Effect of Adult Experience on in-Flight Orientation to Plant and Plant–Host Complex Volatiles inAphidius erviHaliday (Hymenoptera, Braconidae)

The effect of adult experience on in-flight orientation to plant–host complex volatiles byAphidius erviHaliday was studied in a wind tunnel bioassay, usingAcyrthosiphon pisum(Harris), maintained on broad bean plants (Vicia faba) as host. A short oviposition experience (15 s) on the plant–host complex (PHC) was sufficient to induce a drastic decrease of flight propensity and stimulated a foraging behavior characterized by intense walking activity. However, flight activity resumed to normal levels 1 h after the oviposition experience on the PHC occurred. For parasitoids conditioned on the PHC for at least 1 min the recorded proportion making oriented flights to the PHC was significantly higher than that for naive females. In contrast, oviposition experience in the absence of plant material did not influence theA. erviflight response. Oviposition attempts on aphid dummies without egg release did not reduce flight activity. WhenA. ervifemales were exposed to glass beads coated withAc. pisumcornicle secretion, a priming effect was observed, resulting, compared with naive females, in a significantly higher rate of oriented flights to the PHC. In contrast, oviposition attempts visually induced by colored aphid dummies did not influence flight behavior. A strong reaction to volatile cues from uninfested plants was induced by oviposition experience on newly infested broad bean plants. This appears to be a case of associative learning. In fact, uninfested broad bean plants are basically unattractive to naiveA. ervifemales. The results demonstrate that adult experience has a considerable influence onA. ervibehavior and may have important implications for biological control of natural pest aphid populations.     

The influence of prior ovipositional experience on host selection in four species of aphidiid wasps (Hymenoptera: Aphidiidae)

We evaluated the influence of prior ovipositional experience on host selection in four solitary parasitoids, Aphidius erviHaliday, A. pisivorusSmith, A. smithiSharma & Subba Rao, and Praon pequodorumViereck (Hymenoptera: Aphidiidae). When provided simultaneously with equal numbers of two species of aphids, Acyrthosiphon pisum(Harris) and Macrosiphum creeliiDavis (Homoptera: Aphididae), all four parasitoids showed a moderate to strong preference for A. pisum.This preference did not increase after sampling, except in P. pequodorum.Learning did not alter host selection behavior in A. erviand P. pequodorum.However, females of A. pisivorusconditioned on A. pisumselected fewer M. creeliithan unconditioned wasps and wasps conditioned on M. creelii.It is suggested that prior ovipositional experience influences a parasitoid's expectations of the kinds of hosts available, but it does not alter the (innate) rank order of hosts.     

Behavioural differences between Aphidius ervi populations from two tritrophic systems are due to phenotypic plasticity

The Palaeoarctic parasitoid Aphidius ervi Haliday (Hymenoptera, Aphidiidae) parasitises legume aphids in its region of origin. In Chile, it parasitises both legume and cereal aphids. This special situation was studied at two levels: (i) the host searching behaviour of A. ervi from two different tritrophic systems (Acyrthosiphon pisum on alfalfa and Sitobion avenae on wheat) was investigated in dual choice tests in a wind tunnel between odours from both A. pisum-alfalfa host plant complex (HPC) and S. avenae-wheat HPC, and (ii) the genetic structure of A. ervi populations from both sources using molecular markers. Responses of A. ervi females to volatile olfactory cues emanating from A. pisum-alfalfa HPC and S. avenae-wheat HPC were significantly higher towards the HPC on which they were reared during the last generation before experimentation, regardless of the origin of the parasitoid. As previously described for this parasitoid species, oviposition experience was also of major relevance in the preferences of female parasitoids. On the other hand, variation in mitochondrial DNA segments and RAPD-PCR polymorphism using total DNA showed the absence of host-based population structure and a high genetic homogeneity between these A. ervi populations. These results reject the possible existence of different host-strains of this parasitoid in Chile.     

