Predation potential of the earwig Euborellia annulipes on fruit fly larvae and trophic interactions with the parasitoid Diachasmimorpha longicaudata

The earwig Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae), a generalist predator, has been observed in fruits infested with fruit fly larvae, which are frequently parasitized by parasitoid wasps. Neither the capacity of earwigs to predate on fruit flies nor intraguild interactions between earwigs and fruit fly parasitoids have been investigated. Here, we studied in laboratory conditions the predation on the fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) by the earwig E. annulipes, and whether parasitism of fruit fly larvae by the parasitoid wasp Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) influences predation by the earwig. We evaluated the predation capacity, functional response and prey preference of E. annulipes for parasitized and non-parasitized fruit fly larvae in choice and no-choice tests. We found that earwigs prey on second- and third-instar larvae and pupae of C. capitata and consumed larger numbers of second-instar larvae, followed by third-instar larvae and pupae. Females prey on larger numbers of fruit flies than did males, regardless of the prey developmental stage, but both sexes exhibited a type II functional response. Interestingly, males killed but did not consume fruit fly larvae more than did females. In no-choice tests, earwig females consumed equal numbers of parasitized and non-parasitized fruit fly larvae. However, in choice tests, the females avoided feeding on parasitized larvae. Subsequent tests with hexane-washed parasitized and non-parasitized larvae showed that putative chemical markings left on fruit flies by parasitoids did not drive the earwig preference towards non-parasitized larvae. These findings suggest that E. annulipes is a potential biological control agent for C. capitata, and that, because the earwig avoids consuming larvae parasitized by D. longicaudata, a combination of the two natural enemies could have an additive effect on pest mortality.     

Polydnaviral Ankyrin Proteins Aid Parasitic Wasp Survival by Coordinate and Selective Inhibition of Hematopoietic and Immune NF-kappa B Signaling in Insect Hosts

Polydnaviruses are mutualists of their parasitoid wasps and express genes in immune cells of their Lepidopteran hosts. Polydnaviral genomes carry multiple copies of viral ankyrins or vankyrins. Vankyrin proteins are homologous to IκB proteins, but lack sequences for regulated degradation. We tested if Ichnoviral Vankyrins differentially impede Toll-NF-κB-dependent hematopoietic and immune signaling in a heterologous in vivo Drosophila, system. We first show that hematopoiesis and the cellular encapsulation response against parasitoid wasps are tightly-linked via NF-κB signaling. The niche, which neighbors the larval hematopoietic progenitors, responds to parasite infection. Drosophila NF-κB proteins are expressed in the niche, and non cell-autonomously influence fate choice in basal and parasite-activated hematopoiesis. These effects are blocked by the Vankyrin I²-vank-3, but not by P-vank-1, as is the expression of a NF-κB target transgene. I²-vank-3 and P-vank-1 differentially obstruct cellular and humoral inflammation. Additionally, their maternal expression weakens ventral embryonic patterning. We propose that selective perturbation of NF-κB-IκB interactions in natural hosts of parasitic wasps negatively impacts the outcome of hematopoietic and immune signaling and this immune deficit contributes to parasite survival and species success in nature.     

High hemocyte load is associated with increased resistance against parasitoids in Drosophila suzukii, a relative of D. melanogaster

