Mite parasitization on armored scale insects: host suitability

Suitability of three armored scale insects (Homoptera: Diaspididae), their stages, the host plants and their parts, for the parasitic miteHemisarcoptes coccophagus Meyer was investigated. The studied scales were chaff scales,Parlatoria pergandii Comstock andP. cinerea Hadden, infesting grapefruit and orange orchards, latania scale,Hemiberlesia lataniae (Signoret), and oleander scale,Aspidiotus nerii Bouche occurring on acacia (Acacia cyanophylla) branches and leaves. Host suitability was evaluated by mite survival, mean intensity (active mites/attacked scales) and fecundity. It was highest on latania scale, lowest on oleander scale and intermediate on chaff scales. Results of laboratory experiments were consistent with these field findings: while developing on oleander scale, mites suffered 3× higher mortality, and their generation time was twice as long as on latania scale; the full egg clutch on the latter was high whereas on oleander scale it was negligible. Our data suggest that ovipositing females of latania and oleander scales, but not of chaff scales, are more suitable hosts than young scale females and the latter more suitable than immature stages. No direct effects of acacia plant parts on host suitability were detected. Citrus species affected chaff scale suitability for the mite through their effect on host diaspidid size. The ability ofH. coccophagus to control pestiferous diaspidids may depend on the specific host scale, its age structure and the specific host plant. The implications of these findings for the biological control of armored scale insects are discussed.     

Population dynamics ofHemisarcoptes coccophagus Meyer (Astigmata: Hemisarcoptidae) attacking three species of armored scale insects (Homoptera: Diaspididae)

The parasitic miteHemisarcoptes coccophagus Meyer (Astigmata: Hemisarcoptidae) feeds on armored scale insects (Homoptera: Diaspididae). Three host scales were studied: chaff scale (Parlatoria pergandii Comstock andP. cinerea Hadden) infesting citrus (grapefruit and orange) orchards, latania scale [Hemiberlesia lataniae (Signoret)] and olcander scale (Aspidiotus nerii Bouche) occurring on acacia (Acacia cyanophylla) branches and leaves. The overall dynamics of mite populations were similar on the different scale species, although the level of attack on each sampled diaspidid host was different. Rates of young mite stages were highly variable on all three hosts throughout the sampling period while adult rates were stable, constituting around 20% of all mites. No oviposition by mites occurred in mid-winter on any scale species, on any plants or parts, at all sampling sites. Of the three diaspidids, highest mite prevalence (parasitization rate) occurred on latania scale. Mite prevalence on oleander scale infesting mature leaves was higher than on young leaves.H. coccophagus apparently first attacks latania scale on branches, it then disperses to the same host inhabiting mature leaves, from there to co-occurring oleander scale and finally to oleander scale infesting young leaves. Mite prevalence on chaff scale was influenced by host plants and climatic regions: in the Negev it was higher than in the coastal plain, with scales on grapefruit being more heavily attacked at the former site but not on coastal plain. Our data suggest thatH. coccophagus prefers ovipositing scale females. The decreasing order of preference for other scale stages was: young females immature males, 2nd-instar nymphs females.     

Priming Aphytis: behavioral modification of host selection by exposure to a synthetic contact kairomone

The introduced parasitoid, Aphytis melinus DeBach (Hymenoptera: Aphelinidae), utilizes a kairomone, O-caffeoyltyrosine, to recognize California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae). Wasps used in augmentative release programs for California red scale on California citrus are reared on oleander scale, Aspidiotus nerii Bouché (Homoptera: Diaspididae), themselves reared on squash. The goals of this study were: 1) to determine if long-term rearing on oleander scale had caused A. melinus to develop a preference for oleander scale, and 2) to determine if the preference of oleander-reared A. melinus for California red scale might be enhanced by exposing them to synthetic O-caffeoyltyrosine prior to release. Wasps that were removed from their hosts as pupae and allowed to emerge as adults isolated from their hosts retained a strong preference for California red scale regardless of rearing host. This preference was reduced if wasps were allowed to emerge from oleander scale, thus acquiring early adult experience with oleander scale. The preference for California red scale was restored, however, by exposing wasps reared on oleander scale to synthetic O-caffeoyltyrosine prior to bioassay. Exposure of A. melinus reared in commercial insectaries to O-caffeoyltyrosine prior to release may be a means to improve the effectiveness of such wasps in augmentative release programs to control California red scale.     

Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America

The introduced parasitoid,Aphytis melinusDeBach (Hymenoptera: Aphelinidae), is used for augmentative biological control of California red scale,Aonidiella aurantii(Maskell) (Homoptera: Diaspididae). Commercially reared wasps are reared on oleander scale,Aspidiotus neriiBouché (Homoptera: Diaspididae). Oleander scale covers lack the chemical,O-caffeoyltyrosine, a kairomone mediating host selection byA. melinus.Wasps reared on oleander scale but individually exposed, or primed, toO-caffeoyltyrosine more readily accepted California red scale for probing in laboratory bioassays and parasitized a greater proportion of available California red scale in the field than wasps reared similarly but not exposed toO-caffeoyltyrosine. Thus, it may be possible to improve host recognition of commercial, insectary-rearedA. melinusby exposing them toO-caffeoyltyrosine prior to release. The goal of this study was to develop an inexpensive but effective means of priming thousands of wasps simultaneously toO-caffeoyltyrosine. The most effective method, but potentially the most expensive, was simply to spray parasitized oleander scale on their host plant with diluteO-caffeoyltyrosine prior to wasp emergence. In additional experiments, using controlled doses of syntheticO-caffeoyltyrosine applied to scale covers, we showed that primed wasps require both a lower minimum dose ofO-caffeoyltyrosine for recognition and also respond to measuredO-caffeoyltyrosine doses more consistently than unprimed wasps. The ability to mass-prime thousands of wasps prior to release is a crucial step toward realizing the concept of behavioral improvement of host selection of parasitoids on a commercial scale.     

Evasion of the insect immune response by Moniliformis dubius (Acanthocephala): further observations on the origin of the envelope

The envelope around larvae of Moniliformis dubius appears to protect the parasite against immune recognition and encapsulation by the insect host's haemocytes. The origin of this envelope has been the subject of controversy although most evidence suggests it is parasite-derived. If host-derived, the envelope would be expected to share surface properties with host tissue. Thus, experiments were undertaken, transplanting parasites and host tissue to other insects and using haemocytic encapsulation as an assay for immune recognition, in order to compare the response to host tissue and to the parasite's envelope. Parasites without their envelopes, and pieces of tissue (ventral nerve cord) from the experimental host (the locust Schistocerca gregaria) were recognized as foreign and encapsulated in the cockroach, Periplaneta americana. The majority of parasites with their envelopes were unencapsulated or only partially encapsulated on transfer to their normal host, P. americana, indicating that the envelope does not have surface similarity to locust tissue. Cockroach-derived parasites with or without envelopes were not encapsulated in S. gregaria, suggesting that the larva itself can evade or inhibit the locust's recognition mechanism. However, since larvae which develop in S. gregaria are enclosed in an envelope, the formation of the envelope would seem to be an inherent feature of the parasite's development.     

Increased parasitization of California red scale in the field after exposing its parasitoid, Aphytis melinus, to a synthetic kairomone

The introduced parasitoid, Aphytis melinus DeBach (Hymenoptera: Aphelinidae), utilizes a kairomone, O-caffeoyltyrosine, to recognize California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae). Wasps used in augmentative release programs for California red scale on California citrus are reared on oleander scale, Aspidiotus nerii Bouché (Homoptera: Diaspididae), themselves reared on squash. Previous laboratory studies demonstrated that wasps reared on oleander scale but exposed to O-caffeoyltyrosine before bioassay more readily accepted California red scale covers for probing than wasps reared similarly but not exposed to O-caffeoyltyrosine. The goal of this study was to determine if wasps reared on oleander scale but exposed to O-caffeoyltyrosine actually parasitized a greater number of field-reared California red scale than unconditioned wasps. In order to distinguish between progeny of conditioned and unconditioned wasps in the field, we utilized two A.melinus strains that carried different electrophoretically distinguishable alleles. In half of the releases, Strain A was conditioned to O-caffeoyltyrosine and Strain B served as the control, while this pattern of conditioning was reversed for the remaining releases. Although there were differences in the rates of population growth of the two strains, whichever strain was the conditioned strain left a greater than expected number of progeny compared to the control strain in all comparisons. Exposure of A. melinus reared in commercial insectaries to O-caffeoyltyrosine prior to release may be a means to improve the effectiveness of such wasps in augmentative release programs to control California red scale.     