Effects of honeydew and insecticide residues on the distribution of foraging aphid parasitoids under glasshouse and field conditions

The opposing effects of attraction to host-derived kairomones and repellency from the pyrethroid insecticide deltamethrin were investigated with aphid parasitoids from the genus Aphidius (Hymenoptera: Aphidiinae). The spatial distribution of female parasitoids was recorded in a series of experiments conducted in a small glasshouse containing wheat plants either infested with cereal aphids, Sitobion avenae (F.) (Homoptera: Aphididae), uninfested or treated with the recommended field concentration of deltamethrin. The number of parasitoids per plant were counted at 0.5 h, 1 h and then at one hourly intervals up to 8 h after release. Parasitoids showed a strong aggregation response to aphid-infested plants compared to adjacent uninfested plants. With the introduction of insecticidetreated plants around the aphid-infested plants, parasitoids showed a greater tendency to disperse away, resulting in fewer parasitoids on plants and significantly lower rates of aphid parasitism. The degree of aphid fall-off from plants was a good indicator of parasitoid foraging activity. In field studies, using sticky traps to measure the activity of parasitoids in plots sprayed with water, deltamethrin and/or an artificial honeydew solution, repellent properties were evident for up to 2 days after application. The attraction/arrestment stimuli associated with the honeydew solution were sufficient for parasitoids to continue searching insecticide-treated areas. The implications of these findings for parasitoids searching crops contaminated with aphid-derived kairomones and insecticides are discussed.     

Impact of environmental stress on aphid clonal resistance to parasitoids: Role of Hamiltonella defensa bacterial symbiosis in association with a new facultative symbiont of the pea aphid

Resistance to endoparasitoids in aphids involves complex interactions between insect and microbial players. It is now generally accepted that the facultative bacterial symbiont Hamiltonella defensa of the pea aphid Acyrthosiphon pisum is implicated in its resistance to the parasitoid Aphidius ervi. It has also been shown that heat negatively affects pea aphid resistance, suggesting the thermosensitivity of its defensive symbiosis. Here we examined the effects of heat and UV-B on the resistance of A. pisum to A. ervi and we relate its stability under heat stress to different facultative bacterial symbionts hosted by the aphid. For six A. pisum clones harboring four different facultative symbiont associations, the impact of heat and UV-B was measured on their ability to resist A. ervi parasitism under controlled conditions. The results revealed that temperature strongly affected resistance, while UV-B did not. As previously shown, highly resistant A. pisum clones singly infected with H. defensa became more susceptible to parasitism after exposure to heat. Interestingly, clones that were superinfected with H. defensa in association with a newly discovered facultative symbiont, referred to as PAXS (pea aphid X-type symbiont), not only remained highly resistant under heat stress, but also expressed previously unknown, very precocious resistance to A. ervi compared to clones with H. defensa alone. The prevalence of dual symbiosis involving PAXS and H. defensa in local aphid populations suggests its importance in protecting aphid immunity to parasitoids under abiotic stress.     

Beneficial arthropod behavior mediated by airborne semiochemicals. VIII. Learning of host-related odors induced by a brief contact experience with host by-products inCotesia marginiventris (Cresson), a generalist larval parasitoid

Females of Cotesia marginiventris(Cresson), a generalist larval parasitoid, were observed to respond to host related odors in a four-arm olfactometer. The females were significantly more responsive to the odors after a brief contact experience with host-damaged leaves contaminated with host by products. During the experience, actual encounters with hosts were not required to improve subsequent responses to host-related odors. The response to odors of the plant-host complex with which parasitoids had experience was significantly higher than the response to odors of an alternative plant-host complex. This suggests that the experience effect is due, at least partly, to associative learning. We suspect that females of this generalist parasitoid, as was recently found for those of a specialist, recognize specific semiochemicals when they contact frass of suitable host larvae. The parasitoids, subsequently, associate the surrounding odors with the possible presence of hosts, and use these odors as cues in their search for more hosts. This could be an important component in the host-searching behavior of many parasitoids.Mention of a proprietary product does not constitute an endorsement or the recommendation for its use by the USDA.     

