Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Size and sex matter: reproductive biology and determinants of offspring survival in Gazella marica

For prey, many behavioural traits are constrained by the risk of predation. Therefore, shifts between warning and cryptic coloration have been suggested to result in parallel changes in several behaviours. In the present study, we tested whether changes in chromatic contrast among eight populations of the strawberry poison‐dart frog, Dendrobates pumilio, co‐vary with behaviour, as expected if selection is imposed by predators relying on visual detection of prey. These eight populations are geographically isolated on different island in the Bocas del Toro region of Panama and have recently diverged morphologically and genetically. We found that aggression and explorative behaviour were strongly correlated and also that males tended to be more aggressive and explorative if they belonged to populations with conspicuously coloured individuals. We discuss how evolutionary switches between predator avoidance strategies and associated behavioural divergence between populations may affect reproductive isolation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Mating or ovipositing? A crucial decision in the life history of the cabbage aphid parasitoid Diaeretiella rapae (M’Intosh)

1. The reproductive fitness of a parasitoid depends on its mating and ovipositing success. Virgin haplodiploid females can reproduce, but produce only males, and may diminish fitness by producing more male offspring than required. Therefore, females must decide on whether to mate or oviposit first. 2. This study was conducted to assess the mating versus ovipositing decision and its impact on the reproductive fitness of Diaeretiella rapae (Hymenoptera: Aphididae), an endoparasitoid of the cabbage aphid Brevicoryne brassicae (Hemiptera: Aphididae). 3. When newly emerged females were given a choice between mating and ovipositing, about 62% of D. rapae females preferred to mate before ovipositing. Those females who oviposited before mating parasitised only 10% of the available aphids. After mating, females superparasitised their hosts with fertilised eggs, which resulted in a highly female‐biased sex ratio in the offspring. 4. Mating success was very high (91%) in the presence of hosts (cabbage aphid nymphs) compared with that in the absence of aphids. However, mating success was not influenced by the quality (size) of the hosts present in the mating arena, despite a parasitoid preference for larger hosts during oviposition. The time between pairing and mating was also shorter in the presence of host aphids. The mean number of aphids parasitised and the parasitism rate were significantly greater after mating.     

Egg load dynamics and the risk of egg and time limitation experienced by an aphid parasitoid in the field

Insect parasitoids and herbivores must balance the risk of egg limitation and time limitation in order to maximize reproductive success. Egg and time limitation are mediated by oviposition and egg maturation rates as well as by starvation risk and other determinants of adult lifespan. Here, we assessed egg load and nutritional state in the soybean aphid parasitoid Binodoxys communis under field conditions to estimate its risk of becoming either egg‐ or time‐limited. The majority of female B. communis showed no signs of egg limitation. Experimental field manipulations of B. communis females suggested that an average of 4–8 eggs were matured per hour over the course of a day. Regardless, egg loads remained constant over the course of the day at approximately 80 eggs, suggesting that egg maturation compensates for oviposition. This is the first case of such “egg load buffering” documented for a parasitoid in the field. Despite this buffering, egg loads dropped slightly with increasing host (aphid) density. This suggests that egg limitation could occur at very high host densities as experienced in outbreak years in some locations in the Midwestern USA. Biochemical analyses of sugar profiles showed that parasitoids fed upon sugar in the field at a remarkably high rate. Time limitation through starvation thus seems to be very low and aphid honeydew is most likely a source of dietary sugar for these parasitoids. This latter supposition is supported by the fact that body sugar levels increase with host (aphid) density. Together, these results suggest that fecundity of B. communis benefits from both dynamic egg maturation strategies and sugar‐feeding.     