Among the most common parasites of Drosophila in nature are parasitoid wasps, which lay their eggs in fly larvae and pupae. D. melanogaster larvae can mount a cellular immune response against wasp eggs, but female wasps inject venom along with their eggs to block this immune response. Genetic variation in flies for immune resistance against wasps and genetic variation in wasps for virulence against flies largely determines the outcome of any fly-wasp interaction. Interestingly, up to 90% of the variation in fly resistance against wasp parasitism has been linked to a very simple mechanism: flies with increased constitutive blood cell (hemocyte) production are more resistant. However, this relationship has not been tested for Drosophila hosts outside of the melanogaster subgroup, nor has it been tested across a diversity of parasitoid wasp species and strains. We compared hemocyte levels in two fly species from different subgroups, D. melanogaster and D. suzukii, and found that D. suzukii constitutively produces up to five times more hemocytes than D. melanogaster. Using a panel of 24 parasitoid wasp strains representing fifteen species, four families, and multiple virulence strategies, we found that D. suzukii was significantly more resistant to wasp parasitism than D. melanogaster. Thus, our data suggest that the relationship between hemocyte production and wasp resistance is general. However, at least one sympatric wasp species was a highly successful infector of D. suzukii, suggesting specialists can overcome the general resistance afforded to hosts by excessive hemocyte production. Given that D. suzukii is an emerging agricultural pest, identification of the few parasitoid wasps that successfully infect D. suzukii may have value for biocontrol.     

Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

Drosophila Avoids Parasitoids by Sensing Their Semiochemicals via a Dedicated Olfactory Circuit

Detecting danger is one of the foremost tasks for a neural system. Larval parasitoids constitute clear danger to Drosophila, as up to 80% of fly larvae become parasitized in nature. We show that Drosophila melanogaster larvae and adults avoid sites smelling of the main parasitoid enemies, Leptopilina wasps. This avoidance is mediated via a highly specific olfactory sensory neuron (OSN) type. While the larval OSN expresses the olfactory receptor Or49a and is tuned to the Leptopilina odor iridomyrmecin, the adult expresses both Or49a and Or85f and in addition detects the wasp odors actinidine and nepetalactol. The information is transferred via projection neurons to a specific part of the lateral horn known to be involved in mediating avoidance. Drosophila has thus developed a dedicated circuit to detect a life-threatening enemy based on the smell of its semiochemicals. Such an enemy-detecting olfactory circuit has earlier only been characterized in mice and nematodes.     

Host searching behavior and potential of an aquatic ichneumonid pupal parasitoid of rice caseworm (Parapoynx stagnalis) in an upland rice paddy agro-ecosystem of the Western Ghats,India

Parapoynx stagnalis (Zeller) (=Nymphula depunctalis (Guenée), is a sporadic rice paddy pest of south and southeast Asian rice paddy fields, and widely known as rice caseworm. A recent outbreak of the pest was observed in the upland rice paddy agro-ecosystem of the central Western Ghats, India. No potential natural enemies of this pest have so far been reported, mainly due to the semi-aquatic behavior of the larvae and pupae. However, an undescribed aquatic ichneumonid wasp species (belonging to a genus near Litochila) is reported as a potentially useful early stage pupal parasitoid of this pest in our study. The host searching behavior and potential of the parasitoid wasp was studied in detail using both field and laboratory experiments. The adult female wasps usually dive into the water in search of the host pupae, remaining under water for a maximum of 90.2s, while searching for rice caseworm pupae. In field and laboratory experiments, we found 73% of the parasitized pupae of P. stagnalis kept under water yielded adult parasitic wasps. However, no parasitoids emerged from parasitized host pupae maintained in dry terrariums. Sex ratio was 2.8:1. The present discovery of the semi-aquatic parasitoid wasp could aid in better management of the rice caseworm population in rice paddy fields.     

Linear Dispersal of the Filth Fly Parasitoid Spalangia cameroni (Hymenoptera: Pteromalidae) and Parasitism of Hosts at Increasing Distances

Release of parasitic wasps (Hymenoptera: Pteromalidae) as biological control agents for house flies and stable flies in livestock confinements has had variable success. In part, this may reflect a lack of knowledge regarding the optimal distance to be used between parasitoid release stations. In the current study, we assessed the effect of linear distance on host parasitism by the wasp Spalangia cameroni Perkins. In open fields at distances ranging from 1 m to 60 m from a central point, house fly puparia were placed in a mixture of pine shavings soiled with equine manure, urine, and alfalfa hay. Releases of S. cameroni then were made using a 5:1 host: parasitoid ratio. Host pupae were parasitized at all distances, with the highest rate of total parasitism (68.9%) recorded ≤ 5 m from the release site. Analyses of results using non-linear and linear models suggest that S. cameroni should be released in close proximity to host development areas. Additionally, releases may not be suitable in pasture situations where long-distance flight is required for control. However, further testing is needed to examine the effect of density-dependent dispersal and diffusion of S. cameroni.     