Host-locating response by the aphid parasitoid Aphidius ervi to tomato plant volatiles

Abstract

The blend of volatile compounds emitted by tomato plants (Solanum lycopersicum) infested with the potato aphid (Macrosiphum euphorbiae) has been studied comparatively with undamaged plants and aphids themselves. Aphid-infested plants were significantly more attractive towards Aphidius ervi than undamaged plants and aphids themselves. Oriented response towards host-damaged plant, from which aphids were removed just before running the bioassay, did not differ from that recorded for infested plants. Collection of the volatiles and analysis by gas chromatography revealed only quantitative differences between uninfested and aphid-infested plants. Nine compounds, α-pinene, (Z)-3-hexen-1-ol, α-phellandrene, limonene, (E)-β-ocimene, p-cymene, methyl salicylate, (E)-β-caryophyllene and an unknown compound, were emitted at higher levels from aphid-infested plants than from undamaged control plants, whilst no differences were noted for hexanal, 6-methyl-5-hepten-2-one, and humulene (=α-caryophyllene). Synthetic standards of these compounds were tested in wind tunnel bioassays and all elicited a significant increase in oriented flight and landings on the target by the aphid parasitoid Aphidius ervi. (E)-β-caryophyllene resulted the most attractive towards female wasps. These results corroborate the hypothesis that the volatiles produced by the plant in response to aphid attack derive from both jasmonic and salicylic acid pathways, and are exploited by A. ervi as olfactory cues to locate its hosts.     

Effects of honeydew sugar composition on the longevity of Aphidius ervi

Feeding on sugar‐rich foods such as nectar and honeydew is important for survival of many adult parasitoids. Especially in agricultural systems, honeydew is often the most prevalent carbohydrate source. However, relative to plant nectar, honeydew may be relatively unsuitable, as a result of an unfavourable sugar composition or the presence of secondary plant compounds. We studied survival of the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) on honeydew collected from various aphid species feeding on potato (Solanum tuberosum L., cv. Desiree) (Solanaceae), wheat (Triticum aestivum L., cv. Bobwhite) (Poaceae), or artificial diet, as well as the sugar composition of the different honeydews. Honeydews from the tested aphid species on potato, wheat, or artificial diet were found to be relatively suitable food sources for adult A. ervi, although not always as suitable as a 2 M sucrose solution. There were differences in honeydew sugar composition among the different aphid species on the various host plants. Multivariate statistics showed that the factor ‘aphid species’ had a significant influence on the sugar composition of the honeydew, explaining 27% of the variation in the potato system and 89% in the wheat system. When exploring the relationship between carbohydrate composition of the honeydews from aphids on potato and wheat plants, and their nutritional value for A. ervi, data revealed that differences in parasitoid longevity can to some extent be explained by carbohydrate composition. Furthermore, our results confirm that sucrose and its hexose components glucose and fructose are very suitable carbohydrate sources for hymenopteran parasitoids and show that parasitoid survival on an equimolar solution of the two monosaccharides glucose and fructose does not exceed performance on the disaccharide sucrose.     

Population level effects of cadmium and the insecticide imidacloprid to the parasitoid, Aphidius ervi after exposure through its host, the pea aphid, Acyrthosiphon pisum (Harris)

The objective of this study was to assess the influence of toxic substances with different modes of action on a two-species system: an aphid-specific parasitoid, Aphidius ervi Haliday, feeding on the pea aphid, Acyrthosiphon pisum (Harris). The instantaneous rate of population increase (r_i) was used as a measure of population level toxic effect in this study. The toxicants evaluated were imidacloprid, a nonpersistent neurotoxic insecticide, and cadmium, a chronic pollutant with a tendency to accumulate. We evaluated the effects of cadmium and imidacloprid on A. pisum and A. ervi because both toxicants can occur together in polluted areas where crops are grown. Cadmium (200 or 400 mg kg⁻¹ dry weight soil) and imidacloprid (4 or 40 g a.i. ha⁻¹) were applied to soil contained in plastic pots in which broad bean plants, Vicia faba L., were grown. Results of this study indicated that cadmium at the concentrations tested, reduced population growth rate of the pea aphid. Imidacloprid also reduced aphid growth rate, but only at the highest concentration tested (40 g a.i. ha⁻¹). Combinations of cadmium and imidacloprid had the greatest impact on aphid growth rate. Imidacloprid alone had no effect on population growth rate of the parasitoid. However, cadmium alone or in combination with imidacloprid had a negative impact on A. ervi by reducing population growth rate 77%. These results indicate that negative impacts on parasitoids may occur in areas where cadmium contamination is present and imidacloprid is used to control aphids.     

