Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Host selection by virgin and inseminated females of the parasitic wasp, Dinarmus basalis (Pteromalidae, Hymenoptera)

1. Fitness is related to reproduction and survival. There apparently exists a negative correlation between the numbers of male and female offspring. There also exists a trade-off between survival and reproduction. This paper investigates optimal decisions with the reproduction and survival trade-off in host selection by wasps.
2. Whereas inseminated female wasps could manipulate the sex of their offspring, virgin females produced only male offspring. I surveyed behavioural differences and the consequences of oviposition by inseminated and virgin females of a solitary parasitic wasp in host choice situations.
3. Two host types were available at the same time to both inseminated and virgin female wasps: one (a 17-day-old host in one bean) presenting difficulties for the laying of eggs, but more benefits for the offspring and the other (five 12- or 13-day-old hosts in one bean) easier for the female wasp for laying of eggs but less beneficial for the offspring.
4. Inseminated female wasps chose more 17-day-old hosts than 12-day-old hosts, but more 13-day-old hosts than 17-day-old hosts in each pair-wise choice. Virgin females chose the smaller hosts in both situations.
5. Virgin females, having greater longevity than inseminated females, laid larger numbers of eggs than the inseminated females during their lifetime by adopting an energy-saving host choice that had little effect on male offspring fitness.     

Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size

Adaptiveness of sex ratio control by the solitary parasitoid wasp Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size was studied, by examining whether differential effects of host size on the fitness of resulting wasps are to be found between males and females. The offspring sex ratio (male ratio) decreased with increasing host size. Larger hosts yielded larger wasps. Male larvae were less efficient in consuming larger hosts than female larvae. No significant interaction in development time was found between parasitoid sex and host size. Larger female wasps lived longer than smaller females, while longevity of male wasps did not increase with increasing wasp size. Smaller males were able to mate either with small or with large females, while larger males failed to mate with small females. Larger female wasps had a greater number of ovarioles and mature eggs at any one time than smaller females, although the number of eggs produced per host-feeding was not influenced by female wasps. Thus, the differential effect of host size on the fitness of males and females exists in I. naranyae. The basic assumption of the host-size model was therefore satisfied, demonstrating that sex ratio control by I. naranyae in response to host size is adaptive.     

Evolution of behavioural and cellular defences against parasitoid wasps in the Drosophila melanogaster subgroup

It may be intuitive to predict that host immune systems will evolve to counter a broad range of potential challenges through simultaneous investment in multiple defences. However, this would require diversion of resources from other traits, such as growth, survival and fecundity. Therefore, ecological immunology theory predicts that hosts will specialize in only a subset of possible defences. We tested this hypothesis through a comparative study of a cellular immune response and a putative behavioural defence used by eight fruit fly species against two parasitoid wasp species (one generalist and one specialist). Fly larvae can survive infection by melanotically encapsulating wasp eggs, and female flies can potentially reduce infection rates in their offspring by laying fewer eggs when wasps are present. The strengths of both defences varied significantly but were not negatively correlated across our chosen host species; thus, we found no evidence for a trade‐off between behavioural and cellular immunity. Instead, cellular defences were significantly weaker against the generalist wasp, whereas behavioural defences were similar in strength against both wasps and positively correlated between wasps. We investigated the adaptive significance of wasp‐induced oviposition reduction behaviour by testing whether wasp‐exposed parents produce offspring with stronger cellular defences, but we found no support for this hypothesis. We further investigated the sensory basis of this behaviour by testing mutants deficient in either vision or olfaction, both of which failed to reduce their oviposition rates in the presence of wasps, suggesting that both senses are necessary for detecting and responding to wasps.     

The relationship between host selection behaviour and offspring fitness in a koinobiont parasitoid

1. When host quality varies, optimal foraging theory assumes that parasitic wasps select hosts in a manner that increases their individual fitness. In koinobiont parasitoids, where the hosts continue developing for a certain period of time after parasitisation, host selection may not reflect current host quality but may be based on an assessment of future growth rates and resources available for the developing larvae. 2. When presented with hosts of uniform quality, the koinobiont parasitoid Leptomastix dactylopii exhibits a characteristic host‐selection behaviour: some hosts are accepted for oviposition on first encounter, while others are rejected several times before an egg is laid in them, a behaviour that is commonly associated with a changing host acceptance threshold during the course of a foraging bout. 3. The fitness of the offspring that emerged from hosts accepted immediately upon encounter was compared with the fitness of offspring emerged from hosts rejected several times before being accepted for oviposition. 4. The pattern of host acceptance and rejection was not related to any of the measured fitness parameters of the offspring emerging from these hosts (development time, size at emergence, sex ratio at emergence, and female offspring egg load). 5. While complex post facto adaptive explanations can be devised to explain the nature of such a time and energy consuming host selection process, it is suggested that physiological constraints on egg production or oviposition may provide an alternative, purely mechanistic, explanation for the results obtained.     