Deadly venom of Asobara japonica parasitoid needs ovarian antidote to regulate host physiology

Asobara japonica (Braconidae) is an endophagous parasitoid developing in Drosophila larvae. The present study shows that A. japonica was never encapsulated in Drosophila melanogaster, and that it caused an overall inhibition of the host encapsulation reaction since injected foreign bodies were never encapsulated in parasitized hosts. Both the number of circulating hemocytes and the phenoloxidase activity decreased in parasitized larvae, and the hematopoietic organ appeared highly disrupted. We also found that A. japonica venom secretions had atypical effects on hosts compared to other braconid wasps. A. japonica venom secretions induced permanent paralysis followed by death of D. melanogaster larvae, whether injected by the female wasp during an interrupted oviposition, or manually injected into unparasitized larvae. More remarkably, these effects could be reversed by injection of ovarian extracts from female wasps. This is the first report that the venom of an endophagous braconid parasitoid can have a deadly effect on hosts, and moreover, that ovarian extracts can act as an antidote to reverse the effects of the wasp's venom. These results also demonstrate that A. japonica secretions from both venom gland and ovary are required to regulate synergistically the host physiology for the success of the parasitoid.     

Identification of seven species of hymenopteran parasitoids of Spodoptera frugiperda, using polymerase chain reaction amplification and restriction enzyme digestion

1 The fall armyworm Spodoptera frugiperda is a voracious pest of numerous crops of economic importance throughout the New World. In its native Mexico, larvae can be attacked by several species of parasitic wasps, which are candidate biological control agents against this and other lepidopteran pests. 2 We attempted to survey the parasitoid fauna on S. frugiperda in maize and sorghum fields throughout Mexico. However, our efforts have been hampered by the incomplete development of parasitoid larvae emerging from collected Spodoptera caterpillars. 3 This problem was solved by developing a method to identify seven species of parasitic wasps using polymerase chain reaction amplification and restriction enzyme digestion. This enables the precise determination of the species of those parasitoid larvae that are usually not morphologically identifiable.     

Bioinformatic analysis suggests potential mechanisms underlying parasitoid venom evolution and function

Hymenopteran parasitoid wasps are a diverse collection of species that infect arthropod hosts and use factors found in their venoms to manipulate host immune responses, physiology, and behaviour. Whole parasitoid venoms have been profiled using proteomic approaches, and here we present a bioinformatic characterization of the venom protein content from Ganaspis sp. 1, a parasitoid that infects flies of the genus Drosophila. We find evidence that diverse evolutionary processes including multifunctionalization, co-option, gene duplication, and horizontal gene transfer may be acting in concert to drive venom gene evolution in Ganaspis sp.1. One major role of parasitoid wasp venom is host immune evasion. We previously demonstrated that Ganaspis sp. 1 venom inhibits immune cell activation in infected Drosophila melanogaster hosts, and our current analysis has uncovered additional predicted virulence functions. Overall, this analysis represents an important step towards understanding the composition and activity of parasitoid wasp venoms.     

