Is the host or the parasitoid in control?: effects of host age and temperature on pseudoparasitization by Microplitis rufiventris in Spodoptera littoralis

We compared the production of pseudoparasitization by Microplitis rufiventris females in most (third) and less (fourth) preferred instars of Spodoptera littoralis larvae at 20+/-1 and 27+/-1 degrees C. The parasitized hosts were classified into hosts producing parasitoids (type A hosts) and hosts producing no parasitoids, i.e., pseudoparasitized hosts (type B hosts). The latter were further classified into: (a) pseudoparasitized hosts with "well" arrested development (type B1 hosts); (b) pseudoparasitized hosts with partially arrested development (type B2 hosts); and (c) pseudoparasitized hosts that successfully pupated to apparently normal host pupae (type B3 hosts). The present series of experiments showed that parasitization by M. rufiventris was clearly affected by host instar, age within an instar and rearing temperature. Production of type B hosts was less when third instar S. littoralis larvae were exposed to the wasp females than when the host larvae were in fourth instar. The production of type A hosts was much greater when early or mid ages of an instar was stung by the wasp females comparing with stung late age of the same instar. Production of type B hosts may be due to one or overall of the following: (a) dosage dilution of M. rufiventris female's factors in the different age classes of the instar; (b) endocrine system (physiological state) at parasitization time, i.e., early vs late age of the instar; (c) growth rate of host larvae. The lowest production of type B hosts was at highest growth rate; and (d) temperature, larger proportions of type B hosts were produced at 27+/-1 than at 20+/-1degrees C. The three types host development (B1, B2 and B3) are possibly representing three levels of host resistance (host control) resulting in partial or complete failure of parasitoid control. Type A hosts represent complete success of parasitoid control. The results suggest that the impact of parasitoid factor(s) on developmental arrest is affected by host age at the time of parasitism and/or by temperature.     

Inhibition of testicular growth and development in Manduca sexta larvae parasitized by the braconid wasp Cotesia congregata

Tobacco hornworm larvae parasitized by the gregarious larval endoparasitoid Cotesia congregata exhibited an inhibition in testicular growth and development, the extent of which was determined by the age and developmental stage of the host at the time of parasitization. The degree of parasitic castration, as assessed by measurements of testicular volume, was correlated with the stadium in which parasitization occurred. A mathematical formula requiring the measurement of testicular length, width and depth was used to calculate testicular volume. The use of the depth parameter revealed a negative correlation between host weight and testicular volume in parasitized larvae. Testicular volumes of fifth instar hosts, which had been parasitized in the first stadium, were significantly smaller than those originally parasitized as fourth or fifth instar larvae and were not correlated with parasitoid load. Effects of natural parasitism were not duplicated by injections of C. congregata polydnavirus and venom, topical treatment with the juvenile hormone analog methoprene, or starvation of nonparasitized larvae. Larvae receiving virus plus venom or methoprene grew larger due to delayed wandering and had larger testes than controls. Deleterious effects on host testes may be due to the effects of nutrient competition between the developing parasitoid progeny and the gonads, combined with the juvenilizing effects believed to be caused by the polydnavirus.     

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

THE PARASITOID,Cotesia flavipes (CAMERON) (HYMENOPTERA: BRACONIDAE), INFLUENCES FOOD CONSUMPTION AND UTILIZATION BY LARVAL Diatraea saccharalis (F.) (LEPIDOPTERA: CRAMBIDAE)

Parasitoids exploit host insects for food and other resources; they alter host development and physiology to optimize conditions to favor parasitoid development. Parasitoids influence their hosts by injecting eggs, along with a variety of substances, including venoms, polydnaviruses, ovarian fluids, and other maternal factors, into hosts. These factors induce profound changes in hosts, such as behavior, metabolism, endocrine events, and immune defense. Because endoparasitoids develop and consume tissues from within their hosts, it is reasonable to suggest that internal parasitization would also influence host food consumption and metabolism. We report on the effects of parasitism by Cotesia flavipes on the food consumption and utilization of its host, Diatraea saccharalis. Cotesia flavipes reduces the host food consumption, but parasitized larvae considered a unit with their parasitoid's attained the same final weight as the nonparasitized larvae. Nutritional indices, midgut activities of carbohydrases, and trypsin of parasitized and nonparasitized D. saccharalis were assessed. Parasitized larvae had reduced relative food consumption, metabolic and growth rates, coupled with higher efficiency for conversion of the digested, but not ingested, food into body mass. Parasitism also affected food flux through the gut and protein contents in the midgut of parasitized larvae. The activity of α‐amylase and trehalase in parasitized host was enhanced in the first day after parasitism relative to control larvae. Saccharase activity remained unchanged during larval development. Trypsin activity was reduced from the fifth to ninth day after parasitism. We argue on the mechanisms involved in host food processing after parasitism.     

