Oviposition strategy of the parasitic waspDinarmus basalis (Hymenoptera,Pteromalidae)

Summary Host type choice and sex allocation were examined using the solitary parasitic waspDinarmus basalis (Pteromalidae, Hymenoptera) parasitizing larvae or pupae of the bean weevilCallosobruchus chinensis (Bruchidae, Coleoptera) within azuki beans (Vigna angularis). The wasps were offered two types of host; one was hard for the mother to lay eggs in, but was more beneficial for the offspring; the other was easy for the mother to lay eggs in, but was less beneficial for the offspring. The two types of host were one large host (17-day old host) in one bean and 6 small hosts (12-, or 13-day old hosts) in one bean. The same number of each host was presented at the same time to female wasps. The wasps accepted more 17-day old hosts than 12-day old hosts, and more 13-day old hosts than 17-day old hosts in each pair-wise choice experiment. The proportions of accepted host types were different from the proportions predicted by optimization models of random prey encounter with known or unknown prey densities. The wasps showed partial preference of host types. Incomplete information about prey densities, and about the costs and benefits of the two types of host may have generated the partial preference. Two predictions of host sizemodels, that (1) there should be a negative relationship between host size and offspring sex ratio (proportion of male offsprings), and (2) the sex ratio in each size host changes with the relative frequency of each size host utilized, were qualitatively supported.     

Reproductive capacity of females Eupelmus vuilleti (Eupelmidae) inseminated by hyperparasitoid males developed upon the primary parasitoid Dinarmus basalis (Pteromalidae)

Eupelmus vuilleti is a primary and solitary ectoparasitoid of the larval stages of Bruchids (Callosobruchus maculatus, Bruchidius atrolineatus). In a context of intense competition for healthy hosts, E. vuilleti displays ovicide and larvicide behaviours towards the Pteromalid D. basalis during its development (kleptoparasitism), and in an extreme expression of kleptoparasitism the E. vuilleti females hyperparasitize the final larval stage (L5 stage) of D. basalis. In this study, we compared the variability of reproductive success in males that had developed in the context of hyperparasitism to that in males that had developed on primary hosts. The adaptation capacity of the males when 24 h old was analysed in terms of their weight, of the quantity of spermatozoids stored in the seminal vesicles, of the quality of insemination determined from the quantity of spermatozoids stored in the spermatheca of the females after the first mating, and of the number of daughters produced. Adults of E. vuilleti, the larvae of which had developed as hyperparasitoids, are smaller than those that have developed on primary hosts, but they keep all the abilities required to parasite a population of primary hosts once the competitive pressure is reduced.     

Sperm Storage and Use After Multiple Mating in Dinarmus basalis (Hymenoptera: Pteromalidae)

Multiple mating and its effects on the sex ratio in Dinarmus basalis (Hymenoptera: Chalcidoidea, Pteromalidae), an ectoparasitoid of Callosobruchus maculatus (Coleoptera: Bruchidae), were investigated under controlled conditions. Once-mated females suffer a sperm depletion about 21 days after mating and thereafter are constrained to produce only haploid males. On the other hand, three-times-mated females store more sperm in their spermathecae (335 vs 147) and produce daughters during the major part of their reproductive life. Consequently, once-mated females showed a male-biased life time sex ratio (mean = 0.31) as opposed to three-times-mated females (mean = 0.63). Females can copulate only at the beginning of their reproductive life, and multiple mating must occur before egg-laying activity. This behavioral strategy could be an avoidance of consanguinity in a promiscuous environment. This ectoparasitoid species reveals a reproductive strategy which promotes polyandry and a very high sperm efficiency.     

Seasonal variation in the reproductive strategy of the parasitic wasp Eulophus larvarum (Hymenoptera: Chalcidoidea: Eulophidae)

ABSTRACT. 1. Eulophus larvarum (L.) (Hymenoptera: Eulophidae) is a gregarious parasitoid of lepidopterous larvae feeding on broad-leaved trees. Normally there are two generations a year.
2. Sex ratio in the spring generation of larvae is female biased. The bias is probably due to local mate competition as progeny from one brood emerge from their pupae in close proximity to each other.
3. Sex ratio in the summer generation of larvae is near equality. Local mate competition is absent as individuals from the same brood become separated during the winter.
4. Variation in sex-ratio in the spring generation is consistent with a binomial model. Variation in sex ratio in the summer generation is much greater than expected from a binomial model and there is a large proportion of single-sex broods.
5. Two hypotheses are put forward to explain the summer generation pattern: virgin oviposition, and strong intersexual competition.
6. No differences in clutch size were found between the two generations.     