Competition, resource partitioning and coexistence of an endoparasitoid Encarsia perniciosi and an ectoparasitoid Aphytis melinus of the California red scale

Abstract. 1. Laboratory experiments and field studies were conducted to explain the coexistence of an endoparasitoid, Encarsia perniciosi Tower, and an ectoparasitoid, Aphytis melinus DeBach, both of which were introduced into California to control the California red scale, Aonidiella aurantii (Mask.).
2. Encarsia parasitized all scale stages but it preferred first and second instar scales. This is in contrast to Aphytis melinus , in which previous studies have shown that it parasitizes second and third instar females and second instar males but prefers third instar female scales. Encarsia developed most rapidly when it parasitized an early second instar and slowest when it parasitized the mature female scale. However, on early second instar scales it was about 80% as fecund as a wasp that emerged from a mature female scale.
3. Second instar scales parasitized by Encarsia were accepted by Aphytis as readily as unparasitized scales.
4. Encarsia did not distinguish between unparasitized hosts and those previously parasitized by Aphytis.
5. Encarsia always outcompeted by Aphytis when both species parasitize the same host.
6. Encarsia prefers scale on stems whereas Aphytis prefers those on leaves and fruits. This, too, may be a result of interspecific competition with Aphytis.
7. The partitioning of the scale resource by the two species explains why they coexist in coastal southern California but it does not explain why Encarsia disappeared from citrus groves in the inland valleys coincident with the introduction of Aphytis melinus into southern California.     

Seasonal flight activity of two Homalodisca species (Homoptera: Cicadellidae) that spread Xylella fastidiosa in southern California.

Homalodisca coagulata (Say) and Homalodisca lacerta (Fowler) are vectors of a new bacterial disease of oleander in California known as oleander leaf scorch, induced by the bacterium Xylella fastidiosa. H. coagulata also has been implicated in the spread of the strain of X. fastidiosa that induces Pierce's disease of grapevines in California. We monitored the flight activity of H. coagulata and H. lacerta in oleander and citrus by using yellow sticky cards at three southern California locations where outbreaks of oleander leaf scorch have been documented, and where vector compliments are different. Areas sampled included a mesic coastal area (Irvine, CA) that supports predominantly H. coagulata and few H. lacerta, a dry inland location (Palm Desert, CA) that supports predominantly H. lacerta and few H. coagulata, and an intermediate area (Riverside, CA) supporting both Homalodisca species. From November 1996 to October 1999 peak catches of both Homalodisca species occurred during the midsummer at all locations. H. coagulata was trapped in greater numbers in citrus than in oleander at both the Riverside and the Irvine sites. Likewise, H. lacerta in Riverside was more associated with citrus than oleander, yet H. lacerta in Palm Desert was trapped in greater numbers in oleander than citrus.     

Disruption of parasitism of the California red scale (Homoptera: Diaspididae) by three ant species (Hymenoptera: Formicidae)

The relative influence of the southern fire ant, Solenopsis xyloni McCook, Argentine ant, Linepithema humile (Mayr), and native gray ant, Formica aerata (Francoeur), on parasitism of California red scale, Aonidiella aurantii Maskell, was studied in the laboratory for two parasitoids, Comperiella bifasciata Howard and Aphytis melinus DeBach. All three ant species reduced percentage parasitism by C. bifasciata and both percentage parasitism and host mutilation by A. melinus. Southern fire ant was the least disruptive and native gray ant the most disruptive. Southern fire ant removed 12% of scale from the lemons, presumably to feed on them, while the other ant species did not exhibit significant removal of scale compared to the controls. Percentage parasitism of California red scale exhibited by C. bifasciata was more than 2-fold the level exhibited by A. melinus. Percentage mutilation of California red scale, including probing and host feeding, was nearly 5-fold higher for A. melinus than C. bifasciata. Because A. melinus required a longer total host examination + oviposition period in the absence of ants than C. bifasciata and because oviposition occurs as the last act in a sequence of behaviors, disruption by the ants had a more significant negative effect on oviposition by A. melinus.     

Oppositional responses to host density in Aphytis holoxanthus (Hymenoptera: Aphefinidae), an efficient gregarious parasite

Abstract. 1. Ovipositional responses of an efficient gregarious ectoparasite ( Aphytis holoxanthus ) to increasing densities of its host (the Florida red scale, Chrysomphalus aonidum , Homoptera: Diaspididae) were studied in a laboratory system.
2. The female parasite may lay one to several eggs at each encounter with a host.
3. As host density increased, the number of eggs laid by the female parasite during the experimental period increased at a decreasing rate. This type of functional response resulted in less parasitism at higher host densities.
4. As host density increased, the number of eggs laid per encounter decreased. This behavioural response partially compensated for the decrease in parasitism.     