Stage dependent influences of polydnaviruses and the parasitoid larva on host ecdysteroids

In the solitary egg-larval parasitoid Chelonus inanitus (Braconidae) both polydnavirus and the parasitoid larva manipulate host development. Parasitization leads to a premature drop in juvenile hormone titre and a precocious onset of metamorphosis in the 5th larval instar. The C. inanitus bracovirus (CiBV) alone causes a reduction in host ecdysteroid titres at the pupal cell formation stage and prevents pupation. Here we report three new findings. (1) We show that parasitization causes a reduction in haemolymph ecdysteroid titre immediately after the moult to the 5th instar; similarly low values were seen in nonparasitized larvae after the moult to the 6th instar. These data along with parasitoid removal experiments indicate that the low ecdysteroid titre after the moult is a very early sign of the upcoming metamorphosis. (2) In vitro experiments with prothoracic glands and brain extracts showed that CiBV affects both prothoracic glands and prothoracicotropic hormone after the stage of pupal cell formation. (3) In the haemolymph of parasitized larvae the ecdysteroid titre increased in the late cell formation stage, i.e. immediately before egression of the parasitoid. In vitro experiments showed that late 2nd instar parasitoids release ecdysteroids and are thus very likely responsible for the rise in host ecdysteroids.     

Varying degree of physiological integration among host instars and their endoparasitoid affects stress‐induced mortality

In natural populations of insect herbivores, genetic differentiation is likely to occur due to variation in host plant utilization and selection by the local community of organisms with which they interact. In parasitoids, engaging in intimate associations with their host during immature development, local variation may exist in host quality for parasitoid development. We compared the development of a gregarious endoparasitoid, Cotesia glomerata L. (Hymenoptera: Braconidae), collected in The Netherlands, in three strains and three caterpillar instars (L1–L3) of its main host, Pieris brassicae L. (Lepidoptera: Pieridae). Hosts had been collected in The Netherlands and France, and were reared in the laboratory for one generation. We also used an established Dutch laboratory strain that had not been exposed to parasitoids for at least 24 generations. Parasitoid survival to adulthood was inversely correlated with host instar at parasitism. Adult parasitoid body mass was largest when hosts were parasitized as L1 and smallest when hosts were parasitized as L3, whereas egg‐to‐adult development time was quickest on L3 hosts and slowest on L1 hosts. Higher survival and faster development of C. glomerata on French L2 hosts also showed that there is variation in host‐instar‐related suitability. Many L2 and most L3 caterpillars that were parasitized exhibited signs of pathogen infection and perished within a few days of parasitism, whereas this never happened when hosts were parasitized as L1 or in non‐parasitized control caterpillars. Our results reveal that, irrespective of the host strain, L1 hosts are optimally synchronized with C. glomerata development. By contrast, the high precocious mortality of L3 larvae may be due to stress‐induced regulation by the parasitoid in order to ‘force’ its developmental program into synchrony with the developing parasitoid larvae. Our results underscore a potentially important role played by pathogens in mediating herbivore–parasitoid interactions that are host‐instar‐dependent in their expression.     

