A comparative study of body size and clutch size across the parasitoid Hymenoptera

Across animal species, body size and clutch size often form part of a suite of associated life history traits, exemplified by the "fast-slow continuum" in mammals. Across the parasitoid Hymenoptera however, a major axis of life history variation is the development mode of the larva (koinobiosis versus idiobiosis), and body size and clutch size do not seem to form clear associations with this major axis. Here we use a large comparative data set and the latest phylogenetic information to explore hypotheses that might explain the variation in body size and clutch size across species in parasitoids. We find evidence for three novel evolutionary correlations: changes in the stage of host attacked by the parasitoid (i.e. egg, larva, pupa) significantly predict changes in both body size and clutch size, whilst in gregarious species changes to higher latitudes are associated with reduced clutch size. We also find a number of hypothesized cross-species (phenotypic) associations that, however, we cannot demonstrate are the result of evolutionary correlations: large bodied species in our data tend to lay small clutches; koinobionts are larger than idiobionts attacking the same host stage; tropical species are smaller than temperate species (Bergmann's rule). Our results provide support for theoretical models of trait evolution in parasitoids, whilst the associations between latitude and life history may help explain why species richness in the family Ichneumonidae peaks at intermediate latitudes. Our results also show the continuing value of phylogenetically-based comparative analyses and demonstrate that recent work on parasitoid phylogenetics has produced significant benefits for our understanding of life history evolution.     

Does parasitoid attack strategy influence host specificity? A test with New World braconids

Abstract.  1. Parasitoid attack strategy has been divided into two broad categories, koinobiosis and idiobiosis, based on the arrest of host development and the intimacy of larval contact. Koinobionts allow the host to continue developing and larvae usually feed within the host body, whereas idiobionts stop host development and larvae usually feed externally.
2. Comparisons of host ranges from rearings of parasitoids from specific host communities have shown that koinobionts are more host specific than idiobionts. These tests suggested that parasitoid attack strategy influenced specialisation in parasitoid–host interactions within certain host communities.
3. To determine whether this pattern was consistent within a single parasitoid lineage that utilises hosts from many different communities, the host ranges of koinobiont and idiobiont braconid genera of the New World were compared. Koinobiont genera utilised fewer host families than idiobionts, suggesting that parasitoid attack strategy may direct the evolution of host specificity throughout the evolutionary history of parasitoid lineages.     

Comparative life‐history traits in a fig wasp community: implications for community structure

1. Whether life‐history traits can determine community composition and structure is an important question that has been well explored theoretically, but has received scant empirical attention. Life‐history traits of a seven‐member community of galler and parasitoid fig wasp species (Chalcidoidea), developing within the inflorescences (syconia) of Ficus racemosa (Moraceae) in India, were determined and used to examine community structure and ecology. 2. Gallers were pro‐ovigenic (all eggs are mature upon adult emergence) whereas parasitoids were synovigenic (eggs mature progressively during adult lifespan). Initial egg load was correlated with body size for some species, and there was a trade‐off between egg number and egg size across all species. Although all species completed their development and left the syconium concurrently, they differed in their adult and pre‐adult lifespans. Providing sucrose solutions increased parasitoid lifespan but had no effect on the longevity of some galler species. While feeding regimes and body size affected longevity in most species, an interaction effect between these variables was detected for only one species. 3. Life‐history traits of wasp species exhibited a continuum in relation to their arrival sequence at syconia for oviposition during syconium development, and therefore reflected their ecology. The largest number of eggs, smallest egg sizes, and shortest longevities were characteristic of the earliest‐arriving galling wasps at the smallest, immature syconia; the converse characterised the later‐arriving parasitoids at the larger, already parasitised syconia. Thus life history is an important correlate of community resource partitioning and can be used to understand community structure. 4. This is the first comprehensive study of life‐history traits in a fig wasp community. The comparative approach revealed constraints and flexibility in trait evolution.     

