. gossypii increased with fertilization levels from 0 to 38 ppm N and reached a plateau from 38 to 488 ppm N. Increased fertilization beyond 38 ppm N, 10% of the commercial standard, did not result in higher aphid number. Aphids responded to nutrient availability of plants by distributing themselves in areas with higher level of nitrogen. More aphids were found in the apical and middle strata of the plants than the basal stratum, which had the lowest nitrogen content. Leaf nitrogen content increased with increased fertilization level and was consistently higher in the apical and middle strata than the basal stratum. Increased fertilization from 0 to 375 ppm N did not result in higher number of insecticide applications. All three insecticides (bifenthrin, kinoprene or pymetrozine) were effective in keeping the aphid infestation below a pre‐determined level, five aphids per plant, but pymetrozine required the least number of applications. For chrysanthemum, a fast‐growing crop and heavy utilizer of nitrogen, increased fertilization shortened the time to flowering, which would allow growers to harvest their crop sooner and reduce the time for aphid population growth. Reduction in time to harvest could result in significant reduction of insecticide usage by reducing the time for aphid population growth. As a result, high fertilization together with minimal runoff may be a useful tactic to an integrated pest management (IPM) programme for managing A. gossypii on potted chrysanthemums.     

Biology and trophic interactions of lucerne aphids

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The interaction of soybean aphids and soybean cyst nematodes on selected resistant and susceptible soybean lines

Soybean aphids, Aphis glycines Matsumura, and soybean cyst nematodes, Heterodera glycines Ichinohe, are economic pests of soybean, Glycine max (L.) Merr., in the north‐central United States. Combined, these pests may account for 20–50% of yield reductions in a soybean crop. Only limited information is available concerning the interaction of these two pests on soybean production. During the summers of 2006 and 2007, we conducted a field‐experiment near Urbana, IL, to evaluate the effect of resistant and susceptible soybean lines on the development and reproduction of both pests in combination with each other. We also examined how each pest, as well as their interaction, affected the yield of susceptible and resistant soybean lines. Soybean plants grown within caged plots were infested with soybean aphids and soybean cyst nematodes; cumulative aphid days and soybean cyst nematode egg densities were determined at the end of each growing season. Soybean aphids were able to survive on all four soybean lines in both years of this study; however, aphid‐resistant lines generally had fewer cumulative aphid days than aphid‐susceptible lines. Likewise, nematode‐resistant lines typically had fewer eggs than nematode‐susceptible lines. During both years, we failed to observe a significant interaction between these two pests on the reproduction of one another. Yield data from 2006 was inconclusive; however, results from 2007 suggest that yield‐loss when soybean aphids and soybean cyst nematodes occur jointly is not significantly greater than when these two pests occur independently. The relationship between these two pests, and our inability to observe an interaction, are discussed.     

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.     

Predator avoidance behavior in the pea aphid: costs,frequency, and population consequences

Induced prey defenses can be costly. These costs have the potential to reduce prey survival or reproduction and, therefore, prey population growth. I estimated the potential for predators to suppress populations of pea aphids (Acyrthosiphon pisum) in alfalfa fields through the induction of pea aphid predator avoidance behavior. I quantified (1) the period of non-feeding activity that follows a disturbance event, (2) the effect of frequent disturbance on aphid reproduction, and (3) the frequency at which aphids are disturbed by predators. In combination, these three values predict that the disturbances induced by predators can substantially reduce aphid population growth. This result stems from the high frequency of predator-induced disturbance, and the observation that even brief disturbances reduce aphid reproduction. The potential for predators to suppress prey populations through induction of prey defenses may be strongest in systems where (1) predators frequently induce prey defensive responses, and (2) prey defenses incur acute survival or reproductive costs. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Genetic variation in the effect of a facultative symbiont on host-plant use by pea aphids

