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1. Several studies have shown that above- and belowground insects can interact by influencing each others growth, development, and survival when they feed on the same host-plant. In natural systems, however, insects can make choices on which plants to oviposit and feed. A field experiment was carried out to determine if root-feeding insects can influence feeding and oviposition preferences and decisions of naturally colonising foliar-feeding insects. 2. Using the wild cruciferous plant Brassica nigra and larvae of the cabbage root fly Delia radicum as the belowground root-feeding insect, naturally colonising populations of foliar-feeding insects were monitored over the course of a summer season. 3. Groups of root-infested and root-uninfested B. nigra plants were placed in a meadow during June, July, and August of 2006 for periods of 3 days. The root-infested and the root-uninfested plants were either dispersed evenly or placed in clusters. Once daily, all leaves of each plant were carefully inspected and insects were removed and collected for identification. 4. The flea beetles Phyllotreta spp. and the aphid Brevicoryne brassicae were significantly more abundant on root-uninfested (control) than on root-infested plants. However, for B. brassicae this was only apparent when the plants were placed in clusters. Host-plant selection by the generalist aphid M. persicae and oviposition preference by the specialist butterfly P. rapae, however, were not significantly influenced by root herbivory. 5. The results of this study show that the presence of root-feeding insects can affect feeding and oviposition preferences of foliar-feeding insects, even under natural conditions where many other interactions occur simultaneously. The results suggest that root-feeding insects play a role in the structuring of aboveground communities of insects, but these effects depend on the insect species as well as on the spatial distribution of the root-feeding insects., 1. Several studies have shown that above- and belowground insects can interact by influencing each others growth, development, and survival when they feed on the same host-plant. In natural systems, however, insects can make choices on which plants to oviposit and feed. A field experiment was carried out to determine if root-feeding insects can influence feeding and oviposition preferences and decisions of naturally colonising foliar-feeding insects. 2. Using the wild cruciferous plant Brassica nigra and larvae of the cabbage root fly Delia radicum as the belowground root-feeding insect, naturally colonising populations of foliar-feeding insects were monitored over the course of a summer season. 3. Groups of root-infested and root-uninfested B. nigra plants were placed in a meadow during June, July, and August of 2006 for periods of 3 days. The root-infested and the root-uninfested plants were either dispersed evenly or placed in clusters. Once daily, all leaves of each plant were carefully inspected and insects were removed and collected for identification. 4. The flea beetles Phyllotreta spp. and the aphid Brevicoryne brassicae were significantly more abundant on root-uninfested (control) than on root-infested plants. However, for B. brassicae this was only apparent when the plants were placed in clusters. Host-plant selection by the generalist aphid M. persicae and oviposition preference by the specialist butterfly P. rapae, however, were not significantly influenced by root herbivory. 5. The results of this study show that the presence of root-feeding insects can affect feeding and oviposition preferences of foliar-feeding insects, even under natural conditions where many other interactions occur simultaneously. The results suggest that root-feeding insects play a role in the structuring of aboveground communities of insects, but these effects depend on the insect species as well as on the spatial distribution of the root-feeding insects.

In natural systems, pre-adult stages of some insect herbivores are known to be attacked by several species of parasitoids. Under certain conditions, hosts may be simultaneously parasitized by more than one parasitoid species (= multiparasitism), even though only one parasitoid species can successfully develop in an individual host. Here, we compared development, survival, and intrinsic competitive interactions among three species of solitary larval endoparasitoids, Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae), Microplitis demolitor Wilkinson, and Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), in singly parasitized and multiparasitized hosts. The three species differed in certain traits, such as in host usage strategies and adult body size. Campoletis sonorensis and M. demolitor survived equally well to eclosion in two host species that differed profoundly in size, Pseudoplusia includens (Walker) and the larger Heliothis virescens (Fabricius) (both Lepidoptera: Noctuidae). Egg-to-adult development time in C. sonorensis and M. demolitor also differed in the two hosts. Moreover, adult body mass in C. sonorensis (and not M. demolitor) was greater when developing in H. virescens larvae. We then monitored the outcome of competitive interactions in host larvae that were parasitized by one parasitoid species and subsequently multiparasitized by another species at various time intervals (0, 6, 24, and 48 h) after the initial parasitism. These experiments revealed that M. croceipes was generally a superior competitor to the other two species, whereas M. demolitor was the poorest competitor, with C. sonorensis being intermediate in this capacity. However, competition sometimes incurred fitness costs in M. croceipes and C. sonorensis, with longer development time and/or smaller adult mass observed in surviving wasps emerging from multiparasitized hosts. Our results suggest that rapid growth and large size relative to competitors of a similar age may, In natural systems, pre-adult stages of some insect herbivores are known to be attacked by several species of parasitoids. Under certain conditions, hosts may be simultaneously parasitized by more than one parasitoid species (= multiparasitism), even though only one parasitoid species can successfully develop in an individual host. Here, we compared development, survival, and intrinsic competitive interactions among three species of solitary larval endoparasitoids, Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae), Microplitis demolitor Wilkinson, and Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), in singly parasitized and multiparasitized hosts. The three species differed in certain traits, such as in host usage strategies and adult body size. Campoletis sonorensis and M. demolitor survived equally well to eclosion in two host species that differed profoundly in size, Pseudoplusia includens (Walker) and the larger Heliothis virescens (Fabricius) (both Lepidoptera: Noctuidae). Egg-to-adult development time in C. sonorensis and M. demolitor also differed in the two hosts. Moreover, adult body mass in C. sonorensis (and not M. demolitor) was greater when developing in H. virescens larvae. We then monitored the outcome of competitive interactions in host larvae that were parasitized by one parasitoid species and subsequently multiparasitized by another species at various time intervals (0, 6, 24, and 48 h) after the initial parasitism. These experiments revealed that M. croceipes was generally a superior competitor to the other two species, whereas M. demolitor was the poorest competitor, with C. sonorensis being intermediate in this capacity. However, competition sometimes incurred fitness costs in M. croceipes and C. sonorensis, with longer development time and/or smaller adult mass observed in surviving wasps emerging from multiparasitized hosts. Our results suggest that rapid growth and large size relative to competitors of a similar age may

1. Parasitoid wasps are highly efficient organisms at utilizing and assimilating limited resources from their hosts. This study explores interactions over three trophic levels, from the third (primary parasitoid) to the fourth (secondary parasitoid) and terminating in the fifth (tertiary parasitoid). 2. Host utilization and adult body mass of the secondary and tertiary parasitoid Gelis agilis was determined when developing on pre-pupae of its primary parasitoid host, Cotesia glomerata, and from pre-pupae of another secondary parasitoid, Lysibia nana that had developed initially on pre-pupae of C. glomerata. 3. In both C. glomerata and G. agilis, the body mass of emerging adult parasitoids was strongly positively correlated with initial cocoon mass. For a given cocoon mass at parasitism, emerging adult G. agilis wasps were almost 90% as large as C. glomerata adults developing in healthy cocoons of comparable mass. Furthermore, G. agilis adults were still 75% as large as C. glomerata adults even when developing on L. nana that in turn had developed on C. glomerata. Otherwise, in terms of adult body mass 4. Analyses of carbon and nitrogen in body tissues of the parasitoids over the third to the fifth trophic level revealed that percentage nitrogen was higher and carbon lower in G. agilis and L. nana than in C. glomerata. Furthermore, percentage carbon was lower in adult G. agilis wasps that had developed from L. nana than from C. glomerata. 5. We argue that the remarkable efficiency which characterizes the development of secondary and tertiary parasitoids is based on the very high nutritional quality of resources that increase in quality up the food chain and rigid selection optimizing allocation of limited host resources. Consequently, food webs involving secondary and tertiary parasitoids can go to levels hitherto unexplored thus far in empirical studies. The use of molecular markers in future studies may reveal just how long food c, 1. Parasitoid wasps are highly efficient organisms at utilizing and assimilating limited resources from their hosts. This study explores interactions over three trophic levels, from the third (primary parasitoid) to the fourth (secondary parasitoid) and terminating in the fifth (tertiary parasitoid). 2. Host utilization and adult body mass of the secondary and tertiary parasitoid Gelis agilis was determined when developing on pre-pupae of its primary parasitoid host, Cotesia glomerata, and from pre-pupae of another secondary parasitoid, Lysibia nana that had developed initially on pre-pupae of C. glomerata. 3. In both C. glomerata and G. agilis, the body mass of emerging adult parasitoids was strongly positively correlated with initial cocoon mass. For a given cocoon mass at parasitism, emerging adult G. agilis wasps were almost 90% as large as C. glomerata adults developing in healthy cocoons of comparable mass. Furthermore, G. agilis adults were still 75% as large as C. glomerata adults even when developing on L. nana that in turn had developed on C. glomerata. Otherwise, in terms of adult body mass 4. Analyses of carbon and nitrogen in body tissues of the parasitoids over the third to the fifth trophic level revealed that percentage nitrogen was higher and carbon lower in G. agilis and L. nana than in C. glomerata. Furthermore, percentage carbon was lower in adult G. agilis wasps that had developed from L. nana than from C. glomerata. 5. We argue that the remarkable efficiency which characterizes the development of secondary and tertiary parasitoids is based on the very high nutritional quality of resources that increase in quality up the food chain and rigid selection optimizing allocation of limited host resources. Consequently, food webs involving secondary and tertiary parasitoids can go to levels hitherto unexplored thus far in empirical studies. The use of molecular markers in future studies may reveal just how long food c

