Presence of a novel small RNA-containing virus in a laboratory culture of the endoparasitic wasp Venturia canescens (Hymenoptera: Ichneumonidae)

Parasitoid wasps use a variety of mechanisms to alter their host's physiology to the benefit of the developing endoparasite inside the host larva. Association of certain wasps with viruses and virus-like particles (VLPs) that contribute to their success in parasitism is one of the fascinating evolutionary adaptations conferring active or passive protection for the endoparasite from the host immune system. Venturia canescens has been shown to produce VLPs that provide protection for the developing parasitoid egg inside the host, Ephestia kuehniella. Here, we report on the presence of a novel small RNA-containing virus from V. canescens, designated as VcSRV, occurring in the ovaries of the wasp. The virus particles are found together with VcVLPs in the lumen of the calyx region of the ovaries and are injected together with the egg and VcVLPs into E. kuehniella larvae where they enter hemocytes. Alignment of the RNA-dependent RNA polymerase gene of VcSRV indicates that the virus most likely belongs to the recently described genus Iflavirus.     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

Differential host growth regulation by the solitary endoparasitoid, Meteorus pulchricornis in two hosts of greatly differing mass

Solitary koinobiont endoparasitoids generally reduce the growth of their hosts by a significant amount compared with healthy larvae. Here, we compared the development and host usage strategies of the solitary koinobiont endoparasitoid, Meteorus pulchricornis, when developing in larvae of a large host species (Mythimna separata) and a much smaller host species (Plutella xylostella). Caterpillars of M. separata were parasitized as L2 and P. xylostella as L3, when they weighed approximately 2 mg. The growth of parasitized M. separata larvae was reduced by almost 95% compared with controls, whereas parasitized P. xylostella larvae grew some 30% larger than controls. Still, adult wasps emerging from M. separata larvae were almost twice as large as wasps emerging from P. xylostella larvae, had larger egg loads after 5 days and produced more progeny. Survival to eclosion was also higher on M. separata than on P. xylostella, although parasitoids developed significantly faster when developing on P. xylostella. Our results provide evidence that koinobionts are able to differentially regulate the growth of different host species. However, there are clearly also limitations in the ability of parasitoids to regulate phenotypic host traits when size differences between different host species are as extreme as demonstrated here.     

Inhibition of juvenile hormone esterase activity in Lymantria dispar (Lepidoptera, Lymantriidae) larvae parasitized by Glyptapanteles liparidis (Hymenoptera, Braconidae)

Glyptapanteles liparidis is a gregarious, polydnavirus (PDV)-carrying braconid wasp that parasitizes larval stages of Lymantria dispar. In previous studies we showed that parasitized hosts dramatically increase juvenile hormone (JH) titers, whereas JH degradation is significantly inhibited in the hemolymph. Here we (i) quantified the effects of parasitism on JH esterase (JHE) activity in hemolymph and fat body of penultimate and final instars of L. dispar hosts and (ii) assessed the relative contribution of individual and combined wasp factors (PDV/venom, teratocytes, and wasp larvae) to the inhibition of host JHE activity. The effects of PDV/venom was investigated through the use of gamma-irradiated wasps, which lay non-viable eggs (leading to pseudoparasitization), while the effects of teratocytes and wasp larvae were examined by injection or insertion of these two components in either control or pseudoparasitized L. dispar larvae. Parasitism strongly suppressed host JHE activity in both hemolymph and fat body irrespective of whether the host was parasitized early (premolt-third instar) or late (mid-fourth instar). Down-regulation of JHE activity is primarily due to the injection of PDV/venom at the time of oviposition, with only very small additive effects of teratocytes and wasp larvae under certain experimental conditions. We compare the results with those reported earlier for L. dispar larvae parasitized by G. liparidis and discuss the possible role of JH alterations in host development disruption.     