The endoparasitoid Cotesia kariyai (Ck) regulates the growth and metabolic efficiency of Pseudaletia separata larvae by venom and Ck polydnavirus

It was previously demonstrated that parasitization by Cotesia kariyai caused a decrease in weight gain and food consumption in host larvae, resulting in a lower final weight for parasitized hosts. It is predicted that C. kariyai regulates the physiological condition of the host to obtain maximum food under restricted nutritional conditions. Approximate digestibility (AD) was higher following parasitization but the efficiency of conversion of digested food (ECD) of the parasitized hosts was lower. This suggests that resources available to the parasitoid larvae are enhanced in the parasitized hosts. We evaluated the physiological changes caused by injection of calyx fluid (polydnavirus) plus venom (C+V) in nonparasitized hosts. Injection of C+V into the nonparasitized hosts duplicated the effects of parasitism, namely it increased the AD and decreased the ECD. Furthermore, C+V injections elevated trehalose concentrations in nonparasitized host 7 to 10 d after injection (2nd stadium of the parasitoid larva). Protein content also increased on days 9 and 10 after C+V injection. These results suggest that the nutrients that parasitoid larvae require for their growth increase in the hemolymph of the host during the 2nd stadium of the parasitoid larva.     

Development of Meteorus pulchricornis and regulation of its noctuid host, Pseudaletia separata

The solitary endoparasitoid Meteorus pulchricornis can parasitize many lepidopteran host species successfully. In the case of parasitization of Pseudaletia separata, developmental duration of M. pulchricornis was 8-9 days from egg to larval emergence and 6 days from prepupa to adult emergence. Successful parasitism by M. pulchricornis decreased with host age. Following parasitization of day-0 4th host instar, the parasitoid embryo, whilst still enclosed in serosal cell membrane, hatched out of the egg chorion 2 days after oviposition. Subsequently, the 1st instar parasitoid emerged from the surrounding serosal cell membrane. Serosal cells dissociated and developed as teratocytes 3.5 days after oviposition. One embryo of M. pulchricornis gave rise to approximately 1200 teratocytes, a number that remained constant until 6 days after parasitization, but decreased drastically to 200 at 7 days post-oviposition. The teratocytes of M. pulchricornis were round- or oval-shaped and grew from 65 microm at 4 days to 200 microm in the long axis at 6 days post-parasitization. At 4 days post-parasitization, many cells or cell clusters with lipid particles were observed in the hemocoels of parasitized hosts. In addition, paraffin sections of parasitized hosts revealed that many teratocytes were attached to the host's fat body and contributed to disrupting the fat body tissue. Further, examination of the total hemocyte count (THC) during parasitization revealed that THC was maintained at low levels. Surprisingly, a temporal decrease followed by restoration of THC was observed in hosts injected with virus-like particles of M. pulchricornis (MpVLPs) plus venom, which contrasts with the constant THC suppression seen in parasitized hosts. This indicates that MpVLP function is temporal and is involved in regulation of the host during early parasitism. Therefore, teratocytes, a host regulation factor in late parasitism, could be involved in keeping THC at a low level.     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.     

Transient host paralysis as a means of reducing self-superparasitism in koinobiont endoparasitoids

The term ‘idiobiont’ refers to those parasitoid species that permanently paralyse their hosts during parasitism, causing the cessation of host growth and development. This is in contrast to koinobiont parasitoids, which allow their hosts to continue developing after being parasitized. While no koinobiont species induce permanent paralysis in their hosts, a minority of koinobionts induce a temporary paralysis that does not interfere with overall host growth and development. We characterized transient paralysis induction in two koinobiont aphid parasitoids in the genus Binodoxys (Hymenoptera: Aphidiinae). Both Binodoxys species induced transient paralysis in Aphis glycines, with paralysis time ranging between 4.5 and 8 min (depending upon parasitoid species and host instar). In a separate experiment, B. communis was capable of inducing transient paralysis in nine aphid species. We addressed two hypotheses potentially explaining the adaptive value of temporary host paralysis in experiments using A. nerii, which is readily accepted but engages in strong defensive behaviour. The first hypothesis is that paralysis increases oviposition success by interfering with host defences and the second is that it aids in the avoidance of self-superparasitism. Paralysed aphids were more likely to be rejected by B. communis than were aphids that had never been stung or that had recovered from paralysis. This result supports the avoidance-of-self-superparasitism hypothesis and is inconsistent with the hypothesis that transient paralysis increases oviposition success of B. communis.     

