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Abstract Background Cotesia marginiventris (Cresson) and Microplitis rufiventris Kok. (Hymenoptera: Braconidae) are larval endoparasitoids of Spodoptera frugiperda (Smith) and S. littoralis (Boisd.), respectively. Cotesia marginiventris was introduced in Egypt to control the recent invasive pest, S. frugiperda and local one, S. littoralis. Therefore, competition between the two parasitoid species may occur. Results When single females of C. marginiventris (Exp. 1), single females of M. rufiventris (Exp. 2), and one female each of C. marginiventris and M. rufiventris (Exp. 3) were offered a choice of 60 1st, 60 2nd, and 60 3rd instars, of the tested host species, S. littoralis and different parasitization levels were obtained. In the non-competitive context (Exps. 1, 2), all instars were parasitized, but C. marginiventris and M. rufiventris parasitized primarily 2nd and 3rd instar host larvae, respectively. In the competitive context (Exp. 3), the same preference pattern emerged but the percentage of parasitism was lower than in each of the two non-competitive contexts. Conclusions M. rufiventris appears to be a superior competitor when simultaneously exploiting the host with the other parasitoid species. The knowledge of this work can be used to predict and/or understand possible outcomes of a biological control program using the two biocontrol agents against one or the two pests species, and to gain insight into possible biotic interference between species at the same trophic level exploiting the same host. Graphical abstract

The vast spectrum of inducible plant defenses can have direct negative effects on herbivores, or indirect effects, for instance in the form of herbivore-induced plant volatiles (HIPVs) that attract natural enemies. Various arthropods have evolved ways to suppress plant defenses. To test whether this is the case for caterpillar-induced HIPVs, we compared the volatile induction by Spodoptera frugiperda (Lepidoptera: Noctuidae), which is particularly well adapted to feed on maize (Zea mays), with the induction by three more generalist noctuid larvae. We tested the hypothesis that S. frugiperda suppresses HIPV emissions in maize, and thereby reduces attractiveness to natural enemies. HIPV emissions triggered by S. frugiperda when feeding on maize were indeed found to be significantly weaker than by Spodoptera littoralis, Spodoptera exigua, and Helicoverpa armigera. The suppression seems specific for maize, as we found no evidence for this when S. frugiperda caterpillars fed on cotton (Gossypium herbaceum). Artificially damaged maize plants treated with larval regurgitant revealed that HIPV suppression may be related to factors in the caterpillars' oral secretions. We also found evidence that differential physical damage that the caterpillars inflict on maize leaves may play a role. The suppressed induction of HIPVs had no apparent consequences for the attraction of a common parasitoid of S. frugiperda, Cotesia marginiventris (Hymenoptera: Braconidae). Nevertheless, the ability to manipulate the defenses of its main host plant may have contributed to the success of S. frugiperda as a major pest of maize, especially in Africa and Asia, which it has recently invaded. [ABSTRACT FROM AUTHOR]

Carnivorous animals put selection pressure on their herbivorous preys, leading to the development of defensive traits in prey species and counter defensive traits in carnivores—an evolutionary arms race for survival. For instance, larvae of the moth Heliothis virescens use defensive oral secretions to deter attacking parasitic wasps (parasitoids). However, it is not clear whether parasitoids also use the same secretion for host location. If carnivores follow the herbivore model, evolution should favor specialist over generalist parasitoid species in their ability to manipulate host defensive chemicals. To test this prediction, Microplitis croceipes and Cotesia marginiventris, both larval endoparasitoids of H. virescens, were used as a study system. Microplitis croceipes is relatively specialized on Heliothis/Helicoverpa species while C. marginiventris is a generalist parasitoid on several noctuid host species. Innate behavioral responses of parasitoids to host defensive secretion were tested in Petri dish arena and Y-tube olfactometer bioassays. Heliothis virescens defensive secretion was manually applied to a spot in a sector (secretion zone) of the arena. Tracking data suggests that both parasitoid species are likely to locate the secretion spot once they enter the secretion zone. Comparing the two species, however, M. croceipes (specialist) spent significantly more time in the secretion zone and spot than C. marginiventris (generalist). In Y-tube olfactometer bioassays, the secretion elicited a strong attraction in M. croceipes but not in C. marginiventris, supporting our prediction. We discussed chemical-mediated counter defense in natural enemies of pest insects and the significance of the results to parasitoid-host coevolution. [ABSTRACT FROM AUTHOR]

