Effect of parasitism on flight behavior of the soybean aphid, Aphis glycines

Many aphid species possess wingless (apterous) and winged (alate) stages, both of which can harbor parasitoids at various developmental stages. Alates can either be parasitized directly or can bear parasitoids eggs or larvae resulting from prior parasitism of alatoid nymphs. Winged aphids bearing parasitoid eggs or young larvae eventually still engage in long-distance flights, thereby facilitating parasitoid dispersal. This may have a number of important implications for biological control of aphids by parasitoids. In this study, we determined the effect of parasitism by Aphelinus varipes (Hymenoptera: Aphelinidae) on wing development and flight of the soybean aphid, Aphis glycines (Hemiptera: Aphididae). We also quantified the influence of aphid flight distance on subsequent A. varipes development. Parasitism by A. varipes was allowed at different A. glycines developmental stages (i.e., alatoid 3rd and 4th-instar nymphs, alates) and subsequent aphid flight was measured using a computer-monitored flight mill. Only 35% of aphids parasitized as L3 alatoid nymphs produced normal winged adults compared to 100% of L4 alatoids. Flight performance of aphids parasitized as 4th-instar alatoid nymphs 24 or 48 h prior to testing was similar to that of un-parasitized alates of identical age, but declined sharply for alates that had been parasitized as 4th-instar alatoid nymphs 72 and 96 h prior to testing. Flight performance of aphids parasitized as alate adults for 24 h was not significantly different from un-parasitized alates of comparable ages. Flight distance did not affect parasitoid larval or pupal development times, or the percent mummification of parasitized aphids. Our results have implications for natural biological control of A. glycines in Asia and classical biological control of the soybean aphid in North America.     

Cucurbit aphid‐borne yellows virus (CABYV) modifies the alighting,settling and probing behaviour of its vector Aphis gossypii favouring its own spread

Plant pathogens are able to influence the behaviour and fitness of their vectors in such a way that changes in plant–pathogen–vector interactions can affect their transmission. Such influence can be direct or indirect, depending on whether it is mediated by the presence of the pathogen in the vector's body or by host changes as a consequence of pathogen infection. We report the effect that the persistently aphid‐transmitted Cucurbit aphid‐borne yellows virus (CABYV, Polerovirus) can induce on the alighting, settling and probing behaviour activities of its vector, the cotton aphid Aphis gossypii. Only minor direct changes on aphid feeding behaviour were observed when viruliferous aphids fed on non‐infected plants. However, the feeding behaviour of non‐viruliferous aphids was very different on CABYV‐infected than on non‐infected plants. Non‐viruliferous aphids spent longer time feeding from the phloem in CABYV‐infected plants compared to non‐infected plants, suggesting that CABYV indirectly manipulates aphid feeding behaviour through its shared host plant in order to favour viral acquisition. Viruliferous aphids showed a clear preference for non‐infected over CABYV‐infected plants at short and long time, while such behaviour was not observed for non‐viruliferous aphids. Overall, our results indicate that CABYV induces changes in its host plant that modifies aphid feeding behaviour in a way that virus acquisition from infected plants is enhanced. Once the aphids become viruliferous they prefer to settle on healthy plants, leading to optimise the transmission and spread of this phloem‐limited virus.     

Complex interactions between a plant pathogen and insect parasitoid via the shared vector-host: consequences for host plant infection

Plant viruses modify the development of their aphid vectors by inducing physiological changes in the shared host plant. The performance of hymenopterous parasitoids exploiting these aphids can also be modified by the presence of the plant pathogen. We used laboratory and glasshouse microcosms containing beans (Vicia faba) as the host plant to examine the interactions between a plant virus (pea enation mosaic virus; PEMV) and a hymenopterous parasitoid (Aphidius ervi) that share the aphid vector/host Acyrthosiphon pisum. Neither PEMV-infection of V. faba, nor the carriage of PEMV virions by A. pisum, affected the growth or morphology of the aphid, or the oviposition behaviour and development of A. ervi. The presence of developing Aphidius ervi larvae within Acyrthosiphon pisum did not affect the ability of the aphids to transmit PEMV. However, by reducing their longevity, parasitism ultimately decreased the time viruliferous aphids were able to inoculate plants. In terms of virus dispersal, parasitized aphids exhibited more movement around experimental arenas than unparasitized controls, causing a slight increase in the proportion of beans infected with PEMV. Exposure to adult Aphidius ervi caused Acyrthosiphon pisum to rapidly drop off bean plants and disperse to new hosts, resulting in considerably higher plant infection rates (70%) than that seen in control arenas (25%). The results of this investigation demonstrate that when parasitoids are added to a plant-pathogen-vector system, benefits to the host plant due to reduced herbivore infestation must be balanced against the consequences of parasitoid-induced aphid dispersal and a subsequent increase in the level of plant infection.     