Influence of adult experience on host microhabitat location by the generalist parasitoid,Campoletis sonorensis (Hymenoptera: Ichneumonidae)

The effect of adult experience on microhabitat location behavior of the generalist parasitoid Campoletis sonorensis (Cameron)was examined in a wind tunnel bioassay. Responses were tested to the odors of two host plants (cotton and sesame) of Heliothis virescens (F.) or a nonhost plant (potato), either damaged and infested with host larvae and host products (host/plant complex) or intact, clean and uninfested. Parasitoid females remained naive or were allowed one oviposition experience on either of the plants, 1 min, 2 h, or 24 h prior their introduction into the wind tunnel. In a no-choice test, parasitoids experienced 1 min prior to bioassay completed significantly more flights to sesame and potato host/plant complexes than did naive parasitoids. However, 24 h after experience, only females experienced on potato completed more flights to the host/ plant complex than did naive females. Parasitoids experienced 1 min prior to flight to undamaged plants showed a slight increase in flight response (significant only for potato) but, after 24 h, completed only as many flights as naive parasitoids. In a dual-choice situation, parasitoids did not show a preference for either of the two host plants but did prefer a host to a nonhost plant. This innate plant preference was not changed by a single oviposition experience. The potential significance of these results to the microhabitat location behavior of C. sonorensisin the field is discussed.     

Identifying bird cherry-oat aphid Rhopalosiphum padi emigrants, alate exules and gynoparae: application of multivariate methods to morphometric and anatomical features

The host microhabitat location behavior of females of the generalist parasitoid Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae) was studied in a wind tunnel. Visual cues associated with the host plant cotton, Gossypium hirsutum L., were important and significantly more parasitoids completed flights to a damaged 4-leaf cotton plant bearing a Heliothis virescens (F.) (Lepidoptera: Noctuidae) larva and frass than to a similarly damaged single leaf with frass and a larva. This difference in completed flights was not due to differences in amounts of volatiles released by the two stimuli. Both naive and experienced parasitoids responded differently to an undamaged cotton leaf, a mechanially damaged leaf, a naturally damaged leaf with the host removed and a naturally damaged leaf with a host larva. Parasitoids completed significantly fewer flights to the undamaged sources of volatiles than to damaged sources of volatiles. Experienced females responded strongly to all types of damage. The number of flights completed by naive females to the three types of damage differed but not significantly and was less than the number completed by experienced females. Components of the preflight experience were varied to determine which factors were responsible for the higher response of experienced females to the host/plant complex. Oviposition was the most important component of this experience. Contact with host frass or plant damage followed by oviposition did not increase the response over that exhibited by females allowed oviposition only. When frass or damaged plant material were contacted without subsequent oviposition, females completed fewer flights than naive females.     

Arrestment responses ofEretmocerus species andEncarsia deserti (Hymenoptera: Aphelinidae) toBemisia tabaci honeydew

The responses of Eretmocerussp. and Encarsia deserti(Gerling & Rivnay), parasitoids of Bemisia tabaci(Gennadius) to host secretions were investigated. Upon contact with honeydew excreted from the host, the parasitoids' walking speed decreased, whereas the rate of angular turning increased (the walking speed of Eretmocerussp. and Encarsia desertidecreased from 2.0 to 0.8 and from 0.8 to 0.3 mm/s, respectively, and the respective angular turning rates increased from 123.3 to 489.6 and from 267.9 to 979.2 deg/cm). Moreover, the wasps generally turned back toward the secretion as soon as they lost contact with it. This induced arrestment responses resulted in increased searching time by the parasitoids. Laboratory bioassays were conducted in which paper disks in petri dishes containing different amounts of honeydew, equivalent to the secretions of 1, 5, or 50 larvae, were offered to female wasps. The wasps responded by arrestment even to honeydew produced by a single larva. While the response of Eretmocerussp. was quantity dependent, being more intense at higher amounts of honeydew, that of E. desertiwas of equal intensity at all honeydew quantities assayed.     

Comparative Prey Suitability ofOstrinia nubilalisEggs andAcyrthosiphon pisumforColeomegilla maculata