Interspecific discrimination and larval competition amongMicroplitis croceipes,Microplitis demolitor,Cotesia kazak (Hym.: Braconidae), andHyposoter didymator (Hym.: Ichneumonidae), parasitoids ofHeliothis virescens (Lep.: Noctuidae)

Interspecific host discrimination by adults, and larval competition among the endoparasitoidsMicroplitis croceipes (Cresson),Microplitis demolitor Wilkinson,Cotesia kazak (Telenga) andHyposoter didymator (Thunberg) were investigated usingHeliothis virescens (F.) as the host. In ovipositional choice tests, the mean number of encounters and ovipositions for unparasitized hosts was not significantly different from the mean number of encounters and ovipositions for parasitized hosts for each treatment combination (P>0.05). Thus, none of the parasitoid species discriminated between host larvae recently parasitized once by a female of another species und unparasitized hosts. However, in all but two cases, females did discriminate between unparasitized hosts and hosts in which an early first instar of the first-attacking species was developing.Cotesia kazak andH. didymator did not discriminate between unparasitized hosts and hosts parasitized by an early first instar ofM. demolitor. Larval competition among these parasitoid species was studied for three time intervals between the first and second species parasitization: 1) second species attack immediately (5–15 sec) after the first; 2) second species attack 24 h after the first; and 3) second species attack 48 h after the first. Time until egg eclosion was shortest forM. demolitor, thenC. kazak, thenM. croceipes, and longest forH. didymator. When the second parasitoid species attacked a host immediately after the first species, the species in which egg eclosion occurred first was the victor more frequently, except whenM. demolitor competed withC. kazak andH. didymator. With a 24 h delay between the first and second species to attack, the older first instar from the first parasitization usually outcompeted the younger first instar from the second attack. A first instar from the second species to attack generally outcompeted the second instar of the first species when the second parasitization had been delayed 48 h. Competiors were eliminated mainly by physical attack, butC. kazak andM. croceipes apparently also killedH. didymator eggs by physiological processes.     

Survival of the parasitoidEncarsia formosa after treatment of parasitized greenhouse whitefly larvae with fungal spores ofAschersonia aleyrodis

The interaction between the entomopathogenic fungusAschersonia aleyrodis and the parasitoidEncarsia formosa on greenhouse whitefly as a host organism was studied, in particular, the survival of the parasitoid after treatment of parasitized hosts with fungal spores. The mean number of parasitized black pupae per parasitoid produced at 25°C was significantly reduced after spore treatment in the first three days following parasitization. Spore treatment four, seven or ten days after parasitization resulted in a mean number of parasitized pupae not significantly different from the number of black pupae in the control. The rather sudden change from low to high survival of parasitized hosts when treated with spores four days after parasitization in spite of high numbers of infected unparasitized larvae, coincided with the hatching of the parasitoid larva from the egg inside the host. Possible reasons for this decrease in susceptibility to infection after parasitoid egg hatch, such as induced changes in host cuticle or haemolymph, are discussed. Parasitoids emerged from treated hosts did not show differences in reproduction compared with parasitoids emerging from untreated hosts. Both natural enemeies of whitefly are compatible to a great extent.     

Parasitism of Pieris rapae (Lepidoptera: Pieridae) by a pupal endoparasitoid, Pteromalus puparum (Hymenoptera: Pteromalidae): effects of parasitization and venom on host hemocytes

In contrast to the situation with egg-larval and larval endoparasitic wasps, little is known about the effects of pupal endoparasitoids and their secretions on the hemocytes of their insect hosts. This study focuses on the pupal endoparasitoid, Pteromalus puparum, and its host, the small white butterfly, Pieris rapae. Parasitism by P. puparum, resulted in a significant increase in the total number of host hemocytes up to day five after parasitization. From day one to day four after parasitization, the percentage of plasmatocytes significantly decreased, and the proportion of granular cells increased. Moreover, from 12 h to day three after parasitization, hemocyte mortality in parasitized pupae was noticeably higher. When P. rapae pupae were parasitized by adult females of P. puparum irradiated by gamma-ray (pseudoparasitization), it was clear that the treated wasps could induce similar hemocyte changes. However, such phenomena did not occur in punctured host pupae (mimic-parasitization). After treatment with P. puparum venom, both the percentages of spreading plasmatocytes and encapsulated Sephadex G-25 beads were lessened significantly in vitro. Electron microscopy analysis and visualization of hemocyte F-actin with phalloidin-FITC showed that hemocytes treated with venom had a rounded configuration and neither spread nor extended pseudopods, while there was no marked alteration of hemocyte cytoskeletons after venom treatment. The results suggested that venom of P. puparum could actively suppress the hemocyte immune response of its host, but not by destroying the host hemocyte cytoskeleton.     