Tritrophic effects of organic and conventional fertilisers on a cereal‐aphid‐parasitoid system

The impact of parasitoids on pests varies between conventional and low‐intensity agricultural systems. Although the impacts on parasitoid natural enemies of many practices within these agricultural systems are well understood, the role of fertilisers has been less well studied. The effects of organic‐based and conventional fertilisers on Hordeum vulgare L. (Poaceae), the aphid Metopolophium dirhodum Walker (Hemiptera: Aphididae), and its parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) was investigated using cage release experiments and measures of aphid and parasitoid fitness were taken. Barley tiller number and aphid weight were increased by fertilisers, particularly under conventional treatments. Adult parasitoid size correlated positively with that of the host, M. dirhodum, whereas percentage parasitism was not affected by fertiliser treatment or host size. The results suggest that the increased parasitoid impact observed in some low‐intensity or organic systems is not a direct result of fertiliser treatment. Our results indicate that fertiliser treatments that improve cereal‐aphid fitness will improve parasitoid fitness as measured by parasitoid size but may not influence percentage parasitism.     

Within-patch foraging behaviour of the aphid parasitoidAphidius funebris: plant architecture, host behaviour, and individual variation

The influence of plant architecture, host colony size, and host colony structure on the foraging behaviour of the aphid parasitoidAphidius funebris Mackauer (Hymenoptera: Aphidiidae) was investigated using a factorial experimental design. The factorial design involved releasing individual parasitoid females in aphid colonies consisting of either 10 or 20 individuals ofUroleucon jaceae L. (Homoptera: Aphididae) of either only larval instar L3 or a mixture of host instars, both on unmanipulated plants and on plants that had all leaves adjacent to the colony removed. Interactions between the parasitoid and its host were recorded until the parasitoid had left the plant. The time females spent on the host plant and the number of eggs laid varied greatly among females. Host colony size significantly affected patch residence time and the number of contacts between parasitoids and aphids. Plant architecture influenced the time-budget of the parasitoids which used leaves adjacent to the aphid colony for attacking aphids. Female oviposition rate was higher on unmanipulated plants than on manipulated plants. No further significant treatment effects on patch residence time, the number of contacts, attacks or ovipositions were found. Oviposition success ofA. funebris was influenced by instar-specific host behaviour. Several rules-of-thumb proposed by foraging theory did not account for parasitoid patch-leaving behaviour.     

Larvicidal activity of lectins onLucilia cuprina: mechanism of action

Foraging behaviour and host-instar preference of young and old females of the solitary aphid parasitoid,Lysiphlebus cardui Marshall (Hymenoptera: Aphidiidae), were studied in the laboratory. The analysis of interactions between parasitoids and different stages ofAphis fabae cirsiiacanthoidis Scop. (Homoptera: Aphididae) revealed that encounter rates between aphids and parasitoid females and defence reactions of the aphids influenced the degree to which a particular aphid age class is parasitized. Encounter rates between hosts and parasitoid females depended on the foraging pattern of the parasitoid, which varied with age. In mixed aphid colonies patch residence time increased with parasitoid age. Furthermore, younger parasitoids (≦1 day old) laid more eggs into second and third instars, while older parasitoids (≧4 days old) did not show distinct host instar preferences. It is suggested that the oviposition behaviour ofL. cardui is influenced by the physiological state, i.e. the age of the wasp.     

Comparing and contrasting life history variation in four aphid hyperparasitoids

1. In primary parasitoids, significant differences in life history and reproductive traits are observed among parasitoids attacking different stages of the same host species. Much less is known about hyperparasitoids, which attack different stages of primary parasitoids. 2. Parasitoids exploit hosts in two different ways. Koinobionts attack hosts that continue feeding and growing during parasitism, whereas idiobionts paralyse hosts before oviposition or attack non‐growing host stages, e.g. eggs or pupae. 3. Koino‐/idiobiosis in primary parasitoids are often associated with different expression of life history trade‐offs, e.g. endo‐ versus ectoparasitism, high versus low fecundity and short versus long life span. 4. In the present study, life history parameters of two koinobiont endoparasitic species (Alloxysta victrix; Syrphophagus aphidivorus), and two idiobiont ectoparasitic species (Asaphes suspensus; Dendrocerus carpenteri) of aphid hyperparasitoids were compared. These hyperparasitoids attack either the parasitoid larva in the aphid before it is killed and mummified by the primary parasitoid or the parasitoid prepupa or pupa in the dead aphid mummy. 5. There was considerable variation in reproductive success and longevity in the four species. The idiobiont A. suspensus produced the most progeny by far and had the longest lifespan. In contrast, the koinobiont A. victrix had the lowest fecundity. Other developments and life history parameters in the different species were variable. 6. The present results reveal that there was significant overlap in life history and reproductive traits among hyperparasitoid koinobionts and idiobionts, even when attacking the same host species, suggesting that selection for expression of these traits is largely association specific.     