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In most animals, the optimal phenotype is determined by trade-offs in life-history traits. Here, I compare development and reproductive strategies in two species of solitary secondary hyperparasitoids, Lysibia nana and Gelis agilis, attacking pre-pupae of their primary parasitoid host, Cotesia glomerata. Parasitoid larvae of both species exploit a given amount of host resources with similar efficiency. However, adults exhibit quite different reproductive strategies. Both species are synovigenic, and female wasps emerge with no mature eggs. However, G. agilis must first host-feed to produce eggs, while L. nana does not host-feed but mobilizes internal resources carried over from larval feeding to initiate oogenesis. Further, G. agilis is wingless, produces large eggs, has a long life-span, and generates only small numbers of progeny per day, whereas these traits are reversed in L. nana. Given unlimited hosts, the fecundity curve in L. nana was “front-loaded,” whereas in G. agilis it was depressed and extended over much of adult life. In L. nana (but not G. agilis), wasps provided with honey but no hosts lived significantly longer than wasps provided with both honey and hosts. Differences in the fecundity curves of the two hyperparasitoids are probably based on differing costs of reproduction between them, with the wingless G. agilis much more constrained in finding hosts than the winged L. nana. Importantly, L. nana is known to be a specialist hyperparasitoid of gregarious Cotesia species that pupate in exposed locations on the food plant, whereas Gelis sp. attack and develop in divergent hosts such as parasitoid cocoons, moth pupae and spider egg sacs. Consequently, there is a strong match between brood size in C. glomerata and egg production in L. nana, but a mismatch between these parameters in G. agilis., In most animals, the optimal phenotype is determined by trade-offs in life-history traits. Here, I compare development and reproductive strategies in two species of solitary secondary hyperparasitoids, Lysibia nana and Gelis agilis, attacking pre-pupae of their primary parasitoid host, Cotesia glomerata. Parasitoid larvae of both species exploit a given amount of host resources with similar efficiency. However, adults exhibit quite different reproductive strategies. Both species are synovigenic, and female wasps emerge with no mature eggs. However, G. agilis must first host-feed to produce eggs, while L. nana does not host-feed but mobilizes internal resources carried over from larval feeding to initiate oogenesis. Further, G. agilis is wingless, produces large eggs, has a long life-span, and generates only small numbers of progeny per day, whereas these traits are reversed in L. nana. Given unlimited hosts, the fecundity curve in L. nana was “front-loaded,” whereas in G. agilis it was depressed and extended over much of adult life. In L. nana (but not G. agilis), wasps provided with honey but no hosts lived significantly longer than wasps provided with both honey and hosts. Differences in the fecundity curves of the two hyperparasitoids are probably based on differing costs of reproduction between them, with the wingless G. agilis much more constrained in finding hosts than the winged L. nana. Importantly, L. nana is known to be a specialist hyperparasitoid of gregarious Cotesia species that pupate in exposed locations on the food plant, whereas Gelis sp. attack and develop in divergent hosts such as parasitoid cocoons, moth pupae and spider egg sacs. Consequently, there is a strong match between brood size in C. glomerata and egg production in L. nana, but a mismatch between these parameters in G. agilis.

Caterpillars that are attacked by some species of parasitoid wasps are known to survive for several days after the parasitoid larvae emerge and pupate. It has been argued that the behaviour of the parasitized larva is ‘usurped’ by the parasitoid and that it ‘guards’ the parasitoid cocoons against their own natural enemies such as hyperparasitoids (the ‘usurpation hypothesis'). We tested this hypothesis in the association involving a gregarious endoparasitoid, the wasp Cotesia glomerata; caterpillars of its host, the large cabbage white butterfly Pieris brassicae; and a pupal hyperparasitoid, the wasp Lysibia nana. In laboratory experiments, we presented cocoon broods of C. glomerata to single females of L. nana in arenas for 6 h. We tested several treatments for rates of primary parasitoid survival, including variation in the position of the caterpillar and the presence or absence of an additional silk web spun by parasitized caterpillars. Parasitized P. brassicae larvae survived longer than the period necessary for C. glomerata adults to emerge. Rates of parasitoid survival were, however, unaffected by the presence of a P. brassicae larva on the cocoon brood, although significantly more parasitoids emerged when the silk web was present. Analyses of the foraging behaviour of individual L. nana females in arenas, performed using Observer software, revealed that the wasps showed a greater tendency to leave cocoons when caterpillars and silk were present. The laboratory experiments only partially support the usurpation hypothesis. In nature, usurpation of the host of the primary parasitoid may be a more effective strategy against generalist predators than against more specialized and better-adapted hyperparasitoids., Caterpillars that are attacked by some species of parasitoid wasps are known to survive for several days after the parasitoid larvae emerge and pupate. It has been argued that the behaviour of the parasitized larva is ‘usurped’ by the parasitoid and that it ‘guards’ the parasitoid cocoons against their own natural enemies such as hyperparasitoids (the ‘usurpation hypothesis'). We tested this hypothesis in the association involving a gregarious endoparasitoid, the wasp Cotesia glomerata; caterpillars of its host, the large cabbage white butterfly Pieris brassicae; and a pupal hyperparasitoid, the wasp Lysibia nana. In laboratory experiments, we presented cocoon broods of C. glomerata to single females of L. nana in arenas for 6 h. We tested several treatments for rates of primary parasitoid survival, including variation in the position of the caterpillar and the presence or absence of an additional silk web spun by parasitized caterpillars. Parasitized P. brassicae larvae survived longer than the period necessary for C. glomerata adults to emerge. Rates of parasitoid survival were, however, unaffected by the presence of a P. brassicae larva on the cocoon brood, although significantly more parasitoids emerged when the silk web was present. Analyses of the foraging behaviour of individual L. nana females in arenas, performed using Observer software, revealed that the wasps showed a greater tendency to leave cocoons when caterpillars and silk were present. The laboratory experiments only partially support the usurpation hypothesis. In nature, usurpation of the host of the primary parasitoid may be a more effective strategy against generalist predators than against more specialized and better-adapted hyperparasitoids.

Theoretical models predict that brood guarding may evolve in situations where eggs are costly to produce or when handling times are long. This study reveals that females of the secondary hyperparasitoid Trichomalopsis apanteloctena guarded cocoon broods of Cotesia kariyai, a gregarious endoparasitoid. Hyperparasitoid females also monopolized host resources and protected their offspring by driving away other conspecific hyperparasitoid females. The females exhibited antagonistic behavior towards competitors through threatening body postures, biting and chasing. Using a video camera to determine how long a hyperparasitoid female attended and parasitized cocoons within a single host brood, it was found that after about 4 days, cocoon guarding behavior became much less apparent. Moreover, more than 90% of hosts were typically parasitized by a hyperparasitoid female over the course of 4 days after she commenced brood guarding. Observations of egg production during a female’s lifetime revealed a physiological interval rhythm that typically lasted 3–4 days, which correlates almost exactly with the period during which the cocoons were guarded. To confirm the giving-up time for a host cocoon brood, hyperparasitoid females were given access to 24 h-old cocoon clusters, each containing 60–100 individual cocoons. Ninety percent of the females remained on cocoons for approximately 72 h. Furthermore, twenty-five percent of wasps continued attending and presumably guarding host cocoon broods for more than 138 h after the female first attended the brood. C. kariyai larvae pupate within a few hours of egression from their host and emerge as adults about 5 days (120 h) later. Therefore, many hyperparasitoid females continued to guard older host cocoons of greatly reduced quality as a resource for their progeny and some even after eclosion of the primary parasitoid. Late-brood guarding enabled a hyperparasitoid female to protect her own progeny from other hyperparasitoid females that, Theoretical models predict that brood guarding may evolve in situations where eggs are costly to produce or when handling times are long. This study reveals that females of the secondary hyperparasitoid Trichomalopsis apanteloctena guarded cocoon broods of Cotesia kariyai, a gregarious endoparasitoid. Hyperparasitoid females also monopolized host resources and protected their offspring by driving away other conspecific hyperparasitoid females. The females exhibited antagonistic behavior towards competitors through threatening body postures, biting and chasing. Using a video camera to determine how long a hyperparasitoid female attended and parasitized cocoons within a single host brood, it was found that after about 4 days, cocoon guarding behavior became much less apparent. Moreover, more than 90% of hosts were typically parasitized by a hyperparasitoid female over the course of 4 days after she commenced brood guarding. Observations of egg production during a female’s lifetime revealed a physiological interval rhythm that typically lasted 3–4 days, which correlates almost exactly with the period during which the cocoons were guarded. To confirm the giving-up time for a host cocoon brood, hyperparasitoid females were given access to 24 h-old cocoon clusters, each containing 60–100 individual cocoons. Ninety percent of the females remained on cocoons for approximately 72 h. Furthermore, twenty-five percent of wasps continued attending and presumably guarding host cocoon broods for more than 138 h after the female first attended the brood. C. kariyai larvae pupate within a few hours of egression from their host and emerge as adults about 5 days (120 h) later. Therefore, many hyperparasitoid females continued to guard older host cocoons of greatly reduced quality as a resource for their progeny and some even after eclosion of the primary parasitoid. Late-brood guarding enabled a hyperparasitoid female to protect her own progeny from other hyperparasitoid females that

The oviposition choice of an insect herbivore is based on a complex set of stimuli and responses. In this study, we examined the effect of plant secondary chemistry (the iridoid glycosides aucubin and catalpol) and aspects of size of the plant Plantago lanceolata, on the oviposition behavior of the specialist butterfly Melitaea cinxia. Iridoid glycosides are known to deter feeding or decrease the growth rate of generalist insect herbivores, but can act as oviposition cues and feeding stimulants for specialized herbivores. In a previous observational study of M. cinxia in the field, oviposition was associated with high levels of aucubin. However, this association could have been the cause (butterfly choice) or consequence (plant induction) of oviposition. We conducted a set of dual- and multiple-choice experiments in cages and in the field. In the cages, we found a positive association between the pre-oviposition level of aucubin and the number of ovipositions. The association reflects the butterfly oviposition selection rather than plant induction that follows oviposition. Our results also suggest a threshold concentration below which females do not distinguish between levels of iridoid glycosides. In the field, the size of the plant appeared to be a more important stimulus than iridoid glycoside content, with bigger plants receiving more oviposition than smaller plants, regardless of their secondary chemistry. Our results illustrate that the rank of a cue used for oviposition may be dependent on environmental context, The oviposition choice of an insect herbivore is based on a complex set of stimuli and responses. In this study, we examined the effect of plant secondary chemistry (the iridoid glycosides aucubin and catalpol) and aspects of size of the plant Plantago lanceolata, on the oviposition behavior of the specialist butterfly Melitaea cinxia. Iridoid glycosides are known to deter feeding or decrease the growth rate of generalist insect herbivores, but can act as oviposition cues and feeding stimulants for specialized herbivores. In a previous observational study of M. cinxia in the field, oviposition was associated with high levels of aucubin. However, this association could have been the cause (butterfly choice) or consequence (plant induction) of oviposition. We conducted a set of dual- and multiple-choice experiments in cages and in the field. In the cages, we found a positive association between the pre-oviposition level of aucubin and the number of ovipositions. The association reflects the butterfly oviposition selection rather than plant induction that follows oviposition. Our results also suggest a threshold concentration below which females do not distinguish between levels of iridoid glycosides. In the field, the size of the plant appeared to be a more important stimulus than iridoid glycoside content, with bigger plants receiving more oviposition than smaller plants, regardless of their secondary chemistry. Our results illustrate that the rank of a cue used for oviposition may be dependent on environmental context