Body size and the timing of egg production in parasitoid wasps

In insects several key fitness-related variables are positively correlated with intraspecific variation in body size, but little is known about size-related variation in the timing of egg production within species. Female insects are known to vary in the degree to which they concentrate egg production into the early part of life. This variation has been quantified as the ovigeny index, defined as the proportion of the maximum potential lifetime complement of eggs that is mature following emergence from the pupa. We tested the hypothesis that the timing of egg production depends both on body size and on host availability, by means of a dynamic programming model that predicted optimal resource allocation to reproduction and survival together with the resulting ovigeny index, in non-feeding synovigenic parasitoids of different sizes. As body size increases, the proportionate increase in resource allocation to initial egg load is less than the proportionate increase in allocation to lifetime fecundity and potential life span, leading to a deferred investment in reproduction as shown by a decrease in ovigeny index. High habitat quality and high habitat stochasticity in reproductive opportunities have a significant effect on the optimal allocation of resources to reproduction and survival, and thus select for early reproduction, i.e. an increased ovigeny index. The ovigeny concept – ovigeny index together with its life-history correlates – enables understanding of the general occurrence of size-related deferment of reproductive investment in parasitoid wasps and also helps explain a significant part of the considerable life-history variation found among such insects.     

Influence of aphid size,age and behaviour on host choice by the parasitoid wasp Ephedrus californicus: a test of host-size models

Summary When host quality varies, parasitoid wasps are expected to oviposit selectively in high-quality hosts. We tested the assumption underlying host-size models that, for solitary species of wasps, quality is based on host size. Using Ephedrus californicus, a solitary endoparasitoid of the pea aphid, we evaluated the influence of aphid size (= mass), age and defensive behaviours on host selection. Experienced parasitoid females were given a choice among three classes of 5-day-old apterous nymphs: small aphids that had been starved daily for 4 h (S4) and 6 h (S6) respectively, and large aphids permitted to feed (F) normally. Wasps attacked more, and laid more eggs in, small than large aphids (S6>S4>F). This rank-order for attack did not change when females could choose among aphids of the same size that differed in age; however, wasps oviposited in all attacked aphids with equal probability. Host size did not influence parasitoid attack rates when aphids were anaesthetized so that they could not escape or defend themselves. As predicted by host-size models, wasp size increased with host size (F>S4; S6), but large wasps required longer to complete development than their smaller counterparts (S4E. californicus reflects a trade-off between maximization of fitness gains per egg and the economics of search-time allocation. Because large aphids are more likely to escape parasitization, a wasp must balance her potential gain in fitness by ovipositinng in a high-quality (large) aphid against her potential cost in terms of lost opportunity time if the attack fails.     

Small body size in an insect shifts development, prior to adult eclosion, towards early reproduction

Life-history theory has suggested that individual body size can strongly affect the allocation of resources to reproduction and away from other traits such as survival. In many insects, adults eclose with a proportion of their potential lifetime egg production that is already mature (the ovigeny index). We establish for the solitary parasitoid wasp Aphaereta genevensis that the ovigeny index decreases with adult body size, despite both initial egg load and potential lifetime fecundity increasing with body size. This outcome is predicted by adaptive models and is the first unequivocal intraspecific demonstration. Evidence suggests that a high ovigeny index carries a cost of reduced longevity in insects. Our results therefore contribute to the emerging evidence that small body size can favour a developmental shift in juveniles that favours early reproduction, but which has adverse late-life consequences. These findings are likely to have important implications for developmental biologists and population biologists.     

Intrinsic, inter-specific competition between egg, egg–larval, and larval parasitoids of plusiine loopers

Abstract.  1. Intrinsic, inter-specific competition between parasitoid wasp species is a key factor in ecological community dynamics and is particularly important for application in biological control. Here three parasitoid wasp species with overlapping host ranges and differing life history strategies were chosen to examine parasitoid–parasitoid interactions: the egg parasitoid Trichogramma pretiosum, the egg–larval, polyembryonic parasitoid wasp Copidosoma floridanum, and the gregarious larval parasitoid Glyptapanteles pallipes , with the plusiine loopers Acanthoplusia agnata and Trichoplusia ni as hosts.
2.  Copidosoma floridanum has been shown to be an intrinsically superior competitor against larval parasitoids because of their production and increased investment in a soldier larval caste during development, but little is known of their interactions with egg parasitoid species. Trichogramma pretiosum completely dominated intrinsic competition with C. floridanum regardless of oviposition order or sex of the C. floridanum egg.
3. Competition between C. floridanum and G. pallipes , however, depended on the host stage at which parasitism occurred, the sex of the C. floridanum egg, and parasitoid development time. Copidosoma floridanum outcompeted G. pallipes overall, despite the fact that G. pallipes injects a polyDNA virus into the host.
4. The sex of the C. floridanum egg was a significant factor in its ability to shift caste ratios to produce more soldiers in response to G. pallipes competition.
5. Only developing female C. floridanum responded to competition with G. pallipes by increasing the ratio of soldier to reproductive larvae, and this happened only when multiparasitism occurred in the host's 1st and 2nd instar.     