Ecological specialisation on different host plants occurs frequently among phytophagous insects and is normally assumed to have a genetic basis. However, insects often carry microbial symbionts, which may play a role in the evolution of specialisation. The bacterium Regiella insecticola is a facultative symbiont of pea aphids (Acyrthosiphon pisum) where it is found most frequently in aphid clones feeding on Trifolium giving rise to the hypothesis that it may improve aphid performance on this plant. A study in which R. insecticola was eliminated from a single naturally infected aphid clone supported the hypothesis, but a second involving two aphid clones did not find the same effect. We created a series of new pea aphid–R. insecticola associations by injecting different strains of bacteria into five aphid clones uninfected by symbionts. For all aphid clones, the bacteria decreased the rate at which aphids accepted Vicia faba as a food plant and reduced performance on this plant. Their effect on aphids given Trifolium pratense was more complex: R. insecticola negatively affected acceptance by all aphid clones, had no effect on the performance of four aphid clones, but increased performance of a fifth, thus demonstrating genetic variation in the effect of R. insecticola on pea aphid host use. We discuss how these results may explain the distribution and frequency of this symbiont across different aphid populations. Julia Ferrari and Claire L. Scarborough contributed equally to the work.     

Fitness effects of two facultative endosymbiotic bacteria on the pea aphid,Acyrthosiphon pisum,and the blue alfalfa aphid,A. kondoi

The effects of two bacterial endosymbionts, designated PASS and PAR, were evaluated on the pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera:Aphididae), in which they occur facultatively, and on the blue alfalfa aphid, A. kondoi Shinji, in which these bacteria have not been found in natural populations. Subclones of pea aphids and blue alfalfa aphids, derived from parent aphid clones that did not contain PASS or PAR, were infected with one or both bacteria, generating PASS- and/or PAR-positive subclones with minimal genetic differences from the parent clones. Under laboratory conditions at 20 °C, PAR consistently reduced the fecundity (by between 19 and 60%) of subclones derived from three different parent pea aphid clones on bur clover, Medicago hispida Gaertn. PAR had intermediate effects on pea aphids reared on sweet pea, Lathyrus odoratus L., and had no significant effect on pea aphids on alfalfa, Medicago sativa L. The effect of PASS was either neutral or negative, depending on parent clone as well as host plant. Also at 20 °C, PASS reduced fecundity (70–77%) and longevity (40–48%), and increased the age of first reproduction (by up to 1.5 days) of blue alfalfa aphid reared on alfalfa and clover. PAR had a less dramatic effect (e.g., 30–39% reduction in fecundity) on these traits of blue alfalfa aphid. In contrast, PAR and PASS increased the fitness of pea aphid subclones of one parent clone reared for three generations at 25 °C on each of the three test plants. Without facultative bacteria, fecundity of the parent clone was reduced to a mean total of < 6 offspring per adult at this elevated temperature, but with PASS or PAR, mean total fecundity of its subclones was > 35. However, this ameliorative effect of facultative bacteria at 25 °C was not found for two other sets of parent clones and their derived subclones. Alate production in pea aphids was significantly increased in large populations of two PASS- and PAR-positive subclones relative to their parent clones. Attempts to transmit PASS or PAR horizontally, i.e., from aphid to aphid via feeding on host plants (bur clover), were unsuccessful.     

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.     

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.     

Causes of regional and yearly variation in pea aphid numbers in eastern England

Regional variation in the number of pea aphids caught in the suction traps of the Rothamsted Insect Survey (RIS) was associated with the proportion of each region under pea crops. The degree of infestation of crops was similar in areas of high and low pea production as the mean annual abundance of aphids per hectare of crop remained constant. Yearly variation in abundance was loosely associated with temperature from January to July. Cold weather in January and February resulted in large numbers of aphids. Warm weather in February led to early colonization, and emigration from, pea crops as well as making early sowing of the crop more likely. A forecast of the time of first appearance of Acyrthosiphon pisum in the aerial plankton can be made, based on February temperature. Populations of A. pisum on peas appear to be regulated by alata production. High densities of aphids resulted in almost all the nymphs developing into alatae which, on reaching maturity, emigrated, causing populations on the crop to decline. This explains population crashes of the pea aphid observed at early growth stages of the crop, on vining and combining peas. Late sowing of peas, a probable effect of cold winters, results in higher aphid densities at flowering. The probable explanation for this is that late-sown crops are colonized at an earlier growth stage, so that the aphid population has a longer period of time in which to develop.