Plants attacked by herbivorous insects emit volatile compounds that attract predators or parasitoids of the herbivores. Plant fitness increases when these herbivorous insects are parasitized by solitary parasitoids, but whether gregarious koinobiont parasitoids also confer a benefit to plant fitness has been disputed. We investigated the relationship between parasitoid load of the gregarious Cotesia glomerata (L.) (Hymenoptera: Braconidae), food consumption by larvae of their host Pieris brassicae L. (Lepidoptera: Pieridae), and seed production in a host plant, Brassica nigra L. (Brassicaceae), in a greenhouse experiment. Plants damaged by caterpillars containing single parasitoid broods produced a similar amount of seeds as undamaged control plants and produced significantly more seeds than plants with unparasitized caterpillars feeding on them. Increasing the parasitoid load to levels likely resulting from superparasitization, feeding by parasitized caterpillars was significantly negatively correlated with plant seed production. Higher parasitoid brood sizes were negatively correlated with pupal weight of Cotesia glomerata, revealing scramble competition leading to a fitness trade-off for the parasitoid. Our results suggest that in this tritrophic system plant fitness is higher when the gregarious parasitoid deposits a single brood into its herbivorous host. A prediction following from these results is that plants benefit from recruiting parasitoids when superparasitization is prevented. This is supported by our previous results on down-regulation of synomone production when Brassica oleracea was fed on by parasitized caterpillars of P. brassicae. We conclude that variable parasitoid loads in gregarious koinobiont parasitoids largely explain existing controversies about the putative benefit of recruiting these parasitoids for plant reproduction., Plants attacked by herbivorous insects emit volatile compounds that attract predators or parasitoids of the herbivores. Plant fitness increases when these herbivorous insects are parasitized by solitary parasitoids, but whether gregarious koinobiont parasitoids also confer a benefit to plant fitness has been disputed. We investigated the relationship between parasitoid load of the gregarious Cotesia glomerata (L.) (Hymenoptera: Braconidae), food consumption by larvae of their host Pieris brassicae L. (Lepidoptera: Pieridae), and seed production in a host plant, Brassica nigra L. (Brassicaceae), in a greenhouse experiment. Plants damaged by caterpillars containing single parasitoid broods produced a similar amount of seeds as undamaged control plants and produced significantly more seeds than plants with unparasitized caterpillars feeding on them. Increasing the parasitoid load to levels likely resulting from superparasitization, feeding by parasitized caterpillars was significantly negatively correlated with plant seed production. Higher parasitoid brood sizes were negatively correlated with pupal weight of Cotesia glomerata, revealing scramble competition leading to a fitness trade-off for the parasitoid. Our results suggest that in this tritrophic system plant fitness is higher when the gregarious parasitoid deposits a single brood into its herbivorous host. A prediction following from these results is that plants benefit from recruiting parasitoids when superparasitization is prevented. This is supported by our previous results on down-regulation of synomone production when Brassica oleracea was fed on by parasitized caterpillars of P. brassicae. We conclude that variable parasitoid loads in gregarious koinobiont parasitoids largely explain existing controversies about the putative benefit of recruiting these parasitoids for plant reproduction.

Parasitoids display remarkable inter- and intraspecific variation in their reproductive and associated traits. Adaptive explanations have been proposed for many of the between-trait relationships. We present an overview of the current knowledge of parasitoid reproductive biology, focusing on egg production strategies in females, by placing parasitoid reproduction within physiological and ecological contexts. Thus, we relate parasitoid reproduction both to inter- and intraspecific patterns of nutrient allocation, utilization, and acquisition, and to key aspects of host ecology, specifically abundance and dispersion pattern. We review the evidence that resource trade-offs underlie several key intertrait correlations and that reproductive and feeding strategies are closely integrated at both the physiological and the behavioral levels. The idea that parasitoids can be divided into capital-breeders or income-breeders is no longer tenable; such terminology is best restricted to the females’ utilization of particular nutrients., Parasitoids display remarkable inter- and intraspecific variation in their reproductive and associated traits. Adaptive explanations have been proposed for many of the between-trait relationships. We present an overview of the current knowledge of parasitoid reproductive biology, focusing on egg production strategies in females, by placing parasitoid reproduction within physiological and ecological contexts. Thus, we relate parasitoid reproduction both to inter- and intraspecific patterns of nutrient allocation, utilization, and acquisition, and to key aspects of host ecology, specifically abundance and dispersion pattern. We review the evidence that resource trade-offs underlie several key intertrait correlations and that reproductive and feeding strategies are closely integrated at both the physiological and the behavioral levels. The idea that parasitoids can be divided into capital-breeders or income-breeders is no longer tenable; such terminology is best restricted to the females’ utilization of particular nutrients.

can act as vertical communication channels or ‘green phones’ linking soil-dwelling insects and insects in the aboveground ecosystem. When root-feeding insects attack a plant, the direct defense system of the shoot is activated, leading to an accumulation of phytotoxins in the leaves. The protection of the plant shoot elicited by root damage can impair the survival, growth and development of aboveground insect herbivores, thereby creating plant-based functional links between soil-dwelling insects and insects that develop in the aboveground ecosystem. The interactions between spatially separated insects below- and aboveground are not restricted to root and foliar plant-feeding insects, but can be extended to higher trophic levels such as insect parasitoids. Here we discuss some implications of plants acting as communication channels or ‘green phones’ between root and foliar-feeding insects and their parasitoids, focusing on recent findings that plants attacked by root-feeding insects are significantly less attractive for the parasitoids of foliar-feeding insects., can act as vertical communication channels or ‘green phones’ linking soil-dwelling insects and insects in the aboveground ecosystem. When root-feeding insects attack a plant, the direct defense system of the shoot is activated, leading to an accumulation of phytotoxins in the leaves. The protection of the plant shoot elicited by root damage can impair the survival, growth and development of aboveground insect herbivores, thereby creating plant-based functional links between soil-dwelling insects and insects that develop in the aboveground ecosystem. The interactions between spatially separated insects below- and aboveground are not restricted to root and foliar plant-feeding insects, but can be extended to higher trophic levels such as insect parasitoids. Here we discuss some implications of plants acting as communication channels or ‘green phones’ between root and foliar-feeding insects and their parasitoids, focusing on recent findings that plants attacked by root-feeding insects are significantly less attractive for the parasitoids of foliar-feeding insects.

Many species are currently moving to higher latitudes and altitudes1, 2, 3. However, little is known about the factors that influence the future performance of range-expanding species in their new habitats. Here we show that range-expanding plant species from a riverine area were better defended against shoot and root enemies than were related native plant species growing in the same area. We grew fifteen plant species with and without non-coevolved polyphagous locusts and cosmopolitan, polyphagous aphids. Contrary to our expectations, the locusts performed more poorly on the range-expanding plant species than on the congeneric native plant species, whereas the aphids showed no difference. The shoot herbivores reduced the biomass of the native plants more than they did that of the congeneric range expanders. Also, the range-expanding plants developed fewer pathogenic effects4, 5 in their root-zone soil than did the related native species. Current predictions forecast biodiversity loss due to limitations in the ability of species to adjust to climate warming conditions in their range6, 7, 8. Our results strongly suggest that the plants that shift ranges towards higher latitudes and altitudes may include potential invaders, as the successful range expanders may experience less control by above-ground or below-ground enemies than the natives., Many species are currently moving to higher latitudes and altitudes1, 2, 3. However, little is known about the factors that influence the future performance of range-expanding species in their new habitats. Here we show that range-expanding plant species from a riverine area were better defended against shoot and root enemies than were related native plant species growing in the same area. We grew fifteen plant species with and without non-coevolved polyphagous locusts and cosmopolitan, polyphagous aphids. Contrary to our expectations, the locusts performed more poorly on the range-expanding plant species than on the congeneric native plant species, whereas the aphids showed no difference. The shoot herbivores reduced the biomass of the native plants more than they did that of the congeneric range expanders. Also, the range-expanding plants developed fewer pathogenic effects4, 5 in their root-zone soil than did the related native species. Current predictions forecast biodiversity loss due to limitations in the ability of species to adjust to climate warming conditions in their range6, 7, 8. Our results strongly suggest that the plants that shift ranges towards higher latitudes and altitudes may include potential invaders, as the successful range expanders may experience less control by above-ground or below-ground enemies than the natives.

Most studies on plant defenses against insect herbivores investigate direct and indirect plant defenses independently. However, these defenses are not necessarily mutually exclusive. Plant metabolites can be transmitted through the food chain and can also affect the herbivore's natural enemies. A conflict may arise when a natural enemy is attracted to a plant that is suboptimal in terms of its own fitness. In addition, plant defenses are often studied in cultivated plant species in which artificial selection may have resulted in reduced resistance against insect herbivores. In this study, we investigated both direct and indirect plant defenses in two closely related wild brassicaceous plant species, Brassica nigra L. and Sinapis arvensis L. The herbivore Pieris brassicae L. (Lepidoptera: Pieridae), which is specialized on brassicaceous plant species, developed faster and attained higher pupal mass when reared on B. nigra than on S. arvensis. In contrast, Cotesia glomerata L. (Hymenoptera: Braconidae), which is a gregarious endoparasitoid of P. brassicae caterpillars, developed equally well on P. brassicae irrespective of the food plant on which its host had been reared. The feeding strategy of the parasitoid larvae, that is, selectively feeding on hemolymph and fat body, is likely to allow for a much wider host-size range without affecting the size or development time of the emerging parasitoids. In flight chamber experiments, C. glomerata, which had an oviposition experience in a host that fed on Brussels sprout, exhibited significant preference for host-damaged B. nigra over host-damaged S. arvensis plants. Headspace analysis revealed quantitative and qualitative differences in volatile emissions between the two plant species. This parasitoid species may use a range of cues associated with the host and the host's food plant in order to recognize the different plant species on which the host can feed. These results show that there is no conflict between direct and, Most studies on plant defenses against insect herbivores investigate direct and indirect plant defenses independently. However, these defenses are not necessarily mutually exclusive. Plant metabolites can be transmitted through the food chain and can also affect the herbivore's natural enemies. A conflict may arise when a natural enemy is attracted to a plant that is suboptimal in terms of its own fitness. In addition, plant defenses are often studied in cultivated plant species in which artificial selection may have resulted in reduced resistance against insect herbivores. In this study, we investigated both direct and indirect plant defenses in two closely related wild brassicaceous plant species, Brassica nigra L. and Sinapis arvensis L. The herbivore Pieris brassicae L. (Lepidoptera: Pieridae), which is specialized on brassicaceous plant species, developed faster and attained higher pupal mass when reared on B. nigra than on S. arvensis. In contrast, Cotesia glomerata L. (Hymenoptera: Braconidae), which is a gregarious endoparasitoid of P. brassicae caterpillars, developed equally well on P. brassicae irrespective of the food plant on which its host had been reared. The feeding strategy of the parasitoid larvae, that is, selectively feeding on hemolymph and fat body, is likely to allow for a much wider host-size range without affecting the size or development time of the emerging parasitoids. In flight chamber experiments, C. glomerata, which had an oviposition experience in a host that fed on Brussels sprout, exhibited significant preference for host-damaged B. nigra over host-damaged S. arvensis plants. Headspace analysis revealed quantitative and qualitative differences in volatile emissions between the two plant species. This parasitoid species may use a range of cues associated with the host and the host's food plant in order to recognize the different plant species on which the host can feed. These results show that there is no conflict between direct and

