Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

The influence of the rearing host on the response of the parasitoid Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae) to odours from Ephestia kuehniella and Plodia interpunctella in a Y-tube olfactometer

Venturia canescens (Gravenhorst) is an ichneumonid generalist parasitoid that successfully attacks the larvae of different lepidopteran pests that infest stored products. These pest species include Plodia interpunctella and Ephestia kuehniella. In this study, we aimed to evaluate the influence of the rearing host on the parasitoid’s ability to detect and respond to a new host different from the rearing species. For this reason, the trials tested the preference of parasitoids reared on P. interpunctella or E. kuehniella for products that were or were not infested with larvae of these hosts. The trials were conducted in a Y-tube olfactometer. Regardless of the rearing host species, the parasitoids showed no preference for uninfested products. The parasitoids were attracted to products infested with larvae of their rearing host in preference to uninfested products. They also showed preferential attraction to products infested with the new host over uninfested products. E. kuehniella was the preferred host, irrespectively of the parasitoid host rearing species. The results are discussed to develop a better understanding of the ecology of V. canescens for its application in biological control.     

Development and encapsulation of the endoparasitoid,Microplitis croceipes (Hym.: Braconidae), in six candidate host species (Lep.)

Encapsulation and development of the endoparasitoid,Microplitis croceipes (Cresson), were studied in six atypical lepidopteran host species whose usual host isHelicoverpa zea (Boddie). The candidate hosts examined were: the fall armywormSpodoptera frugiperda (J. E. Smith); the beet armyworm,Spodoptera exigua (Hübner); the cabbage looper,Trichoplusia ni (Hübner); the greater wax moth,Galleria mellonella (L.); the Indian meal moth,Plodia interpunctella (Hübner); and the diamondback moth,Plutella xylostella (L.). BothS. exigua andT. ni were completely unsuitable forM. croceipes development due to the high rate of eggs that were encapsulated within three days after parasitism. Encapsulation inS. frugiperda included mainly parasitoid eggs and was first detected six days after parasitization at 25°C and two days at 30°C. Encapsulation inG. mellonella occurred only in the larval stage of the parasitoid. InP. interpunctella, parasitoid larvae reached the 3rd stadium, but none of them pupated. OnlyS. frugiperda andG. mellonella supported successful development ofM. croceipes from egg to adult. The percentage of parasitoids reaching the adult stage in these hosts was higher at 30°C than at 25°C (13% vs. 4% inS. frugiperda, and 21% vs. 3% inG. mellonella, respectively). However, these percentages were too low to substitute them as a more economical host for rearingM. croceipes. This biological information will be useful in additional laboratory studies directed toward reducing the rate of encapsulation (e.g., manipulation of host rearing temperature) to increase production ofM. croceipes on these hosts.     

The effect of host nutrition on growth and development of the parasitoid waspVenturia canescens

We investigated the effect of host (Plodia interpunctella; Lepidoptera: Pyralidae) nutritional status on development of the solitary endoparasitoid,Venturia canescens (Hymenoptera: Ichneumonidae). Parasitoids from 3rd (L3) instars reared on a deficient diet during early parasitism took longer to develop and suffered higher mortality than those reared from hosts fedad libitum although there was not a significant difference in the size of eclosing wasps from the two groups. L5 hosts reared at high density produced smaller parasitoids, which developed more rapidly than those reared from hosts from low density containers, although mortality was higher in the latter. In a separate experiment we starved groups of 10–20 hosts (parasitized as L3) daily beginning on the 4th day after parasitism, to determine the host developmental stage required for successful parasitoid development to eclosion. Parasitoid survivorship increased with length of host access to food, while the egg-to-adult parasitoid development time increased throughout the experiment. Parasitoid size decreased with increasing periods of host starvation. The successful emergence ofVenturia depends uponPlodia reaching the size normally attained in the mid-5th instar, or 50–70% of the mass of healthy late 5th instars. Our results show that when earlier instars are parasitized, host growth is essential for successful parasitoid development to eclosion. Furthermore, they suggest that, for many koinobionts, host suitability may be greatly influenced by feeding rate and food quality.     