Comparing and contrasting life history variation in four aphid hyperparasitoids

1. In primary parasitoids, significant differences in life history and reproductive traits are observed among parasitoids attacking different stages of the same host species. Much less is known about hyperparasitoids, which attack different stages of primary parasitoids. 2. Parasitoids exploit hosts in two different ways. Koinobionts attack hosts that continue feeding and growing during parasitism, whereas idiobionts paralyse hosts before oviposition or attack non‐growing host stages, e.g. eggs or pupae. 3. Koino‐/idiobiosis in primary parasitoids are often associated with different expression of life history trade‐offs, e.g. endo‐ versus ectoparasitism, high versus low fecundity and short versus long life span. 4. In the present study, life history parameters of two koinobiont endoparasitic species (Alloxysta victrix; Syrphophagus aphidivorus), and two idiobiont ectoparasitic species (Asaphes suspensus; Dendrocerus carpenteri) of aphid hyperparasitoids were compared. These hyperparasitoids attack either the parasitoid larva in the aphid before it is killed and mummified by the primary parasitoid or the parasitoid prepupa or pupa in the dead aphid mummy. 5. There was considerable variation in reproductive success and longevity in the four species. The idiobiont A. suspensus produced the most progeny by far and had the longest lifespan. In contrast, the koinobiont A. victrix had the lowest fecundity. Other developments and life history parameters in the different species were variable. 6. The present results reveal that there was significant overlap in life history and reproductive traits among hyperparasitoid koinobionts and idiobionts, even when attacking the same host species, suggesting that selection for expression of these traits is largely association specific.     

Does synovigeny confer reproductive plasticity upon a parasitoid wasp that is faced with variability in habitat richness?

Understanding the factors that constrain the reproductive success of animals and their demographics requires detailed insight into the processes of resource acquisition and allocation in relation to habitat richness. Parasitoid wasp females are valuable models in this respect because their lifetime reproductive success is closely tied to host availability. Parasitoids that manufacture eggs throughout adult life (i.e. ‘synovigenic’ species) and characteristically acquire nutrients via feeding are predicted to be plastic in their allocation to egg manufacture. Using the synovigenic parasitoid wasp Venturia canescens, we tested whether this prediction holds when females are faced with variation in the availability of both hosts and food. Laboratory experiments were conducted to determine how environmental variation affects parasitoid reproductive success and the lifetime dynamics of egg load and of major nutrient types. Our results, surprisingly, show that female V. canescens lacks a significant degree of reproductive plasticity under our experimental conditions. In particular, allocation of resources to reproduction was high irrespective of host availability. We attribute this lack of flexibility to the low energy content of V. canescens' eggs and to features peculiar to the ecology of this species. Our findings shed new light on the physiological factors that constrain parasitoid lifetime reproductive success. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 621–632.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.     

Ontogeny and reproductive biology of Diadegma semiclausum (Hym.: Ichneumonidae), a larval endoparasitoid of Diamondback Moth,Plutella xylostella (Lep.: Plutellidae)

In this article, the ontogeny and reproductive biology of Diadegma semiclausum (Hym.: Ichneumonidae), an important parasitoid of Plutella xylostella (Lep.: Plutellidae) are described in detail. We did dissect parasitized P. xylostella larvae in phosphate-buffered saline and determine the external morphology of its parasitoid at all developmental stages. The developmental duration of its immature stages, adult longevity, total oviposition period and fecundity of the parasitoid are determined at 24 ± 1 °C, 65  ±  5% R.H., and a photoperiod of 16:8 h L:D. The mean duration of egg and larval stages is 9.56 days and the prepupa and pupa stages last for 8.27 days. In average, female longevity is 1.31 times longer than that of males, and females lay 300 eggs in total. The peak of D. semiclausum oviposition is on the eighth day after mating. The egg loading pattern of D. semiclausum was investigated to determine the parasitoid ‘ovigeny index’ throughout the female's parasitoid lifetime. Initial egg load in immature females (<2 h age) is 1.45 per female and the mean lifetime potential fecundity (total immature and mature oocytes), at four interval ages, is 34. With an ovigeny index value of 0.038, D. semiclausum is considered moderately to strongly synovigenic. In the absence of the host, after 3 days, the number of eggs is decreased in D. semiclausum. Our results demonstrated that there is a negative relation between the ovigeny index and egg resorption in this parasitoid.     