Development of the parasitoid,Cotesia rubecula (Hymenoptera: Braconidae) in Pieris rapae and Pieris brassicae (Lepidoptera: Pieridae): evidence for host regulation

Several recent models examining the developmental strategies of parasitoids attacking hosts which continue feeding and growing after parasitism (=koinobiont parasitoids) assume that host quality is a non-linear function of host size at oviposition. We tested this assumption by comparing the growth and development of males of the solitary koinobiont endoparasitoid, Cotesia rubecula, in first (L1) to third (L3) larval instars of its preferred host, Pieris rapae and in a less preferred host, Pieris brassicae. Beginning 3 days after parasitism, hosts were dissected daily, and both host and parasitoid dry mass was determined. Using data on parasitoid dry mass, we measured the mean relative growth rate of C. rubecula, and compared the trajectories of larval growth of the parasitoid during the larval and pupal stages using non-linear equations. Parasitoids generally survived better, completed development faster, and grew larger in earlier than in later instars of both host species, and adult wasps emerging from P. rapae were significantly larger than wasps emerging from all corresponding instars of P. brassicae. During their early larval stages, parasitoids grew most slowly in L1 P. rapae, whereas in all other host classes of both host species growth to pupation proceeded fairly uniformly. The growth of both host species was markedly reduced after parasitism compared with controls, with the development of P. brassicae arrested at an earlier stage, and at a smaller body mass, than P. rapae. Our results suggest that C. rubecula regulates certain biochemical processes more effectively in P. rapae than in P. brassicae, in accordance with its own nutritional and physiological requirements. Furthermore, we propose that, for parasitoids such as C. rubecula, which do not consume all host tissues prior to pupation, that parasitoid size and host quality may vary independently of host size at oviposition and at larval parasitoid egression.     

Transient host paralysis as a means of reducing self-superparasitism in koinobiont endoparasitoids

The term ‘idiobiont’ refers to those parasitoid species that permanently paralyse their hosts during parasitism, causing the cessation of host growth and development. This is in contrast to koinobiont parasitoids, which allow their hosts to continue developing after being parasitized. While no koinobiont species induce permanent paralysis in their hosts, a minority of koinobionts induce a temporary paralysis that does not interfere with overall host growth and development. We characterized transient paralysis induction in two koinobiont aphid parasitoids in the genus Binodoxys (Hymenoptera: Aphidiinae). Both Binodoxys species induced transient paralysis in Aphis glycines, with paralysis time ranging between 4.5 and 8 min (depending upon parasitoid species and host instar). In a separate experiment, B. communis was capable of inducing transient paralysis in nine aphid species. We addressed two hypotheses potentially explaining the adaptive value of temporary host paralysis in experiments using A. nerii, which is readily accepted but engages in strong defensive behaviour. The first hypothesis is that paralysis increases oviposition success by interfering with host defences and the second is that it aids in the avoidance of self-superparasitism. Paralysed aphids were more likely to be rejected by B. communis than were aphids that had never been stung or that had recovered from paralysis. This result supports the avoidance-of-self-superparasitism hypothesis and is inconsistent with the hypothesis that transient paralysis increases oviposition success of B. communis.     

Factors influencing parasitism of Sirex noctilio (Hymenoptera: Siricidae) by the nematode Deladenus siricidicola (Nematoda: Neotylenchidae) in summer rainfall areas of South Africa

Control of the invasive wasp, Sirex noctilio Fabricius using the parasitic nematode Deladenus siricidicola Bedding is a well known example of a successful classical biological control program. Despite its wide-scale success, this control method has recently had poor success in the summer rainfall areas of South Africa. Data from previous studies showed variation in nematode parasitism from inoculated trees (inoculation success) between different tree sections and among inoculation times. They also pointed to moisture content of the wood or virulence of the nematode as the most likely underlying factors influencing variations in inoculation success. The results from our study showed that the highest levels of parasitism was obtained from early inoculations and from the bottom sections of trees, where moisture content of the wood was highest, supporting the hypothesis that moisture content influences parasitism. However, even when moisture content was adequate, average inoculation success remained below 25% and was often 0%, suggesting that there are other barriers to inoculation success. Different sources from which the nematodes were produced did not influence inoculation success, indicating that nematode virulence is most likely not the cause of the low success. Another interesting finding was that parasitized wasps were larger than unparasitized wasps. Background parasitism was present despite the poor success with past inoculations, but the data also suggest that the natural build-up of this population could be constrained by the same factors that influence inoculations.     