Behaviour of roach (Rutilus rutilus L.) altered by Ligula intestinalis (Cestoda: Pseudophyllidea): a field demonstration

1. We studied the influence of a cestode parasite, the tapeworm Ligula intestinalis (L.) on roach ( Rutilus rutilus L.) spatial occupancy in a French reservoir (Lake Pareloup, South-west of France).
2. Fish host age, habitat use and parasite occurrence and abundance were determined during a 1 year cycle using monthly gill-net catches. Multivariate analysis [generalized linear models (GLIM)], revealed significant relationships ( P  < 0.05) between roach age, its spatial occupancy and parasite occurrence and abundance.
3. Three-year-old roach were found to be heavily parasitized and their location toward the bank was significantly linked to parasite occurrence and abundance. Parasitized fish, considering both parasite occurrence and abundance, tended to occur close to the bank between July and December. On the contrary, between January and June no significant relationship was found.
4. These behavioural changes induced by the parasite may increase piscivorous bird encounter rate and predation efficiency on parasitized roach and therefore facilitate completion of the parasite's life cycle.     

Effects of pyriproxyfen on California red scale (Hemiptera: Diaspididae) development and reproduction

Two life stages of a laboratory colony of California red scale, Aonidiella aurantii (Maskell) (Hemiptera: Diaspididae), were exposed to 10 concentrations of pyriproxyfen to evaluate the effect of this insect growth regulator (IGR) on scale development and fecundity. First instars exposed to pyriproxyfen responded with mortality during the first and second molts. Second instars were more tolerant of pyriproxyfen than were first instars, indicating that growers should treat California red scale in the field before the first molt to achieve maximum efficacy. Male scales responded with an LC50 12-fold lower and an LC90 47-fold lower than was observed for female scales. Female scales that survived first instar exposure to pyriproxyfen experienced reduced fecundity with increasing pyriproxyfen concentration. Reduced fecundity was due to reduced survival of the females rather than sterility. Results from this study provide baseline California red scale susceptibility data for pyriproxyfen, and they suggest that 1 ppm pyriproxyfen can be used as a concentration that would discriminate between susceptible and resistant populations of scales. Field monitoring for incipient pyriproxyfen resistance in California red scale should be implemented in the San Joaquin Valley of California, and if resistance is detected, integrated resistance management strategies should be used to slow the progression of resistance.     

Volatile cues used by the parasitoid, Aphytis melinus, for host location: California red scale revisited

Aphytis melinus DeBach (Hymenoptera: Aphelinidae) is a biological control agent of diaspidid scale insects. The parasitoid has a narrow host range but its hosts are polyphagous. We determined the source of volatile cues the wasp uses to locate its few host species when those hosts occur on more than one host plant species. We addressed four questions in regard to the use of volatile cues in host location of California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae): (1) Does A. melinus use volatile cues to assist in host location? (2) Are these cues innately recognized or learned? (3) Are cues produced by female California red scale, or from other sources? (4) Are the cues specific to the host or host plant? These questions were tested through the use of a Y-tube olfactometer. Female A. melinus used volatile cues to orient toward both infested and uninfested host plant material. Wasps learned these cues by associating odors from the host plant with host presence. They had no innate preferences for scale insect or host plant volatile stimuli. Contrary to previous studies, we found no evidence of orientation toward the female-produced sex pheromone of California red scale, nor to volatile cues from the attacked host stage. Wasps given experience with scale insects growing on lemon fruit subsequently oriented toward lemon and orange fruit and leaves. The scale species with which the wasp was given experience did not affect this preference. Wasps given experience with California red scale growing on squash did not orient toward infested lemon fruit. The host ranges of the parasitoid and its hosts are used to explain the adaptive value for the evolution of learned rather than fixed responses to cues used in foraging behavior.     

