Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

Host discrimination modulates brood guarding behaviour and the adaptive superparasitism in the parasitoid wasp Trissolcus semistriatus (Hymenoptera: Scelionidae)

Because hosts utilized by parasitoids are vulnerable to further oviposition by conspecifics, host guarding benefits female wasps. The present study aims to test whether female adults regulate brood guarding behaviour by host discrimination in a solitary parasitoid Trissolcus semistriatus by presenting an intact or parasitized host egg mass to a female adult. Virgin females without oviposition experience have host discrimination ability, which enables them to adjust the number of eggs laid in the hosts. Mating experience increases superparasitism by female adults, whereas mated females achieve a higher discrimination ability as a result of oviposition experience and show a lower superparasitism rate. As expected, females exhibit brood guard after parasitizing an intact host egg mass, whereas those females visiting a previously parasitized host egg mass, do not. Because the survival of eggs in superparasitized hosts is relatively low, regulating brood guarding behaviour by host discrimination is adaptive for female wasps.     

Endophyte-mediated tritrophic interactions between a grass-feeding caterpillar and two parasitoid species with different life histories

Plant secondary chemicals can alter herbivore suitability for parasitoids by weakening or stunting the host, delaying its development, or when larval parasitoids encounter ingested phytotoxins in the body of their host. Experiments with different parasitoids that exploit the same host species feeding on the same plant may provide insight about how parasitoid life history affects the strength of such interactions. The encyrtid wasp Copidosoma bakeri, a slow-developing polyembryonic egg-larval parasitoid, and the tachinid fly Linnaemya comta, a fast-developing solitary species, both parasitize Agrotis ipsilon, a generalist noctuid. We tested the hypothesis that of the two parasitoid species, the encyrtid, because of its more prolonged developmental association with the host, would suffer greater fitness costs when A. ipsilon feeds on perennial ryegrass containing an alkaloid-producing fungal endophyte. Indeed, fewer parasitized cutworms yielded C. bakeri broods, and those host mummies were smaller, formed more slowly, and contained fewer adults when the hosts fed on endophytic as opposed to endophyte-free grass. In contrast, L. comta fitness parameters were similar regardless of the type of grass upon which their host fed. Our results highlight that the outcome of endophyte-mediated tritrophic interactions may differ for different parasitoid species. Implications for integrating the use of endophytic grasses and biological control are discussed.     

RETRACTED ARTICLE: Foraging of Host-Habitat and Superparasitism in <Emphasis Type="Italic">Cotesia glomerata</Emphasis>: A Gregarious Parasitoid of <Emphasis Type="Italic">Pieris brassicae</Emphasis>

The study investigates differences in the oviposition pattern of a braconid parasitoid, Cotesia glomerata (Linn.) in Pieris brassicae (Linn.) in relation to their use of different cruciferous food plants. The response of P. brassicae to superparasitism and consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour. Female parasitoid located various crucifers and searched for host more frequently almost on all the host plants tested i.e. cabbage, cauliflower, Chinese cabbage, broccoli and radish. According to the estimated relative number of female locating hosts, cabbage was the most attractive plant for C. glomerata and total number of eggs laid in host larvae feeding on it was higher than in larvae feeding on other plants. Laboratory experiments demonstrated that superparasitism reduced survivorship of P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae. Sex ratio and the body weight of emergent wasps correlated negatively with brood size. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater female dry mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitized hosts are rare in a habitat.     

Determination of the optimal parasitoid‐to‐host ratio for efficient mass‐rearing of the parasitoid,Sclerodermus pupariae (Hymenoptera: Bethylidae)

Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae) is used as a potential biocontrol agent for several buprestid and cerambycid larvae. This study aimed to enhance the efficiency of mass‐rearing of this parasitoid by investigating the fitness gain of this bethylid wasp, including the proportion of successful parasitism and development, brood size, sex ratio, proportion of winged female offspring, body size and longevity of female offspring, under eight different maternal parasitoid density treatments using Thyestilla gebleri Faldermann as host in the laboratory. The results indicated that the foundress densities did not affect the parasitism or emergence rate of this parasitoid. Brood size of the parasitoids increased significantly when the number of maternal wasps ranged from one to four. However, further increases in foundress number did not affect the parasitoid brood size. The sex ratios of S. pupariae were always female‐biased. The proportions of male in the progeny colonies were <10% throughout all experimental treatments. The percentage of winged female progeny was not significantly influenced by the density of adult maternal parasitoids. Body sizes of parasitoids significantly declined with increasing maternal parasitoid densities. Although the parasitoid body size reduced when maternal wasp number was higher, it could be compromised by the relatively higher number of female offspring produced. Further, more than 70% of the parasitoids remained alive when they were stored at 12°C for four months throughout the experiments. These findings suggest that exposure of four female wasps to a single host larva would result in the highest fitness of S. pupariae. Our findings might provide a new approach to enhance the efficiency of mass‐rearing of this bethylid wasp.     

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.     

Influence of nutrient deficiency caused by host developmental arrest on the growth and development of a koinobiont parasitoid

Koinobiont parasitoids utilize nutrients obtained from hosts that contine to feed and grow after parasitization. However, if the ecdysis of early host instars is prevented, parasitized larvae will fail to grow large enough to support the development of the parasitoid brood and both organisms will perish. When L5 instar larvae (the penultimate stage) of Pseudaletia separata were parasitized by Cotesia kariyai and injected with Euplectrus separatae venom (5PV), the development of these hosts was arrested before molting to the next stage and the caterpillars thus failed to gain weight. These hosts remained at approximately 300 mg until parasitoid emergence. In contrast, hosts parasitized as L5 but without the injection of venom (5P) exhibited an increase in weight after molting to the next stage and ultimately grew to approximately 700 mg. The inhibition of ecdysis reduced the amount of food resource (e.g. fat body) for the parasitoid larvae. On the other hand, when final (= L6) host instars were parasitized and injected with E. separatae venom (6PV), the maximum weight attained by these larvae was about 710 mg, although weight gain was depressed compared to hosts parasitized without the injection of E. separatae venom (6P). The adult weight of C. kariyai that emerged from 5PV hosts was less than conspecifics that emerged from 5P, 6P, and 6PV respectively, although the egg-pupal period of the parasitoid from 5PV hosts was extended. The offspring sex ratio (percentage males) of adult wasps did not vary significantly with treatment. Female parasitoids that eclosed from 5PV hosts laid almost the same number of eggs in day 0-6th host instars as those emerging from 5P, 6P, 6PV hosts. Their egg-pupal period was extended and the cocoon cluster mass and the parasitoid body mass on subsequent generations was lighter than those reared from 5P, 6P, 6PV hosts. The sex ratio of F2 C. kariyai wasps that eclosed from 5PV increased more than in wasps that eclosed from the other host treatments (5P, 6P, 6PV). Our results reveal that a reduction in host quality and offspring fitness in the first generation negatively impacted female fitness in the second generation. An early arrestment of host growth, mediated by the addition of E. separatae venom, has severe implications on parasitoid fitness by reducing host quality, especially in smaller hosts.     

Brood Guarding by an Adult Parasitoid Reduces Cannibalism of Parasitoid-Attacked Conspecifics by a Caterpillar Host

Adult female bethylid parasitoids (Hymenoptera: Bethylidae) commonly guard their offspring until they develop into later immature stages to protect them from competing parasitoids and predators. Another possible mortality factor is from caterpillar larvae that cannibalize parasitized conspecific larvae. This study determined the effect of brood guarding on the occurrence of host cannibalism by using a brood guarding bethylid parasitoid Goniozus legneri and a non-guarding braconid parasitoid Habrobracon gelechiae (Hymenoptera: Braconidae), both are gregarious ectoprasitoids that attack the obliquebanded leafroller Choristoneura rosaceana (Lepidoptera: Tortricidae). When a C. rosaceana larva parasitized by G. legneri was presented to a live conspecific larva, the frequency of cannibalism on the parasitized conspecific was lower in the presence than in the absence of a guarding female G. legneri. In contrast, when a C. rosaceana larva parasitized by H. gelechiae was presented to a live conspecific larva, the presence or absence of H. gelechiae did not affect cannibalism frequency. Cannibals did not gain a fitness advantage in terms of larval survival, developmental time, or body mass of developed pupae; therefore, cannibalism by the host larva may be a group defense against increased parasitism in subsequent population generations. We suggest that brood guarding by parasitoids can act as a broad defense strategy to protect offspring from the cannibalism of parasitized conspecifics by the host larvae.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