EVOLUTION OF AN APHID-PARASITOID INTERACTION: VARIATION IN RESISTANCE TO PARASITISM AMONG APHID POPULATIONS SPECIALIZED ON DIFFERENT PLANTS

The evolution of associations between herbivorous insects and their parasitoids is likely to be influenced by the relationship between the herbivore and its host plants. If populations of specialized herbivorous insects are structured by their host plants such that populations on different hosts are genetically differentiated, then the traits affecting insect-parasitoid interactions may exhibit an associated structure. The pea aphid (Acyrthosiphon pisum) is a herbivorous insect species comprised of genetically distinct groups that are specialized on different host plants (Via 1991a, 1994). Here, we examine how the genetic differentiation of pea aphid populations on different host plants affects their interaction with a parasitoid wasp, Aphidius ervi. We performed four experiments. (1) By exposing pea aphids from both alfalfa and clover to parasitoids from both crops, we demonstrate that pea aphid populations that are specialized on alfalfa are successfully parasitized less often than are populations specialized on clover. This difference in parasitism rate does not depend upon whether the wasps were collected from alfalfa or clover fields. (2) When we controlled for potential differences in aphid and parasitoid behavior between the two host plants and ensured that aphids were attacked, we found that pea aphids from alfalfa were still parasitized less often than pea aphids from clover. Thus, the difference in parasitism rates is not due to behavior of either aphids or wasps, but appears to be a physiologically based difference in resistance to parasitism. (3) Replicates of pea aphid clones reared on their own host plant and on a common host plant, fava bean, exhibited the same pattern of resistance as above. Thus, there do not appear to be nutritional or secondary chemical effects on the level of physiological resistance in the aphids due to feeding on clover or alfalfa, and therefore the difference in resistance on the two crops appears to be genetically based. (4) We assayed for genetic variation in resistance among individual pea aphid clones collected from clover fields and found no detectable genetic variation for resistance to parasitism within two populations sampled from clover. This is in contrast to Henter and Via's (1995) report of abundant genetic variation in resistance to this parasitoid within a pea aphid population on alfalfa. Low levels of genetic variation may be one factor that constrains the evolution of resistance to parasitism in the populations of pea aphids from clover, leading them to remain more susceptible than populations of the same species from alfalfa.     

A negative effect of a pathogen on its vector? A plant pathogen increases the vulnerability of its vector to attack by natural enemies

Plant pathogens that are dependent on arthropod vectors for transmission from host to host may enhance their own success by promoting vector survival and/or performance. The effect of pathogens on vectors may be direct or indirect, with indirect effects mediated by increases in host quality or reductions in the vulnerability of vectors to natural enemies. We investigated whether the bird cherry-oat aphid Rhopalosiphum padi, a vector of cereal yellow dwarf virus (CYDV) in wheat, experiences a reduction in rates of attack by the parasitoid wasp Aphidius colemani when actively harboring the plant pathogen. We manipulated the vector status of aphids (virus carrying or virus free) and evaluated the impact on the rate of attack by wasps. We found that vector status did not influence the survival or fecundity of aphids in the absence of parasitoids. However, virus-carrying aphids experienced higher rates of parasitism and greater overall population suppression by parasitoid wasps than virus-free aphids. Moreover, virus-carrying aphids were accepted as hosts by wasps more often than virus-free aphids, with a greater number of wasps stinging virus-carrying aphids following assessment by antennal palpations than virus-free aphids. Therefore, counter to the prevailing idea that persistent vector-borne pathogens enhance the performance of their vectors, we found that infectious aphids actively carrying a plant pathogen experience greater vulnerability to natural enemies. Our results suggest that parasitoids may contribute to the successful biological control of CYDV by disproportionately impacting virus-carrying vectors, and thus reducing the proportion of vectors in the population that are infectious.     