Edin Expression in the Fat Body Is Required in the Defense Against Parasitic Wasps in Drosophila melanogaster

The cellular immune response against parasitoid wasps in Drosophila involves the activation, mobilization, proliferation and differentiation of different blood cell types. Here, we have assessed the role of Edin (elevated during infection) in the immune response against the parasitoid wasp Leptopilina boulardi in Drosophila melanogaster larvae. The expression of edin was induced within hours after a wasp infection in larval fat bodies. Using tissue-specific RNAi, we show that Edin is an important determinant of the encapsulation response. Although edin expression in the fat body was required for the larvae to mount a normal encapsulation response, it was dispensable in hemocytes. Edin expression in the fat body was not required for lamellocyte differentiation, but it was needed for the increase in plasmatocyte numbers and for the release of sessile hemocytes into the hemolymph. We conclude that edin expression in the fat body affects the outcome of a wasp infection by regulating the increase of plasmatocyte numbers and the mobilization of sessile hemocytes in Drosophila larvae.     

Survey of the native insect natural enemies of Hyphantria cunea (Drury) (Lepidoptera: Arctiidae) in China

The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae), is an invasive and important pest in China. Investigations on insect natural enemies have been conducted from 1996 to 1999 in five provinces and one municipality of China in order to select effective species for biological control. Two carabid predators (Coleoptera) and 25 parasitoid species were found, among which 23 were parasitic wasps (Hymenoptera), including five hyperparasitic species and two tachinid flies (Diptera). The two carabids preyed on young larvae inside webs, two braconid wasps parasitized larvae, and 18 parasitoid species attacked the fall webworm during the pupal and/or 'larval-pupal' stages. Among these parasitoids, there were one genus and nine species that are new to science and four species new to China, which were described and published by the senior author Yang. The average parasitism rates of fall webworm pupae were 25.8% and 16.1% in the overwintering generation and the first generation (summer generation), respectively. These findings reveal that these natural enemies play an important role in the natural control of the pest. Chouioia cunea Yang (Hymenoptera: Eulophidae), a gregarious pupal endo-parasitoid, was recommended as a promising biological control agent against the fall webworm in China.     

Tissue communication in a systemic immune response of Drosophila

Several signaling pathways, including the JAK/STAT and Toll pathways, are known to activate blood cells (hemocytes) in Drosophila melanogaster larvae. They are believed to regulate the immune response against infections by parasitoid wasps, such as Leptopilina boulardi, but how these pathways control the hemocytes is not well understood. Here, we discuss the recent discovery that both muscles and fat body take an active part in this response. Parasitoid wasp infection induces Upd2 and Upd3 secretion from hemocytes, leading to JAK/STAT activation mainly in hemocytes and in skeletal muscles. JAK/STAT activation in muscles, but not in hemocytes, is required for an efficient encapsulation of wasp eggs. This suggests that Upd2 and Upd3 are important cytokines, coordinating different tissues for the cellular immune response in Drosophila. In the fat body, Toll signaling initiates a systemic response in which hemocytes are mobilized and activated hemocytes (lamellocytes) are generated. However, the contribution of Toll signaling to the defense against wasps is limited, probably because the wasps inject inhibitors that prevent the activation of the Toll pathway. In conclusion, parasite infection induces a systemic response in Drosophila larvae involving major organ systems and probably the physiology of the entire organism.     

Aggressive Behavior in Olive Fruit Fly Females: Oviposition Site Guarding against Parasitic Wasps

Among Tephritidae flies, the females display agonistic behavior to maintain single oviposition sites and reduce larval competition for food. In the olive fruit fly, Bactrocera oleae, female-female aggressive interactions are characterized by reciprocal wing waving, chasing, head butting and boxing with forelegs. Little is known on tephritid aggressive behaviors directed towards natural enemies, with special reference to parasitoids attacking their young instars. In this study, we quantified the aggressive behavior of B. oleae females guarding their oviposition site against the braconid parasitoid Psyttalia concolor. The fly aggressive behavior displayed against the parasitoids was compared that directed towards paper dummies mimicking P. concolor adults. When a P. concolor female came close (<20 mm) to a B. oleae female guarding the oviposition site, the 91.67% of the flies displayed wing-waving, the 63.34% chased the parasitoid, the 45% showed head-butting, while boxing was observed only in the 26.67% of the aggressions. When paper dummies were tested, only the 66.67% of the flies displayed wing-waving, the 8.33% performed chasing, followed by head butting (5.00%) and boxing (3.33%). B. oleae displayed longer aggression bouts towards live wasps over dummies. Overall, this is the first evidence pointing out that tephritid aggressive acts, besides their role in intraspecific interactions, were also highly effective to displace parasitic wasps from the fly oviposition site. Further research on potential consequences on fitness traits arising from the above-discussed behaviors, as well as on parasitoid learning-mediated responses to tephritid aggressions, is urgently required.     