Trade-offs between developmental parameters of two endoparasitoids developing in different instars of the same host species

Trade-offs amongst life history traits is a major theme in evolutionary biology. Parasitoid wasps are important biological control agents and make excellent organisms to examine trade-offs in fitness related traits such as size, development rate and survival. Here, we examined trait-related trade-offs in 2 solitary endoparasitoids developing in different stages (or instars) of the same caterpillar host, the cabbage moth Mamestra brassicae. Microplitis mediator is a small specialist parasitoid that attacks first (L1) to third (L3) instars of M. brassicae; Meteorus pulchricornis is a larger highly generalized parasitoid that attacks L1–L4 instars of the same host species. When developing in early host instars (e.g. L1–L2), both parasitoids differently traded-off size against development time. In M. mediator, adult body mass was smaller in wasps developing in L1 than in L2 and L3 hosts, whereas development time was unaffected by instar. By contrast, adult body mass in M. pulchricornis was smaller and development time longer when developing in L1 and L2 than in L3 and L4 instars. Periodic starvation of M. brassicae caterpillars parasitized by M. pulchricornis further reduced adult mass and extended development time of wasps in L2 (but not L4) hosts. Maximum egg load in M. pulchricornis (but not M. mediator) was correlated with adult female body size. Our results imply that rapid development time is more important than body size for fitness in both species, although in M. pulchricornis both development time and adult size are traded off in determining the optimal phenotype. Developing a better understanding of association-specific patterns of development in parasitoids can assist in the optimization of mass rearing of these insects for biological control.     

The effect of host age ofLymantria dispar larvae [Lep.: Lymantriidae] on the development ofGlyptapanteles liparidis [Hym.: Braconidae]

The endoparasitic development ofG. liparidis was examined in 3 different host stages of gypsy moth larvae. Hatching ofG. liparidis-larvae occurred 3 to 5 days after oviposition in hosts parasitized during their premoulting period, and after 5 to 7 days in those parasitized in the 3^rd midinstar state. The parasites generally moulted to the 2^nd larval instar between the 11^th and 13^th day in the first group, and between the 13^th and 15^th day in the latter, when they had reached a volume of 0.04–0.05 mm³. The positive correlation between host ecdysis and the ecdysis of 1^st stadium larvae to L₂ suggested that host moulting influenced the development of the parasitoid larvae. Emergence from the host larvae occurred at 20°C after 27 days on average, and coincided with the parasites moulting to the 3^rd instar. Five to 7 days after spinning their cocoons near the developmentally arrested host larva, the male, and 1 to 2 days later the female wasps eclosed. Due to the variation in the number of parasites per host, no difference was observed between the hosts parasitized at various stages; however, a tendency for later parasitized hosts to contain more parasite larvae was evident. The nutritional conditions of the moth parental generation influenced both host and parasite development. On the other hand no influence of host age was observed on emergence dates of larvae and wasps.      

Parasitism-induced Effects on Host Growth and Metabolic Efficiency in Tobacco Hornworm Larvae Parasitized by Cotesia congregata