Sex pheromones have rarely been studied in parasitoids, and it remains largely unknown how male and female parasitoids locate each other. We investigated possible attraction (and repellency) between the sexes of two braconid wasps belonging to the same genus, the gregarious parasitoid, Cotesia glomerata (L.), and the solitary parasitoid, Cotesia marginiventris (Cresson). Males of both species were strongly attracted to conspecific virgin females. Interestingly, in C. glomerata, the males were repelled by mated females, as well as by males of their own species. This repellency of mated females was only evident hours after mating, implying a change in pheromone composition. Males of C. marginiventris were also no longer attracted, but not repelled, by mated females. Females of both species showed no attraction to the odors of conspecific individuals, male or female, and C. glomerata females even appeared to be repelled by mated males. Moreover, the pheromones were found to be highly specific, as males were not attracted by females of the other species. Males of Cotesia glomerata even avoided the pheromones of female Cotesia marginiventris, indicating the recognition of non-conspecific pheromones. We discuss these unique responses in the context of optimal mate finding strategies in parasitoids.

Microplitis croceipes and Cotesia marginiventris (Hymenoptera: Braconidae) are parasitoids of lepidopteran larvae with different degrees of host specificity. Both parasitoid species rely on host-related plant volatiles as odor cues to locate their herbivore hosts. To better understand mechanisms of odor processing in parasitoids, we tested responses of olfactory sensory neurons (OSNs) in the antennal sensilla placodea of female parasitoids to select plant volatiles and mixtures . The compounds tested include two green leaf volatiles (i.e., cis -3-hexenol and hexanal) and three herbivore-induced plant volatiles (i.e., cis -3-hexenyl butyrate, cis- 3-hexenyl acetate and linalool). Single-sensillum recording showed that the test compounds elicited activity in large and small amplitude neurons housed in the short sensilla placodea of both parasitoid species. In general, C. marginiventris showed greater OSN responses to a low dose while M. croceipes showed greater responses to a high dose of test compounds. Binary mixtures of cis -3-hexenol and linalool inhibited OSN activity in M. croceipes , but not in C. marginiventris . These differences may have implications for odor discrimination in the two parasitoid species. In addition, anterograde neurobiotin stainings were performed to map glomerular projections of OSNs in the antennal lobe of the parasitoids. In M. croceipes , a mixture of cis -3-hexenol and linalool inhibited activity of the glomerulus activated by cis -3-hexenol alone. In C. marginiventris, a mixture of cis -3-hexenol and cis -3-hexenyl acetate showed intense labeling in their respective glomeruli, possibly suggesting a synergistic interaction. These differences in detection and coding of single compounds and mixtures may impact host location strategies in the two parasitoid species. [ABSTRACT FROM AUTHOR]

Fall armyworm larvae, Spodoptera frugiperda (J. E. Smith), were collected from sweet corn plants ( Zea mays L.) in fields located in three south Florida counties. Fields were sampled from 2010 to 2015 during the fall and spring seasons. Larvae were transferred to the laboratory to complete development. The objective of the study was to identify the common parasitoids emerging from larvae that are present in sweet corn habitats where insecticides are traditionally used. A total of 8353 fall armyworm larvae were collected, of which 60.6% (5062) developed into moths after feeding on corn tissue and artificial diet. Parasitoids emerged from 2365 larvae (28.3%), and parasitism ranged from 1% to 91.7%, depending on site. Parasitism was higher at the University of Florida Everglades Research and Education (EREC) in Belle Glade (50.4 ± 11.8%) than at other locations in south Florida. Parasitism was comparable between fall and spring seasons, but was much higher in unsprayed fields (44.0 ± 9.6%) than in the sprayed fields (15.0 ± 2.5%). The two most common parasitoids that emerged from larvae were the solitary endoparasitoids Cotesia marginiventris (Cresson), found in 23 of the 25 sites sampled, and Chelonus insularis Cresson, found in 18 of the 25 sites sampled. Other parasitoid species that emerged from fall armyworm larvae were Aleiodes laphygmae (Viereck), Euplectrus platyhypenae Howard, Meteorus spp., Ophion flavidus Brullé, and unidentified species of Tachinidae. Techniques to improve the management of fall armyworm in overwintering areas of south Florida using conservation biological control are discussed. [ABSTRACT FROM AUTHOR]