Sperm production and mating potential of males after a cold shock on pupae of the parasitoid wasp Dinarmus basalis (Hymenoptera: Pteromalidae)

For ectothermic species, temperature is a key environmental factor influencing several aspects of their physiology and ecology, acting particularly on reproduction. To measure the consequences of a severe thermal stress during development on male reproduction, a cold shock (1h at -18 degrees C) was tested on Dinarmus basalis pupae. D. basalis (Hymenoptera: Pteromalidae) is a parasitoid wasp in which sperm management in both male and female is of prime importance. After a cold shock, developmental success was reduced, with a quarter of cold-shocked males not emerging correctly. The stress effects were estimated at the level of sperm stock in seminal vesicles of males at different ages and on the ability of 2-day-old males to access females in single and multiple mating and in male-male competition. Cold-shocked males had a reduced sperm stock compared to control males and this difference persisted with age. The rate of sperm production was similar in both groups. The consequences of a cold shock on male reproductive ability were perceptible in multiple mating and male-male competition but not in single mating. Cold-shocked males were at a disadvantage, inseminating fewer females and copulating less frequently. Finally, male pupae of D. basalis were able to withstand severe temperature stresses and their reproductive functions were partially preserved.     

Effect of wasp size, physiological state, and prior host experience on host-searching behavior in a parasitoid wasp (Hymenoptera: Ichneumonidae)

Host-searching behavior in insects generally varies among individuals. A number of physiological and environmental factors can be involved in such individual variation. Here, a series of behavioral observations were made to highlight the importance of physiological state (i.e., number of mature eggs a female carries, amount of nutrient reserves, etc.) and learning state (i.e., prior host experience) on host-approaching behavior of parasitoids. Itoplectis naranyae (Hymenoptera: Ichneumonidae), a solitary endoparasitoid wasp attacking lepidopteran pupae and prepupae, was used as a test insect. The results show that female wasps with experience ovipositing on hosts 2 days before the test found hosts more quickly than did naïve wasps. Prior experience of host odor itself did not affect host-finding behavior, however. A single oviposition was enough for wasps to shorten time to find a host; additional experience had no significant effect on the efficiency of searching. The number of mature eggs a female carried had no effect on the time required to find a host regardless of prior host experience. The size of wasps, instead, was a significant factor when wasps had no prior host experience, and larger wasps found hosts more rapidly than did smaller wasps. Searching activity was not affected by how many hosts a female wasp had fed on before testing.     

First report of chromosomes of the parasitoid wasp Trissolcus basalis (Wollaston) (Hymenoptera: Platygastridae: Telenominae)

Here we present the first data on the chromosome complement of Trissolcus basalis gathered in conjunction with the ongoing whole genome sequencing efforts for this species. The cytogenetic investigation revealed a diploid karyotype with 20 chromosomes, comprising one pair of large metacentrics and nine pairs of acrocentrics that gradually decrease in length.     

Effect of host size on male fitness in the parasitoid wasp Dinarmus basalis

The effect of host size on male fitness was tested in the parasitoid wasp Dinarmus basalis (Hymenoptera, Pteromalidae) using hosts of different fresh weight. Fitness was measured as the sperm stock in seminal vesicles, and the ability to access females in single or competition situations. Both body size and sperm in seminal vesicles increased with host fresh weight. Males from small hosts had a reduced size and sperm stock compared to those from larger hosts. In single situations, males from both small and large hosts had similar reproductive capacities, whereas in multiple mating or competition situations, males from small hosts were at a disadvantage, inseminating fewer females and copulating less frequently. However, females did not appear to choose between males, and no effect on sperm stored in the spermatheca was observed. Being small does not prevent a D. basalis male mating and producing progeny in single situations, although more offspring could be expected from larger males because of their better competitive abilities.     

Characterization and biochemical analyses of venom from the ectoparasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae)

During parasitism, the ectoparasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) induces a developmental arrest in host pupae that is sustained until the fly is either consumed by developing larvae or the onset of death. Bioassays using fluids collected from the female reproductive system (calyx, alkaline gland, acid gland, and venom reservoir) indicated that the venom gland and venom reservoir are the sources of the arrestant and inducer(s) of death. Infrared spectroscopic analyses revealed that crude venom is acidic and composed of amines, peptides, and proteins, which apparently are not glycosylated. Reversed phase high performance liquid chromatography (HPLC) and sodium dodecyl polyacrylamide gel electrophoresis (SDS-PAGE) confirmed the proteinaceous nature of venom and that it is composed mostly of mid to high molecular weight proteins in the range of 13 to 200.5 kilodaltons (kDa). Ammonium sulfate precipitation and centrifugal size exclusion membranes were used to isolate venom proteins. SDS-PAGE protein profiles of the isolated venom fractions displaying biological activity suggest that multiple proteins contribute to arresting host development and eliciting death. Additionally, HPLC fractionation coupled with use of several internal standards implied that two of the low molecular weight proteins were apamin and histamine. However, in vitro assays using BTI-TN-5B1-4 cells contradict the presence of these agents.     