Linking reproductive and feeding strategies in the parasitoid Ibalia leucospoides: does feeding always imply profit?

Resource acquisition and allocation to different biological functions over the course of life have strong implications for animal reproductive success. Animals can experience different environmental conditions during their lifetime, and this may play an important role in shaping their life-history and resource allocation strategies. In this study we investigate larval and adult resource allocation to reproductive and survival functions in the parasitoid wasp Ibalia leucospoides (family Ibaliidae). The pattern of larval resource allocation was inferred from the relationship between adult body size and ovigeny index (OI; a relative measure of investment in early reproduction determined as the ratio between the initial egg load and the potential lifetime fecundity); and adult resource allocation was explored through the influence of adult feeding on reproduction, maintenance and metabolism, in laboratory experiments. Food acquisition by this parasitoid in the wild was also examined. The relationship between size and OI was constant, suggesting no differential resource allocation to initial egg load and potential lifetime fecundity with size. This finding is in line with that predicted by adaptive models for the proovigenic egg maturation strategy (OI = 1). Despite of this, I. leucospoides showed a high OI of 0.77, which places this species among the weakly synovigenic ones (OI < 1). Adult feeding had no effect on post-emergence egg maturation. However, wasps extended their lifespan through feeding albeit only when food was provided ad libitum. Although the information we obtained on the feeding behaviour of free-foraging wasps is limited, our results suggest that food intake in the wild, while possible, may not be frequent in this parasitoid. We discuss the results relative to the environmental factors, such as reproductive opportunities and food availability, which may have driven the evolution of larval and adult pattern of resource allocation in parasitoids.     

Latitudinal gradients and the shaping of life‐history traits in a gregarious caterpillar

The present study aimed to investigate how the impact of several factors linked to geography would shape life‐history traits in a gregarious species, using the pine processionary moth (PPM) Thaumetopoea pityocampa as a model system. PPM has a wide geographical distribution over the Mediterranean Basin, and it is a strictly gregarious species throughout larval development, where the total reproductive output of each female forms a colony. We reviewed both published and unpublished data on PPM from all over its distribution in the Mediterranean Basin and extracted data on fecundity, egg size, egg parasitoid mortality, flight period, and development time. These life‐history traits were then related to location, expressed as latitude and altitude, local average temperatures, and host tree species. We found that PPM fecundity increaseed with latitude, concomitant with an increase in the length of development and an earlier onset of adult flight. These results are the opposite of that found in other Lepidoptera species with a wide geographical distribution, as well as in insects in general. We propose that a large colony size in PPM is important at higher latitudes because this confers an advantage for thermoregulation and tent building in areas where larvae have to face harsher conditions during the winter, thus shifting the optimal trade‐off between the number and size of eggs with latitude. However, host tree species also affected the relationship between egg number and size and the optimal outcome of these traits is likely a compromise between different selection pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 224–236.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

白蜡吉丁柄腹茧蜂的寄生和繁殖生物学

白蜡吉丁柄腹茧蜂Spathius agrili Yang是我国天津地区寄生白蜡窄吉丁幼虫的一种重要天敌。本研究通过林间调查并结合室内观察,测定了该寄生蜂在不同营养条件下的存活时间和不同时期的产卵量,调查了不同寄主大小对寄生蜂后代生长发育的影响,以及寄生蜂对不同深度树皮下的寄主利用情况。结果表明：不同营养条件下白蜡吉丁柄腹茧蜂的存活时间差异很大,补充20％蜂蜜水的成蜂存活时间明显延长,最长可达3个月。相同条件下雌蜂的存活时间比雄蜂略长。在补充营养的条件下,茧蜂的产卵时间最长可达8周,产卵的高蜂在羽化后第2周前后,平均每雌每周产卵量在整个产卵期内基本保持在8～12粒上下,没有大的波动。茧蜂低龄幼虫也可能分泌毒素主动攻击寄主,提高了寄生蜂后代的生存能力。寄主个体大,营养量也大,寄生其上的茧蜂幼虫成功完成发育的数量随之增加,表明寄主营养资源的多少直接影响到寄生蜂后代的生长发育和存活。白蜡吉丁柄腹茧蜂对1～6.5 mm树皮深度范围内的适龄寄主幼虫均能够寄生利用,约92％的被寄生白蜡窄吉丁幼虫个体分布于1～4 mm深度的树皮下,在3 mm厚的树皮下分布的数量最多。茧蜂数量与树皮深度大致上呈抛物线关系,理论上在树皮深度为3.97 mm处茧蜂分布数量最多。     