Experiments were conducted between 2001 and 2003 in constructed plant communities that were set up in 1996 on abandoned agricultural land. The primary aim of the experiment was to study how different secondary vegetation succession scenarios influence community development of invertebrates in different trophic levels. The succession scenarios were obtained by sowing high diversity or low diversity seed mixtures of mid-successional plant species in 1996 in comparison with unsown plots where agriculture ceased in 1996 or 1999. Carnivorous carabid species generally preferred plots characterized by open vegetation, whereas herbivorous carabids generally favored plots associated with high plant diversity. However, carabid community composition was affected most by sampling year and there was also a dramatic shift over the three years in the relative proportion of the different trophic groups. Irrespective of treatment, the proportion of carnivores in the community declined with time, whereas phytophages increased over the course of the three years. By contrast, the proportion of omnivores peaked during the second year. These long-term changes were, at least to some extent, related to short-term changes in the structure and composition of the plant communities. The importance of local variation and temporal changes in plant species richness on carabids and other insect fauna are discussed., Experiments were conducted between 2001 and 2003 in constructed plant communities that were set up in 1996 on abandoned agricultural land. The primary aim of the experiment was to study how different secondary vegetation succession scenarios influence community development of invertebrates in different trophic levels. The succession scenarios were obtained by sowing high diversity or low diversity seed mixtures of mid-successional plant species in 1996 in comparison with unsown plots where agriculture ceased in 1996 or 1999. Carnivorous carabid species generally preferred plots characterized by open vegetation, whereas herbivorous carabids generally favored plots associated with high plant diversity. However, carabid community composition was affected most by sampling year and there was also a dramatic shift over the three years in the relative proportion of the different trophic groups. Irrespective of treatment, the proportion of carnivores in the community declined with time, whereas phytophages increased over the course of the three years. By contrast, the proportion of omnivores peaked during the second year. These long-term changes were, at least to some extent, related to short-term changes in the structure and composition of the plant communities. The importance of local variation and temporal changes in plant species richness on carabids and other insect fauna are discussed.

Black mustard, Brassica nigra (L.) Koch, is a wild annual species found throughout Europe and fed on by larvae of the large cabbage-white butterfly, Pieris brassicae L. We examined the impact of herbivory from P. brassicae, a gregarious herbivore, on B. nigra grown from wild seed collected locally. In greenhouse studies, the response of B. nigra to four herbivore densities in two developmental stages of the plant was quantified by measuring leaf damage, plant height, days to flowering, silique number, and seed production. Pieris brassicae readily attacked B. nigra leaves, although the timing of the attack did not affect seed production; attacked plants produced as many seeds as as nonattacked plants. Plant height was affected when plants were attacked early, but not later, in development, suggesting a connection between their belowground zone of influence and ability to regain biomass. These results demonstrate that at the herbivore densities and timing of damage studied, B. nigra tolerates folivory from Pieris brassicae through compensation., Black mustard, Brassica nigra (L.) Koch, is a wild annual species found throughout Europe and fed on by larvae of the large cabbage-white butterfly, Pieris brassicae L. We examined the impact of herbivory from P. brassicae, a gregarious herbivore, on B. nigra grown from wild seed collected locally. In greenhouse studies, the response of B. nigra to four herbivore densities in two developmental stages of the plant was quantified by measuring leaf damage, plant height, days to flowering, silique number, and seed production. Pieris brassicae readily attacked B. nigra leaves, although the timing of the attack did not affect seed production; attacked plants produced as many seeds as as nonattacked plants. Plant height was affected when plants were attacked early, but not later, in development, suggesting a connection between their belowground zone of influence and ability to regain biomass. These results demonstrate that at the herbivore densities and timing of damage studied, B. nigra tolerates folivory from Pieris brassicae through compensation.

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 para, 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 para

Through artificial selection, domesticated plants often contain modified levels of primary and secondary metabolites compared to their wild progenitors. It is hypothesized that the changed chemistry of cultivated plants will affect the performance of insects associated with these plants. In this paper, the development of several specialist and generalist herbivores and their endoparasitoids were compared when reared on a wild and cultivated population of cabbage, Brassica oleracea, and a recently established feral Brassica species. Irrespective of insect species or the degree of dietary specialization, herbivores and parasitoids developed most poorly on the wild population. For the specialists, plant population influenced only development time and adult body mass, whereas for the generalists, plant populations also affected egg-to-adult survival. Two parasitoid species, a generalist (Diadegma fenestrale) and a specialist (D. semiclausum), were reared from the same host (Plutella xylostella). Performance of D. semiclausum was closely linked to that of its host, whereas the correlation between survival of D. fenestrale and host performance was less clear. Plants in the Brassicaceae characteristically produce defense-related glucosinolates (GS). Levels of GS in leaves of undamaged plants were significantly higher in plants from the wild population than from the domesticated populations. Moreover, total GS concentrations increased significantly in wild plants after herbivory, but not in domesticated or feral plants. The results of this study reveal that a cabbage cultivar and plants from a wild cabbage population exhibit significant differences in quality in terms of their effects on the growth and development of insect herbivores and their natural enemies. Although cultivated plants have proved to be model systems in agroecology, we argue that some caution should be applied to evolutionary explanations derived from studies on domesticated plants, unless some knowledge

In nature, plants defend themselves by production of allelochemicals that are toxic to herbivores. There may be considerable genetic variation in the expression of chemical defenses because of various selection pressures. In this study, we examined the development of the small cabbage butterfly, Pieris rapae, and its gregarious pupal ectoparasitoid, Pteromalus puparum, when reared on three wild populations (Kimmeridge, Old Harry, Winspit) of cabbage, Brassica oleracea, and a Brussels sprout cultivar. Wild plant populations were obtained from seeds of plants that grow naturally along the south coast of Dorset, England. Significant differences in concentrations of allelochemicals (glucosinolates) were found in leaves of plants damaged by P. rapae. Total glucosinolate concentrations in Winspit plants, the population with the highest total glucosinolate concentration, were approximately four times higher than in the cultivar, the strain with the lowest total glucosinolate concentration. Pupal mass of P. rapae and adult body mass of Pt. puparum were highest when reared on the cultivar and lowest when developing on Kimmeridge plants, the wild strain with the lowest total glucosinolate concentration. Development of male parasitoids was also more negatively affected than female parasitoids. Our results reveal that plant quality, at least for the development of ‘adapted’ oligophagous herbivores, such as P. rapae, is not based on total glucosinolate content. The only glucosinolate compound that corresponded with the performance of P. rapae was the indole glucosinolate, neoglucobrassicin. Our results show that performance of ectoparasitoids may closely reflect constraints on the development of the host., In nature, plants defend themselves by production of allelochemicals that are toxic to herbivores. There may be considerable genetic variation in the expression of chemical defenses because of various selection pressures. In this study, we examined the development of the small cabbage butterfly, Pieris rapae, and its gregarious pupal ectoparasitoid, Pteromalus puparum, when reared on three wild populations (Kimmeridge, Old Harry, Winspit) of cabbage, Brassica oleracea, and a Brussels sprout cultivar. Wild plant populations were obtained from seeds of plants that grow naturally along the south coast of Dorset, England. Significant differences in concentrations of allelochemicals (glucosinolates) were found in leaves of plants damaged by P. rapae. Total glucosinolate concentrations in Winspit plants, the population with the highest total glucosinolate concentration, were approximately four times higher than in the cultivar, the strain with the lowest total glucosinolate concentration. Pupal mass of P. rapae and adult body mass of Pt. puparum were highest when reared on the cultivar and lowest when developing on Kimmeridge plants, the wild strain with the lowest total glucosinolate concentration. Development of male parasitoids was also more negatively affected than female parasitoids. Our results reveal that plant quality, at least for the development of ‘adapted’ oligophagous herbivores, such as P. rapae, is not based on total glucosinolate content. The only glucosinolate compound that corresponded with the performance of P. rapae was the indole glucosinolate, neoglucobrassicin. Our results show that performance of ectoparasitoids may closely reflect constraints on the development of the host.

It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata, a parasitoid of the leaf herbivore Pieris brassicae, mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities.

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Contains fulltext : 90882.pdf (publisher's version ) (Closed access)

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Interactions between butterflies and caterpillars in the genus Pieris and plants in the family Brassicaceae are among the best explored in the field of insect–plant biology. However, we report here for the first time that Pieris brassicae, commonly assumed to be a typical folivore, actually prefers to feed on flowers of three Brassica nigra genotypes rather than on their leaves. First- and second-instar caterpillars were observed to feed primarily on leaves, whereas late second and early third instars migrated via the small leaves of the flower branches to the flower buds and flowers. Once flower feeding began, no further leaf feeding was observed. We investigated growth rates of caterpillars having access exclusively to either leaves of flowering plants or flowers. In addition, we analyzed glucosinolate concentrations in leaves and flowers. Late-second- and early-third-instar P. brassicae caterpillars moved upward into the inflorescences of B. nigra and fed on buds and flowers until the end of the final (fifth) instar, after which they entered into the wandering stage, leaving the plant in search of a pupation site. Flower feeding sustained a significantly higher growth rate than leaf feeding. Flowers contained levels of glucosinolates up to five times higher than those of leaves. Five glucosinolates were identified: the aliphatic sinigrin, the aromatic phenyethylglucosinolate, and three indole glucosinolates: glucobrassicin, 4-methoxyglucobrassicin, and 4-hydroxyglucobrassicin. Tissue type and genotype were the most important factors affecting levels of identified glucosinolates. Sinigrin was by far the most abundant compound in all three genotypes. Sinigrin, 4-hydroxyglucobrassicin, and phenylethylglucosinolate were present at significantly higher levels in flowers than in leaves. In response to caterpillar feeding, sinigrin levels in both leaves and flowers were significantly higher than in undamaged plants, whereas 4-hydroxyglucobrassicin leaf levels were lower, Interactions between butterflies and caterpillars in the genus Pieris and plants in the family Brassicaceae are among the best explored in the field of insect–plant biology. However, we report here for the first time that Pieris brassicae, commonly assumed to be a typical folivore, actually prefers to feed on flowers of three Brassica nigra genotypes rather than on their leaves. First- and second-instar caterpillars were observed to feed primarily on leaves, whereas late second and early third instars migrated via the small leaves of the flower branches to the flower buds and flowers. Once flower feeding began, no further leaf feeding was observed. We investigated growth rates of caterpillars having access exclusively to either leaves of flowering plants or flowers. In addition, we analyzed glucosinolate concentrations in leaves and flowers. Late-second- and early-third-instar P. brassicae caterpillars moved upward into the inflorescences of B. nigra and fed on buds and flowers until the end of the final (fifth) instar, after which they entered into the wandering stage, leaving the plant in search of a pupation site. Flower feeding sustained a significantly higher growth rate than leaf feeding. Flowers contained levels of glucosinolates up to five times higher than those of leaves. Five glucosinolates were identified: the aliphatic sinigrin, the aromatic phenyethylglucosinolate, and three indole glucosinolates: glucobrassicin, 4-methoxyglucobrassicin, and 4-hydroxyglucobrassicin. Tissue type and genotype were the most important factors affecting levels of identified glucosinolates. Sinigrin was by far the most abundant compound in all three genotypes. Sinigrin, 4-hydroxyglucobrassicin, and phenylethylglucosinolate were present at significantly higher levels in flowers than in leaves. In response to caterpillar feeding, sinigrin levels in both leaves and flowers were significantly higher than in undamaged plants, whereas 4-hydroxyglucobrassicin leaf levels were lower