Raspberry viruses affect the behavior and performance of Amphorophora agathonica in single and mixed infections

Pathogens may alter their hosts, which consequently increases transmission efficiency by vectors. We examined the effects of Raspberry leaf mottle virus [RLMV; Closterovirus (Closteroviridae)] and Raspberry latent virus [RpLV; Reovirus (Reoviridae)], alone and in a co‐infection in raspberry, Rubus idaeus L. (Rosaceae) cv. Meeker, on the behavior and performance of its vector, Amphorophora agathonica Hottes (Hemiptera: Aphididae). Longevity was increased in aphids feeding on all infected‐plant treatments compared with healthy plants, but aphid fecundity only increased in the co‐infection treatment. In a two‐way choice study between infected and healthy plants, aphids showed no difference in preference between plants after 30 min of exposure. After 24 h, aphids significantly preferred to settle on plants infected with RLMV over healthy, but healthy plants over plants infected with RpLV. There were no differences in settling preferences between healthy and co‐infected plants. An electrical penetration graph study showed no differences in aphid feeding behavior on plants infected with RLMV and RLMV+RpLV when compared with healthy controls. Our results are consistent with past findings that infected plant's impact vector performance and behavior, but also highlight the need to further investigate greater virus diversity and effects of mixed infections.     

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.     

Aphelinus albipodus(Hymenoptera: Aphelinidae) andDiaeretiella rapae(Hymenoptera: Braconidae) Parasitism onDiuraphis noxia(Homoptera: Aphididae) Infesting Barley Plants Differing in Plant Resistance to Aphids

Russian wheat aphid,Diuraphis noxia(Mordvilko), as a pest of small grains, has prompted research into biological control and host plant resistance. In the presence of Russian wheat aphid, leaves of a susceptible barley (Morex) are curled and chlorotic and sustain large densities of this aphid, while leaves of a resistant barley (STARS-9301B) remain flat and green and sustain fewer aphids. Might parasitism of Russian wheat aphid byAphelinus albipodusHayat & Fatima andDiaeretiella rapaeMcIntosh be affected differently by these plant types? When presented the plants separately and based on parasitism rate relative to aphid density, the largerD. rapaewas more effective in parasitizing relatively high densities of aphids within curled leaves of Morex than relatively low densities of aphids on uncurled leaves of STARS-9301B. Parasitism byA. albipodusdid not significantly differ among the plants. When given a choice of plants, approximately equal rates of parasitism occurred on the two plant lines for both parasitoid species, and parasitism byD. rapaewas greater thanA. albipodus.These data indicate that using parasitoid size as an indicator of success in a physically restricted environment may be misleading, when considered in a plant environment responsive in several manners to aphids (chlorosis, curling, and ability to sustain Russian wheat aphid). We expect that use of resistant barley will result in decreased parasitoid abundance as aphid densities decrease. However, parasitism rates are expected to be approximately equal on resistant and susceptible barley. In this system, plant resistance and biocontrol are compatible management strategies.     

Plant quality vs. risk of parasitism: within-plant distribution and performance of the corn leaf aphid, Rhopalosiphum maidis

1 Colonies of the aphid Rhopalosiphum maidis on Johnsongrass, Sorghum halepense, usually occur inside the whorl. The present work assessed the role of two biotic factors, plant quality and parasitism by Lysiphlebus testaceipes, in determining within‐plant distribution and performance of R. maidis. The mean relative growth rate of aphids inside the whorl and on a mature leaf was compared, and the concentration of hydroxamic acids in those tissues determined as an indicator of plant quality. Parasitism effectiveness and parasitoid behaviour were evaluated using three treatments: aphid colonies placed (1) wholly inside the whorl, (2) on the inner and outer surfaces of the whorl, and (3) on a mature leaf. 2 The mean relative growth rate of aphids was lower on the whorl than on the mature leaf, and hydroxamic acid concentration in the whorl was higher than in the mature leaf. 3 The number of parasitized dead aphids (mummies) inside the whorl was considerably lower than in the other two treatments. Mummies were present in 80% of the replicates in the whorl, whereas mummies were present in all replicates of the other treatments. 4 Host location time of the parasitoid was increased on the mature leaf compared with the other treatments. No significant differences in the residence time of the parasitoid in the aphid colony occurred between treatments. 5 Host location time showed a negative correlation, and residence time of the parasitoid a positive correlation with the number of aphids on the outer surface of the whorl.     