Parasitism by the braconid wasp Cotesia congregata affects the growth of Manduca sexta larvae in a parasitoid 'dose-dependent' fashion. Following parasitization of fourth-instar larvae, more heavily parasitized larvae grew larger compared to those containing fewer parasitoids due to an increase in host dry weight. The differences in host mass appeared to arise after oviposition. A 'dose-dependent' enhancement of host dry weight would appear nutritionally beneficial for the parasitoids developing in more 'crowded' hosts. The efficiencies of conversion of ingested and digested food to body mass and the approximate digestibility of the diet ingested by the host caterpillar did not vary significantly with clutch size although parasitoids took slightly longer to develop in the more heavily parasitized hosts. Larval parasitoids developing in the presence of many competitors weighed up to 50% less than those developing in hosts with fewer endoparasitoids, although the weight of adult female parasitoids did not vary significantly with wasp clutch size. The maximum number of emerging wasps was 200 parasitoids, possibly representing the host's 'carrying capacity' for larvae parasitized in the fourth-instar. The ratio of emerging to non-emerging parasitoids decreased as parasitoid clutch size increased, with few or none emerging from very heavily parasitized hosts containing more than 400 parasitoids. Copyright 1997 Elsevier Science Ltd. All right reserved     

Does the Polyphagous Egg Parasitoid <Emphasis Type="Italic">Trichogramma platneri</Emphasis> Nagarkatti (Hymenoptera: Trichogrammatidae) Display Behavioral Plasticity When Parasitizing Different Hosts?

Trichogramma platneri oviposition behavior on Amorbia cuneana egg masses was investigated under laboratory conditions. No relationship was detected between host surface area, number of edge turning on host eggs, and parasitoid clutch size. It was also observed that females parasitized the egg mass randomly by drilling into the egg mass and parasitizing individual eggs without using stereotypical behaviors to assess individual hosts (i.e. drumming and turning). However, clutch size did change due to ovipositional experience as naïve wasps with little or no ovipositional experience (<6 eggs parasitized), allocated significantly more progeny per host than wasps with longer ovipositional experience (24-h oviposition experience on a single egg mass). Moreover, naïve wasps parasitized significantly more eggs on the outer edge of the egg mass than experienced wasps. We contend that the physical characteristics of A. cuneana eggs and egg masses preclude T. platneri from completely discriminating between individual eggs. However, because T. platneri may be using kairomones from the egg mass, this described oviposition strategy remains effective.     

Growth rate adjustment of two Drosophila parasitoids in response to the developmental stage of hosts

1. Generalist koinobiont parasitoids often exhibit high flexibility in their development; their larvae shorten or prolong the developmental period depending on the host quality at parasitisation. However, flexibility of the growth rate of parasitoid larvae has rarely been investigated so far. 2. This study investigated how the koinobiont parasitoid wasps Asobara japonica and Leptopilina ryukyuensis regulate their larval growth when they parasitise host Drosophila larvae with varying larval periods. 3. In both parasitoid species, the preimaginal period was longer when they parasitised 1‐day‐old larvae of Drosophila rufa than when they parasitised older larvae of D. rufa or when they parasitised larvae of Drosophila simulans, a species with a shorter larval period than D. rufa. After host pupariation, A. japonica accelerated its growth, thereby showing a biphasic growth curve. On the other hand, L. ryukyuensis did not accelerate its growth after host pupariation. 4. Growth retardation of parasitoid larvae in 1‐day‐old D. rufa larvae would contribute to avoiding excess growth before host pupariation, because the excess growth of parasitoid larvae would have negative effects on host growth. The growth rate acceleration of A. japonica after host pupariation suggests that they enhance resource utilisation in a host that has reached maximum body mass. It remains uncertain as to why L. ryukuensis does not show clear accelerated growth after host pupariation. Nonetheless, these results suggest that parasitoid larvae have the ability to detect the developmental stage of hosts in a species‐specific manner.     