Development of the braconid wasp Cotesia flavipes in two Crambids, Diatraea saccharalis and Eoreuma loftini: Evidence of host developmental disruption

Cotesia flavipes is an important gregarious larval endoparasitoid of several crambid stem borers, including Diatraea saccharalis. The suitability of two crambid species, Eoreuma loftini and D. saccharalis, pests of sugarcane and rice in Texas, for C. flavipes development was tested. The effect of parasitization by C. flavipes on encapsulation response was assessed in vivo in both D. saccharalis and E. loftini. The results indicated that the parasitoid developed and emerged successfully in D. saccharalis larvae. Although E. loftini larvae were readily parasitized by C. flavipes parasitoids, no wasp larvae hatched from the eggs in this host because eggs were encapsulated by the host's hemocytes. The developmental fate of the E. loftini larvae with encapsulated parasitoids was variable. Most died as abnormal fifth instars or as post-wandering prepupae, while a few developed normally to the pupal stage. In vivo experiments, there was a significant reduction in the percent of beads encapsulated in parasitized larvae in both hosts. However, the percent of beads showing melanization decreased significantly in parasitized D. saccharalis larvae but did not differ significantly in parasitized or unparasitized E. loftini larvae. Our results showed that D. saccharalis is a suitable host for C. flavipes whereas E. loftini is an unsuitable host. This study indicated that lepidopteran stem borers that are taxonomically, behaviorally, and ecologically very similar can differ in their ability to encapsulate a parasitoid species.     

Performance of Microplitis tuberculifer (Hymenoptera: Braconidae) parasitizing Mythimna separata (Lepidoptera: Noctuidae) in different larval instars

The solitary parasitoid Microplitis tuberculifer (Wesmael) is an important biological control agent of various lepidopteran pests in Asia. We examined the preference of M. tuberculifer for different instars of its common host, Mythimna separata (Walker), host instar effects on parasitoid development, and the consequences of parasitism in different stages for growth and consumption of host larvae. The wasp successfully parasitized the first four larval instars of M. separata, but not the fifth, which appeared to be behaviorally resistant. First and second instars were parasitized at higher rates compared to thirds and fourths in no-choice situations, ostensibly due to longer handling times for the latter, but second instars were most preferred in a choice test that presented all stages simultaneously. Although later instar hosts yielded heavier cocoons, the fastest parasitoid development was obtained in second instars. Lower sex ratios were obtained from first instars as females appeared to lay a smaller proportion of fertilized eggs in small hosts. Both weight gain and food consumption of parasitized larvae were reduced significantly within 24 h of parasitism, regardless of the stage parasitized, and final body weights were less than 10% those of unparasitized larvae. Thus, M. tuberculifer has good potential as a biological control agent of M. separata, successfully parasitizing the first four larval instars and dramatically reducing plant consumption by the host in all cases.     

Developmental rates of two congeneric parasitoids,Encarsia formosa and E. pergandiella (Hymenoptera: Aphelinidae), utilizing different egg provisioning strategies

Endoparasitic Hymenoptera vary in the extent to which they provision their eggs and thus in the degree to which they appear to rely on their hosts for resources during embryonic development. In this study, developmental rates were examined in two congeneric parasitoid species, Encarsia formosa and E. pergandiella, that provision their eggs to different degrees. E. formosa eggs are much larger than E. pergandiella eggs. E. formosa eggs hatch significantly earlier than the eggs of E. pergandiella when deposited in 1st or 4th instar nymphs of a common whitefly host, Bemisia tabaci. Both species hatch earlier in 4th instar nymphs, but the delay in hatching in hosts parasitized as 1st instars is much greater in E. pergandiella. While E. formosa develops more rapidly to the 1st larval instar, E. pergandiella emerge as adults significantly earlier, though smaller, than E. formosa adults regardless of the host instar parasitized. These findings show that the extent of provisioning in the eggs of these wasps does not strictly determine their order of progression through different stages of development.     