Relations of carrot red leaf and carrot mottle viruses with their aphid vector, Cavariella aegopodii

Studies with Scottish isolates of carrot red leaf (CRLV) and carrot mottle (CMotV) viruses confirmed the dependency of CMotV on CRLV for transmission by the aphid Cavariella aegopodii. CMotV was transmitted by aphids only when the two viruses were present in the same source plant, and its transmission was not assisted by anthriscus yellows virus, which acts as a helper for parsnip yellow fleck virus. Some test plants became infected with CRLV alone, and a few with CMotV alone. In winter, aphid transmission of CRLV and CMotV was greatly increased when the source plants received supplementary lighting whereas the CMotV infectivity of sap was not increased. C. aegopodii acquired CRLV and CMotV after minimum acquisition access times of 30 min and inoculated them after minimum inoculation access times of 2 min. There was a minimum latent period of 7–18 h. The viruses were retained by the aphid after moulting and are therefore circulative in the vector, but were not transmitted to progeny insects. Aphids allowed 24 h to acquire the viruses continued to transmit them for at least 12 days, but some aphids allowed 6 h or less for virus acquisition ceased to transmit after 3 or 4 days. CRLV is considered a tentative member of the luteovirus group.     

Host nutritional status mediates degree of parasitoid virulence

Parasitic organisms rely on the resources of their hosts to obtain nutrients essential for growth and reproduction. Insect parasitoids constitute an extreme condition since they develop in a single host from which they typically consume all available resources. As a result, the host is killed following parasitism. However, a few intriguing cases of host survival have been reported wherein hosts resume foraging and may even reproduce following parasitoid emergence. Yet, the ultimate and proximate mechanisms responsible for host recovery remain unresolved. We tested the impact of host nutrition on host fate and parasitoid fitness, using the association between Dinocampus coccinellae and the spotted lady beetle Coleomegilla maculata. Under laboratory conditions, we fed parasitized ladybirds on different aphid diets, with or without pollen. In the field, we followed the fate of parasitized ladybirds during seasonal variations in pollen and aphid abundance. We found that ladybirds fed on aphids or a combination of aphids and pollen recovered more frequently from parasitism (from 65 to 81%) than those eating only pollen (48%). Field data suggest that the fate of parasitized ladybirds is also related to food availability. On the other hand, when hosts fed on a combination of aphids and pollen, consequences for parasitoid fitness were often ‘all‐or‐nothing’: parasitoid emergence rate was the lowest of all host nutrition regimes (～50%), but parasitoids that did emerge were larger than individuals emerging from other host nutrition regimes. Laboratory and field results concur to show that host nutritional status during parasitoid development significantly influences both host fate and parasitoid fitness.     

Blockage of stylet tips as the mechanism of resistance to virus transmission by Aphis gossypii in melon lines bearing the Vat gene

Aphis gossypii is the main virus vector in muskmelon crops. The melon gene Vat confers resistance to non‐persistent virus transmission by this aphid. The mechanism of this resistance is not well understood, but no relationship has been detected between resistance and the probing behaviour of aphids on resistant plants. Results presented here suggest that temporary blockage of aphid stylet tips preventing virus particle release may explain the resistance conferred by Vat gene. We performed experiments in which viruliferous aphids were allowed to probe different sequences of resistant (Vat‐bearing) and/or susceptible melon plants. The results demonstrated that A. gossypii inoculates Cucumber mosaic virus (CMV) efficiently in susceptible plants having previously probed resistant plants, showing that the resistance mechanism is reversible. Furthermore, the infection rate obtained for susceptible plants was the same (25%) regardless of whether the transmitting aphid had come directly from the CMV source or had subsequently probed on resistant plants. This result suggests that virus is not lost from stylet to plant during probing of resistant plants, supporting the temporary blockage hypothesis. We also found that the ability of Myzus persicae to transmit CMV is noticeably reduced after probing on resistant plants, providing evidence that this aphid species also responds to the presence of the Vat gene. Finally, we also found that in probes immediately after virus acquisition M. persicae inoculates resistant plants with CMV more efficiently than susceptible plants, perhaps because the Vat gene product induces increased salivation by this aphid.     

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.     