Oviposition behaviour, influence of experience on host size selection, and niche overlap of the solitary Leptomastix epona and the gregarious Pseudaphycus flavidulus, two endoparasitoids of the mealybug Pseudococcus viburni

Oviposition behaviour and host size selection of the solitary parasitoid Leptomastix epona(Walker) and the gregarious Pseudaphycus flavidulus(Brèthes) [both Hymenoptera: Encyrtidae] were examined on five size classes of the mealybug Pseudococcus viburni(Signoret) [Hemiptera: Pseudococcidae]. The host size classes mostly consisted of one stage (first, second, third instar nymph, young adult and preovipositing adult) and were presented together to wasps of either parasitoid species. Both parasitoid species locate the host by drumming the surface of the patch with the antennae. Leptomastix eponaseems to use mainly the antennae to examine the host but P. flavidulusmay accept or reject a host for oviposition after antennation or insertion of the ovipositor. Leptomastix eponaattempts oviposition in all the host stages from second instar nymphs but P. flavidulusincludes first instar. Both parasitoid species select mainly larger hosts (>1 mm, third instar nymphs) to oviposit but P. flavidulusis able to parasitize more second instar nymphs compared to L. epona. Female wasps of L. eponamay host feed on small mealybugs (second and third instar nymphs) that they do not use for oviposition. Oviposition experience of either parasitoid species for 24 hours does not influence host size selection on patches with hosts of similar mixed sizes. Oviposition decisions are independent of the host sizes of the preceding ovipositions. Implications about stability of a single parasitoid – host system and the success of biological control of the mealybug were discussed in respect of the developmental refugia of the two parasitoid species. Niche overlap of the two parasitoid species was discussed with a view to giving an insight into a single or multiple introduction.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Lifetime Reproductive Success in the Solitary Endoparasitoid, Venturia canescens

Parasitoid wasps have long been considered excellent organisms in studies examining the evolution of reproductive and life-history strategies. In examining the lifetime reproductive success of parasitoids in the laboratory, most investigations have provided the insects with excess hosts and food, where they exist in a relatively constraint-free environment. Importantly, these conditions may not accurately reflect the true heterogeneity of natural systems, where suitable hosts and food sources are likely to be limiting. This study examines the influence of differences in host and food availability on reproductive and life-history parameters in an asexual strain of the solitary endoparasitoid, Venturia canescens (Hymenoptera: Ichneumonidae). Lifetime reproductive success in V. canescens was measured in response to temporal variations in host and food (honey solution) access. Cohorts of parasitoids were provided with 200 fifth-instar larvae of the Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae), and food for variable periods daily after eclosion. V. canescens is synovigenic, and host-deprived wasps continued to mature eggs over the first few days after eclosion until the egg storage capacity was reached in the oviducts. When these wasps were subsequently provided with hosts, oogenesis resumed and continued until later in adult life. Constantly fed wasps lived longer and produced more progeny than wasps from cohorts which were alternately fed and starved or were starved from eclosion. Moreover, wasps with constant host and food access produced most progeny early in life and usually experienced prolonged periods of postreproductive survival. In contrast, the reproductive period of wasps with limited host access was more evenly distributed throughout the adult life. Consequently, the cumulative progeny production by V. canescens with constant food access was fairly uniform irrespective of host availability. Longevity and fecundity in V. canescens were positively correlated with adult size. However, variable host access had little effect on the longevity of wasps which were constantly supplied with honey. Over the first 2 days of adult life, variation in food access also had no effect on progeny production by V. canescens. We argue that manipulating temporal host and food access to parasitoids in the laboratory more closely approximates natural conditions, where these resources are likely to be spatially separated. Moreover, our findings suggest that many highly synovigenic parasitoids like V. canescens, which produce microtype (=hydropic) eggs, have a considerably higher reproductive potential than ovary dissections have revealed. Our findings are discussed in relation to life-history evolution in the parasitic Hymenoptera.     