The thale cress, Arabidopsis thaliana, is considered to be an important model species in studying a suite of evolutionary processes. However, the species has been criticized on the basis of its comparatively small size at maturity (and consequent limitations in the amount of available biomass for herbivores) and on the duration and timing of its life cycle in nature. In the laboratory, we studied interactions between A. thaliana and the cabbage butterfly, Pieris rapae, in order to determine if plants are able to support the complete development of the herbivore. Plants were grown in pots from seedlings in densities of one, two, or four per pot. In each treatment, one, two, or five newly hatched larvae of P. rapae were placed on fully developed rosettes of A. thaliana. In a separate experiment, the same densities of P. rapae larvae were reared from hatching on single mature cabbage (Brassica oleracea) plants. Pupal fresh mass and survival of P. rapae declined with larval density when reared on A. thaliana but not on B. oleracea. However, irrespective of larval density and plant number, some P. rapae were always able to complete development on A. thaliana plants. A comparison of the dry mass of plants in different treatments with controls (= no larvae) revealed that A. thaliana partially compensated for plant damage when larval densities of P. rapae were low. By contrast, single cress plants with 5 larvae generally suffered extensive damage, whereas damage to B. oleracea plants was negligible. Rosettes of plants that were monitored in spring, when A. thaliana naturally grows, were not attacked by any insect herbivores, but there was often extensive damage from pulmonates (slugs and snails). Heavily damaged plants flowered less successfully than lightly damaged plants. Small numbers of generalist plant-parasitic nematodes were also recovered in roots and root soil. By contrast, plants monitored in a sewn summer plot were heavily attacked by insect herbivores, primarily, The thale cress, Arabidopsis thaliana, is considered to be an important model species in studying a suite of evolutionary processes. However, the species has been criticized on the basis of its comparatively small size at maturity (and consequent limitations in the amount of available biomass for herbivores) and on the duration and timing of its life cycle in nature. In the laboratory, we studied interactions between A. thaliana and the cabbage butterfly, Pieris rapae, in order to determine if plants are able to support the complete development of the herbivore. Plants were grown in pots from seedlings in densities of one, two, or four per pot. In each treatment, one, two, or five newly hatched larvae of P. rapae were placed on fully developed rosettes of A. thaliana. In a separate experiment, the same densities of P. rapae larvae were reared from hatching on single mature cabbage (Brassica oleracea) plants. Pupal fresh mass and survival of P. rapae declined with larval density when reared on A. thaliana but not on B. oleracea. However, irrespective of larval density and plant number, some P. rapae were always able to complete development on A. thaliana plants. A comparison of the dry mass of plants in different treatments with controls (= no larvae) revealed that A. thaliana partially compensated for plant damage when larval densities of P. rapae were low. By contrast, single cress plants with 5 larvae generally suffered extensive damage, whereas damage to B. oleracea plants was negligible. Rosettes of plants that were monitored in spring, when A. thaliana naturally grows, were not attacked by any insect herbivores, but there was often extensive damage from pulmonates (slugs and snails). Heavily damaged plants flowered less successfully than lightly damaged plants. Small numbers of generalist plant-parasitic nematodes were also recovered in roots and root soil. By contrast, plants monitored in a sewn summer plot were heavily attacked by insect herbivores, primarily

Larvae of the moth, Hadena bicruris, constitute the most important predispersal seed predator on Silene latifolia (Caryophyllaceae). Parasitoids attacking the larvae of this specialist noctuid can potentially decrease the amount of damage to the plant. This paper describes and quantifies the parasitoid complex associated with H. bicruris in the Netherlands, and documents life history characters of its species. Forty-four percent of larval H. bicruris were parasitized by at least 13 species of parasitoids. The most prevalent of these were the braconids Microplitis tristis (22.9%) and Bracon variator (4.9%) and the ichneumonids Eurylabus tristis (11.7%) and Ophion pteridis (3.4%). Other species occurring sporadically were: the ichneumonids Scambus brevicornis, S. buolianae, Erigorgus cerinops, and Hyposoter sp.; the tachinids Phryxe vulgaris, P. nemea, Blondelia nigripes and Siphona geniculata; and a Mermithidae sp. The ichneumonid hyperparasitoid Mesochorus lanceolatus was fo! und occasionally in larvae of M. tristis. The hyperparasitoid ichneumonids Gelis agilis, G. hortensis and the chalcids Barvscapus endemus, Pteromalus chrysos and P. vibulenus were found in cocoons of both M. tristis and B. variator. The primary parasitoids M. tristis, E. tristis, and to a lesser extent, O. pteridis, are believed to be specialized on Hadena. The effect of the parasitoids on herbivory by their host is probably small because the most common parasitoid species are koinobionts that attack large, late instar hosts. Only B. variator and Scambus spp., which are ectoparasitoids, arrest host development immediately upon parasitism, but like the other parasitoids they kill mainly large L4 or L5 hosts. The gregarious M. tristis and B. variator produce clutches with a female-biased sex ratio, in contrast to the solitary E. tristis where the sex ratio approaches equality. The ectoparasitoid B. variator produces mostly single-sex clutches, probably avoiding deleterious ef! fects of inbreeding, Larvae of the moth, Hadena bicruris, constitute the most important predispersal seed predator on Silene latifolia (Caryophyllaceae). Parasitoids attacking the larvae of this specialist noctuid can potentially decrease the amount of damage to the plant. This paper describes and quantifies the parasitoid complex associated with H. bicruris in the Netherlands, and documents life history characters of its species. Forty-four percent of larval H. bicruris were parasitized by at least 13 species of parasitoids. The most prevalent of these were the braconids Microplitis tristis (22.9%) and Bracon variator (4.9%) and the ichneumonids Eurylabus tristis (11.7%) and Ophion pteridis (3.4%). Other species occurring sporadically were: the ichneumonids Scambus brevicornis, S. buolianae, Erigorgus cerinops, and Hyposoter sp.; the tachinids Phryxe vulgaris, P. nemea, Blondelia nigripes and Siphona geniculata; and a Mermithidae sp. The ichneumonid hyperparasitoid Mesochorus lanceolatus was fo! und occasionally in larvae of M. tristis. The hyperparasitoid ichneumonids Gelis agilis, G. hortensis and the chalcids Barvscapus endemus, Pteromalus chrysos and P. vibulenus were found in cocoons of both M. tristis and B. variator. The primary parasitoids M. tristis, E. tristis, and to a lesser extent, O. pteridis, are believed to be specialized on Hadena. The effect of the parasitoids on herbivory by their host is probably small because the most common parasitoid species are koinobionts that attack large, late instar hosts. Only B. variator and Scambus spp., which are ectoparasitoids, arrest host development immediately upon parasitism, but like the other parasitoids they kill mainly large L4 or L5 hosts. The gregarious M. tristis and B. variator produce clutches with a female-biased sex ratio, in contrast to the solitary E. tristis where the sex ratio approaches equality. The ectoparasitoid B. variator produces mostly single-sex clutches, probably avoiding deleterious ef! fects of inbreeding

1. Root feeding insects can influence foliar quality of the host plant, which can affect the development and behaviour of leaf herbivores and parasitoids. Thus far, such interactions have been reported in situations where root and leaf associated organisms share a host-plant. We tested whether root herbivory influences searching behaviour of an above-ground parasitoid when the foliar feeding host of the parasitoid and the root herbivore are feeding on different plants. 2. We manipulated the proportion of 25 plants (ranging from 0 to 1) exposed to root herbivory by Delia radicum (neighbouring-plants). Five additional plants were infested above-ground with Pieris brassicae larvae (host-infested plants) and were placed in-between the neighbouring plants. We then released females of the parasitoid Cotesia glomerata which attacks P. brassicae and studied foraging efficiency of the parasitoid. 3. Overall, parasitoids located more host-infested plants during the maximum allowed searching time, and found their hosts about three times faster when neighbouring plants were exposed to root herbivory, than when neighbouring plants were not infested with D. radicum. Similar results were found when the host-infested plants were also exposed to root herbivory. 4. Our results show that the interaction between an above-ground foliar feeding insect and its parasitoid can be influenced by the presence of non-host herbivores feeding on the roots of neighbouring conspecific plants., 1. Root feeding insects can influence foliar quality of the host plant, which can affect the development and behaviour of leaf herbivores and parasitoids. Thus far, such interactions have been reported in situations where root and leaf associated organisms share a host-plant. We tested whether root herbivory influences searching behaviour of an above-ground parasitoid when the foliar feeding host of the parasitoid and the root herbivore are feeding on different plants. 2. We manipulated the proportion of 25 plants (ranging from 0 to 1) exposed to root herbivory by Delia radicum (neighbouring-plants). Five additional plants were infested above-ground with Pieris brassicae larvae (host-infested plants) and were placed in-between the neighbouring plants. We then released females of the parasitoid Cotesia glomerata which attacks P. brassicae and studied foraging efficiency of the parasitoid. 3. Overall, parasitoids located more host-infested plants during the maximum allowed searching time, and found their hosts about three times faster when neighbouring plants were exposed to root herbivory, than when neighbouring plants were not infested with D. radicum. Similar results were found when the host-infested plants were also exposed to root herbivory. 4. Our results show that the interaction between an above-ground foliar feeding insect and its parasitoid can be influenced by the presence of non-host herbivores feeding on the roots of neighbouring conspecific plants.