Artificial selection of crop plants for desired traits such as increased yield and improved seed or fruit quality has been hypothesized to have had a cost for other potentially useful traits, including resistance to herbivores. Besides direct defences, such as the production of toxins, plants may also indirectly protect themselves by emitting volatile organic compounds (VOCs) that attract the natural enemies of herbivores. Parasitoid wasps are known to use these VOCs to localize hosts for their offspring. However, domestication and selective breeding of crop plants have reportedly led to the loss of such signals. The aim of this study was to identify possible differences in the attraction of parasitoid wasps by modern maize and its wild ancestors, the teosintes. In a six-arm olfactometer, we compared the capacity of teosintes and maize to attract the parasitoid wasps Cotesia marginiventris (Hymenoptera: Braconidae) and Campoletis sonorensis (Hymenoptera: Ichneumonidae). We studied the attractiveness of plants in which VOC emission was induced by the application of artificial damage and caterpillar regurgitant, as well as the attractiveness of extracts of volatiles that we collected from plants exposed to herbivory. C. sonorensis did not distinguish between the odours of maize and teosintes, whereas C. marginiventris showed a significant preference for the odours of teosintes over the odours of maize. The fact that we obtained very similar results with extracts of collected volatiles implies that we could use these extracts to identify the key compounds that are responsible for wasp attraction. Restoring and/or enhancing such key parasitoid attractants in cultivated plants could be an effective way to increase natural pest control. [ABSTRACT FROM AUTHOR]

The relationship between the degree of specialization of parasitoids and their responses to host-related volatiles is an important and current evolutionary question. Specialist parasitoids which have evolved to attack fewer host species are predicted to be more responsive to host-related volatiles than generalists. We tested the above prediction by comparing behavioural responses of both sexes of two parasitoids (Hymenoptera: Braconidae) with different degrees of host specificity, Microplitis croceipes (Cresson) (specialist) and Cotesia marginiventris (generalist), to different suites of synthetic host-related volatile compounds. The compounds tested at two doses (1 and 100μg) include two green leaf volatiles (GLVs: hexanal and (Z)-3-hexen-1-ol) and four herbivore-induced plant volatiles (HIPVs: (Z)-3-hexenyl acetate, linalool, (Z)-3-hexenyl butyrate and (E,E)-α-farnesene). Two hypotheses were tested: (i) M. croceipes (specialist) would show relatively greater behavioural responses to the HIPVs, whereas C. marginiventris (generalist) would show greater behavioural responses to the GLVs, and (ii) females of both species would show greater responses than conspecific males to the host-related volatiles. At the low dose (1μg), females of the specialist showed significantly greater responses than females of the generalist to three of the tested HIPVs, (Z)-3-hexenyl acetate, linalool and (Z)-3-hexenyl butyrate. In contrast, females of the generalist showed relatively greater responses to the GLVs. The same trends were recorded at the high dose but fewer significant differences were detected. In general, similar results were recorded for males, with the exception of linalool (an HIPV) which elicited significantly greater response in the generalist than the specialist. Comparing the sexes, females of both species showed greater responses than conspecific males to most of the tested volatiles. The ecological significance of these findings is discussed. [ABSTRACT FROM PUBLISHER]

The ability of parasitoids to learn a wide range of volatiles, including ecologically relevant and novel odors, and respond favorably to the learned stimuli has been documented for various species. Comparison of odor learning in closely related species can elucidate adaptive differences in species responses to infochemicals. The present study used a Y-tube olfactometer to compare odor learning and priming of behavioral responses of two parasitoids, Cotesia marginiventris (Cresson) and Microplitis croceipes (Cresson), to four selected host-related volatile compounds which have been previously shown to elicit behavioral response in both species; trans-2-hexanal, α-pinene, cis-3-hexenyl butyrate, and ( E,E)-α-farnesene. The two parasitoid species differ in their degree of specialization at the host level with M. croceipes having a more specialized host range, but otherwise represent infochemical generalist parasitoid species at the plant level. Naïve females as well as females that had been trained to associate sugar water with the test compounds were tested. Compared to naïve females, trained females of C. marginiventris showed a significant increase in behavioral response to all four tested compounds. In contrast, trained females of M. croceipes showed a significant increase in behavioral response only to α-pinene, and ( E,E)-α-farnesene. Overall, the species with a wider host range, C. marginiventris showed a greater learning-induced increase in response than the species with a relatively specialized host range, M. croceipes. The results imply that animals with different ecological constraints may show variations in learning. The adaptive significance of behavioral responses as related to dietary specializations of the parasitoids is discussed. [ABSTRACT FROM AUTHOR]

Insect herbivores often induce plant volatile compounds that can attract natural enemies. Cotesia marginiventris (Hymenoptera: Braconidae) is a generalist parasitoid wasp of noctuid caterpillars and is highly attracted to Spodoptera exigua-induced plant volatiles. The plasticity of C. marginiventris associative learning to volatile blends of various stimuli, such as host presence, also has been shown, but little is known about how this generalist parasitoid distinguishes between host species of varying suitability. Spodoptera exigua is an excellent host that yields high parasitoid emergence, while Trichoplusia ni serves as a sub-optimal host species due to high pre-imaginal wasp mortality. We have found that S. exigua and T. ni induce different volatile blends while feeding on cotton. Here, wind tunnel flight assays were used to determine the importance of differentially induced volatiles in host-finding by C. marginiventris. We found that, while this generalist parasitoid wasp can distinguish between the two discrete volatile blends when presented concurrently, a positive oviposition experience on the preferred host species ( S. exigua) is more important than host-specific volatile cues in eliciting flight behavior towards plants damaged by either host species. Furthermore, wasps with oviposition experience on both host species did not exhibit a deterioration in positive flight behavior, suggesting that oviposition in the sub-optimal host species ( T. ni) does not cause aversive odor association. [ABSTRACT FROM AUTHOR]