Development of a solitary koinobiont hyperparasitoid in different instars of its primary and secondary hosts

Parasitoid wasps are excellent organisms for studying the allocation of host resources to different fitness functions such as adult body mass and development time. Koinobiont parasitoids attack hosts that continue feeding and growing during parasitism, whereas idiobiont parasitoids attack non-growing host stages or paralyzed hosts. Many adult female koinobionts attack a broad range of host stages and are therefore faced with a different set of dynamic challenges compared with idiobionts, where host resources are largely static. Thus far studies on solitary koinobionts have been almost exclusively based on primary parasitoids, yet it is known that many of these are in turn attacked by both koinobiont and idiobiont hyperparasitoids. Here we compare parasitism and development of a primary koinobiont hyperparasitoid, Mesochorus gemellus (Hymenoptera: Ichneumonidae) in larvae of the gregarious primary koinobiont parasitoid, Cotesia glomerata (Hymenoptera: Braconidae) developing in the secondary herbivore host, Pieris brassicae (Lepidoptera: Pieridae). As far as we know this is the first study to examine development of a solitary primary hyperparasitoid in different stages of its secondary herbivore host. Pieris brassicae caterpillars were parasitized as L1 by C. glomerata and then these parasitized caterpillars were presented in separate cohorts to M. gemellus as L3, L4 or L5 instar P. brassicae. Different instars of the secondary hosts were used as proxies for different developmental stages of the primary host, C. glomerata. Larvae of C. glomerata in L5 P. brassicae were significantly longer than those in L3 and L4 caterpillars. Irrespective of secondary host instar, every parasitoid cluster was hyperparasitized by M. gemellus but all only produced male progeny. Male development time decreased with host stage attacked, whereas adult male body mass did not, which shows that M. gemellus is able to optimally exploit older host larvae in terms of adult size despite their decreasing mass during the pupal stage. Across a range of cocoon masses, hyperparasitoid adult male body mass was approximately 84% as large as primary parasitoids, revealing that M. gemellus is almost as efficient at exploiting host resources as secondary (pupal) hyperparasitoids.     

Effects of the gregarious parasitoids,Apanteles ruficrus and A. Kariyai,on host growth and development

Two gregarious parasitoids, Apanteles ruficrus and A. kariyai attack larvae of the common armyworm, Pseudaletia separata. Their growth pattern and growth rate of the parasitoids were not affected by host age at the time of oviposition, even though host weight increased exponentially with age. Consequently, the maximal weight of a single parasitoid larva was nearly constant regardless of host instar parasitized. Parasitoid females laid more eggs in later-instar hosts than in earlier-instar hosts. When parasitized at the same age, heavily parasitized hosts attained a larger mass than lightly parasitized larvae. Therefore, the ratio of the maximal weight of the host to the parasitoid mass was nearly constant. These results indicate that host growth is regulated by the parasitoids.     

螟蛉盘绒茧蜂的过寄生行为及其对子代生长发育的影响

【目的】过寄生现象普遍存在于寄生蜂寄生过程中。本研究旨在探究螟蛉盘绒茧蜂Cotesia ruficrus的过寄生对子代发育的影响及影响过寄生行为的主要因素。【方法】室内研究了螟蛉盘绒茧蜂在其寄主稻纵卷叶螟Cnaphalocrocis medinalis3龄幼虫上过寄生行为的发生,不同产卵次数对寄主存活及子代蜂生长发育的影响,研究了不同接蜂时间和不同接蜂密度对过寄生发生的影响。【结果】螟蛉盘绒茧蜂存在过寄生行为,无论雌蜂有无产卵寄生经历,均能在被自身寄生过和同种不同个体寄生过的寄主内产卵。寄生蜂茧量随着被产卵次数的增加而增加,被产卵3~5次的寄主体内死亡的寄生蜂幼虫数随着增大。寄主在育出蜂前的死亡率随着被产卵寄生次数的增加而增加,被产卵5次时,寄主育出蜂前死亡率达50%。过寄生使螟蛉盘绒茧蜂子代蜂卵-蛹的历期延长,羽化率和雌雄性比下降,雌蜂体型随寄生次数的增加显著变小。过寄生率随着接蜂密度及接蜂时间的增加而增加。【结论】在寄主上产卵2次对螟蛉盘绒茧蜂子代发育是最适合的,产卵3次及以上为过寄生。过寄生使蜂子代发育适合度降低,不利于子代生长发育。在室内大量繁殖螟蛉盘绒茧蜂时,应减少接蜂密度和接蜂时间,从而减少过寄生的发生。     