Influence of cornicle droplet secretions of the cabbage aphid,Brevicoryne brassicae,on parasitism behavior of na＇fve and experienced Diaeretiella rapae

Insects have evolved amazing methods of defense to ward off enemies. Many aphids release cornicle secretions when attacked by predators and parasitoids. These se cretions contain an alarm pheromone that alerts other colony members of danger, thereby providing indirect fitness benefits to the releaser. In addition, contact with cornicle se cretions could also threaten an attacker and could provide direct fitness to the releaser. However, cornicle secretions may also be recruited as a kairomonal cue by aphid natural enemies. In this study, we investigated the effect of the cornicle droplet volatiles of the cabbage aphid, Brevicoryne brassicae （L.）, on the hostsearching behavior of naive and experienced female Diaeretiella rapae （M＇Intosh） parasitoids in olfactometer studies. In addition, we evaluated the role ofB. brassicae cornicle droplets on the oviposition prefer ence of the parasitoid in a twochoice bioassay. Naive females did not exhibit any preference between volatiles from aphids secreting cornicle droplets over nonsecreting aphids, while experienced parasitoids exploited the secretions in their host location. Experienced females were also able to choose volatiles from both secreting and nonsecreting aphids over clean air, while this ability was not observed in naive females. Although secretion of cornicle droplets did not influence the percentage of first attack in either naive or experienced females, the success of attack （i.e. resulting in a larva） was significantly different between secreting and nonsecreting aphids in the case of experienced parasitoids.     

Host genotype affects the relative success of competing lines of aphid parasitoids under superparasitism

1. In solitary parasitoids, only one individual can complete development in a given host. Therefore, solitary parasitoids tend to prefer unparasitised hosts for oviposition, yet under high parasitoid densities, superparasitism is frequent and results in fierce competition for the host's limited resources. This may lead to selection for the best intra‐host competitors. 2. Increased intra‐host competitive ability may evolve under a high risk of superparasitism if this trait exhibits genetic variation, and if competitive differences among parasitoid genotypes are consistent across environments, e.g. different host genotypes. 3. These assumptions were addressed in the aphid parasitoid Lysiphlebus fabarum (Hymenoptera: Braconidae: Aphidiinae) and its main host, the black bean aphid, Aphis fabae (Scopoli) (Hemiptera: Aphididae). Three parthenogenetic lines of L. fabarum were allowed to parasitise three aphid clones singly and in all pairwise combinations (superparasitism). The winning parasitoid in superparasitised aphids was determined by microsatellite analysis. 4. The proportions of singly parasitised aphids that were mummified were similar for the three parasitoid lines and did not differ significantly among host clones. 5. Under superparasitism, significant biases in favour of one parasitoid line were observed for some combinations, indicating that there is genetic variation for intra‐host competitive ability. However, the outcome of superparasitism was inconsistent across aphid clones and thus influenced significantly by the host clone in which parasitoids competed. 6. Overall, this study shows that the fitness of aphid parasitoids under superparasitism is determined by complex interactions with competitors as well as hosts, possibly hampering the evolution of improved intra‐host competitive ability.     