Abstract: Synthetic elicitors can be used to induce resistance in plants against pathogens and arthropod herbivores. Such compounds may also change the emission of herbivore-induced plant volatiles, which serve as important cues for parasitic wasps to locate their hosts. Therefore, the use of elicitors in the field may affect biological control of insect pests. To test this, we treated maize seedlings growing in a subtropical field in Mexico with methyl jasmonate (MeJA), an elicitor of defense responses against many insects, and benzo-(1,2,3)-thiadiazole-7-carbothioic acid S-methyl ester (BTH), an elicitor of resistance against certain pathogens. Volatile emission, herbivore infestation, pathogen infection, and plant performance (growth and grain yield) of treated and untreated maize plants were measured. Application of BTH slightly reduced volatile emission in maize, while MeJA increased the emission compared to control treatments. Despite the apparent changes in volatile emissions, the elicitor application did not consistently affect infestation by Spodoptera frugiperda larvae, the main insect pest found on the maize seedlings, and had only marginal effects on parasitism rates. Similarly, there were no treatment effects on infestation by other herbivores and pathogens. Results for the six replications that stretched over one summer and one winter season were highly variable, with parasitism rates and the species composition of the parasitoids differing significantly between seasons. This variability, as well as the severe biotic and abiotic stresses on young seedlings might explain why we measured only slight effects of elicitor application on pest incidence and biological control in this specific field study. Indeed, an additional field experiment under milder and more standardized conditions revealed that BTH induced significant resistance against Bipolaris maydis, a major pathogen in the experimental maize fields. Similar affects can be expected for herbivory and parasitism rates. [Copyright &y& Elsevier]

The ability to cope with plant defense chemicals differs between specialist and generalist species. In this study, we examined the effects of the concentration of the two main iridoid glycosides (IGs) in Plantago lanceolata, aucubin and catalpol, on the performance of a specialist and two generalist herbivores and their respective endoparasitoids. Development of the specialist herbivore Melitaea cinxia was unaffected by the total leaf IG concentration in its host plant. By contrast, the generalist herbivores Spodoptera exigua and Chrysodeixis chalcites showed delayed larval and pupal development on plant genotypes with high leaf IG concentrations, respectively. This result is in line with the idea that specialist herbivores are better adapted to allelochemicals in host plants on which they are specialized. Melitaea cinxia experienced less post-diapause larval and pupal mortality on its local Finnish P. lanceolata than on Dutch genotypes. This could not be explained by differences in IG profiles, suggesting that M. cinxia has adapted in response to attributes of its local host plants other than to IG chemistry. Development of the specialist parasitoid Cotesia melitaearum was unaffected by IG variation in the diet of its host M. cinxia, a response that was concordant with that of its host. By contrast, the development time responses of the generalist parasitoids Hyposoter didymator and Cotesia marginiventris differed from those of their generalist hosts, S. exigua and C. chalcites. While their hosts developed slowly on high-IG genotypes, development time of H. didymator was unaffected. Cotesia marginiventris actually developed faster on hosts fed high-IG genotypes, although they then had short adult longevity. The faster development of C. marginiventris on hosts that ate high-IG genotypes is in line with the 'immunocompromized host' hypothesis, emphasizing the potential negative effects of toxic allelochemicals on the host's immune response. [ABSTRACT FROM AUTHOR]

Plant volatile compounds induced by herbivore attack have been demonstrated to provide a signal to herbivore enemies such as parasitic wasps that use these volatiles to locate their hosts. However, in addition to herbivore-induced volatiles, plants often release volatiles constitutively. We assessed the interaction between herbivore-induced and constitutively released volatiles of maize in the attraction of the wasp Cotesia marginiventris that parasitizes herbivorous lepidopteran larvae feeding on maize. Experiments were carried out with olfactometers in which the sources of volatiles were transgenic Arabidopsis thaliana plants overexpressing maize sesquiterpene synthases that produce blends of herbivore-induced or constitutive compounds. We found that the constitutive volatiles of maize terpene synthase 8 (TPS8) were attractive to C. marginiventris, just like the herbivore-induced volatiles of TPS10 studied earlier. A mixture of both the TPS8 and TPS10 volatile blends, however, was more effective in parasitoid attraction, indicating that constitutively released sesquiterpenes enhance the attraction of those induced by herbivores. While C. marginiventris did not distinguish among the volatiles of TPS8, TPS10, nor those of another maize sesquiterpene synthase (TPS5), when these blends were combined, their attractiveness to the wasp appeared to increase with the complexity of the blend. [ABSTRACT FROM AUTHOR]