Development of the solitary endoparasitoid Microplitis demolitor: host quality does not increase with host age and size

Abstract.  1. Many studies examining the relationship between host size, an index of host quality, and parasitoid fitness use development time and/or adult parasitoid size as currencies of fitness, while ignoring pre-adult mortality. Because the physiological suitability of the host may vary in different stages, sizes, or ages of hosts, a misleading picture of host quality may therefore be obtained in cases where fitness is based on only one or two developmental traits.
2. The development of the solitary koinobiont endoparasitoid Microplitis demolitor is examined in different larval age-classes of its host the soybean looper Pseudoplusia includens . Hosts were parasitised on days 1–8 after hatching from the egg, and development time, adult body size, and mortality of the parasitoid were compared.
3. A comparison of larval growth trajectories (using dry body mass) of M. demolitor revealed that parasitoid larvae attained over twice as much body mass in old hosts than in younger hosts. Similarly, adult parasitoid size at eclosion generally increased with host size, although parasitoids developing in smaller hosts lost a much lower proportion of mass between pupation and eclosion.
4. Overall egg-to-adult development was most rapid in intermediate-aged hosts, and longer in hosts at opposite ends of the age continuum. Moreover, parasitoid mortality varied non-linearly with host stage, and was generally higher in very young and older hosts.
5. Based on these results and other empirical data for koinobionts, it is argued that fitness functions in this group of parasitoids are not simply a positive function of host size or age, but instead may be distinctly dome-shaped, both patterns reflecting the degree of physiological and nutritional compatibility between the two organisms.     

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

Dose-dependent influence of Microplitis rufiventris factors on developmental rate and growth of last-instar Spodoptera littoralis larvae

The solitary endoparasitoid, Microplitis rufiventris, attacks and can develop in earlier instars of Spodoptera littoralis larvae with preference to third‐instar larvae. We used the last stadium (sixth instar), a stage which is not naturally parasitized. The newly moulted larvae (0–3 h old) of this stadium were more acceptable for parasitization by the wasp females than the older ones (24 h old). Parasitization by M. rufiventris wasp of last instar S. littoralis larvae leads to dose (no. of eggs + parasitoid factors)‐dependent effects which were more pronounced at 20°C than at 27°C. A single oviposition into a sixth instar host larva resulted in normal development of the host. However, superparasitization increased the proportions of developmentally arrested hosts and number of live wasp larvae. Development of supernumerary individuals of the parasitoid in the host larva leads to dose‐related adverse effects on host growth and development. The present study may provide interesting opportunities for studying the physiological bases of host–parasitoid interactions and parasitoid intra‐specific competition in the biological system considered.     

Endogenous and exogenous factors affecting parasitism of gypsy moth egg masses by shape Ooencyrtus kuvanae

Factors affecting the orientation, reproduction, and sex ratio of the egg parasitoid Ooencyrtus kuvanae Howard were examined. Adult females were attracted to airborne volatiles from the egg mass and accessory gland of the primary host, the gypsy moth Lymantria dispar L. Visual cues also affected host selection. Background colors against which egg masses were placed affected oviposition preference. In the absence of egg masses, color variation did not affect wasp behavior. Light is required for parasitism by O. kuvanae. The age and density of both the host and parasitoid affected wasp reproduction and sex ratios. Older egg masses issued relatively fewer wasps and higher proportions of males than did young egg masses. Likewise, wasp reproduction and the proportion of females declined with wasp age. Larger egg masses produced more wasps and lower proportions of males than did smaller egg masses. The number of offspring per female, and the proportion of female offspring, were inversely related to wasp density. Implications to biological control of the gypsy moth and parasitoid ecology are discussed.