Behavioral Attraction of Two Parasitoids,Venturia canescens and Bracon hebetor,to Silk Extracts of a Host Plodia interpunctella

Kairomonal activities of silk extracts of host Plodia interpunctella were determined by measuring the rates of behavioral responses of two parasitic wasps, Venturia canescens and Bracon hebetor. Silk of P. interpunctella larvae attracted both parasitic wasps but the cocoon silk of silkworm, Bombyx mori and the web silk of two-spotted spider mite, Tetranychus urticae did not. Silk components of P. interpunctella were extracted by using either hexane or methanol, and tested the rates of three serial responses of wasps; host location, antennal drumming and ovipositor probing behaviors. The patterns of each behavioral response were similar in two wasps. The rates of each response were increased at the higher concentrations of both extracts. Antennal drumming behavior was much more responsive to lower concentrations of both extracts than ovipositor probing behavior was. Furthermore, the rate of antennal drumming response was higher in hexane-extracts rather than methanol-extract in both wasps; V. canescens and B. hebetor for 20 and 17 times, respectively. However, ovipositor probing response was similar in two different extracts. Both extracts elicited 100% of antennal drumming response but ovipositor probing response was only 60 to 80% of all tested individuals. Our results were shown that silk extracts of host larvae elicited strong behavioral responses of two parasitic wasps and could be applied for practical application of parasitoids attraction in the biological control of agricultural pests.     

蝇蛹俑小蜂对寄主及果实的趋性反应

【背景】蝇蛹俑小蜂是实蝇类害虫蛹期重要的寄生蜂之一。有关植物果实和寄主挥发物在蝇蛹俑小蜂寻找寄主过程中的作用尚不明确。【方法】采用"Y"型嗅觉仪测试了寄主和果实(健康果实和虫害诱导果实)对蝇蛹俑小蜂的引诱作用。【结果】虫伤1 d的番石榴和杨桃果实对蝇蛹佣小蜂雌蜂具有显著的引诱作用;寄主蛹——3日龄橘小实蝇蛹和4日龄瓜实蝇蛹对蝇蛹俑小蜂雌蜂也具有显著的引诱作用。蝇蛹俑小蜂雌蜂对雄蜂有显著的吸引作用。橘小实蝇性诱剂甲基丁香酚对蝇蛹俑小蜂的搜索行为则无显著影响。【结论与意义】寄主蛹和幼虫危害1 d的果实能释放对蝇蛹俑小蜂有引诱作用的物质,这可为利用蝇蛹俑小蜂防治实蝇类害虫提供基础信息。     

Entomological and functional role of floral strips in an organic apple orchard: Hymenopteran parasitoids as a case study

Habitat manipulation techniques improve the availability of resources required by natural enemies to increase their effectiveness. This study focused on the effects of floral strips on Hymenopteran parasitoid presence. The experiments were conducted during spring 2007 in one organic low-input apple orchard located in south-eastern France. The density and the diversity of parasitic wasps collected from sown floral strips were higher than those from mown plants. The family of parasitic wasps of Braconidae was strongly dominant, followed by Mymaridae and Pteromalidae. By studying 26 flowering species, the greatest diversity and density of parasitic wasps were collected from Potentilla reptans, Achillea millefolium, Trifolium repens and Torilis arvensis. In terms of the early flowering plants, the most important results were observed in Euphorbia helioscopia, Senecio vulgaris and Veronica persica. To give an idea of the functional role of these plants, we studied the parasitic wasps of the diapausing larvae (cocoon) of codling moth Cydia pomonella. We recorded three emerged species: Ascogaster quadridentata, Pristomerus vulnerator and the hyperparasite Perilampus fulvicornis. However, none of these species have been observed on the 26 studied plants. Hence, this result may be suggesting that the studied plants do not have a functional role concerning these parasitoids. These studies may be advantageous for biological control programs in order to select flowering plant species attracting parasitic wasps specific to fruit pests.     