Abstract: In order to test whether the electroantennogram (EAG) response spectrum of an insect correlates to its degree of host specificity, we recorded EAG responses of two parasitoid species with different degrees of host specificity, Microplitis croceipes (specialist) and Cotesia marginiventris (generalist), to a wide array of odor stimuli including compounds representing green leaf volatiles (GLVs), herbivore-induced plant volatiles (HIPV), ecologically irrelevant (not used by the parasitoid species and their hosts for host location) plant volatiles, and host-specific odor stimuli (host sex pheromones, and extracts of host caterpillar body and frass). We also tested the EAG responses of female moths of the caterpillar hosts of the parasitoids, Heliothis virescens and Spodoptera exigua, to some of the odor stimuli. We hypothesized that the specialist parasitoid will have a narrower EAG response spectrum than the generalist, and that the two lepidopteran species, which are similar in their host plant use, will show similar EAG response spectra to plant volatiles. As predicted, the specialist parasitoid showed greater EAG responses than the generalist to host-specific odor and one HIPV (cis-3-hexenyl butyrate), whereas the generalist showed relatively greater EAG responses to the GLVs and unrelated plant volatiles. We detected no differences in the EAG responses of H. virescens and S. exigua to any of the tested odor. [Copyright &y& Elsevier]

Associative learning is known to modify foraging behavior in numerous parasitic wasps. This is in agreement with optimal foraging theory, which predicts that the wasps will adapt their responses to specific cues in accordance with the rewards they receive while perceiving these cues. Indeed, the generalist parasitoid Cotesia marginiventris shows increased attraction to a specific plant odor after perceiving this odor during contact with hosts. This positive associative learning is common among many parasitoids, but little is known about the effects of unrewarding host searching events on the attractiveness of odors. To study this, preferences of female C. marginiventris for herbivore-induced odors of three plant species were tested in a six-arm olfactometer after the wasps perceived one of these odors either i) without contacting any caterpillars, ii) while contacting the host caterpillar Spodoptera littoralis, or iii) while contacting the non-host caterpillar Pieris rapae. The results confirm the effects of positive associative learning, but showed no changes in innate responses to the host-induced odors after “negative” experiences. Hence, a positive association is made during an encounter with hosts, but unsuccessful host-foraging experiences do not necessarily lead to avoidance learning in this generalist parasitoid. [ABSTRACT FROM AUTHOR]

Parasitoids employ different types of host-related volatile signals for foraging and host-location. Host-related volatile signals can be plant-based, originate from the herbivore host or produced from an interaction between herbivores and their plant host. In order to investigate potential sex- and species-related differences in the antennal response of parasitoids to different host-related volatiles, we compared the electroantennogram (EAG) responses of both sexes of the specialist parasitoid, Microplitis croceipes (Cresson), and the generalist, Cotesia marginiventris (Cresson), to varying doses of selected plant-based host-related volatiles: two green leaf volatiles (cis-3-hexenol and hexanal) and three inducible compounds (cis-3-hexenyl acetate, linalool, and (E,E)-??-farnesene). Mating had no significant effect on EAG response. Females of both species showed significantly greater EAG responses than conspecific males to green leaf volatiles, which are released immediately after initiation of herbivore feeding damage. In contrast, males showed greater responses than conspecific females to inducible compounds released much later after initial damage. Cotesia marginiventris females and males showed greater EAG responses than counterpart M. croceipes to the tested compounds at various doses, suggesting that the generalist parasitoid shows greater antennal sensitivity than the specialist to the tested host-plant volatiles. These results are discussed in relation to the possible roles of green leaf volatiles and inducible compounds in the ecology of female and male parasitoids. [ABSTRACT FROM AUTHOR]