Defence strategies against a parasitoid wasp in Drosophila: fight or flight?

Hosts may defend themselves against parasitism through a wide variety of defence mechanisms, but due to finite resources, investment in one defence mechanism may trade-off with investment in another mechanism. We studied resistance strategies against the parasitoid wasp Leptopilina boulardi in two Drosophila species. We found that D. melanogaster had significantly lower physiological resistance against L. boulardi than D. simulans, and hypothesized that D. melanogaster might instead invest more heavily in other forms of defence, such as behavioural defence. We found that when given a choice between clean oviposition sites and sites infested with wasps, both D. melanogaster and D. simulans detected and avoided infested sites, which presumably limits later exposure of their offspring to infection. Unlike D. simulans, however, D. melanogaster laid significantly fewer eggs than controls in the forced presence of wasps. Our findings suggest that D. melanogaster relies more heavily on behavioural avoidance as defence against wasp parasitism than D. simulans, and that this may compensate for a lack of physiological defence.     

Nutritional ecology of the interaction between larvae of the gregarious ectoparasitoid, Muscidifurax raptorellus (Hymenoptera: Pteromalidae), and their pupal host, Musca domestica (Diptera: Muscidae)

In this study we examined the relationship between clutch size and parasitoid development of Muscidifurax raptorellus (Hymenoptera: Pteromalidae), a gregarious idiobiont attacking pupae of the housefly, Musca domestica (Diptera: Muscidae). Host quality was controlled in the experiments by presenting female parasitoids with hosts of similar size and age. This is the first study to monitor the development of a gregarious idiobiont parasitoid throughout the course of parasitism. Most female wasps laid clutches of one to four eggs per host, although some hosts contained eight or more parasitoid larvae. In both sexes, parasitoids completed development more rapidly, but emerging adult wasp size decreased as parasitoid load increased. Furthermore, the size variability of eclosing parasitoid siblings of the same sex increased with clutch size. Irrespective of clutch size, parasitoids began feeding and growing rapidly soon after eclosion from the egg and this continued until pupation. However, parasitoids in hosts containing five or more parasitoid larvae pupated one day earlier than hosts containing one to four larvae. The results are discussed in relation to adaptive patterns of host utilization by gregarious idiobiont and koinobiont parasitoids.     

A parasitoid wasp uses landmarks while monitoring potential resources

Social insects and insects that provision nests are well known to have complex foraging behaviour involving repeated visits to learned locations. Other insects do not forage from a central location and are generally assumed to respond to resources by simple attraction without spatial memory. This simple response to resource cues is generally taken as giving rise to patterns of resource use that correspond directly to resource distribution. By contrast, the solitary parasitoid wasp Hyposoter horticola monitors the locations of multiple potential hosts (butterfly eggs) for up to several weeks, until the hosts become susceptible to parasitism. Essentially all hosts in the landscape are found, and one-third of them are parasitized, independent of host density. Here, we show that the wasps do not relocate hosts using odour markers previously left by themselves or other foragers, nor do they find the eggs anew repeatedly. Instead, the wasps relocate host eggs by learning the position of the eggs relative to visual landmarks. The anticipatory foraging behaviour presented here is a key to the wasp's exceptionally stable population dynamics.     