Foraging for solitarily and gregariously feeding caterpillars: A comparison of two related parasitoid species (Hymenoptera: Braconidae)

In the present study we apply a comparative approach, in combination with experimentation, to study behavior of two parasitoid species that attack caterpillar hosts with different feeding strategies (gregarious or solitary). In a semifield setup, consisting of clean cabbage plants and plants infested with one of two host species, the foraging behavior of the specialistCotesia rubecula, on obligate parasitoid of solitarily feedingPieris rapae larvae, was compared to that of the generalistCotesia glomerata, a polyphagous parasitoid of several Pieridae species (mainly the gregariously feedingPieris brassicae).Cotesia glomerata displayed equal propensity to search for and parasitize larvae of both host species. AlthoughC. glomerata exhibited a relatively plastic foraging behavior in that it searched differently under different host distribution conditions, its behavior seems more adapted to search for gregariously feeding hosts. Females exhibited a clear area-restricted search pattern and were more successful in finding the gregariously feeding caterpillars.Cotesia rubecula showed a higher propensity to search forP. rapae than forP. brassicae, i.e., females left the foraging setup significantly earlier when their natural hostP. rapae was not present.C. rubecula showed a more fixed foraging behavior, which seems adapted to foraging for solitarily feeding host larvae. In a setup with onlyP. rapae larvae, the foraging strategies of the two parasitoid species were quite similar. In a choice situationC. glomerata did not show a preference for one of the host species, whileCotesia rubecula showed a clear preference for its natural host species. The latter was shown by several behavioral parameters such as the number of first landings, allocation of search time, and percentage parasitization.     

Influence of laboratory hosts on the biological attributes ofChelonus blackburni [Hym.: Braconidae]

Among 6 laboratory hosts tested, the egg-larval parasitoidChelonus blackburni Cameron completed its development in only 3;Corcyra cephalonica Stainton,Phthorimaea operculella (Zeller) andAchroia grisella (Fabricius). The parasitoid failed to develop inSpodoptera litura (Fabricius),Galleria mellonella L. andSitotroga cerealella (Olivier). The development period of the parasitoids obtained from permissive hosts correspond to the development period of their hosts. Development period ofP. operculella, A grisella andC. cephalonica were 24.7±2.0, 39.5±2.1 and 50.5±4.1 days respectively while the development period of the parasitoid reared from these hosts were 25.8±1.6, 36.4±3.5 and 42.5±3.5 days respectively. The fecundity of the parasitoids reared onA. grisella, P. operculella andC. cephalonica were 365.2±52.8, 287.9±101.9 and 248.7±50.8 respectively. The size of the parasitoids reared from the above 3 hosts also followed the similar trend. However, no significant difference was observed on the percent parasitism and the longevity of the parasitoids reared on different hosts. This study was conducted at a temperature of 24±2°C and 60±5% RH. Contribution No 46001 of Biological Control Centre (NCIPM), Bangalore.     

Consequences of mixed species infestation on the searching behavior and parasitism success of a larval parasitoid

When two herbivore pest species are potential hosts of a single parasitoid species, two questions arise. Firstly, which host is preferable for mass rearing in terms of later parasitoid performance, and secondly, how do parasitoids perform in mixed herbivore situations after colony establishment? We tested Hyssopus pallidus, a gregarious parasitoid of two major pests of apple, Cydia (Grapholita) molesta and Cydia pomonella, before and after landing on apples infested by one of the two Cydia species. Pre-alighting host preference was tested in a Y-tube olfactometer setup, and parasitism success in a contact bioassay. To gain information on parasitoid performance throughout the growing season, different fruit growth stages were used. Irrespective of the host they had developed on, the parasitoids showed similar olfactory preferences when given a dual choice between infested and healthy fruits, and they did not discriminate between fruits infested by C. molesta and C. pomonella. Responsiveness was generally high, especially late in the season close to harvest. Both hosts are parasitized regardless of the host the parasitoid female had developed on, and no differences in parasitism rates or number of offspring were noted for the two hosts offered. Results were consistent for all apple growth stages tested. In conclusion, mass rearing of this parasitoid can be carried out on either host, without limiting the future efficacy of the bio-control agent. Similarly, established colonies are expected to develop further on both hosts without any bias in host preference.     

Flexible larval development and the timing of destructive feeding by a solitary endoparasitoid: an optimal foraging problem in evolutionary perspective

1. In studying the evolution of life-history strategies in parasitoids, considerable attention has been paid to the relationship between host quality and parasitoid fitness. Various workers have reported that host quality influences parasitoid size, development time, and survival. Because body size is frequently correlated with fecundity, longevity, and host-finding ability in parasitoids, this parameter is usually considered to be the main target of selection. 2. In koinobiont parasitoids that consume the entire host before pupation, adult parasitoid size and development time are often strongly correlated with host size at the time when it is developmentally arrested through destructive feeding by the parasitoid larva. 3. Here, a mathematical model is proposed to describe the larval feeding behaviour of the solitary koinobiont endoparasitoid Venturia canescens in four larval stadia of its host Plodia interpunctella. In particular, the model describes how adult size, represented by an exponential growth rate, and development time are traded off when the parasitoid develops in nutritionally suboptimal second stadium hosts. 4. Using a graphical model, the different conditions faced by V. canescens during development in various host species of greatly differing mass are illustrated. 5. It is argued that the relative importance of size and development time on parasitoid fitness is determined by ecological and biological characteristics of both host and parasitoid, and it is suggested that there may be correlations between life-history traits and host-utilisation strategies among koinobionts.     