Plants counteract attack by herbivorous insects using a variety of inducible defence mechanisms. The production of toxic proteins and metabolites that instantly affect the herbivore's development are examples of direct induced defence. In addition, plants may release mixtures of volatile organic compounds (VOCs) that indirectly protect the plant by attracting natural enemies of the herbivore. Recent studies suggest that these VOCs can also prime nearby plants for enhanced induction of defence upon future insect attack. However, evidence that this defence priming causes reduced vulnerability to insects is sparse. Here we present molecular, chemical and behavioural evidence that VOC-induced priming leads to improved direct and indirect resistance in maize. A differential hybridization screen for inducible genes upon attack by Spodoptera littoralis caterpillars identified 10 defence-related genes that are responsive to wounding, jasmonic acid (JA), or caterpillar regurgitant. Exposure to VOCs from caterpillar-infested plants did not activate these genes directly, but primed a subset of them for earlier and/or stronger induction upon subsequent defence elicitation. This priming for defence-related gene expression correlated with reduced caterpillar feeding and development. Furthermore, exposure to caterpillar-induced VOCs primed for enhanced emissions of aromatic and terpenoid compounds. At the peak of this VOC emission, primed plants were significantly more attractive to parasitic Cotesia marginiventris waSPS. This study shows that VOC-induced priming targets a specific subset of JA-inducible genes, and links these responses at the molecular level to enhanced levels of direct and indirect resistance against insect attack. [ABSTRACT FROM AUTHOR]

After herbivore attack, plants release a plethora of different volatile organic compounds (VOCs), which results in odor blends that are attractive to predators and parasitoids of these herbivores. VOCs in the odor blends emitted by maize plants ( Zea mays) infested by lepidopteran larvae are well characterized. They are derived from at least three different biochemical pathways, but the relative importance of each pathway for the production of VOCs that attract parasitic wasps is unknown. Here, we studied the importance of shikimic acid derived VOCs for the attraction of females of the parasitoids Cotesia marginiventris and Microplitis rufiventris. By incubating caterpillar-infested maize plants in glyphosate, an inhibitor of the 5-enolpyruvylshikimate-3-phospate (EPSP) synthase, we obtained induced odor blends with only minute amounts of shikimic acid derived VOCs. In olfactometer bioassays, the inhibited plants were as attractive to naive C. marginiventris females as control plants that released normal amounts of shikimic acid derived VOCs, whereas naive M. rufiventris females preferred inhibited plants to control plants. By adding back synthetic indole, the quantitatively most important shikimic acid derived VOC in induced maize odors, to inhibited plants, we showed that indole had no effect on the attraction of C. marginiventris and that M. rufiventris preferred blends without synthetic indole. Exposing C. marginiventris females either to odor blends of inhibited or control plants during oviposition experiences shifted their preference in subsequent olfactometer tests in favor of the experienced odor. Further learning experiments with synthetic indole showed that C. marginiventris can learn to respond to this compound, but that this does not affect its choices between natural induced blends with or without indole. We hypothesize that for naïve wasps the attractiveness of an herbivore-induced odor blend is reduced due to masking by nonattractive compounds, and that during oviposition experiences in the presence of complex odor blends, parasitoids strongly associate some compounds, whereas others are largely ignored. [ABSTRACT FROM AUTHOR]

1. Interspecific competition among parasitoids may play a key role in the community dynamics of tritrophic plant–herbivore–parasitoid systems and has important implications for management of herbivorous insect pests. 2. A model system was used to explore the outcome of interspecific competition between parasitoids that differ in host specificity. The system included the lepidopteran pest Heliothis virescens, the generalist parasitoid Cotesia marginiventris, and two specialist parasitoids, Microplitis croceipes and Cardiochiles nigriceps. 3. The generalist, C. marginiventris, dominated intrinsic competition when given an 8-h developmental head start over C. nigriceps or when its oviposition was simultaneous with that of M. croceipes. Microplitis croceipes and especially C. nigriceps larvae prevailed when they were allowed to oviposit prior to C. marginiventris. 4. Rates of host mortality prior to parasitoid emergence varied with parasitoid species composition and with the order of oviposition. 5. Implications for integrated pest management and the adaptive significance of competition as related to host specialisation are discussed. [ABSTRACT FROM AUTHOR]

Abstract: A recombinant baculovirus insecticide, AcMLF9.ScathL, has been constructed that expresses a cathepsin L-like protease. This protease degrades the basement membrane of infected-lepidopteran larvae. AcMLF9.ScathL kills larvae significantly faster than the wild type baculovirus. Before field trials can be conducted with this virus, risk assessment studies are required to examine the potential impact of the virus on nontarget organisms. We examined the impact of AcMLF9.ScathL on the survival, development, and behavior of the parasitoid Cotesia marginiventris (Cresson) that parasitizes infected second instar larvae. Larvae of Heliothis virescens were exposed to the parasitoid and infected with the wild type or recombinant virus at >LD99 at 72, 96, or 120h after parasitism. Control larvae were mock infected. Choice tests showed that infection with AcMLF9.ScathL did not affect the host preference of the parasitoid. At 72h post-parasitism, the survival of parasitoids emerging from hosts infected with AcMLF9.ScathL was lower than for parasitoids emerging from hosts infected with wild type virus or uninfected controls. There were no significant differences between AcMLF9.ScathL and wild type virus treatments for the larval emergence time, adult emergence time, sex ratio, size, and fecundity of C. marginiventris. These results indicate that use of AcMLF9.ScathL in insect pest management poses a slightly greater risk to the parasitoid at early stages of parasitoid development than use of the wild type virus at >LD99 dose. [Copyright &y& Elsevier]