The adaptive significance of host location by vibrational sounding in parasitoid wasps

Vibrational sounding, which is a form of echolocation, is a means of host location by some parasitoid wasps. The wasp taps the substrate (wood, stem or soil) and detects the position of a potential host through the returning 'echoes'. The deployment of vibrational sounding is inferred through the form of the subgenual organ in the female tibia in combination with the presence of modifications to the female antenna used for tapping the substrate. Vibrational sounding and its associated modifications were found in two families. The use of vibrational sounding by parasitoid wasps was positively correlated with the depth of the host in the substrate relative to the size of the parasitoid. There were also significant correlations between the use of vibrational sounding and parasitism of immobile and concealed hosts and between vibrational sounding and idiobiosis. The data suggested that vibrational sounding evolved under a variety of ecological conditions, being employed in the location of wood-boring, stem-boring, soil-dwelling and cocooned hosts and stem-nesting aculeates, often in situations in which the host does not produce vibrations itself.     

Evolutionary patterns of host utilization by ichneumonoid parasitoids (Hymenoptera: Ichneumonidae and Braconidae)*

The four major biological strategies of ichneumonoid parasitoids, koinobiont and idiobiont, ecto-and endoparasitism, are discussed and the evolutionary radiations of the two families Ichneumonidae and Braconidae compared in an attempt to relate differences in patterns of host utilization to differences in evolutionary history. The most primitive members of both families are idiobiont ectoparasitoids of hosts concealed in plant tissue. Idiobiont ectoparasitic braconids are all still primarily associated with such hosts, but idiobiont ectoparasitic ichneumonids have radiated to attack hosts in other situations, such as in aculeate nests or in cocoons. A shift in emphasis between the behavioural steps, host habitat location and host location, is envisaged as being important in such evolutionary change. Idiobiont endoparasitism is postulated as having arisen amongst ectoparasitoids attacking cocooned hosts, as an adaptation that allows them to exploit pupae and puparia in relatively exposed positions; it is a fairly common strategy in the Ichneumonidae, but virtually unknown in the Braconidae. Koinobiosis is perceived as having evolved in association with hosts which feed in a relatively weakly concealed position, but pupate in a more secluded and safe location. The strategy is advantageous as it allows a parasitoid to oviposit on an easily discoverable host, but to use the host's pupation concealment to complete its own development. The evolution of koinobiosis has allowed parasitoids to exploit hosts that feed in exposed positions, and to attack hosts at a younger and numerically more common stage in the host's life cycle. Koinobiont ectoparasitism is envisaged, in some braconid and ichneumonid groups, to occupy an evolutionary transitional position between idiobiosis and endoparasitic koinobiosis; only in the Ichneumonidae have large radiations of koinobiont ectoparasitoids occurred. Endoparasitic koinobiosis is hypothesized as having arisen in the Braconidae in association with lepidopterous/coleopterous hosts, whilst in the major lineage of endoparasitic koinobiont ichneumonids, this habit is hypothesized as having arisen in association with symphytan hosts. The great majority of braconids are koinobiont endoparasitoids, but only about 50% of the Ichneumonidae have this habit. Very few koinobiont braconids develop as endoparasitoids of hymenopterous hosts, although many endoparasitic ichneumonids attack Hymenoptera. However, lineages of the Braconidae have radiated to exploit adult insects and exopterygote nymphs; ichneumonids do not utilize such hosts.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