Studies on the characteristics of parasitism ofPlutella xylostella (Lep.: Plutellidae) by a larval parasiteDiadegma semiclausum (Hym.: Ichneumonidae)

Laboratory experiments were conducted to study the characteristics of parasitism of diamondback moth,Plutella xylostella (L.), a worldwide pest of crucifers, by a larval parasite,Diadegma semiclausum Hellén. Results showed that the greater the host larval density, the lesser was the percentage of larvae parasitized. The area of discovery decreased with increasing parasite density, but k-value remained practically unchanged. Parasitism reduced food consumption in parasitised larvae. No difference was observed in duration of larval instars between the parasitized and healthy larvae. As the frequency of superparasitism increased, the proportion of production of female progeny also increased. Presence of parasite larva within the host larva deterred superparasitism greater than the presence of parasite egg. Presence of larva of eitherD. semiclausum or another larval parasite,Cotesia plutellae Kurdjumov, within host larva deterred multiparasitism by either species greater than the presence of egg of either parasite species.Diadegma semiclausum was much more aggressive thanC. plutellae in parasitisingP. xylostella larvae.     

Preferential infectivity of entomopathogenic nematodes in an envenomed host

Entomopathogenic nematodes and parasitoid wasps are used as biological control agents for management of insect pests such as the Indian meal moth, Plodia interpunctella. The parasitoid wasp Habrobracon hebetor injects a paralytic venom into P. interpunctella larvae before laying eggs. A previous study reported that the entomopathogenic nematode Heterorhabditis indica preferentially infects P. interpunctella that have been envenomed by H. hebetor while results in this study showed a similar preference by the entomopathogenic nematode, Steinernema glaseri. We therefore tested four hypotheses for why nematode infection rates are higher in envenomed hosts: (1) elevated CO₂ emission from envenomed hosts attracts nematodes, (2) paralysis prevents hosts from escaping nematodes, (3) volatile chemicals emitted from envenomed hosts attract nematodes and increase infection, and (4) reduced immune defenses in envenomed hosts increase nematode survival. Results showed that envenomed P. interpunctella larvae emitted lower amounts of CO₂ than non-envenomed larvae. Physical immobilization of P. interpunctella larvae did not increase infection rates by S. glaseri but did increase infection rates by H. indica. Emissions from envenomed hosts were collected and analyzed by thermal desorption gas chromatography/mass spectrometry. The most abundant compound, 3-methyl-3-buten-1-ol, was found to be an effective cue for S. glaseri attraction and infection but was not an effective stimulus for H. indica. Envenomed P. interpunctella exhibited a stronger immune response toward nematodes than non-envenomed hosts. Altogether, we conclude that different mechanisms underlie preferential infection in the two nematode species: host immobilization for H. indica and chemical cues for S. glaseri.     

Nondevelopment ofCotesia kariyai(Hymenoptera: Braconidae) in Entomopoxvirus-Infected Larvae ofPseudaletia separata(Lepidoptera: Noctuidae)

Interaction between an entomopoxvirus (PsEPV) and a gregarious braconid endoparasitoid,Cotesia kariyai,inPseudaletia separatalarvae showed that infection of larvae with PsEPV was deleterious to the development and survival ofC. kariyai.The survival and development ofC. kariyaiin PsEPV-infectedP. separatalarvae depended on the length of time between parasitization and viral infection. No parasitoid larvae emerged from PsEPV-infected hosts when host larvae were exposed simultaneously to parasitization and PsEPV inoculation whereas more than 80% of the hosts produced parasitoids when PsEPV was administered 5 days postparasitization.C. kariyailarvae in PsEPV-infected hosts showed a retarded development, shrank, and died about 8 days after viral exposure. Virion-free plasma from PsEPV-infectedP. separatalarvae was toxic to the parasitoid larvae even up to a dilution level of 32 when it was injected intrahemocoelically into the host larvae. Development of parasitoids in hosts that were simultaneously parasitized and injected with the virion-free plasm never progressed beyond the egg stage. The parasitizedP. separatalarvae injected with the virion-free plasma did not pupate and died within 30 days after injection.     

