Summary 1.Ecologists concerned with life-history strategies of parasitoid wasps have recently focused on interspecific variation in the fraction of the maximum potential lifetime egg complement that is mature when the female emerges into the environment. Species that have all of this complement mature upon emergence are termed 'pro-ovigenic', while those that do not are termed 'synovigenic'. We document and quantify the diversity of egg maturation patterns among 638 species of parasitoid wasps from 28 families. 2. We test a series of hypotheses concerning variation in 'ovigeny' and likely life-history correlates by devising a quantitative index -the proportion of the maximum potential lifetime complement that is mature upon female emergence. 3. Synovigeny, which we define as emerging with at least some immature eggs, was found to be by far the predominant egg maturation pattern (98·12% of species). Even allowing for some taxonomic bias in our sample of species, pro-ovigeny is rare among parasitoid wasps. 4. There is strong evidence for a predicted continuum in ovigeny index among parasitoid wasps, from pro-ovigenic (ovigeny index = 1) to extremely synovigenic species (ovigeny index = 0). 5. As predicted, synovigenic species are longer-lived than pro-ovigenic ones, and ovigeny index and life span are negatively correlated across parasitoid taxa, suggesting a life span cost of concentrating reproductive effort early in adult life. 6. There is equivocal evidence that host feeding (i.e. consumption of host haemolymph and /or tissues by adult wasps) is confined to synovigenic parasitoid wasps. It is also not certain from our analyses whether host feeding is associated with a relatively low ovigeny index. 7. As predicted, egg resorption capability is concentrated among producers of yolkrich eggs. Also, the hypothesis that it is associated with a tendency towards a low ovigeny index is supported. Parasitoid species that produce yolk-rich eggs also exhibit a lower ovigeny index than species that produce yolk-deficient eggs. 8. Ovigeny index appears to be linked to parasitoid development mode (koinobiosisidiobiosis). 9. We conclude that 'ovigeny' is a concept applicable to insects generally.

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.

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Immature development of parasitoid wasps is restricted to resources found in a single host that is often similar in size to the adult parasitoid. When two or more parasitoids of the same or different species attack the same host, there is competition for monopolization of host resources. The success of intrinsic competition differs between parasitoids attacking growing hosts and parasitoids attacking paralyzed hosts. Furthermore, the evolution of gregarious development in parasitoids reflects differences in various developmental and behavioral traits, as these influence antagonistic encounters among immature parasitoids. Fitness-related costs (or benefits) of competition for the winning parasitoid reveal that time lags between successive attacks influence the outcome of competition. Physiological mechanisms used to exclude competitors include physical and biochemical factors that originate with the ovipositing female wasp or her progeny. In a broader multitrophic framework, indirect factors, such as plant quality, may affect parasitoids through effects on immunity and nutrition.

Many species are currently moving to higher latitudes and altitudes. 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 effects 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 range. 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.

Summary1. 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 belowground 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.

Parasitoid wasps have long been considered as model organisms for examining optimal resource allocation to different fitness functions, such as body size and development time. Unlike insect predators, which may need to consume many prey items to attain maturity, parasitoids generally rely on a limited amount of resources that are obtained from a single source (the host). This review discusses a range of ecophysiological constraints that affect host quality and concomitantly the evolution of development strategies in parasitoids. Two macroevolutionary differences in host usage strategies (idiobiosis, koinobiosis) are initially described. Over many years, particular attention has been paid in examining a range of quantitative host attributes such as size, age, or stage, as these affect idiobiont and koinobiont parasitoid development. Parasitoids and their hosts, however, constitute only a small part of an ecological community. Consequently, host quality may be affected by a broad range of factors that may operate over variable spatial and temporal scales. Intimate factors include aggressive competition with other parasitoids and pathogens for access to host resources, whereas less intimate factors include the effects of toxic plant compounds (allelochemicals) on parasitoid performance as mediated through primary and/or secondary hosts. It is suggested that future experiments should increase the levels of trophic complexity as these influence the evolution of life history and development strategies in parasitoids. This includes integration of a suite of direct and indirect mechanisms, including biological processes occurring in different ecological realms, such as above-ground and belowground interactions.

Abstract. Populations of wild Brassica oleracea L. grow naturally along the Atlantic coastlines of the United Kingdom and France. Over a very small spatial scale (i.e., ,15 km) these populations differ in the expression of the defensive compounds, glucosinolates (GS). Thus far, very few studies have examined interactions between genetically distinct populations of a wild plant species and associated consumers in a multitrophic framework. Here, we compared the development of a specialist (Pieris rapae) and a generalist (Mamestra brassicae) insect herbivore and their endoparasitoids (Cotesia rubecula and Microplitis mediator, respectively) on three wild populations and one cultivar of B. oleracea under controlled greenhouse conditions. Herbivore performance was differentially affected by the plant population on which they were reared. Plant population influenced only development time and pupal mass in P. rapae, whereas plant population also had a dramatic effect on survival of M. brassicae. Prolonged development time in P. rapae corresponded with high levels of the indole GS, neoglucobrassicin, whereas reduced survival in M. brassicae coincided with high levels of the aliphatic GS, gluconapin and sinigrin. The difference between the two species can be explained by the fact that the specialist P. rapae is adapted to feed on plants containing GS and has evolved an effective detoxification system against aliphatic GS. The different B. oleracea populations also affected development of the endoparasitoids. Differences in foodplant quality for the hosts were reflected in adult size in C. rubecula and survival in M. mediator, and further showed that parasitoid performance is also affected by herbivore diet.