Are viviparous lizards from cool climates ever exclusively nocturnal? Evidence for extensive basking in a New Zealand gecko

Viviparity has evolved numerous times among squamate reptiles; however, the combination of viviparity and nocturnality is apparently rare among lizards. We used time‐lapse photography to examine evidence for diurnal activity in a viviparous lizard often described as nocturnal, the gecko Woodworthia ‘Otago/Southland’ from southern New Zealand (family Diplodactylidae). Evidence for diurnal emergence was extensive. Females have a higher incidence of basking compared to males, although no difference was detected between females in different reproductive conditions. Temperature loggers inserted into calibrated copper models were used to compare the body temperatures available to geckos in two basking positions and in two retreat types. Models in basking positions reached higher mean temperatures than models in retreats, although there was no significant effect of basking position or retreat type on model temperatures. Collectively, our results indicate that pregnant geckos that bask consistently could reduce gestation length by at least 14 days compared with females that remain in retreats. Extensive basking in this species adds to the growing evidence of diurno‐nocturnality in many New Zealand lepidosaurs, including other viviparous geckos. Our results lead us to question whether viviparity in lizards is ever compatible with ‘pure’ nocturnality in a cool climate. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Effects of pea aphid secondary endosymbionts on aphid resistance and development of the aphid parasitoid Aphidius ervi: a correlative study

In order to reduce parasite‐induced mortality, hosts may be involved in mutualistic interactions in which the partner contributes to resistance against the parasite. The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae), harbours secondary bacterial endosymbionts, some of which have been reported to confer resistance against aphid parasitoids. Although this resistance often results in death of the developing parasitoid larvae, some parasitoid individuals succeed in developing into adults. Whether these individuals suffer from fitness reduction compared to parasitoids developing in pea aphid clones without symbionts has not been tested so far. Using 30 pea aphid clones that differed in their endosymbiont complement, we studied the effects of these endosymbionts on aphid resistance against the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae: Aphidiinae), host–parasitoid physiological interactions, and fitness of emerging adult parasitoids. The number of symbiont species in an aphid clone was positively correlated with a number of resistance measurements but there were also clear symbiont‐specific effects on the host–parasitoid interaction. As in previous studies, pea aphid clones infected with Hamiltonella defensa Moran et al. showed resistance against the parasitoid. In addition, pea aphid clones infected with Regiella insecticola Moran et al. and co‐infections of H. defensa–Spiroplasma, R. insecticola–Spiroplasma, and R. insecticola–H. defensa showed reduced levels of parasitism and mummification. Parasitoids emerging from symbiont‐infected aphid clones often had a longer developmental time and reduced mass. The number of teratocytes was generally lower when parasitoids oviposited in aphid clones with a symbiont complement. Interestingly, unparasitized aphids infected with Serratia symbiotica Moran et al. and R. insecticola had a higher fecundity than unparasitized aphids of uninfected pea aphid clones. We conclude that in addition to conferring resistance, pea aphid symbionts also negatively affect parasitoids that successfully hatch from aphid mummies. Because of the link between aphid resistance and the number of teratocytes, the mechanism underlying resistance by symbiont infection may involve interference with teratocyte development.     

Host stage selection of the mealybug parasitoid Anagyrus spec. nov near sinope

The mealybug parasitoid Anagyrus spec. nov near sinope (Hymenoptera: Encyrtidae) is an undescribed parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Homoptera: Pseudococcidae). We investigated the preference of Anagyrus spec. nov near sinope for six developmental stadia (first‐ and second‐instar nymphs, third‐instar immature females, third‐ or fourth‐instar immature males, pre‐reproductive adult females, and ovipositing adult females) of P. madeirensis and the fitness consequences of the host stage selection behavior. In the no‐choice test, Anagyrus spec. nov near sinope parasitized and completed development in all host stadia except third‐instar immature males. When all host stadia were offered simultaneously, the parasitoids preferred third‐instar immature and pre‐reproductive adult females. Dissection of the stung mealybugs revealed that the clutch size (number of eggs per host) was approximately four and three in the third‐instar and pre‐reproductive females, respectively, and one egg per first‐instar nymph. Parasitoids emerged from P. madeirensis parasitized at third‐instar or pre‐reproductive adult female completed development in the shortest duration, achieved a higher progeny survival rate, larger brood and body size, and the lowest proportion of males. We showed that the continued development of mealybugs had significant influence on the fitness of the parasitoids. Although deposited as eggs in first‐ or second‐instar nymphs, parasitoids emerged from mummies that had attained third‐instar or adult development achieved similar progeny survival rate, brood size, body size, and sex ratio as those parasitoids deposited and developed in third‐instar or adult mealybugs. By delaying larval development in young mealybugs, Anagyrus spec. nov near sinope achieved higher fitness by allowing the parasitized mealybugs to grow and accumulate body size and resources. We suggest that the fitness consequence of host stage selection of a koinobiont parasitoid should be evaluated on both the time of parasitism and the time of mummification.     