Is the host or the parasitoid in control?: effects of host age and temperature on pseudoparasitization by Microplitis rufiventris in Spodoptera littoralis

We compared the production of pseudoparasitization by Microplitis rufiventris females in most (third) and less (fourth) preferred instars of Spodoptera littoralis larvae at 20+/-1 and 27+/-1 degrees C. The parasitized hosts were classified into hosts producing parasitoids (type A hosts) and hosts producing no parasitoids, i.e., pseudoparasitized hosts (type B hosts). The latter were further classified into: (a) pseudoparasitized hosts with "well" arrested development (type B1 hosts); (b) pseudoparasitized hosts with partially arrested development (type B2 hosts); and (c) pseudoparasitized hosts that successfully pupated to apparently normal host pupae (type B3 hosts). The present series of experiments showed that parasitization by M. rufiventris was clearly affected by host instar, age within an instar and rearing temperature. Production of type B hosts was less when third instar S. littoralis larvae were exposed to the wasp females than when the host larvae were in fourth instar. The production of type A hosts was much greater when early or mid ages of an instar was stung by the wasp females comparing with stung late age of the same instar. Production of type B hosts may be due to one or overall of the following: (a) dosage dilution of M. rufiventris female's factors in the different age classes of the instar; (b) endocrine system (physiological state) at parasitization time, i.e., early vs late age of the instar; (c) growth rate of host larvae. The lowest production of type B hosts was at highest growth rate; and (d) temperature, larger proportions of type B hosts were produced at 27+/-1 than at 20+/-1degrees C. The three types host development (B1, B2 and B3) are possibly representing three levels of host resistance (host control) resulting in partial or complete failure of parasitoid control. Type A hosts represent complete success of parasitoid control. The results suggest that the impact of parasitoid factor(s) on developmental arrest is affected by host age at the time of parasitism and/or by temperature.     

Spatial distribution patterns of three fig wasps on Ficus semicordata: How non-pollinators affect pollinator’s sex ratio

Sex ratio theory is one of the most productive fields in research on evolutionary biology. Pollinating fig wasps, due to their particular natural life history, are considered to be a valuable model for the study of sex ratio evolution. A great deal of research concerning the factors that affect pollinator fig wasp (Agaonidae) progeny sex ratio has been done, and at present three main factors (haplodiploidy, local mate competition and inbreeding) are found to be important at the population level. However, there still exists variation between empirical data and model predictions. Another factor to which little thought has been given before is the effect of non-pollinating fig wasps (NPFWs) which parasitize in the larvae gall of pollinator thus kill pollinators and exploit the fig/fig pollinator mutualistic systems. In this study, we focus on why and how non-pollinating fig wasps distort pollinator fig wasp’s original sex ratio. Through controlling the number of ovipositing foundresses inside a fig, combined with the observation of ovipositing behavior and sequence, we studied three species of wasp in the figs of a dioecious fig Ficus semicordata including the pollinator Ceratosolen gravely and NPFWs Platyneura cunia, Sycoscapter trifemmensis in tropical area of Xishuangbanna from September to December 2009. First, we observed the timing of oviposition of all fig wasps utilizing F. semicordata and found differences when compared to previous studies. Such as P. cunia is the fourth rather then the secondary fig wasps to oviposit on the syconia approximately 10 days after the pollinator. S. trifemmensis oviposits much earlier than previously thought, 14–32 days after the pollinators. We examined the spatial location of male and female progeny of the pollinator. We found foundresses of pollinator prefer to use innermost ovules first. Only at high offspring numbers were the outer ovules used. More male pollinator offspring were developed near the fig cavity, while female pollinator offspring were more evenly distributed among ovule layers. As pollinator offspring numbers increased, this phenomenon became more pronounced. This pattern of segregation of male larvae gall in inner ovules and female larvae gall in outer ovules suggests that female offspring might be more vulnerable to attack by parasitic wasps that oviposit from outside the syconium. Experiments later demonstrated that NPFWs are restricted by their ovipositor length and they prefer to or can only lay their eggs into ovules near the fig wall. Then we examined the spatial location of NPFWs and compared this with the spatial location of male/female progeny of pollinator. NPFWs had a high probability of parasitizing female pollinator larvae. Thus, NPFWs have a substantial effect on the sex ratio of the pollinator, as parasitism risk decreases towards the center of the syconium, where inner ovules provide enemy-free space for most of male pollinator offspring. Partial correlation analyse shows that sex ratio of pollinator progeny has a positive relationship with the number of NPFWs. We suggest that the resulting gradient in offspring viability between male and female contributes to selection on pollinators’ for a less femalebiased sex ratio. When the affect of NPFWs was excluded, the pollinator sex ratio was not in good agreement with local mate competition theory, although it was still female-biased. In addition, the average number of offspring per foundress decreased with increasing foundress number, but pollinator sex ratio was positively related to brood size. Thus, pollinator females do not appear to adjust their sex ratio to foundress density directly, but use brood size and foundress density simultaneously as cues to assess potential LMC.     