寄生性膜翅目昆虫毒素的生态机制及应用前景

膜翅目昆虫利用高效的毒素进行自身防卫、攻击猎物和调节寄主生长发育.从寄生性膜翅目昆虫毒素的产生、类别、组份、性质、毒素的生态功能以及毒素的作用机制等方面综述了寄生性膜翅目昆虫毒素的研究概况.膜翅目的泌毒器官起源于外胚层,由生殖系统的附腺演化而来.毒液由成熟雌蜂的毒腺或酸腺所分泌,并贮于毒囊中.昆虫毒素是成分复杂的混合物,已知膜翅目昆虫毒素中含有烃类、醇类、醛类、酮类、羧酸类、酯类、内酯类、酶类等多种化合物.寄生性膜翅目昆虫的毒素在提高自身适应能力方面的作用是巨大的,如通过麻痹寄主提高产卵成功的概率、通过抑制寄主的生长发育和免疫功能提高后代的存活率、通过干扰寄主的生理活动改善后代的营养需求等.体外寄生蜂毒素可造成寄主幼虫停止发育、永久性的麻痹甚至死亡,这类毒素常为抑性的、广谱的,一般作用于中枢神经系统或神经-肌肉连接点.而体内寄生蜂多为容性寄生,其毒液中含有多分DNA病毒（PDV）,PDV通过抑制寄主免疫系统而巧妙地调节寄主的生理活动和发育,影响寄主的正常变态,大多数种类直到寄主结茧或做好蛹室时才将其杀死在安全的场所,从而使寄生蜂后代能够顺利完成发育.容性寄生蜂毒素对PDV的功能具有显著的增效或协同作用,而不会使寄主产生永久性麻痹.PDV对寄生蜂本身是非致病性的,与寄生蜂是一种分子水平上的共生或依生关系.寄生性膜翅目昆虫毒素显示了良好的应用前景,特别是在开发人类医药和特异性生物杀虫剂方面.但分离和纯化毒液中各个活性成分是应用的前提,也是生化和毒理研究的需要.     

Host Stage Selection and Sex Allocation by Gyranusoidea tebygi (Hymenoptera: Encyrtidae), a Parasitoid of the Mango Mealybug, Rastrococcus invadens (Homoptera: Pseudococcidae)

Host stage selection and sex allocation by Gyranusoidea tebygi Noyes (Hym,, Encyrtidae) were studied in choice and no-choice experiments in the laboratory. The parasitoid reproduced on first, second, and third instars of the mango mealybug, Rastrococcus invadens Williams (Hem., Pseudococcidae), and it avoided hosts that were already parasitized. Host feeding was occasionally observed. Sex ratios of the offspring produced by individual wasps were highly biased in favor of females, whereas the sex ratio of groups of wasps foraging under crowded conditions varied from male biased in smaller hosts to female biased in larger hosts. Females had longer developmental times than males, developed faster in larger mealybugs than in smaller ones, and were always larger than males emerging from the same host instar. Their size increased with the instar of the host at oviposition. About 90% of all ovipositions in second and third instar nymphs resulted from an attack with multiple stings, starting with a sting in the head of the host for the most part. The function of these head stings is either to assess quality of the host or to subdue hosts prior to oviposition. Encounter rates, number of attacks, and number of stings during one attack increased, while ovipositions decreased with host instar. Time investment per oviposition and time spent preening increased with increasing host age because older hosts defended themselves more vigorously than younger ones. Thus, while fitness of the parasitoid increased with host size, fitness returns per time decreased. The implications of this host selection behavior for the biological control of the mango mealybug are discussed.     

Effect of the <Emphasis Type="Italic">Drosophila</Emphasis> endosymbiont <Emphasis Type="Italic">Spiroplasma</Emphasis> on parasitoid wasp development and on the reproductive fitness of wasp-attacked fly survivors

In a previous study, we showed that Spiroplasma, a maternally transmitted endosymbiotic bacterium of Drosophila hydei, enhances larval to adult survival of its host when exposed to oviposition attack by the parasitoid wasp Leptopilina heterotoma. The mechanism by which Spiroplasma enhances host survival has not been elucidated. To better understand this mechanism, we compared the growth of wasp larvae in Spiroplasma-infected and uninfected hosts. Our results indicate that wasp embryos in Spiroplasma-infected hosts hatch and grow normally for ~2 days, after which their growth is severely impaired, compared to wasps developing in uninfected hosts. Thus, despite their reduced ability to complete development in Spiroplasma-infected hosts, developing wasps may exert fitness costs on their hosts that are manifested after host emergence. The severity of these costs will influence the degree to which this protective mechanism contributes to the long-term persistence of Spiroplasma in D. hydei. We therefore examined survival to 10-day-old adult stage and fecundity of Spiroplasma-infected flies surviving a wasp treatment. Our results suggest detrimental effects of wasp attack on longevity of Spiroplasma-infected adult flies. However, compared to Spiroplasma-free flies exposed to wasps, Spiroplasma-infected flies exposed to wasps have ~5 times greater survival from larva to 10 day-adult. The relative fecundity of wasp-attacked Spiroplasma-infected females was ~71% that of un-attacked Spiroplasma-free females. Our combined survival and female fecundity results suggest that under high wasp parasitism, the reproductive fitness of Spiroplasma-infected flies may be ~3.5 times greater than that of uninfected females, so it is potentially relevant to the persistence of Spiroplasma in natural populations of D. hydei. Interestingly, Spiroplasma-infected males surviving a wasp attack were effectively sterile during the 3-day period examined. This observation is consistent with the expectation that, as a maternally transmitted symbiont, there is little selective pressure on Spiroplasma to enhance the reproductive fitness of its male hosts.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Biased oviposition and biased survival together help resolve a fig–wasp conflict