Interactions between host plant resistance and biological control may benefit or hinder pest management efforts. Turfgrass cultivars have rarely been tested for extrinsic resistance characteristics such as occurrence and performance of beneficial arthropods on plant genotypes with resistance to known turf pests. Parasitism of fall armyworm, Spodoptera frugiperda (J.E. Smith), among six turfgrass genotypes was evaluated. The six grasses tested [Sea Isle-1 and 561-79 seashore paspalum, Paspalum vaginatum Swartz; TifSport and TifEagle hybrid Bermuda grass, Cynodon dactylon (L.) x C. transvaalensis (Burtt-Davy); and Cavalier and Palisades zoysiagrass, Zoysia japonica von Steudel and Z. Marcello (L.) Merrill, respectively] represented a range in resistance to S. frugiperda. Differential recovery of larvae released as first instars reflected this gradient in resistance of Cavalier ≥ Palisades ≥ TifSport = TifEagle ≥ 561- = Sea Isle-1 Larval recovery (percentage of initial number released) was greatest in May, less in July and August, and least in October, probably reflecting the increase in activity of on-site predators and disease pressure. Parasitism of the fall armyworm by the braconid Aleiodes laphygmae Viereck varied among turfgrass genotypes. Parasitism was greatest during July. In total, 20,400 first instars were placed in the field; 2,368 were recovered; 468 parasitoids were subsequently reared; 92.2% were A. laphygmae. In the field, the greatest percentage of reduction in S. frugiperda larvae by A. laphygmae occurred on the armyworm-susceptible seashore paspalums (51.9% on Sea Isle-1 in July). Cotesia marginiventris Cresson and Meteorus sp. also were reared from collected larvae. No parasitoids were reared from larvae collected from resistant Cavalier zoysiagrass. A. laphygmae and C. marginiventris were reared from larvae collected from the other five grass cultivars. No parasitoids of older larvae or pupae were observed. [ABSTRACT FROM AUTHOR]

Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed. [ABSTRACT FROM AUTHOR]

Abstract In natural populations, plants demonstrate an array of indirect and direct defence strategies that help to protect them from their herbivores and pathogens. Indirect defences include the release of odours that attract the natural enemies of herbivores, whereas direct defences may include the production of secondary compounds, allelochemicals that impair herbivore development or repel herbivore attack. Although both strategies have been well studied independently, comparatively little attention has been paid to examining the conflict that may arise between indirect and direct defences, such as when the performance of ‘recruited’ parasitoids or predators is negatively affected by plant allelochemicals. Here, we examine the growth and development of polyphagous and oligophagous folivores and their respective endoparasitoids on three crucifer species. One of the species, Brassica oleracea , was recently cultivated, whereas populations of B. nigra and Barbarea vulgaris occur naturally. Additionally, these species possess contrasting life-history patterns and are also known to exhibit differences in secondary chemistry. The development of the generalist herbivore–parasitoid system was much more variable over the three crucifers than that exhibited by the specialists. Moreover, generalist herbivore and/or parasitoid fitness-related traits (survival, development time, pupal, or adult size) were much more negatively affected on the wild crucifers than in the specialist association. Our results suggest that the relative importance of direct and indirect defences in plants may rest on the degree of dietary specialisation exhibited by herbivores and their natural enemies, and on the level of toxicity in the plant species under investigation. [ABSTRACT FROM AUTHOR]