Effects of the endoparasitoid Cotesia chilonis (Hymenoptera: Braconidae) parasitism,venom, and calyx fluid on cellular and humoral immunity of its host Chilo suppressalis (Lepidoptera: Crambidae) larvae

The larval endoparasitoid Cotesia chilonis injects venom and bracoviruses into its host Chilo suppressalis during oviposition. Here we study the effects of the polydnavirus (PDV)-carrying endoparasitoid C. chilonis (Hymenoptera: Braconidae) parasitism, venom and calyx fluid on host cellular and humoral immunity, specifically hemocyte composition, cellular spreading, encapsulation and melanization. Total hemocyte counts (THCs) were higher in parasitized larvae than in unparasitized larvae in the late stages following parasitization. While both plasmatocyte and granulocyte fractions and hemocyte mortality did not differ between parasitized and unparasitized hosts, in vitro spreading behavior of hemocytes was inhibited significantly by parasitism throughout the course of parasitoid development. C. chilonis parasitism suppressed the encapsulation response and melanization in the early stages. Venom alone did not alter cellular immune responses, including effects on THCs, mortality, hemocyte composition, cell spreading and encapsulation, but venom did inhibit humoral immunity by reducing melanization within 6 h after injection. In contrast to venom, calyx fluid had a significant effect on cell spreading, encapsulation and melanization from 6 h after injection. Dose–response injection studies indicated the effects of venom and calyx fluid synergized, showing a stronger and more persistent reduction in immune system responses than the effect of either injected alone.     

Assessing augmentative releases of parasitoids using the «Recruitment method», with reference toEdovum puttleri,a parasitoid of the colorado potato beetle [Coleoptera: chrysomelidae]

Release of approximately 17,700 experienced adult femaleEdovum puttleri Grissell against 1^st generation Colorado potato beetle eggs in 1987 in a 0.4 ha potato field in S. Deerfield, Massachusetts resulted in only 3.6% parasitism as assessed by direct measurement of host and parasitoid recruitment. Levels of non-viability indicated an additional 2.8% of hosts killed by parasitoid hostfeeding, for an overall impact of 6.4%. Release in 1988 of 126,300 parasitoid against 1^st generation hosts in a 0.4 ha potato (Solanum tuberosum L.) field at the same site produced only slightly higher levels of parasitism (10.6%) and host feeding (2.0%). Release in 1987 of 32,800 wasps against 2^nd generation eggs resulted in only 0.7% parasitism and 0.3% host feeding due to the toxicity of fenvalerate (Pydrin^R) residues from a single application applied for control of the potato leafhopper,Empoasca fabae (Harris). Release in 1988 of 47,400 wasps against the 2^nd host generation in the absence of any pesticide applications resulted in 34.4% parasitism and 16.1% host feeding, for a total impact of 50.5%. Difference in parasitization levels between host generations supports the idea thatE. puttleri adults require an in-field carbohydrate source such as aphid honeydew to reproduce. In Massachusetts, aphid populations in potato typically do not develop until the end of the 1^st larval generation. The recruitment method ofVan Driesche & Bellows (1988) proved to be a satisfactory approach for determining results of augmentative parasitoid releases.      

Could hosts considered as low quality for egg-laying be considered as high quality for host-feeding?

When parasitoid females encounter a host, they can either lay an egg and thus invest in current reproduction or feed on the host and thus invest in future reproduction. However, hosts could have different values according to their parasitized status. Whereas already parasitized hosts represent poor quality for egg-laying, they could have a high nutritive value for feeding. Moreover, the optimal strategy adopted generally depends on the females’ physiological state. In this study, the impact of the females’ physiological state on their reproductive strategies was investigated in the solitary parasitoid Anisopteromalus calandrae. We analysed how their age and diet influenced (i) the use of hosts (feeding vs. oviposition) and (ii) host selection (previously parasitized vs. unparasitized). Our results show that both age and diet influence the reproductive strategy of A. calandrae females: old females fed with the poorer diet laid fewer eggs and made more host-feeding than others. Females also showed a preference for already parasitized hosts for feeding. This strategy cannot be explained by the nutritive value of haemolymph, as parasitized hosts carry fewer lipids. However, as parasitized hosts are also paralyzed, it could be less costly to feed on them than on unparasitized hosts.     

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.