We report evidence that helps resolve two competing explanations for stability in the mutualism between Ficus racemosa fig trees and the Ceratosolen fusciceps wasps that pollinate them. The wasps lay eggs in the tree's ovules, with each wasp larva developing at the expense of a fig seed. Upon maturity, the female wasps collect pollen and disperse to a new tree, continuing the cycle. Fig fitness is increased by producing both seeds and female wasps, whereas short‐term wasp fitness increases only with more wasps, thereby resulting in a conflict of interests. We show experimentally that wasps exploit the inner layers of ovules first (the biased oviposition explanation), which is consistent with optimal‐foraging theory. As oviposition increases, seeds in the middle layer are replaced on a one‐to‐one basis by pollinator offspring, which is also consistent with biased oviposition. Finally, in the outer layer of ovules, seeds disappear but are only partially replaced by pollinator offspring, which suggests high wasp mortality (the biased survival or ‘unbeatable seeds’ explanation). Our results therefore suggest that both biased oviposition and biased survival ensure seed production, thereby stabilizing the mutualism. We further argue that biased oviposition can maintain biased survival by selecting against wasp traits to overcome fig defenses. Finally, we report evidence suggesting that F. racemosa balances seed and wasp production at the level of the tree. Because figs are probably selected to allocate equally to male and female function, a 1:1 seed:wasp ratio suggests that fig trees are in control of the mutualism.     

Adult diet affects the life history and host-killing behavior of a host-feeding parasitoid

Neochrysocharis formosa (Westwood), an important biocontrol agent of agromyzid leafminers worldwide, is a host-feeding, idiobiont parasitoid. Female wasps have three types of host-killing behaviors: reproductive (parasitism), non-reproductive host feeding (host feeding), and host stinging without oviposition or feeding (host stinging). In this study, we compared the life history and host-killing behaviors of female parasitoids under four adult diets: starvation, hosts only, hosts plus honey (10% w/v honey solution), and honey only. Furthermore, we analyzed the host-feeding and oviposition preferences of adult females in the hosts-only and hosts-plus-honey treatments. Female parasitoids feeding on hosts had significantly increased longevity, higher fecundity, more host-stinging events, and caused a higher total host mortality than parasitoids in the starvation treatment. The honey supplement significantly increased longevity, fecundity, host-stinging events, and total host mortality, as well as average daily fecundity, but did not alter host-feeding events, host-stinging events, or daily total host mortality. However, the honey supplement did reduce the number of daily host-feeding events and induced a shift toward oviposition. Finally, we found that the non-reproductive host killing caused by host feeding and host stinging enhanced the control potential of N. formosa. These results should contribute to a better understanding of the biocontrol efficiency of destructive host feeders.     

Divvying up an incubator: How parasitic and mutualistic fig wasps use space within their nursery microcosm

Differential occupancy of space can lead to species coexistence. The fig–fig wasp pollination system hosts species-specific pollinating and parasitic wasps that develop within galls in a nursery comprising a closed inflorescence, the syconium. This microcosm affords excellent opportunities for investigating spatial partitioning since it harbours a closed community in which all wasp species are dependent on securing safe sites inside the syconium for their developing offspring while differing in life history, egg deposition strategies and oviposition times relative to nursery development. We determined ontogenetic changes in oviposition sites available to the seven-member fig wasp community of Ficus racemosa comprising pollinators, gallers and parasitoids. We used species distribution models (SDMs) for the first time at a microcosm scale to predict patterns of spatial occurrence of nursery occupants. SDMs gave high true-positive and low false-positive site occupancy rates for most occupants indicating species specificity in oviposition sites. The nursery microcosm itself changed with syconium development and sequential egg-laying by different wasp species. The number of sites occupied by offspring of the different wasp species was negatively related to the risk of syconium abortion by the plant host following oviposition. Since unpollinated syconia are usually aborted, parasitic wasps ovipositing into nurseries at the same time as the pollinator targeted many sites, suggesting response to lower risk of syconium abortion owing to reduced risk of pollination failure compared to those species ovipositing before pollination. Wasp life history and oviposition time relative to nursery development contributed to the co-existence of nursery occupants.     