The majority of studies exploring interactions between above- and below-ground biota have been focused on the effects of root-associated organisms on foliar herbivorous insects. This study examined the effects of foliar herbivory by Pieris brassicae L. (Lepidoptera: Pieridae) on the performance of the root herbivore Delia radicum L. (Diptera: Anthomyiidae) and its parasitoid Trybliographa rapae (Westwood) (Hymenoptera: Figitidae), mediated through a shared host plant Brassica nigra L. (Brassicaceae). In the presence of foliar herbivory, the survival of D. radicum and T. rapae decreased significantly by more than 50%. In addition, newly emerged adults of both root herbivores and parasitoids were significantly smaller on plants that had been exposed to foliar herbivory than on control plants. To determine what factor(s) may have accounted for the observed results, we examined the effects of foliar herbivory on root quantity and quality. No significant differences in root biomass were found between plants with and without shoot herbivore damage. Moreover, concentrations of nitrogen in root tissues were also unaffected by shoot damage by P. brassicae larvae. However, higher levels of indole glucosinolates were measured in roots of plants exposed to foliar herbivory, suggesting that the development of the root herbivore and its parasitoid may be, at least partly, negatively affected by increased levels of these allelochemicals in root tissues. Our results show that foliar herbivores can affect the development not only of root-feeding insects but also their natural enemies. We argue that such indirect interactions between above- and below-ground biota may play an important role in the structuring and functioning of communities., The majority of studies exploring interactions between above- and below-ground biota have been focused on the effects of root-associated organisms on foliar herbivorous insects. This study examined the effects of foliar herbivory by Pieris brassicae L. (Lepidoptera: Pieridae) on the performance of the root herbivore Delia radicum L. (Diptera: Anthomyiidae) and its parasitoid Trybliographa rapae (Westwood) (Hymenoptera: Figitidae), mediated through a shared host plant Brassica nigra L. (Brassicaceae). In the presence of foliar herbivory, the survival of D. radicum and T. rapae decreased significantly by more than 50%. In addition, newly emerged adults of both root herbivores and parasitoids were significantly smaller on plants that had been exposed to foliar herbivory than on control plants. To determine what factor(s) may have accounted for the observed results, we examined the effects of foliar herbivory on root quantity and quality. No significant differences in root biomass were found between plants with and without shoot herbivore damage. Moreover, concentrations of nitrogen in root tissues were also unaffected by shoot damage by P. brassicae larvae. However, higher levels of indole glucosinolates were measured in roots of plants exposed to foliar herbivory, suggesting that the development of the root herbivore and its parasitoid may be, at least partly, negatively affected by increased levels of these allelochemicals in root tissues. Our results show that foliar herbivores can affect the development not only of root-feeding insects but also their natural enemies. We argue that such indirect interactions between above- and below-ground biota may play an important role in the structuring and functioning of communities.

It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata, a parasitoid of the leaf herbivore Pieris brassicae, mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities., It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata, a parasitoid of the leaf herbivore Pieris brassicae, mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities.

Mature larvae of the gregarious endoparasitoid Cotesia kariyai construct cocoons for pupation approximately 10 days after parasitization and emerge from their host Pseudaletia separata under a long day photo-regime (16L8D) at 25 ± 1°C. The parasitoid larvae make capsules in the host hemocoel just prior to their emergence. These capsules function as anchors, which enable them to press against the host integument from inside the host. It was predicted that this anchor might be composed of silk proteins secreted from the parasitoid larvae, because a previous study showed that the anchor was made up of a glycoprotein and that the silk gland of parasitoid larvae developed from 2nd larval stage. Fibroin-like proteins in C. kariyai larva mainly consist of two proteins with molecular masses of the 300.6 and 46.7 kDa estimated by SDS-PAGE. The fibroin-like proteins with the same molecular mass were detected from the anchor proteins just prior to parasitoid emergence. These results indicate that the anchor was assembled with fibrion-like proteins and was formed just before parasitoid emergence while in the host body cavity. Injection of bovine pancreatic trypsin inhibited the emergence of parasitoid larvae from the host because the anchor was decomposed by trypsin. Trypsin activity in the parasitized host hemolymph increased only after parasitoid emergence., Mature larvae of the gregarious endoparasitoid Cotesia kariyai construct cocoons for pupation approximately 10 days after parasitization and emerge from their host Pseudaletia separata under a long day photo-regime (16L8D) at 25 ± 1°C. The parasitoid larvae make capsules in the host hemocoel just prior to their emergence. These capsules function as anchors, which enable them to press against the host integument from inside the host. It was predicted that this anchor might be composed of silk proteins secreted from the parasitoid larvae, because a previous study showed that the anchor was made up of a glycoprotein and that the silk gland of parasitoid larvae developed from 2nd larval stage. Fibroin-like proteins in C. kariyai larva mainly consist of two proteins with molecular masses of the 300.6 and 46.7 kDa estimated by SDS-PAGE. The fibroin-like proteins with the same molecular mass were detected from the anchor proteins just prior to parasitoid emergence. These results indicate that the anchor was assembled with fibrion-like proteins and was formed just before parasitoid emergence while in the host body cavity. Injection of bovine pancreatic trypsin inhibited the emergence of parasitoid larvae from the host because the anchor was decomposed by trypsin. Trypsin activity in the parasitized host hemolymph increased only after parasitoid emergence.

Herbivory can induce changes in plant growth, physiology, morphology, and phenology. Such herbivore-induced changes by insects are commonly observed in many plant species. Changes in the host plant can alter interaction linkages between plants, herbivores and higher trophic levels such as carnivores of the herbivores. These interaction linkages shape community organization and biodiversity in plant-based terrestrial communities. Considering that plants have both aboveground and belowground parts, systemic herbivore-induced changes within the plant could potentially link spatially separated herbivores associated with plants in the soil and in the aboveground (sub) system. Relatively recently, an increasing number of studies have shown that root herbivores, among other soil dwelling organisms, can indeed interact with foliar herbivores. Root herbivores change the quality and biomass of aboveground plant parts, which in turn influences the growth, development and survival of foliar herbivores. Similarly, foliar-feeding insects have been shown to influence root-feeding insects. This thesis explores whether, and mechanistically how, these feed-backs between root and foliar herbivores are transmitted through the soil and the aboveground food chains influencing the performance and behaviour of higher trophic levels. In Chapter 1, I introduce plant-insect interactions in a multitrophic context, briefly describing how the plant mediates these intricate interactions, and I describe the general aim and the outline of the thesis. In Chapter 2 I study the effects of root herbivores, at different densities, on the biomass and quality of the plant shoot, and on the performance of a foliar herbivore, its primary parasitoid and its secondary parasitoid (hyperparasitoid) sharing the host plant. In Chapter 3 I study the reverse effect; the effects of foliar herbivores on a root herbivore and its parasitoid mediated by the host plant. The results of these Chapters show that the spatial

Because plants cannot run away from their attackers, move to more favourable locations or hide, they have to be able to tolerate, adapt and/or defend themselves. Plants have evolved an enormous array of mechanical and chemical defences against herbivores. One can distinguish three types of defence strategies: direct defences that directly affect the herbivores, indirect defences that attract the enemies of the herbivores and tolerance, which reduces the fitness consequences of herbivore damage. It is unlikely that direct and indirect defences act independently in plants. Natural enemies of the herbivores (the indirect defence) can be negatively affected when attacking larvae which feed on plants defended by high levels of allelochemicals (direct defence), causing potential incompatibility between host plant resistance and biological control. On the other hand, herbivores feeding on plants with high levels of allelochemicals may develop more slowly and have an impaired immune function, leading to longer exposure of vulnerable stages to parasitism and predation and causing potential synergism between host plant resistance and biological control. Studies of interactions between direct and indirect defences are almost exclusively based on studies with crop species. It has frequently been observed that domesticated plants have lower levels of defensive chemicals than their wild relatives.In this study I used a natural plant-herbivore-parasitoid complex. The main focus in this thesis is the effect of direct defence chemicals in the plant on higher trophic levels in the system, to detect if there is potential for a conflict between direct chemical defence and indirect defence of the plant. Natural enemies of herbivores are a potential source of indirect defences for a plant. Plants can attract these enemies by providing shelter (domatia), food or signalling them with volatiles. The efficiency of these indirect defences depends on the effect of the natural enemy on the

The majority of studies exploring interactions between above- and below-ground biota have been focused on the effects of root-associated organisms on foliar herbivorous insects. This study examined the effects of foliar herbivory by Pieris brassicae L. (Lepidoptera: Pieridae) on the performance of the root herbivore Delia radicum L. (Diptera: Anthomyiidae) and its parasitoid Trybliographa rapae (Westwood) (Hymenoptera: Figitidae), mediated through a shared host plant Brassica nigra L. (Brassicaceae). In the presence of foliar herbivory, the survival of D. radicum and T. rapae decreased significantly by more than 50%. In addition, newly emerged adults of both root herbivores and parasitoids were significantly smaller on plants that had been exposed to foliar herbivory than on control plants. To determine what factor(s) may have accounted for the observed results, we examined the effects of foliar herbivory on root quantity and quality. No significant differences in root biomass were found between plants with and without shoot herbivore damage. Moreover, concentrations of nitrogen in root tissues were also unaffected by shoot damage by P. brassicae larvae. However, higher levels of indole glucosinolates were measured in roots of plants exposed to foliar herbivory, suggesting that the development of the root herbivore and its parasitoid may be, at least partly, negatively affected by increased levels of these allelochemicals in root tissues. Our results show that foliar herbivores can affect the development not only of root-feeding insects but also their natural enemies. We argue that such indirect interactions between above- and below-ground biota may play an important role in the structuring and functioning of communities.

A number of studies have reported that consumers affect a range of community-level processes, and in turn their diversity and abundance is influenced by the structure and diversity of the plant community. Although gastropods are important generalist herbivores in many environments, few studies have examined the effects of plant species richness and plant community structure on gastropods. This study investigated gastropod species richness and interactions with various above-ground parameters of the vegetation on an experimental field with four plant treatments: low and high diversity of sown later succession plant species, natural colonization at the start of the experiment and natural colonization after 3 years of continued agricultural practice. The investigated gastropod assemblage contained only seven species and was highly dominated by two of them. Both in pitfalls and with hand-sorting the number of species collected per plot was highest in plots with natural plant colonization. Multivariate analysis revealed that overall gastropod abundance was positively associated with plant height and percentage cover of plants, and negatively with percentage grass cover. The same pattern holds for one of the dominant species-complex (Cochlicopa lubrica/lubricella). The other dominant gastropod species (Deroceras reticulatum) was more abundant in samples with higher percentages of moss cover and higher plant diversity, while less abundant at samples with higher plant cover, indicating that the gastropod species preferences may matter more than just their response to plant diversity. Two plant–gastropod species-level associations were observed: Senecio jacobaea with D. reticulatum and Tanacetum vulgare with Cochlicopa spp. The present study also demonstrated that pitfall-traps are suitable for collecting terrestrial gastropods, at least for species-poor grassland habitats. [KEYWORDS: Cochlicopa spp. ; Deroceras reticulatum ; Plant diversity ; Plant–gastropod interactions, A number of studies have reported that consumers affect a range of community-level processes, and in turn their diversity and abundance is influenced by the structure and diversity of the plant community. Although gastropods are important generalist herbivores in many environments, few studies have examined the effects of plant species richness and plant community structure on gastropods. This study investigated gastropod species richness and interactions with various above-ground parameters of the vegetation on an experimental field with four plant treatments: low and high diversity of sown later succession plant species, natural colonization at the start of the experiment and natural colonization after 3 years of continued agricultural practice. The investigated gastropod assemblage contained only seven species and was highly dominated by two of them. Both in pitfalls and with hand-sorting the number of species collected per plot was highest in plots with natural plant colonization. Multivariate analysis revealed that overall gastropod abundance was positively associated with plant height and percentage cover of plants, and negatively with percentage grass cover. The same pattern holds for one of the dominant species-complex (Cochlicopa lubrica/lubricella). The other dominant gastropod species (Deroceras reticulatum) was more abundant in samples with higher percentages of moss cover and higher plant diversity, while less abundant at samples with higher plant cover, indicating that the gastropod species preferences may matter more than just their response to plant diversity. Two plant–gastropod species-level associations were observed: Senecio jacobaea with D. reticulatum and Tanacetum vulgare with Cochlicopa spp. The present study also demonstrated that pitfall-traps are suitable for collecting terrestrial gastropods, at least for species-poor grassland habitats. [KEYWORDS: Cochlicopa spp. ; Deroceras reticulatum ; Plant diversity ; Plant–gastropod interactions