The effects ofOstrinia nubilalis(Hübner) (Lepidoptera: Pyralidae) eggs andAcyrthosiphon pisum(Harris) (Homoptera: Aphididae), when provided as single prey species and in combination, on life history characteristics ofColeomegilla maculataDeGeer (Coleoptera: Coccinellidae) larvae and adults were quantified. Preimaginal development was not influenced by the larval prey regime; development at 26 ± 1°C was completed in approximately 13.5 days onO. nubilaliseggs,A. pisum,orA. pisumalternated daily withO. nubilaliseggs. The resulting adults weighed 13.0, 10.7, and 12.5 mg when reared onO. nubilaliseggs,A. pisum,andA. pisumalternated daily withO. nubilaliseggs, respectively. Eighteen percent of the individuals died when reared onA. pisum,28% died when reared onO. nubilaliseggs, and 22% died when fedA. pisumalternated daily withO. nubilaliseggs. Seven adult diet combinations, based on diet regimes of larvae and adults, did not cause significant differences in preoviposition period, interoviposition period, and the number of days on which eggs were laid. Total fecundity was influenced both by larval and adult diet. The diet that resulted in highly fecund females wasA. pisumalternated daily withO. nubilaliseggs for larvae andO. nubilaliseggs for adults. FemaleC. maculatafedO. nubilaliseggs had the highest intrinsic rate of increase and net reproductive rate.     

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.     

Effect of honeydew and hosts on plant colonization by the aphid parasitoidEphedrus cerasicola

Females ofEphedrus cerasicola Stary were released into small glasshouses (13.6 m³) or cages (0.125 m³) which contained paprika plants in 2 of the following categories: fresh plants (without honeydew and aphids), honeydew-contaminated plants (aphids removed) and aphid-infested plants (with honeydew). Two hundred females were released into each glasshouse with 20 plants, 10 of each category, while 10 females were released into the cages with 2 plants, one of each category. Control experiments with only fresh plants were conducted in both the glasshouses and the cages. The female parasitoids on each plant were counted at regular intervals for 24 or 48 h in the glasshouse and for 4 h in the cage experiments; this number was used as an indicator of plant preferences. The following preferences were established (preferred plant > less preferred plant): honeydew-plant > fresh plant, aphid-plant > fresh plant, aphid-plant > honeydew-plant. The number of females on the aphid-plants increased throughout the experimental period, whereas the number on the honeydew-plants tended to be more stable.      

Foraging behaviour at the fourth trophic level: a comparative study of host location in aphid hyperparasitoids

In studies of foraging behaviour in a multitrophic context, the fourth trophic level has generally been ignored. We used four aphid hyperparasitoid species: Dendrocerus carpenteri (Curtis) (Hymenoptera: Megaspilidae), Asaphes suspensus Walker (Hymenoptera: Pteromalidae), Alloxysta victrix (Westwood) (Hymenoptera: Alloxystidae) and Syrphophagus aphidivorus (Mayr) (Hymenoptera: Encyrtidae), to correlate their response to different cues with their ecological attributes such as host range and host stage. In addition, we compared our results with studies of primary parasitoids on the same plant–herbivore system. First, the olfactory response of females was tested in a Y‐tube olfactometer (single choice: plant, aphid, honeydew, parasitised aphid, aphid mummy, or virgin female parasitoid; dual choice: clean plant, plant with aphids, or plant–host complex). Second, their foraging behaviour was described on plants with different stimuli (honeydew, aphids, parasitised aphids, and aphid mummies). The results indicated that olfactory cues are probably not essential cues for hyperparasitoid females. In foraging behaviour on the plant, all species prolonged their total visit time and search time as compared to the control treatment (clean plant). Only A. victrix did not react to the honeydew. Oviposition in mummies prolonged the total visit time because of the long handling time, but the effect of this behaviour on search time could not be determined. No clear correlation between foraging behaviour and host stage or host range was found. In contrast to specialised primary aphid parasitoids that have strong fixed responses to specific kairomones and herbivore‐induced synomones, more generalist aphid hyperparasitoids seem to depend less on volatile olfactory stimuli, but show similarities with primary parasitoids in their use of contact cues while searching on a plant.     

<Emphasis Type="Italic">Aphidius ervi</Emphasis> Preferentially Attacks the Green Morph of the Pea Aphid, <Emphasis Type="Italic">Acyrthosiphon pisum</Emphasis>

The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae) is found in red and green color morphs. Previous work has suggested that the aphidiine parasitoid Aphidius ervi Haliday preferentially attacks green pea aphids in the field. It is not clear whether these results reflect a real preference, or some unknown clonal difference, such as in immunity, between the aphids used in the previous studies. We used three susceptibility-matched pairs of red and green morph pea aphid clones to test for preferences. In a no-choice situation, the parasitoids attacked equal proportions of each color morph. When provided with a choice, A. ervi was significantly more likely to oviposit into colonies formed from green morphs when the neighboring colony was formed from red morph aphids. In contrast, red morphs were less likely to be attacked when their neighboring colony was of the green morph. By preferentially attacking green colonies, A. ervi may reduce the likelihood of intraguild predation, as it is suggested that visually foraging predators preferentially attack red aphid colonies. Furthermore, if this host choice behavior is replicated in the field, we speculate that color morphs of the pea aphid may interact indirectly through their shared natural enemies, leading to intraspecific apparent competition.     