The influence of plant species on attraction and host acceptance inCotesia glomerata (Hymenoptera: Braconidae)

Females of the larval parasitoidCotesia glomerata (L.) use plant-associated cues to locate their lepidopteran host,Pieris rapae L. In this study we investigated the influence of four host plant species,Brassica oleracea var.acephala (‘Vates’ kale),Tropaeolum majus (nasturtium),Lunaria annua (honesty), andCleome spinosa (spider flower), on two components of the host selection process inC. glomerata, namely, attraction and host acceptance. Choice tests in a flight tunnel showed that parasitoids were attracted to some host plant species more than to others in the absence of host larvae.B. oleracea was the most attractive plant species, followed byL. annua, T. majus, andC. spinosa. In previous studies it was shown thatB. oleracea carries highly suitable hosts forC. glomerata and that, in the field, parasitization rates on this plant were the highest. When host larvae were reared on the four host plant species and then transferred to a common substrate (B. oleracea var.capitata, cabbage), plant species that had served as diet for the hosts did not have a significant effect on acceptance for parasitization. Thus, parasitoids were attracted to host plant species differentially, but they did not discriminate among host larvae based on the dietary history of their hosts. ForC. glomerata, it appears that phytochemistry mediates host selection more by influencing parasitoid attraction than it does by affecting host acceptance.     

Potential for lab rearing of <Emphasis Type="Italic">Apanteles taragamae</Emphasis>, the larval endoparasitoid of coconut pest <Emphasis Type="Italic">Opisina arenosella</Emphasis>, on the rice moth <Emphasis Type="Italic">Corcyra cephalonica</Emphasis>

A lab rearing technique was standardised for Apanteles taragamae Viereck (Hymenoptera: Braconidae), the early larval parasitoid of the coconut leaf-eating caterpillar, Opisina arenosella Walker on the alternate host Corcyra cephalonica Stainton (Lepidoptera: Pyralidae). The parasitoid took 23.3 ± 3.2 days to complete the egg to adult period. Adult longevity for males and females was 15.3 ± 4.6 and 13.8 ± 4.6 days respectively. Fecundity was 14.8 ± 4.3 eggs per female. The percentage parasitism was 60.6 ± 5.7 on the alternative host C. cephalonica and 64.6 ± 5.5 on the natural host O. arenosella. Eight- to ten-day-old caterpillars were the ideal stage of C. cephalonica for rearing A. taragamae. The results indicated the amenability of rearing A. taragamae on C. cephalonica in the laboratory.     

Bioecology ofLydella thompsoni Herting, [Dip. Tachinidae] within the Rhone Delta in Southern France

Lydella thompsoni Hertin,Tachinidae, is an endoparasitoid frequently associated in southern France with larvae ofOstrinia nubilalis, Pyralidae, Sesamia nonagrioides, Noctuidae, Archanara geminipuncta andA. dissoluta, Noctuidae. The tachinid was reared successfully at 21°C in the laboratory by providing the adult flies with a high humidity, a light intensity of 8,000–10,000 lux for mating and a mixture of casein proteolysate and honey as food. Under such conditions, the flies lived for about 30 days and about half of them mated successfully. Hosts were infested by dissecting mature tachinid females and placing 1–2 of the extracted planidia onto each moth larva. Half of the planidia successfully entered their larval hosts. The biology of the tachinid larvae was studied on the 2 main hosts,O. nubilalis andS. nonagrioides. At 21°C, 25°C and 28°C, larval development took less time onO. nubilalis than onS. nonagrioides. At 25°C, female larvae onO. nubilalis required 9.0±0.5 days and onS. nonagrioides 10.5±0.3 days, male larvae onO. nubilalis required 8.3±0.5 days and onS. nonagrioides 10.6±0.3 days. Pupal duration was also influenced by the larval host. In winter, 2nd instar larvae ofL. thompsoni enter a resting or quiescent condition. This condition is terminated sooner (December–January) in larvae developing onS. nonagrioides than in those developing onO. nubilalis (February–March). The life cycle of the tachinid in the field was studied by trapping flies in water dishes and by collecting parasitized host larvae from various plants. Flies were caught from April to October, mainly in September. Larvae of the spring generation of the parasitoid developed on larvae of species ofArchanara that fed on the reed,Phragmites communis. From 1976 to 1982, parasitism averaged about 16%. Parasitism by summer generation onS. nonagrioides was highest (4–5%) in June. Parasitism ofO. nubilalis did not change by more than 2 fold in either of the 2 summers studied (10–17 % in 1981, 6–10 % in 1982). By September the numbers of host larvae had increased to 10–20 times the number available earlier in the season. Larval populations ofL. thompsoni similarly increased from 3–400 larvae per ha in June to 4–5,000 per ha in September. The stability of the relationship between this parasitoid and its hosts in southeastern France is discussed and compared to relationship described elsewhere.     