Using multiple traits to estimate the effects of heat shock on the fitness of Aphidius colemani

Extreme temperature events are a great challenge for most ectotherms, particularly for the immature stages of parasitoids, as they do not possess the ability to behaviourally thermoregulate. In this study, we measured the effect of an acute heat shock, combined with desiccation stress (34 °C and 35% r.h. during 10 h) during the mummy stage on several fitness‐related traits of emerging adults of the aphid parasitoid Aphidius colemani Viereck (Hymenoptera: Braconidae: Aphidiinae). Our results showed that the emergence rate was strongly affected by the heat shock (63 ± 2.3 vs. 96.1 ± 0.7% for stress vs. control conditions, respectively), and the resulting population was male biased. Heat stress reduced the lifespan of emerged parasitoids but had no effect on fat reserves and female size. Egg load at emergence and parasitism rate were also reduced by heat treatment. The antennal symmetry was disturbed by the heat treatment, and stressed individuals had reduced mating success compared to control parasitoids. Specifically, time before wing fanning, a typical male courtship behaviour, was significantly longer in parasitoid pairs involving females that had emerged from heat‐treated mummies. Additionally, olfactometry tests on experimental females indicated that their response to host odour was similar to that of control individuals, but they required a longer time for perception. These results highlight that exposure to high temperatures during the mummy stage has cumulative detrimental effects that may strongly impact parasitoid populations under natural conditions and hinder the success of biological control programs.     

Food web structure of aphids and their parasitoids in Belgian fruit agroecosystems

Community structures of aphids and their parasitoids were studied in fruit crop habitats of eastern Belgium in 2014 and 2015. Quantitative food webs of these insects were constructed separately for each year, and divided into subwebs on three host‐plant categories, fruit crop plants, non‐crop woody and shrub plants and non‐crop herbaceous plants. The webs were analyzed using the standard food web statistics designed for binary data. During the whole study period, 78 plant species were recorded as host plants of 71 aphid species, from which 48 parasitoid species emerged. The community structure, aphid / parasitoid species‐richness ratio and trophic link number varied between the two years, whereas the realized connectance between parasitoids and aphids was relatively constant. A new plant–aphid–parasitoid association for Europe was recorded. Dominant parasitoid species in the study sites were Ephedrus persicae, Binodoxys angelicae and Praon volucre: the first species was frequently observed on non‐crop trees and shrubs, but the other two on non‐crop herbaceous plants. The potential influence, through indirect interactions, of parasitoids on aphid communities was assessed with quantitative parasitoid‐overlap diagrams. Symmetrical links were uncommon, and abundant aphid species seemed to have large indirect effects on less abundant species. These results show that trophic indirect interactions through parasitoids may govern aphid populations in fruit crop habitats with various non‐crop plants, implying the importance for landscape management and biological control of aphid pests in fruit agroecosystems.     

Thermal change alters the outcome of behavioural interactions between antagonistic partners

1. Concerns about climate change often trigger the question whether physiological and behavioural responses of species will enable them to persist. However, species do not exist alone and are largely dependent on interactions with others within communities. 2. In the present study, a mechanistic approach is used to test the hypothesis that inter‐specific differences in metabolic response to unpredictable short‐term thermal changes can change the outcome of host–parasitoid behavioural interactions. 3. The effect of a drop or a rise of 5 °C on resting metabolic rates (RMR) of the main aphid pest of cereal crops in Western Europe, the host Sitobion avenae Fabricius and its main natural enemy, the parasitoid Aphidius rhopalosiphi De Stefani‐Perez was measured. Also, defence and attack behaviours were measured for host and parasitoid separately as well as in interaction, since behavioural strategies of both species largely determine parasitism success. 4. The results showed that, when no change in temperature occurred, parasitoids had the highest oviposition rate. However, only with a rise of temperature behavioural interactions were disrupted: the parasitoid attack rate decreased whereas the aphid defence rate increased. This alteration in behaviour was associated with a stronger thermal response of RMR in hosts than in parasitoids, suggesting that species‐specific thermal responses of RMR could give valuable information on changes in the outcome of species interactions under warm spells but not under cold ones. 5. It was shown that relatively modest thermal changes with non‐lethal effects can have profound consequences for interacting co‐evolved species which may affect ecosystem services, such as biological control of pest populations.     