Prevalence, intensity, and differential development of Pseudodelphis oligocotti (Nematoda: Dracunculoidea) in sympatric fish hosts of the northeastern Pacific coast

Counter to expectations of coevolved parasite-host relationships, parasites frequently infect hosts that never contribute to their reproduction, making the identification of a parasite's true host-specificity problematic. Pseudodelphis oligocotti (Nematoda: Dracunculoidea) infects several coastal Pacific fishes, but its course of development appears highly variable, suggesting that incidence does not reflect effective host range. To determine the host range of P. oligocotti and describe its relationship to various potential hosts, 24 fish species were examined from several British Columbia localities for prevalence, intensity, and extent and tissue location of parasite development. Pseudodelphis oligocotti infects 9 species of fishes from 5 orders, of which penpoint gunnel, Apodichthys flavidus, showed the highest prevalence and intensity, up to 80% and 19 (+/- 17.1 SD) worms per host, respectively. Although subadult and adult P. oligocotti occurred in all 9 fishes, larvigerous P. oligocotti only occurred in A. flavidus and rarely in the northern clingfish, Gobiesox maeandricus. Infective first-stage larvae were recovered from gill tissue of A. flavidus. Thus, at most only 2 of the 9 host species infected by P. oligocotti actually contribute to its transmission. The occurrence of P. oligocotti in diverse hosts may be accounted for by the parasite's indiscriminant mode of transmission via ingestion of free-living intermediate copepod hosts, where highly exposed or more suitable fishes (or both) are closely related by diet and microhabitat. This study demonstrates how parasite transmission and host ecology can greatly affect observed host range and ultimately its potential for expansion.     

Host-related fitness trade-offs in a presumed generalist parasitoid, Diaeretiella rapae (Hymenoptera: Aphidiidae)

Abstract.  1. Host ranges of parasitoid wasps are mediated by behavioural responses to hosts and their environment (infectivity), and development in hosts (virulence). Determinants of host range were measured in Diaeretiella rapae (Hymenoptera: Aphidiidae), which has been described as a generalist that attacks more than 60 species.
2. In northern Colorado, this wasp mainly attacks two hosts: cabbage aphid ( Brevicoryne brassicae ) and Russian wheat aphid ( Diuraphis noxia ). Here, laboratory experiments are described in which D. rapae originating from these two hosts were offered several hosts for oviposition. Both infectivity and virulence were measured.
3. Infectivity included host acceptance and handling time, while virulence was measured as productivity (number of progeny), survival of immatures within hosts, development time, and sex ratio.
4. Wasps had higher productivity and survival when attacking 'home' hosts than 'alternate' hosts, and trade-offs were found by quantitative genetic analyses to be genetically determined. Sex ratio and development times also showed trade-offs, but mainly related to the host environment in which females were reared.
5. In previous genetic studies in northern Colorado, populations were genetically subdivided on the scale of 1 km. The fitness differences described here could be strong enough to create populations adapted to different hosts, but it appears that gene flow is sufficient to prevent formation of separate lineages on the two hosts.
6. Rather than being a generalist with a broad host range, D. rapae is a serial specialist, attacking particular hosts according to availability in different seasons or in different geographical areas.     

Uncoupling physical and chemical cues: The independent roles of scale cover size and kairomone concentration on host selection byAphytis melinus DeBach (Hymenoptera: Aphelinidae)

The ectoparasitoidAphytis melinus initially selects its host, California red scale,Aonidiella aurantii (Maskell) (Homoptera: Diaspididae), using characteristics of the host’s cover before assessing the quality of the scale body beneath. Host suitability is known to increase with host size until the scale insect reaches maturity, after which it is no longer available for parasitism. The wasp uses a combination of scale cover size and a kairomone,O-caffeoyltyrosine, in the cover for initial assessment. Under natural conditions these two factors are frequently coupled. We quantified the relative importance of cover size and kairomone concentration independently by removing and selectively reapplying controlled doses of syntheticO-caffeoyltyrosine to scale covers of known size. In the absence of the kairomone, wasps did not discriminate among scale covers differing in size. Wasps showed a curvilinear response to kairomone dose for each scale cover age group. Wasps preferred low doses ofO-caffeoyltyrosine on young, small scale covers, and high doses on old, large scale covers. The ability of wasps to respond quantitatively to the kairomone may be used in the field to differentiate small second-instar from larger and more suitable third-instar scale insect larvae.     

Effect of selected insecticides on Homalodisca coagulata (Homoptera: Cicadellidae) and transmission of oleander leaf scorch in a greenhouse study.