Several studies have reported that flowering herbs, which grow naturally or are sown adjacent to agricultural fields, may be an important source of nutrients for natural enemies. Many parasitoids readily feed on plant exudates such as floral nectar, which contain different types of sugars that enable the insects to optimize their longevity, mobility and reproductive success. However, leaf tissues of plants grown in the margins of agricultural fields may also provide food for immature stages of insect herbivores, such as caterpillars, that are in turn attacked by parasitoids. Herbivores and their parasitoids may later disperse into the crop, so the nutritional quality of surrounding plants, as this affects herbivore and parasitoid fitness, may also influence the success of biological control programmes, especially later in the season. Here, we compare the suitability of three species of cruciferous plants (Brassicaceae) on the development of Pieris rapae L. (Lep., Pieridae) and its solitary endoparasitoid, Cotesia rubecula Marshall (Hym., Bracondiae). Insects were reared on a feral population of cabbage, Brassica oleracea, on radish Raphanus sativus, which is widely sown in agricultural margins, and on hedge mustard, Sisymbrium officinale, a wild crucifer which often grows in medium to large stands along road verges and field edges. Development time in both the herbivore and parasitoid were extended on R. sativus, compared with the other two species, whereas C. rubecula completed its development most rapidly on B. oleracea. Moreover, adult butterflies and parasitoids were significantly smaller when reared on R. sativus plants. Our results reveal that differences in the quality of plants growing adjacent to agricultural fields can affect the development of key herbivores and their parasitoids. This should be borne in mind when establishing criteria for the selection of floral biodiversity., Several studies have reported that flowering herbs, which grow naturally or are sown adjacent to agricultural fields, may be an important source of nutrients for natural enemies. Many parasitoids readily feed on plant exudates such as floral nectar, which contain different types of sugars that enable the insects to optimize their longevity, mobility and reproductive success. However, leaf tissues of plants grown in the margins of agricultural fields may also provide food for immature stages of insect herbivores, such as caterpillars, that are in turn attacked by parasitoids. Herbivores and their parasitoids may later disperse into the crop, so the nutritional quality of surrounding plants, as this affects herbivore and parasitoid fitness, may also influence the success of biological control programmes, especially later in the season. Here, we compare the suitability of three species of cruciferous plants (Brassicaceae) on the development of Pieris rapae L. (Lep., Pieridae) and its solitary endoparasitoid, Cotesia rubecula Marshall (Hym., Bracondiae). Insects were reared on a feral population of cabbage, Brassica oleracea, on radish Raphanus sativus, which is widely sown in agricultural margins, and on hedge mustard, Sisymbrium officinale, a wild crucifer which often grows in medium to large stands along road verges and field edges. Development time in both the herbivore and parasitoid were extended on R. sativus, compared with the other two species, whereas C. rubecula completed its development most rapidly on B. oleracea. Moreover, adult butterflies and parasitoids were significantly smaller when reared on R. sativus plants. Our results reveal that differences in the quality of plants growing adjacent to agricultural fields can affect the development of key herbivores and their parasitoids. This should be borne in mind when establishing criteria for the selection of floral biodiversity.

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 a, 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 a

Lysibia nana is a solitary, secondary idiobiont hyperparasitoid that attacks newly cocooned pre-pupae and pupae of several closely related gregarious endoparasitoids in the genus Cotesia, including C. glomerata. Prior to oviposition, the female wasp injects paralysing venom into the host, thus preventing further development. Here, host fate, emerging hyperparasitoid mass, and egg-to-adult development time was compared in hosts parasitized at different ages over 24-h intervals. Cocoons of C. glomerata were parasitized by L. nana at 12, 36, 60, 84, and 108 h post-egression from the secondary host, Pieris brassicae. Hyperparasitoid survival exceeded 80% in hosts parasitized within the first 60 h after pupation, but dropped thereafter, with no hyperparasitoids emerging in hosts aged 108 h. The mass of hyperparasitoids was positively correlated with the mass of the host cocoon, and this relationship remained consistent in hosts up to 60 h old. Within each host age cohort, the mass of male and female wasps was not significantly different. Development time in L. nana was uniform in hosts up to 60 h old, but increased significantly in 84-h-old hosts, and male wasps completed their development earlier than female wasps. Regulation of host growth varied with the age of the host at parasitism, with the early growth of older hosts reduced much more dramatically than young hosts. Unlike most parasitoids, pupal hyperparasitoids do not make cocoons but instead pupate within the already prepared cocoon of the host parasitoid. Consequently, for a given mass of cocoon, newly emerged L. nana adults were remarkably similar in size with male and female adults of C. glomerata. This reveals that L. nana is extremely efficient at exploiting its primary parasitoid host. [KEYWORDS: Cotesia glomerata ; development ; host quality ; hyperparasitoid ; idiobiont ; Lysibia nana], Lysibia nana is a solitary, secondary idiobiont hyperparasitoid that attacks newly cocooned pre-pupae and pupae of several closely related gregarious endoparasitoids in the genus Cotesia, including C. glomerata. Prior to oviposition, the female wasp injects paralysing venom into the host, thus preventing further development. Here, host fate, emerging hyperparasitoid mass, and egg-to-adult development time was compared in hosts parasitized at different ages over 24-h intervals. Cocoons of C. glomerata were parasitized by L. nana at 12, 36, 60, 84, and 108 h post-egression from the secondary host, Pieris brassicae. Hyperparasitoid survival exceeded 80% in hosts parasitized within the first 60 h after pupation, but dropped thereafter, with no hyperparasitoids emerging in hosts aged 108 h. The mass of hyperparasitoids was positively correlated with the mass of the host cocoon, and this relationship remained consistent in hosts up to 60 h old. Within each host age cohort, the mass of male and female wasps was not significantly different. Development time in L. nana was uniform in hosts up to 60 h old, but increased significantly in 84-h-old hosts, and male wasps completed their development earlier than female wasps. Regulation of host growth varied with the age of the host at parasitism, with the early growth of older hosts reduced much more dramatically than young hosts. Unlike most parasitoids, pupal hyperparasitoids do not make cocoons but instead pupate within the already prepared cocoon of the host parasitoid. Consequently, for a given mass of cocoon, newly emerged L. nana adults were remarkably similar in size with male and female adults of C. glomerata. This reveals that L. nana is extremely efficient at exploiting its primary parasitoid host. [KEYWORDS: Cotesia glomerata ; development ; host quality ; hyperparasitoid ; idiobiont ; Lysibia nana]

Since the concept of the 'niche' was proposed by Hutchinson almost 50 years ago, many researchers have investigated factors that enable species within 'guilds' (i.e., species that exploit a common resource and are likely to compete with one another) to co-exist and to persist. Here, development, host utilization, and life-history characteristics are compared in two species of solitary secondary hyperparasitoids, Lysibia nana and Gelis agilis, attacking pre-pupae of their primary parasitoid host, Cotesia glomerata. More than 80% of the host cocoons presented to L. nana successfully produced adult hyperparasitoids compared with only about 20% for G. agilis. Adult hyperparasitoid body mass increased with host mass, but there was little difference in the mean overall body mass of either species. However, egg-to-adult development time for L. nana was significantly less than that for G. agilis. When provided with unlimited food (honey solution), G. agilis had a lifespan that was almost twice that of L. nana. However, dissections of the ovaries of adult wasps at death revealed that L. nana usually had accumulated more than 20 mature eggs, whereas G. agilis was effectively sterile. We suggest that several eco-physiological factors, as well as differing degrees of specialization amongst the guild of secondary hyperparasitoids attacking C glomerata cocoons, enable them to successfully co-exist in nature. [KEYWORDS: competition ; Cotesia glomerata ; guild ; hyperparasitoid ; niche construction]

Soil organisms can influence higher trophic level aboveground organisms, but only very few studies have considered such effects. We manipulated soil community composition of model grassland ecosystems by introducing nematode communities, microorganisms, neither or both groups. Above ground, aphids (Rhopalosiphum padi) and parasitoids (Aphidius colemani) were introduced, and we measured individual performance and population dynamics of plants, aphids and parasitoids. In microcosms with nematode inoculations either with or without microorganism inoculation, aphids offspring production was significantly reduced by 31%. Aphid populations on both host plants Agrostis capillaris and Anthoxanthum odoratum were lowest in microcosms with combined nematode and microorganism inoculations. Opposite results were found for parasitoids. While the number of emerged parasitoids did not differ between treatments, parasitoid mortality and the proportion of males were significantly lower in microcosms with nematode and microorganism inoculations. Parasitized aphids were significantly larger in microcosms with nematodes inoculated. Plant biomass did not differ, but in the preferred host plant A. odoratum, foliar phenolic content was reduced in the presence of nematodes, and also the concentration of amino acids in the phloem. This study shows that the composition of the soil community matters for aboveground multitrophic interactions. [KEYWORDS: Abovegroundbelowground interactions ; aphid ; Aphidius colemani ; fitness ; herbivory ; microcosm ; nematode ; parasitoid ; Rhopalosiphum padi]