Studies onBonnetia comta [Dipt.: Tachinidae] parasitizingAgrotis ipsilon [Lep.: Noctuidae] larvae

Laboratory studies showed that 1st-instarBonnetia comta (Fallén) maggots (planidia) had a significant impact (P<0.05) on mortality of all black cutworm (BCW),Agrotis ipsilon (Hufnagel), instars, either by killing 1st- and 2nd-instar BCWs 2.3 to 9.7 days after parasitization or by producing a puparium from older host instars. Diet consumption and utilization by BCW larvae parasitized byB. comta as 4th instars were similar to those of nonparasitized larvae until 1 to 2 days before the parasitoid emerged. In a 2-year host exposure study in Iowa, it was found thatB. comta primarily parasitized released BCWs in June through September and did not seem to play a role in controlling the damaging 1st generation of BCW larvae. Techniques were developed to produce and store large numbers ofB. comta planidia.Bonnetia comta deposited large numbers of planidia on filter paper treated with a fecal supernatant. These planidia could be stored on filter paper in a covered Petri dish at 4.4°C for 5 days with minimal mortality. Preliminary field data show that planidia placed around corn seedlings infested with 4th-instar BCW larvae do parasitize the pest and reduce the hosts cutting potential.     

Comparing the physiological effects and function of larval feeding in closely‐related endoparasitoids (Braconidae: Microgastrinae)

Abstract The larvae of most endoparasitoid wasps consume virtually all host tissues before pupation. However, in some clades, the parasitoid larvae primarily consume haemolymph and fat body and emerge through the side of the host, which remains alive and active for up to several days. The evolutionary significance of this host‐usage strategy has attracted attention in recent years. Recent empirical studies suggest that the surviving larva guards the parasitoid broods against natural enemies such as predators and hyperparasitoids. Known as the ‘usurpation hypothesis’, the surviving larvae bite, regurgitate fluids from the gut, and thrash the head capsule when disturbed. In the present study, the ‘usurpation hypothesis’ is tested in the association involving Manduca sexta, its parasitoid Cotesia congregata, and a secondary hyperparasitoid Lysibia nana. Percentage parasitoid survival is higher and hyperparasitism lower when cocoons of C. congregata are attached to the dorsum of M. sexta caterpillars. Fat body contents in several associations involving solitary and gregarious parasitoids feeding on haemolymph and fat body are also compared. The amount of fat body retained in parasitized caterpillars varies considerably from one association to another. In M. sexta and Pieris brassicae, considerable amounts of fat body remain after parasitoid emergence whereas, in Cotesia kariyai and Cotesia rufricus, virtually all of the fat body is consumed by the parsasitoid larvae. The length of post‐egression survival of parasitized caterpillars differs considerably in several tested associations. In Pseudeletia separata, most larvae die within a few hours of parasitoid emergence whereas, in M. sexta, parasitized larvae live up to 2 weeks after parasitoid emergence. Larvae in other associations parasitized by gregarious and solitary endoparasitoids live for intermediate periods. The results are discussed in relation to the adaptive significance of different feeding strategies of immature parasitoids and of the costs and benefits of retaining the parasitized caterpillar in close proximity with the parasitoid cocoons.     

Field confirmation of a mechanism causing synergism between Bacillus thuringiensis and the gypsy moth parasitoid, Apanteles melanoscelus

In 16-ha plots aerially sprayed with single and double applications of Bacillus thuringiensis, percentage parasitism by A. melanoscelus and the number of A. melanoscelus cocoons under burlap strips were higher than in comparable untreated plots in the same area. Strong correlations occurred between percentage parasitism and caterpillar size, with plots having the smallest caterpillars being the most heavily parasitized. However, these parameters were also negatively correlated with number of caterpillars per plot. The increased numbers of parasitoid progeny, i.e., cocoons, found in treated plots showed that corresponding increases in percentage parasitism could not be due simply to improved parasitoid: host ratios. Evidence strongly suggests that the retarding effect of B. thuringiensis infection kept gypsy moth larvae small enough in the treated plots to permit A. melanoscelus females to parasitize relatively large numbers of caterpillars.