Homalodisca coagulata (Say) is a recent introduction to California. It is known to spread a strain of the bacterium Xylella fastidiosa Wells, Raju, Hung, Weisberg, Mandelco-Paul & Brenner that induces oleander leaf scorch disease in oleander, Nerium oleander L. Oleander leaf scorch is lethal to oleander and threatens to decimate one of the most important landscape shrubs in California. Towards developing a management strategy for H. coagulata-spread oleander leaf scorch, we documented the affects of selected insecticides on H. coagulata mortality, feeding behavior, and disease transmission in a greenhouse study. Oleanders treated with fenpropathrin, fenpropathrin + acephate, and imidacloprid caused significant mortality to caged H. coagulata within 4 h of exposure. Within 24 h, these pesticides caused nearly 100% mortality 3 wk after treatment. In other experiments, acetamiprid and fenpropathrin treatments reduced time spent feeding and total time on plants. H. coagulata on fenpropathrin-, acetamiprid-, and imidacloprid-treated oleander died in less than 13 min on average. Oleander leaf scorch transmission by H. coagulata was blocked by applications of foliar-applied acetamiprid, and soil-applied imidacloprid and thiamethoxam.     

Infection of the western treehole mosquito,Aedes sierrensis (Diptera: Culicidae), with Lambornella clarki (Ciliophora: Tetrahymenidae)

Lambornella clarki was a common parasite of Aedes sierrensis immatures collected from treeholes in Mendocino County, California, in 1982–1983. The ciliate was not found in mosquitoes from treeholes with water having the most extreme values of electrical conductivity (<0.23 and >1.74 mmhos/cm) and pH (<6.5 and >7.7). Infection rates for individual monthly samples from L. clarki-positive treeholes ranged from 1 to 75%; 67% of all infections were observed in 4th-instar larvae. Infection with pathogens and parasites such as L. clarki, Ascogregarina clarki, Octomyomermis troglodytis, and unidentified bacteria and fungi, appeared responsible for high mortality rates (21–76%). Parasitism with L. clarki did not always result in death of the mosquito host; 7% of adults emerging from samples held in the laboratory were found to be infected. Ciliates were restricted to the host hemocoel except in older females where they invaded the ovaries, resulting in parasitic castration. This phenomenon may be associated with parasite dispersal.     

Absence of intraspecific interference during feeding by the predatory ladybirds Chilocorus spp. (Coleoptera: Coccinellidae)

Abstract. 1. The hypothesis was tested that intraspecific behavioural interference does not adversely affect the feeding behaviour of adults of three predatory coccinellid species, Chilocorus nigritus (F.), C.bipustulatus (L.) and C.infernalis Mulsant, at densities found under field conditions.
2. Feeding rates on mature oleander scale Aspidiotus nerii Bouché were evaluated by two methods at various predator densities. Proportion of the population dispersing was also measured for one of the species.
3. Feeding rate did not decrease and dispersal did not increase with increasing predator density. No significant behavioural interference that might have reduced predatory efficiency was observed, counter to assumptions on which published interference models are based.
4. Results here help to explain the relative importance of parasitoids and predators in the effective control of red scale Aonidiella aurantii (Mask.). The results also provide guidelines for release of these bio-control agents.     

Developmental strategy of the endoparasite Xenos vesparum (strepsiptera, Insecta): host invasion and elusion of its defense reactions

To successfully complete its endoparasitic development, the strepsipteran Xenos vesparum needs to elude the defense mechanisms of its host, the wasp Polistes dominulus. SEM and TEM observations after artificial infections allow us to outline the steps of this intimate host-parasite association. Triungulins, the mobile 1st instar larvae of this parasite, are able to "softly" overcome structural barriers of the larval wasp (cuticle and epidermis) without any traumatic reaction at the entry site, to reach the hemocoel where they settle. The parasite molts 48 h later to a 2nd instar larva, which moves away from the 1st instar exuvium, molts twice more without ecdysis (a feature unique to Strepsiptera) and pupates, if male, or develops into a neotenic female. Host encapsulation involves the abandoned 1st larval exuvium, but not the living parasite. In contrast to the usual process of encapsulation, it occurs only 48 h after host invasion or later, and without any melanization. In further experiments, first, we verified Xenos vesparum's ability to reinfect an already parasitized wasp larva. Second, 2nd instar larvae implanted in a new host did not evoke any response by hemocytes. Third, we tested the efficiency of host defense mechanisms by implanting nylon filaments in control larval wasps, excluding any effect due the dynamic behavior of a living parasite; within a few minutes, we observed the beginning of a typical melanotic encapsulation plus an initial melanization in the wound site. We conclude that the immune response of the wasp is manipulated by the parasite, which is able to delay and redirect encapsulation towards a pseudo-target, the exuvia of triungulins, and to elude hemocyte attack through an active suppression of the immune defense and/or a passive avoidance of encapsulation by peculiar surface chemical properties.