Striking cuticular hydrocarbon dimorphism in the mason wasp Odynerus spinipes and its possible evolutionary cause (Hymenoptera: Chrysididae,Vespidae)

Cleptoparasitic wasps and bees smuggle their eggs into the nest of a host organism. Here the larvae of the cleptoparasite feed upon the food provision intended for the offspring of the host. As cleptoparasitism incurs a loss of fitness for the host organism (offspring of the host fail to develop), hosts of cleptoparasites are expected to exploit cues that alert them to potential cleptoparasite infestation. Cuticular hydrocarbons (CHCs) could serve as such cues, as insects inevitably leave traces of them behind when entering a nest. By mimicking the host''s CHC profile, cleptoparasites can conceal their presence and evade detection by their host. Previous studies have provided evidence of cleptoparasites mimicking their host''s CHC profile. However, the impact of this strategy on the evolution of the host''s CHC profile has remained unexplored. Here, we present results from our investigation of a host–cleptoparasite system consisting of a single mason wasp species that serves syntopically as the host to three cuckoo wasp species. We found that the spiny mason wasp (Odynerus spinipes) is able to express two substantially different CHC profiles, each of which is seemingly mimicked by a cleptoparasitic cuckoo wasp (i.e. Chrysis mediata and Pseudospinolia neglecta). The CHC profile of the third cuckoo wasp (Chrysis viridula), a species not expected to benefit from mimicking its host''s CHC profile because of its particular oviposition strategy, differs from the two CHC profiles of its host. Our results corroborate the idea that the similarity of the CHC profiles between cleptoparasitic cuckoo wasps and their hosts are the result of chemical mimicry. They further suggest that cleptoparasites may represent a hitherto unappreciated force that drives the evolution of their hosts'' CHCs.     

Remarkable similarity in body mass of a secondary hyperparasitoid Lysibia nana and its primary parasitoid host Cotesia glomerata emerging from cocoons of comparable size

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.     

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.     

Adult size and reproduction in the ectoparasitoid Agrothereutes lanceolatus Walker (Hym., Ichneumonidae)

The effects of adult male and female size on reproductive performance in Agrothereutes lanceolatus Walker, a solitary ectoparasitoid of mature larvae and pupae of several pyralids and tortricids, were investigated. Females had a longer mean lifetime than males. Larger females lived longer, whereas the size of males did not influence the longevity. The number of ovarioles per female did not differ among females of different sizes but larger females carried a greater number of mature and immature eggs. When presented with host cocoons, large females parasitized a higher percentage of hosts than small females. This was due to the fact that larger females accepted more hosts for oviposition whereas smaller ones rejected more hosts. Small and large females used equal numbers of hosts for host-feeding, suggesting that large females removed a greater amount of materials from single hosts through host-feeding.     

Venom is beneficial but not essential for development and survival of Nasonia

1. Parasitoid wasps sting and inject venom into arthropod hosts, which alters host metabolism and development while keeping the host alive for several days, presumably to induce benefits for the parasitoid young. 2. This study investigates the consequences of host envenomation on development and fitness of wasp larvae in the ectoparasitoid Nasonia vitripennis, by comparing wasps reared on live unstung, previously stung, and cold‐killed hosts. Developmental arrest and suppression of host response to larvae are major venom effects that occur in both stung and cold‐killed hosts, but not in unstung hosts, whereas cold‐killed hosts lack venom effects that require a living host. Thus, cold‐killed hosts mimic some of the effects of venom, but not others. 3. Eggs placed on live unstung hosts have significantly higher mortality during development; however, successfully developing wasps from these hosts have similar lifetime fecundity to that of wasps from cold‐killed or stung hosts. Therefore, although venom is beneficial, it is not required for wasp survival. 4. While wasps developing on both cold‐killed and stung hosts have similar fitness levels, rearing multiple generations on cold‐killed hosts results in significant fitness reductions of wasps. 5. It is concluded that the largest benefits of venom are induction of host developmental arrest and suppression of host response to larva (e.g. immune responses), although more subtle benefits may accrue across generations or under stressful conditions.