Wasp behavior leads to uniform parasitism of a host available only a few hours per year

The parasitoid wasp, Hyposoter horticola, parasitizes a nearlyfixed fraction of its host butterfly larvae within a host metapopulationof 300–500 local populations in a 50 x 70-km area. Weshow, through laboratory observation, that the wasp lays eggsin fully developed larvae that have not yet hatched from theegg, constraining the period of host vulnerability to severalhours out of the host's one year lifecycle. The parasitoid achievesa persistent high rate of parasitism over the entire host rangedespite the extremely limited period of host vulnerability aswell as a high rate of host population extinctions and colonizationsof new habitat patches every year. It does this in part by beingextremely mobile. In addition, we show by using a field experimentand observation of marked wasps foraging for hosts in naturalpopulations, that the wasp finds virtually all host egg clustersin the weeks before the hosts become vulnerable to parasitism,and then later returns to parasitize them. By locating the hostsbefore their vulnerability, the wasp extends the time availablefor searching from hours to weeks. After parasitizing aboutone-third of the larvae in a host cluster the wasp stops, apparentlyleaving a mark that deters further parasitism by other individuals.The result of this novel combination of mobility and local foragingbehavior is a stable population size despite an unstable hostthat is vulnerable during about one thousandth of its lifecycle.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

高榕榕果内Eupristina属两种榕小蜂的遗传进化关系

对生长在福州地区的高榕进行长期追踪观察,发现高榕榕果内仅生活着Eupristina altissima和Eupristina sp.榕小蜂,前者为高榕的传粉小蜂,后者无传粉行为,两者雌蜂之间在体色、触角、花粉袋和花粉刷等部位存在细微的差异,而两者雄蜂之间无形态差异。通过克隆福建地区5个样地的高榕榕果内收集到的E. altissima和Eupristina sp.榕小蜂,以及细叶榕的传粉小蜂Eupristina verticillata(外群)的Cytb及COI基因,并进行碱基组成及遗传距离分析,用邻接法构建系统发育树,分析两榕小蜂群体之间的遗传进化关系,结果显示:(1)榕小蜂COI及Cytb序列碱基组成中A+T的含量(Cytb序列中A+T=75.3%,COI序列中A+T=75.5%)显著高于G+C,符合膜翅目昆虫线粒体基因碱基组成特征。(2)对两群体小蜂进行遗传距离分析显示,Cytb序列中E. altissima和Eupristina sp. 群体内各样本之间的平均遗传距离分别为0.0092和0.0030,而E. altissima与Eupristina sp. 群体间的平均距离为0.1588;COI序列中E. altissima 与Eupristina sp. 群体内各样本之间的平均遗传距离分别为0.0065和0.0205,而二者群体间的平均遗传距离为0.1043,表明两者群体间的遗传距离明显大于各自群体内各样本间的遗传距离。统计GenBank中下载的6个属34种榕小蜂Cytb序列的种间遗传距离为0.0811-0.1723,6个属28种榕小蜂COI序列的种间遗传距离为0.0939-0.1986。由此认为E. altissima和Eupristina sp.之间的遗传距离差异已经达到了种间水平,即E. altissima与Eupristina sp.为两个不同的种。(3)在形态上,两种小蜂的雌蜂之间有微小差异,而二者雄蜂之间无差异,但Cytb与COI序列分析结果一致表明:E. altissima与Eupristina sp.雄蜂之间,以及二者雌蜂之间的遗传距离均差异显著,表明形态变异滞后于基因变异。雌蜂在表型上进化快于雄蜂,可能是由于雌蜂羽化后从榕果出飞,受到外界环境因素的影响较大,且两种雌蜂在传粉功能上存在差异,故二者之间的形态差异较大,而雄蜂寿命短,又终生生活在黑暗封闭、环境变化相对恒定的榕果内,两种雄蜂在行为上不存在差异,故二者表型变异较为缓慢。E. altissima和Eupristina sp.小蜂对宿主的专一性不强,在榕-蜂协同进化过程中,可能发生过宿主转移事件。     