Larval Defense against Attack from Parasitoid Wasps Requires Nociceptive Neurons

Parasitoid wasps are a fierce predator of Drosophila larvae. Female Leptopilina boulardi (LB) wasps use a sharp ovipositor to inject eggs into the bodies of Drosophila melanogaster larvae. The wasp then eats the Drosophila larva alive from the inside, and an adult wasp ecloses from the Drosophila pupal case instead of a fly. However, the Drosophila larvae are not defenseless as they may resist the attack of the wasps through somatosensory-triggered behavioral responses. Here we describe the full range of behaviors performed by the larval prey in immediate response to attacks by the wasps. Our results suggest that Drosophila larvae primarily sense the wasps using their mechanosensory systems. The range of behavioral responses included both “gentle touch” like responses as well as nociceptive responses. We found that the precise larval response depended on both the somatotopic location of the attack, and whether or not the larval cuticle was successfully penetrated during the course of the attack. Interestingly, nociceptive responses are more likely to be triggered by attacks in which the cuticle had been successfully penetrated by the wasp. Finally, we found that the class IV neurons, which are necessary for mechanical nociception, were also necessary for a nociceptive response to wasp attacks. Thus, the class IV neurons allow for a nociceptive behavioral response to a naturally occurring predator of Drosophila.     

Parasitoid hymenopterans of Costa Rica: Geographical distribution of the species associated with fruit flies [Diptera: Tephritidae]

The geographical distribution of the parasitoid species associated with fruit flies in Costa Rica is presented. In a study of 319 fruit samples infested with larvae of fruit flies, collected from 135 localities, 8 parasitoid species were recovered from 11.0% of the fruit samples. Two species are considered to be indigenous, two were introduced for the biological control ofCeratitis capitata (Wiedemann) in this country, at least 3 unidentified species of eucoiline cynipoids and one species of uncertain origin was found for the first time. The identified parasitoids were:Doryctobracon crawfordi (Viereck) (indigenous),D. areolatus (Szépligeti) (indigenous),Biosteres longicaudatus Ashmead (introduced),Aceratoneuromyia indica (Silvestri) (introduced) andTrichopria sp. (indigenous?). The introduced species were more frequently associated withAnastrepha spp. than withC. capitata. The need for systematic monitoring of populations of released parasitoid species is suggested.     

Cellular and humoral immune interactions between Drosophila and its parasitoids

The immune interactions occurring between parasitoids and their host insects, especially in Drosophila–wasp models, have long been the research focus of insect immunology and parasitology. Parasitoid infestation in Drosophila is counteracted by its multiple natural immune defense systems, which include cellular and humoral immunity. Occurring in the hemocoel, cellular immune responses involve the proliferation, differentiation, migration and spreading of host hemocytes and parasitoid encapsulation by them. Contrastingly, humoral immune responses rely more heavily on melanization and on the Toll, Imd and Jak/Stat immune pathways associated with antimicrobial peptides along with stress factors. On the wasps’ side, successful development is achieved by introducing various virulence factors to counteract immune responses of Drosophila. Some or all of these factors manipulate the host's immunity for successful parasitism. Here we review current knowledge of the cellular and humoral immune interactions between Drosophila and its parasitoids, focusing on the defense mechanisms used by Drosophila and the strategies evolved by parasitic wasps to outwit it.