Plant infection by two different viruses induce contrasting changes of vectors fitness and behavior

Insect-vectored plant viruses can induce changes in plant phenotypes,thus influencing plant-vector interactions in a way that may promote their dispersal according to their mode of transmission (i.e.,circulative vs.noncirculative).This indirect vector manipulation requires host-virus-vector coevolution and would thus be effective solely in very specific plant-virus-vector species associations.Some studies suggest this manipulation may depend on multiple factors relative to various intrinsic characteristics of vectors such as transmission efficiency.In anintegrative study,we tested the effects of infection of the Brassicaceae Camelina sativa with the noncirculative Cauliflower mosaic virus (CaMV)or the circulative Turnip yellows virus (TuYV)on the host-plant colonization of two aphid species differing in their virus transmission efficiency:the polyphagous Myzus persicae,efficient vector of both viruses,and the Brassicaceae specialist Brevicoryne brassicae,poor vector of TuYV and efficient vector of CaMV.Results confirmed the important role of virus mode of transmission as plant-mediated effects of CaMV on the two aphid species induced negative alterations of feeding behavior (i.e.,decreased phloem sap ingestion)and performance that were both conducive for virus fitness by promoting dispersion after a rapid acquisition.In addition,virus transmission efficiency may also play a role in vector manipulation by viruses as only the responses of the efficient vector to plant-mediated effects of TuYV,that is,enhanced feeding behavior and performances,were favorable to their acquisition and further dispersal.Altogether,this work demonstrated that vector transmission efficiency also has to be considered when studying the mechanisms underlying vector manipulation by viruses.Our results also re- inforce the idea that vector manipulation requires coevolution between plant,virus and vector.     

Spread of barley yellow dwarf virus and relative importance of local aphid vectors in central Washington

Data from bioassays of field collected aphids, barley indicator plants exposed to natural conditions, and various types of aphid traps were used to describe the spread of barley yellow dwarf virus (BYDV) in wheat and barley near Prosser, Washington. Bioassays were also used to assess the relative importance of local vector species. Of alate aphids collected from grain in the 1982 and 1983 fall migration seasons, 3.4–14–5% transmitted BYDV. Data from concurrent and post-migration assays of resident aphids (apterae and nymphs) reflected an increase in the proportion of infected plants in the field. Maximum increase in the percentage of viruliferous aphids occurred in late November and December of 1982 and November of 1983. The 1982 increase occurred after aphid flights had ceased for the year, suggesting active secondary spread. Collections in pitfall traps and infected trap plants from November to February confirmed aphid activity and virus spread. Rhopalosiphum padi was the most important vector in central Washington in 1982 and 1983 because of its abundance and relative BYDV transmission efficiency. Metopolophium dirhodum was more winter-hardy than R. padi and equal to R. padi in its efficiency as a vector; however, it was not as abundant as R. padi except during the mild winter of 1982–83, when it was a major contributor to secondary spread. Sitobion avenae may be important in years when it is abundant, but it was only a quarter as efficient as R. padi. Rhopalosiphum maidis was a much less efficient vector than R. padi and it only reached high populations in late autumn barley.     

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

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

Infection of potato plants with potato leafroll virus changes attraction and feeding behaviour of Myzus persicae

Potato leafroll virus (PLRV; genus Polerovirus, family Luteoviridae) is a persistently transmitted circulative virus that depends on aphids for spreading. The primary vector of PLRV is the aphid Myzus persicae (Sulzer) (Homoptera: Aphididae). Solanum tuberosum L. potato cv. Kardal (Solanaceae) has a certain degree of resistance to M. persicae: young leaves seem to be resistant, whereas senescent leaves are susceptible. In this study, we investigated whether PLRV‐infection of potato plants affected aphid behaviour. We found that M. persicae's ability to differentiate headspace volatiles emitted from PLRV‐infected and non‐infected potato plants depends on the age of the leaf. In young apical leaves, no difference in aphid attraction was found between PLRV‐infected and non‐infected leaves. In fact, hardly any aphids were attracted. On the contrary, in mature leaves, headspace volatiles from virus infected leaves attracted the aphids. We also studied the effect of PLRV‐infection on probing and feeding behaviour (plant penetration) of M. persicae using the electrical penetration graph technique (DC system). Several differences were observed between plant penetration in PLRV‐infected and non‐infected plants, but only after infected plants showed visual symptoms of PLRV infection. The effects of PLRV‐infection in plants on the behaviour of M. persicae, the vector of the virus, and the implications of these effects on the transmission of the virus are thoroughly discussed.     