Catching Ariadne by her thread: how a parasitoid exploits the herbivore's marking trails to locate its host

Chemical signals that can be associated with the presence of a host insect often work as arrestants in close range host location by parasitoids, leading to longer searching times on patches where such signals are present. Our current view of parasitoid host location is that by prolonging the search times in patches, randomly searching parasitoids enhance their chance of detecting host insects. However, prolonged search times are not necessarily the only modification in parasitoid behaviour. In this study, we examine the exploitation of host-fruit marking pheromone of rose-hip flies, Rhagoletis basiolaOsten-Sacken (Diptera: Tephritidae) by the specialized egg-larval parasitoid Halticoptera rosae Burks (Hymenoptera: Pteromalidae). We provide evidence that the instantaneous probability that a host egg will be located by a searching parasitoid wasp differs markedly between pheromone-marked and unmarked fruits. The arresting response to the marking pheromone, i.e., the prolonged time a wasp is willing to search on marked fruits, can only account for a small fraction of the difference in successful host location on marked and unmarked fruits. We further demonstrate that the time wasps require to locate the host egg is independent of the size of the rose-hip harbouring the fly egg, and thus is independent of the area the wasp potentially has to search. A comparison of our findings with results of different search algorithms for parasitoid wasps suggests that wasps use the fly's pheromone marking trail as a guide way to the fly's oviposition site and thus the host egg.     

Effect of host‐cocoon mass on adult size in the secondary hyperparasitoid wasp,Pteromalus semotus (Hymenoptera: Pteromalidae)

Abstract Parasitoids have long proven to be model organisms in studying resource‐related constraints on immature development. Here we examine the relationship between host cocoon (= pupal) size in the gregarious endoparasitoid wasp, Cotesia glomerata, and development time and adult size in the solitary idiobiont hyperparasitoid, Pteromalus semotus. Little is known about the biology or ecology of this ecto‐hyperparasitoid species, although it is one of the major secondary hyperparasitoids of C. glomerata. The size of the adult wasp covaried with the size of the host cocoon at parasitism. Moreover, female wasps were larger than male wasps for a given cocoon size. Adult wasps have remarkably long life‐spans, 3 months on average. Longevity did not significantly differ with sex. We also examined how larvae of P. semotus exclude other potential competitors. P. semotus is protandrous, with females taking significantly longer to complete their development than males. In experiments where several eggs of P. semotus were placed on individual pupae of C. glomerata, newly hatched hyperparasitoid larvae moved rapidly over the surface of the host and destroyed the eggs of any conspecifics by biting them before they would initiate feeding on host tissues. Our results are discussed in relation to those with other studies with solitary ichneumonid idiobiont hyperparasitoids of C. glomerata.     

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.