We examined the effect of Bt-cotton (Event 531) plants expressing the Bacillus thuringiensis δ-endotoxin CryIA(c) on two hymenopteran endoparasitoids, Cotesia marginiventris and Copidosoma floridanum. In the laboratory, parasitized and unparasitized Pseudoplusia includens larvae were reared on foliage from a conventional soybean cultivar (Pioneer 97B61), a conventional cotton cultivar (DPL 5415), or a Bt-cotton cultivar (NuCotn 33B). C. marginiventris developed significantly faster within P. includens larvae feeding on Pioneer 97B61 and DPL 5415 compared to those feeding on NuCotn 33B. C. marginiventris that developed inside P. includens larvae feeding on NuCotn 33B suffered reduced longevity, and females had fewer ova. NuCotn 33B also affected the growth and development of P. includens parasitized with C. floridanum and life history parameters of adult C. floridanum. Parasitized and unparasitized P. includens developed more slowly when they were fed NuCotn 33B and the prepupae weighed less. Survival of parasitized and unparasitized P. includens was lower when larvae were fed NuCotn 33B and some evidence points to higher susceptibility of parasitized caterpillars to intoxication by NuCotn 33B. Fewer C. floridanum adults emerged from hosts fed NuCotn 33B, but pupal weight and adult longevity were unaffected. Analysis comparing the two experiments conducted with C. floridanum suggests that older NuCotn 33B plants (90–120 days after planting) may affect parasitoid development and adult survival less than younger NuCotn 33B plants (60–90 days after planting). Feeding on NuCotn 33B by P. includens affected the survival and development of the two hymenopteran endoparasitoids studied here, and the degree of the effect was similar to that observed with natural resistance found in soybean plants. It remains to be determined if the effects demonstrated here are less than, equal to, or greater than the impact of conventional insecticide applications used in conventional, non-transgenic, cotton. [Copyright &y& Elsevier]

Herbivore-induced plant volatiles can function as indirect defense signals that attract natural enemies of herbivores. Several parasitoids are known to exploit these plant-provided cues to locate their hosts. One such parasitoid is the generalist Cotesia marginiventris, which is, among others, attracted to maize volatiles induced by caterpillar damage. Maize plants can be induced to produce the same blend of attractive volatiles by treating them with regurgitant of Spodoptera species. We collected and analyzed the regurgitant-induced emissions of two plant species (cowpea and maize) and of eight Mexican maize varieties and found significant differences among their volatile emissions, both in terms of total quantity and the quality of the blends. In a Y-tube olfactometer, the odors of the same artificially induced plant species and Mexican varieties were offered in dual choice experiments to naïve mated females of C. marginiventris. Wasps preferred cowpea over maize odor and, in 3 of 12 combinations with the maize varieties, they showed a preference for the odors of one of the varieties. A comparison of the odor collection with results from the behavioral assays indicates that not only the quantity of the volatile emissions, but also the quality composition of the volatile blends is important for attraction of C. marginiventris. The results are discussed in the context of the possibility of breeding crop varieties that are particularly attractive to parasitoids. [ABSTRACT FROM AUTHOR]

Cotton plants under herbivore attack release volatile semiochemicals that attract natural enemies of the herbivores to the damaged plant. The volatiles released in response to herbivory are not only released from the damaged leaves but from the entire cotton plant. We found that cotton plants that released myrcene, ( Z)-3-hexenyl acetate, ( E)-β-ocimene, linalool, ( E)-4,8-dimethyl-1,3,7-nonatriene, ( E)-β-farnesene, and ( E, E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene systemically from undamaged leaves of caterpillar damaged plants were attractive to the generalist parasitoid Cotesia marginiventris and the specialist parasitoid Microplitis croceipes. Plants from which the caterpillar damaged leaves were removed and that released those compounds systemically were significantly preferred over undamaged control plants in two-choice experiments in a flight tunnel. Artificially damaged cotton plants that released green leafy volatiles and constitutive terpenoids were less attractive for M. croceipes and C. marginiventris. Only C. marginiventris preferred artificially damaged plants over undamaged control plants, whereas M. croceipes showed no preference. The apparent lack of specificity of systemically released compounds in response to different herbivores feeding on the lower leaves is discussed. [ABSTRACT FROM AUTHOR]

In theory, the degree of specificity of the signals a parasitoid species needs to successfully locate its host correlates with its level of specialization. We examined this question by comparing the foraging strategies of two parasitoids that differ in their host ranges. In wind-tunnel experiments, we investigated how systemically released herbivore-induced volatiles were used by the generalist parasitoid, Cotesia marginiventris (Cresson) and the specialist, Microplitis croceipes (Cresson). We determined the relative influence of these volatiles as compared to other signals emitted in the host orientation of the two parasitoids. Both the generalist and the specialist parasitoid strongly preferred Spodoptera exigua (Hübner) leaf-induced systemic plants over undamaged plants when no other information was available. When wasps were given a choice between leaf-induced and undamaged plants carrying other plant- or host-related materials, the responses differed for the two species. C. marginiventris appeared to cue primarily on recent damage volatiles, whereas M. croceipes appeared to cue primarily on host frass volatiles. However, recent damage on previously leaf-induced plants, was strongly preferred to recent damage on plants previously damaged by both species. When plants were induced at the squares by Helicoverpa zea (Boddie), only M. croceipes exhibited a preference for these plants over undamaged plants. The adaptive significance of the behaviors as related to dietary specializations of the parasitoids is discussed. [ABSTRACT FROM AUTHOR]