The role of physical cues in the regulation of host recognition and acceptance behavior ofAphidius ervi Haliday (Hymenoptera: Braconidae)

The role of color and shape in the host recognition and acceptance behavior ofAphidius ervi Haliday was studied. A quantitative analysis of the oviposition behavior ofA. ervi was carried out with a computer-aided analysis of 150 video-recorded oviposition sequences on its natural host,Acyrthosiphon pisum (Harris). The importance of visual stimuli was assessed in a choice condition bioassay, observing the behavioral reaction of female parasitoids to various test materials flame-sealed into glass capillaries. Glass beads 2 and 6 mm in diameter and a flat arena were coated with cornicle secretion ofA. pisum, and their acceptance rates by both naive and experienced female parasitoids were assessed under no-choice conditions. In most cases,A. ervi females switched from random searching to attack position when the host was within a range of 1 cm, suggesting that host recognition is regulated in part by cues acting before physical contact. The glass capillary bioassay indicated that visual cues are important factors in the host recognition and acceptance phases. Pea aphid color alone can elicit the oviposition response of naiveA. ervi females, and this response is enhanced when color is combined with aphid shape. The cornicle secretion ofA. pisum stimulated an oviposition response which was stronger in naive females ofA. ervi than in experienced ones and was not significantly affected by the glass bead size or flat surface. These results, along with those from previous studies, suggest that manipulation of the oviposition behavior ofA. ervi is feasible under laboratory conditions.     

Aphid parasitoid responses to semiochemicals — Genetic, conditioned or learnt?

Parasitoid foraging behaviour is known to be influenced by interactions of genetic, physiological, environmental and experiential factors. Although the role of genetics, learning and conditioning in determining responses to foraging cues has been studied in lepidopteran parasitoids, aphid parasitoids have been less intensively researched. Using the tritrophic system,Vicia faba — Acyrthosiphon pisum — Aphidius ervi, evidence for the role of genetics and learning in parasitoid foraging is presented, and the difficulty of differentiating between genetic responses and those conditioned during parasitoid development is discussed. Aphidius ervi responds to aphid sex pheromones both in the field and in the laboratory. Since laboratory reared individuals have never experienced sexual aphids, the response must be genetic as it cannot have been conditioned during development. An example of a response conditioned during development is the variable response ofA. rhopalosiphi to different wheat cultivars depending upon host feeding. Aphid parasitoids also are adept at learning as shown by their responses to plant-derived cues which are learnt as Conditioned Stimuli (CS). Host products such as honeydew, as well as the host itself, can act as the Unconditioned Stimulus (US) in the learning process. Aphidius ervi offers a good model for investigating the role of these factors in parasitoid foraging behaviour. Finally, the value of such research for biological control programmes involving aphid parasitoids is discussed.     

Oviposition behaviour of the aphid parasitoid,Aphidius ervi: Are wet aphids recognized as host?

Females of the central European population of the aphid parasitoid, Aphidius ervi, did not attack wet pea aphids (Acyrthosiphon pisum) that were washed previously with water. After 1 hour, this phenomenon disappeared and A. ervi attacked washed hosts to the same degree as dry ones. Similarly, A. ervi attacked dead aphids killed in liquid nitrogen readily if they were dry but not if they were wet. This effect was also reversible and disappeared after 1 h. When A. ervi females were foraging on broad beans (Vicia faba), they laid significantly more eggs into dry aphids than into wet aphids. Resource utilization of wet aphids, however, was significantly lower in this design than in Petri dishes, due to a changed drop-off behaviour of the aphid. We conclude that females did not use visual cues for host recognition but instead relied on chemical cues. These cues may be covered by a thin water layer directly after aphids became wet. Our results also demonstrate the importance of abiotic factors for the estimation of the reproductive success of parasitoids in the field.