Studies in crop species show that the effect of plant allelochemicals is not necessarily restricted to herbivores, but can extend to (positive as well as negative) effects on performance at higher trophic levels, including the predators and parasitoids of herbivores. We examined how quantitative variation in allelochemicals (iridoid glycosides) in ribwort plantain, Plantago lanceolata, affects the development of a specialist and a generalist herbivore and their respective specialist and generalist endoparasitoids. Plants were grown from two selection lines that differed ca. 5-fold in the concentration of leaf iridoid glycosides. Development time of the specialist herbivore, Melitaea cinxia, and its solitary endoparasitoid, Hyposoter horticola, proceeded most rapidly when reared on the high iridoid line, whereas pupal mass in M. cinxia and adult mass in H. horticola were unaffected by plant line. Cotesia melitaearum, a gregarious endoparasitoid of M. cinxia, performed equally well on hosts feeding on the two lines of P. lanceolata. In contrast, the pupal mass of the generalist herbivore, Spodoptera exigua, and the emerging adult mass of its solitary endoparasitoid, C. marginiventris, were significantly lower when reared on the high line, whereas development time was unaffected. The results are discussed with regards to (1) differences between specialist and generalist herbivores and their natural enemies to quantitative variation in plant secondary chemistry, and (2) potentially differing selection pressures on plant defense. [KEYWORDS: Chemical defense ; iridoid glycosides ; Melitaea cinxia ; multitrophic interactions ; Plantago lanceolata ; Spodoptera exigua], Studies in crop species show that the effect of plant allelochemicals is not necessarily restricted to herbivores, but can extend to (positive as well as negative) effects on performance at higher trophic levels, including the predators and parasitoids of herbivores. We examined how quantitative variation in allelochemicals (iridoid glycosides) in ribwort plantain, Plantago lanceolata, affects the development of a specialist and a generalist herbivore and their respective specialist and generalist endoparasitoids. Plants were grown from two selection lines that differed ca. 5-fold in the concentration of leaf iridoid glycosides. Development time of the specialist herbivore, Melitaea cinxia, and its solitary endoparasitoid, Hyposoter horticola, proceeded most rapidly when reared on the high iridoid line, whereas pupal mass in M. cinxia and adult mass in H. horticola were unaffected by plant line. Cotesia melitaearum, a gregarious endoparasitoid of M. cinxia, performed equally well on hosts feeding on the two lines of P. lanceolata. In contrast, the pupal mass of the generalist herbivore, Spodoptera exigua, and the emerging adult mass of its solitary endoparasitoid, C. marginiventris, were significantly lower when reared on the high line, whereas development time was unaffected. The results are discussed with regards to (1) differences between specialist and generalist herbivores and their natural enemies to quantitative variation in plant secondary chemistry, and (2) potentially differing selection pressures on plant defense. [KEYWORDS: Chemical defense ; iridoid glycosides ; Melitaea cinxia ; multitrophic interactions ; Plantago lanceolata ; Spodoptera exigua]

1. Habitat complexity may stabilize interactions among species of different trophic levels by providing refuges to organisms of lower trophic levels. 2. Searching behaviour of the parasitoid, Diadegma semiclausum, was followed in different semifield set-ups, a low and high-density monoculture of Brassica oleracea and two intercrops, B. oleracea with Sinapis alba (also a member of the Brassicaceae) and B. oleracea with Hordeum vulgare (Poaceae). 3. When a low-density monocrop of B. oleracea was compared with a high-density monocrop, no differences were found in the ability of the female wasps to locate a host-infested plant, B. oleracea, infested with Plutella xylostella that was placed in the centre of the set-up. 4. The efficiency of the parasitoid to locate the host-infested plant was differentially affected by the species composition of the vegetation. Wasps entered the Sinapis-Brassica set-up faster, but took more time to find the host-infested plant than in the Hordeum-Brassica set-up. 5. The horizontal arrangement, i.e. by mixing S. alba or H. vulgare with, or placing them as rows between B. oleracea, did not affect host-finding efficiency. 6. Plant height did influence host finding. Wasps found the host-infested plants earlier in the set-up with short Sinapis plants compared with tall Sinapis plants. 7. Once the wasps had landed on the host-infested plant, the surrounding vegetation did not affect time needed to parasitize five consecutive hosts on the same infested plant, regardless of the composition or horizontal/vertical arrangement of the set-up. 8. Chemical and structural refuges in complex landscapes may play an important role in the persistence of this system through dampening oscillations of parasitoid and host populations. [KEYWORDS: crucifers ; Diadegma semiclausum ; Plutella xylostella ; proportional hazards model ; refuges], 1. Habitat complexity may stabilize interactions among species of different trophic levels by providing refuges to organisms of lower trophic levels. 2. Searching behaviour of the parasitoid, Diadegma semiclausum, was followed in different semifield set-ups, a low and high-density monoculture of Brassica oleracea and two intercrops, B. oleracea with Sinapis alba (also a member of the Brassicaceae) and B. oleracea with Hordeum vulgare (Poaceae). 3. When a low-density monocrop of B. oleracea was compared with a high-density monocrop, no differences were found in the ability of the female wasps to locate a host-infested plant, B. oleracea, infested with Plutella xylostella that was placed in the centre of the set-up. 4. The efficiency of the parasitoid to locate the host-infested plant was differentially affected by the species composition of the vegetation. Wasps entered the Sinapis-Brassica set-up faster, but took more time to find the host-infested plant than in the Hordeum-Brassica set-up. 5. The horizontal arrangement, i.e. by mixing S. alba or H. vulgare with, or placing them as rows between B. oleracea, did not affect host-finding efficiency. 6. Plant height did influence host finding. Wasps found the host-infested plants earlier in the set-up with short Sinapis plants compared with tall Sinapis plants. 7. Once the wasps had landed on the host-infested plant, the surrounding vegetation did not affect time needed to parasitize five consecutive hosts on the same infested plant, regardless of the composition or horizontal/vertical arrangement of the set-up. 8. Chemical and structural refuges in complex landscapes may play an important role in the persistence of this system through dampening oscillations of parasitoid and host populations. [KEYWORDS: crucifers ; Diadegma semiclausum ; Plutella xylostella ; proportional hazards model ; refuges]

1. Plants and insects are part of a complex multitrophic environment, in which they closely interact. However, most of the studies have been focused mainly on bi-tritrophic above-ground subsystems, hindering our understanding of the processes that affect multitrophic interactions in a more realistic framework. 2. We studied whether root herbivory by the fly Delia radicum can influence the development of the leaf feeder Pieris brassicae, its parasitoid Cotesia glomerata and its hyperparasitoid Lysibia nana, through changes in primary and secondary plant compounds. 3. In the presence of root herbivory, the development time of the leaf herbivore and the parasitoid significantly increased, and the adult size of the parasitoid and the hyperparasitoid were significantly reduced. The effects were stronger at low root fly densities than at high densities. 4. Higher glucosinolate (sinigrin) levels were recorded in plants exposed to below-ground herbivory, suggesting that the reduced performance of the above-ground insects was via reduced plant quality. Sinigrin contents were highest in plants exposed to low root fly densities, intermediate in plants exposed to high root fly densities and lowest in plants that were not exposed to root herbivory. 5. Our results show, for the first time, that root herbivory via changes in plant quality can reduce the performance of an above-ground multitrophic level food chain. This underlines the importance of integrating a broader range of above- and below-ground organisms to facilitate a better understanding of complex multitrophic interactions and interrelationships. [KEYWORDS: above–below-ground interactions ; Cotesia glomerata ; Lysibia nana ; Pieris brassicae ; plant–insect interactions], 1. Plants and insects are part of a complex multitrophic environment, in which they closely interact. However, most of the studies have been focused mainly on bi-tritrophic above-ground subsystems, hindering our understanding of the processes that affect multitrophic interactions in a more realistic framework. 2. We studied whether root herbivory by the fly Delia radicum can influence the development of the leaf feeder Pieris brassicae, its parasitoid Cotesia glomerata and its hyperparasitoid Lysibia nana, through changes in primary and secondary plant compounds. 3. In the presence of root herbivory, the development time of the leaf herbivore and the parasitoid significantly increased, and the adult size of the parasitoid and the hyperparasitoid were significantly reduced. The effects were stronger at low root fly densities than at high densities. 4. Higher glucosinolate (sinigrin) levels were recorded in plants exposed to below-ground herbivory, suggesting that the reduced performance of the above-ground insects was via reduced plant quality. Sinigrin contents were highest in plants exposed to low root fly densities, intermediate in plants exposed to high root fly densities and lowest in plants that were not exposed to root herbivory. 5. Our results show, for the first time, that root herbivory via changes in plant quality can reduce the performance of an above-ground multitrophic level food chain. This underlines the importance of integrating a broader range of above- and below-ground organisms to facilitate a better understanding of complex multitrophic interactions and interrelationships. [KEYWORDS: above–below-ground interactions ; Cotesia glomerata ; Lysibia nana ; Pieris brassicae ; plant–insect interactions]

1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid. 2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals. 3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured. 4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time. 5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size. 6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short. 7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size. [KEYWORDS: Ageing ; clutch size adjustment ; egg limitation ; Hadena bicruris ; host deprivation ; time limitation], 1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid. 2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals. 3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured. 4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time. 5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size. 6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short. 7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size. [KEYWORDS: Ageing ; clutch size adjustment ; egg limitation ; Hadena bicruris ; host deprivation ; time limitation]

1. An important constraint upon life-history evolution in parasitoids is the limit imposed by body size on allocation of limited metabolic resources to different fitness-related physiological functions such as reproduction and survival. 2. The influence of adult nutrition on reproductive and maintenance variables was studied in the synovigenic ectoparasitoid Mastrus ridibundus, and it was determined whether resource allocation to these different functions depends on body size. 3. Over the course of adult life there was a positive relationship between body size and the number of mature eggs in adult females both in the presence and absence of food. However, only in the presence of food did egg maturation rates increase significantly with body size. Starved wasps produced significantly smaller eggs than fed ones, which has not been documented before. Moreover, starved wasps produced fewer offspring than fed wasps, and attacked fewer hosts. 4. The availability of food had a major effect on longevity, with fed females living about 10 times longer than starved ones. There was also a positive relationship between body size and longevity. In starved wasps, this relationship was the same both in the presence and absence of hosts, but in fed wasps there was a positive relationship between body size and longevity in the absence of hosts only. Allocation to initial eggs relative to lifetime progeny production did not decline with body size 5. The data reveal that in M. ridibundus the trade-off between maintenance and reproduction varies with life expectancy. [KEYWORDS: Egg maturation ; egg production ; egg size ; reproductive strategies ; resource allocation ; sizefitness relationship ; synovigeny], 1. An important constraint upon life-history evolution in parasitoids is the limit imposed by body size on allocation of limited metabolic resources to different fitness-related physiological functions such as reproduction and survival. 2. The influence of adult nutrition on reproductive and maintenance variables was studied in the synovigenic ectoparasitoid Mastrus ridibundus, and it was determined whether resource allocation to these different functions depends on body size. 3. Over the course of adult life there was a positive relationship between body size and the number of mature eggs in adult females both in the presence and absence of food. However, only in the presence of food did egg maturation rates increase significantly with body size. Starved wasps produced significantly smaller eggs than fed ones, which has not been documented before. Moreover, starved wasps produced fewer offspring than fed wasps, and attacked fewer hosts. 4. The availability of food had a major effect on longevity, with fed females living about 10 times longer than starved ones. There was also a positive relationship between body size and longevity. In starved wasps, this relationship was the same both in the presence and absence of hosts, but in fed wasps there was a positive relationship between body size and longevity in the absence of hosts only. Allocation to initial eggs relative to lifetime progeny production did not decline with body size 5. The data reveal that in M. ridibundus the trade-off between maintenance and reproduction varies with life expectancy. [KEYWORDS: Egg maturation ; egg production ; egg size ; reproductive strategies ; resource allocation ; sizefitness relationship ; synovigeny]