Comparative transmission of pea mosaic virus by one strain ofMyzus persicae Sulz. and two strains ofAcyrthosiphon pisum Harris

Experiments were conducted to compare the efficiency of transmission of a strain of pea mosaic virus (PMV) isolated in Czechoslovakia by two strains (clones) of the pea aphidAcyrthosiphon pisum Harris (green and red) and one strain ofMyzus persicae Sulz. PMV is a nonpersistent virus and the preliminary fasting of aphids before acquisition feeding increases the efficiency of aphids in transmission of this virus. In our experiments two hour fasted individuals were used and two periods of acquisition feeding on the source (1 and 5 minutes). On the healthy test plants the aphids were left over night. As the source and test plants pea (Pisum sativum L.) of the cv. Raman were used. During the one minute acquisition period on the source of infection the aphids were observed under the stereoscopic microscope. They usually made two to three probes. During the five minute acquisition feeding time the aphids were not observed and they were taken from the source of infection after a lapse of five minutes. To compare the efficiency in transmission of this virus by these aphids only one aphid per tested plant was used and in all trials only two to four day old nymphs were taken. Differences in transmission efficiency between two strains ofAcyrthosiphon pisum Harris were highly significant. The green strain of pea aphid was the less efficient vector in comparison with the green peach aphid and the red strain of the pea aphid, the latter being the most efficient vector of this virus.     

Introduced ants reduce interaction diversity in a multi‐species,ant–aphid mutualism

Mutualisms contribute in fundamental ways to the origin, maintenance and organization of biological diversity. Introduced species commonly participate in mutualisms, but how this phenomenon affects patterns of interactions among native mutualists remains incompletely understood. Here we examine how networks of interactions among aphid‐tending ants, ant‐tended aphids, and aphid‐attacking parasitoid wasps differ between 12 spatially paired riparian study sites with and without the introduced Argentine ant Linepithema humile in southern California. To resolve challenges in species identification, we used DNA barcoding to identify aphids and screen for parasitoid wasps (developing inside their aphid hosts) from 170 aphid aggregations sampled on arroyo willow Salix lasiolepis. Compared to uninvaded sites, invaded sites supported significantly fewer species of aphid‐tending ants and ant‐tended aphids. At invaded sites, for example, we found only two species of ant‐tended aphids, which were exclusively tended by L. humile, whereas at uninvaded sites we found 20 unique ant–aphid interactions involving eight species of ant‐tended aphids and nine species of aphid‐tending ants. Ant–aphid linkage density was thus significantly lower at invaded sites compared to uninvaded sites. We detected aphid parasitoids in 14% (28/198) of all aphid aggregations. Although the level of parasitism did not differ between invaded and uninvaded sites, more species of wasps were detected within uninvaded sites compared to invaded sites. These results provide a striking example of how the assimilation of introduced species into multi‐species mutualisms can reduce interaction diversity with potential consequences for species persistence.     

Parasitoids aid dispersal of a nonpersistently transmitted plant virus by disturbing the aphid vector

• 1 Aphids are the major group of insects that vector plant viruses, and they often display a preference for foliage showing disease symptoms. Although this behaviour will increase the numbers of vectors acquiring the pathogen, it will not in itself result in a greater spread of the disease.
• 2 The present study examined how infection of Vicia faba by the nonpersistently transmitted virus bean yellow mosaic virus (BYMV) affected colonization by pea aphids Acyrthosiphon pisum. We then examined how foraging by the hymenopterous parasitoid Aphidius ervi affected aphid settling/movement behaviour and the consequences for dissemination of the virus.
• 3 In Petri dish arenas, aphids colonized discs from BYMV‐infected leaves more rapidly than discs from uninfected plants. Reflectance from infected foliage was approximately 20% higher than from uninfected leaves in the green–yellow wavelengths, indicating that aphids might be responding to visual cues from the brighter foliage. Settling was reduced by A. ervi, with the foraging wasps preventing the aphids reaching and/or remaining on the leaf tissue.
• 4 In multiple plant arenas, A. ervi caused a reduction in aphid numbers but also a nine‐fold increase in BYMV infection. It is hypothesized that disturbance by the parasitoids resulted in more aphid movement as well as more cases of aphids probing on a BYMV‐infected plant and then a new host within the critical time period for successful inoculation to occur. This effect of parasitoids on virus dispersal should be considered in epidemiological models of insect‐vectored plant diseases, and also when evaluating the use of natural enemies in biocontrol strategies of insect herbivore/vector pests.