Winter host exploitation influences fitness traits in a parasitoid

Organisms can either evade winter's unfavourable conditions by migrating or diapausing, or endure them and maintain their activities. When it comes to foraging during winter, a period of scarce resources, there is strong selective pressure on resource exploitation strategy. Generalist parasitoids are particularly affected by this environmental constraint, as their fitness is deeply linked to the profitability of the available hosts. In this study, we considered a cereal aphid–parasitoid system and investigated (1) the host–parasitoid community structure, host availability, and parasitism rate in winter, (2) the influence of host quality in terms of species and instars on the fitness of the aphid parasitoid Aphidius rhopalosiphi De Stefani‐Perez (Hymenoptera: Braconidae: Aphidiinae), and (3) whether there is a detectable impact of host fidelity on parasitism success of this parasitoid species. Host density was low during winter and the aphid community consisted of the species Rhopalosiphum padi L. and Sitobion avenae Fabricius (both Hemiptera: Aphididae), both parasitized by A. rhopalosiphi at non‐negligible rates. Aphidius rhopalosiphi produced more offspring when parasitizing R. padi compared with S. avenae, whereas bigger offspring were produced when parasitizing S. avenae. Although aphid adults and old larvae were significantly larger hosts than young larvae, the latter resulted in higher emergence rates and larger parasitoids. No impact of host fidelity on emergence rates or offspring size was detected. This study provides some evidence that winter A. rhopalosiphi populations are able to take advantage of an array of host types that vary in profitability, indicating that host selectivity may drop under winter's unfavourable conditions.     

Rapid evolution of parasitoids when faced with the symbiont-mediated resistance of their hosts

Insects harbour a wild diversity of symbionts that can spread and persist within populations by providing benefits to their host. The pea aphid Acyrthosiphon pisum maintains a facultative symbiosis with the bacterium Hamiltonella defensa, which provides enhanced resistance against the aphid parasitoid Aphidius ervi. Although the mechanisms associated with this symbiotic‐mediated protection have been investigated thoroughly, little is known about its evolutionary effects on parasitoid populations. We used an experimental evolution procedure in which parasitoids were exposed either to highly resistant aphids harbouring the symbiont or to low innate resistant hosts free of H. defensa. Parasitoids exposed to H. defensa gained virulence over time, reaching the same parasitism rate as those exposed to low aphid innate resistance only. A fitness reduction was associated with this adaptation as the size of parasitoids exposed to H. defensa decreased through generations. This study highlighted the considerable role of symbionts in host–parasite co‐evolutionary dynamics.     

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.     

Dropping of pea aphids from feeding site: a consequence of parasitism by the wasp, Monoctonus paulensis

We examined the dropping behaviour of the pea aphid, Acyrthosiphon pisum (Harris), feeding on broad-bean plants in the laboratory. Aphid responses to foraging and oviposition by females of Monoctonus paulensis (Ashmead) were instar-specific and included kicking with the hind legs, walking away and dropping from the feeding site. Fourth nymphal instars were most likely to drop, followed by second, third, and first instars, in that order. Compared with first instars, the odds that a fourth-instar aphid will drop were 6-times higher independent of the stimulus, and 16-times higher after parasitoid attack. Dropping from the feeding site increases an aphid's mortality risk. If parasitoid offspring are more likely to survive in small pea aphids, it may be adaptive for M. paulensis to choose smaller hosts, regardless of possible fitness gains due to increased body size from development in larger aphids.