Putative Host Volatiles Used by <Emphasis Type="Italic">Habrobracon hebetor</Emphasis> (Hymenoptera: Braconidae) to Locate Larvae of <Emphasis Type="Italic">Plodia interpunctella</Emphasis> (Lepidoptera: Pyralidae)

Indianmeal moth, Plodia interpunctella (Hubner) (Lepidoptera: Pyralidae), is a post-harvest pest of grains, milled and processed food, processing plants, warehouses and bakeries. The parasitoid, Habrobracon hebetor (Say) is among the most important natural enemies of Pyralidae infesting stored grains and grain products. Many parasitoids use semiochemicals originating from their hosts, or host’s habitat as cues to locate hosts, hosts’ food or habitat. The authors used Y-tube and four-way olfactometers to assay responses to stimuli with the moth host and thereby understand the role of host-associated semiochemicals in host location by H. hebetor. Responses of mated parasitoid females were assayed to the following stimuli: P. interpunctella sex pheromone, female adults, larvae, or hexane extracts of residue of the rearing medium. Generally, host-related odor sources generated stimuli that elicited better responses than those to blank controls. Previous exposure to odor sources from the host shortened latency periods and response times compared to naïve females. Odors emanating from live moth larvae elicited the strongest responses. When responses from the four odor sources were compared in a four-way olfactometer, it was confirmed that volatiles from larval moths elicited the strongest attraction to the parasitoid. The involvement of host-specific chemical cues in both long and short range host location by female parasitoid is discussed.     

Tritrophic Choice Experiments with Bt Plants,the Diamondback Moth (Plutella xylostella) and the parasitoid Cotesia plutellae

Parasitoids are important natural enemies of many pest species and are used extensively in biological and integrated control programmes. Crop plants transformed to express toxin genes derived from Bacillus thuringiensis (Bt) provide high levels of resistance to certain pest species, which is likely to have consequent effects on parasitoids specialising on such pests. A better understanding of the interaction between transgenic plants, pests and parasitoids is important to limit disruption of biological control and to provide background knowledge essential for implementing measures for the conservation of parasitoid populations. It is also essential for investigations into the potential role of parasitoids in delaying the build-up of Bt-resistant pest populations. The diamondback moth (Plutella xylostella), a major pest of brassica crops, is normally highly susceptible to a range of Bt toxins. However, extensive use of microbial Bt sprays has led to the selection of resistance to Bt toxins in P. xylostella. Cotesia plutellae is an important endoparasitoid of P. xylostella larvae. Although unable to survive in Bt-susceptible P. xylostella larvae on highly resistant Bt oilseed rape plants due to premature host mortality, C. plutellae is able to complete its larval development in Bt-resistant P. xylostella larvae. Experiments of parasitoid flight and foraging behaviour presented in this paper showed that adult C. plutellae females do not distinguish between Bt and wildtype oilseed rape plants, and are more attracted to Bt plants damaged by Bt-resistant hosts than by susceptible hosts. This stronger attraction to Bt plants damaged by resistant hosts was due to more extensive feeding damage. Population scale experiments with mixtures of Bt and wildtype plants demonstrated that the parasitoid is as effective in controlling Bt-resistant P. xylostella larvae on Bt plants as on wildtype plants. In these experiments equal or higher numbers of parasitoid adults emerged per transgenic as per wildtype plant. The implications for integrated pest management and the evolution of resistance to Bt in P. xylostella are discussed.