Host Age Response in the Parasitoid Wasp Spalangia cameroni (Hymenoptera: Pteromalidae)

Female Spalangia cameroni produced more offspring from younger house fly pupae, both when given a choice of host ages and when not given a choice. Host age did not affect offspring survivorship. Offspring were larger when they had developed on younger hosts and the effect was independent of offspring sex. Having previously parasitized old hosts versus young hosts did not reduce a female's production of offspring in subsequent hosts. Females distinguished between young and old hosts both in the light and in the dark. Females did not distinguish between host ages prior to physical contact with the host but could distinguish by the time they first began exploring a host by tapping it with their antennae; thus, they could distinguish before drilling into a host.     

Analysis of multiparasitism by Eupelmus vuilleti (Craw) (Eupelmidae) and Dinarmus basalis (Rond) (Pteromalidae) in the presence of one of their common hosts,Bruchidius atrolineatus (Pic) (Coleoptera Bruchidae)

Bruchidius atrolineatus (Pic) is a tropical beetle (Coleoptera Bruchidae) that develops during the larval and pupal stages in the seeds of a legume Vigna unguiculata (Walp). Two species of Hymenoptera, Dinarmus basalis (Rond) and Eupelmus vuilleti (Craw), solitary ectoparasitoids of the larvae and pupae of B. atrolineatus, were introduced successively in the presence of their hosts, varying the interval between the two introductions. When D. basalis females were introduced 24 h, 3 days or 7 days after E. vuilleti, multiparasitism was low. The females had low fecundity, and their eggs were not distributed randomly over the different available hosts. When E. vuilleti females were introduced second, they oviposited on the different hosts availabe and did not avoid multiparasitism. The presence of hosts already parasitised by D. basalis increased the reproduction of E. vuilleti, and the fecundity of the females was higher than in control batches with E. vuilleti alone. E. vuilleti seems capable of detecting the ovipositor shafts drilled by the D. basalis females, and by introducing its own ovipositors killing the D. basalis eggs or larvae. When interspecific competition was occurring the number of E. vuilleti adults emerging from the seeds was no different from that observed in control batches with E. vuilleti alone, and there were always fewer D. basalis adults than in control batches (D. basalis alone). This interspecific competition reduces the influence of the two parasitoids in the biological control of bruchid populations.     

红脉穗螟寄生蜂寄主选择研究

[目的]优化红脉穗螟寄生蜂繁育技术,为该虫的生物防治提供参考。[方法]通过室内饲养研究了红脉穗螟幼虫寄生蜂褐带卷蛾茧蜂和蛹寄生蜂周氏啮小蜂(海南种)对4种昆虫的寄生效果,并对最佳接种比例进行筛选。[结果]褐带卷蛾茧蜂对大蜡螟幼虫寄生效果优于米蛾幼虫,且对黄粉虫和大麦虫幼虫不表现寄生特性,褐带卷蛾茧蜂和大蜡螟幼虫的最佳接种比例为1∶1和2∶1,其中在2∶1接种比例处理中的寄生率和单寄主产蜂量分别为76.67%和34.60头;4种昆虫蛹均可用于繁育周氏啮小蜂,从寄主的繁育成本和寄生效果分析,以黄粉虫蛹效果较最佳,接种蜂虫比以2∶1为宜,此时的单寄主产蜂量为148.60头。[结论]寄主和接种比例不同会影响寄生蜂的寄生效果,本研究中褐带卷蛾茧蜂适宜寄主为大蜡螟,最优蜂虫比为1∶1和2∶1,周氏啮小蜂(海南种)适宜寄主为黄粉虫,最优接种蜂虫比为2∶1。     

Marking Behavior and Discrimination of Concealed Hosts by the Ectoparasitoid, Dinarmus basalis Rond. (Hym. Pteromalidae)

The parasitoid species, Dinarmus basalis, attacks bruchid beetle larvae concealed within hard seeds. Female wasps are able to discriminate the quality of their host but the position in the concealed host system where this discrimination takes place remains unclear. Under laboratory conditions, we investigated whether the cues were detectable: (1) at a medium or short distance from the seed, (2) on the seed, or (3) within the seed, at the point of contact with the host. In particular, we investigated whether internal or external markings are applied by the parasitoid and then recognized in a subsequent encounter or by a conspecific encountering the host. Detailed behavioral observations did not provide evidence of markings being applied onto the seed or used, but showed that host discrimination occurred on the basis of internal cues. The lack of evidence of external markings is unexpected and raises the question as to which circumstances in the parasitoid's ecology and life history could result in such a lack not causing a reduction of fitness.     

Susceptibility of the bruchid Callosobruchus maculatus (Coleoptera: Bruchidae) and its parasitoid Dinarmus basalis (Hymenoptera: Pteromalidae) to three essential oils

The bruchid Callosobruchus maculatus (F.) causes major losses during the storage of seeds of Vigna unguiculata (Walp.) in West Africa. An endemic parasitoid, the pteromalid Dinarmus basalis (Rond.) reduces the increase in bruchid populations in stores and could be used for biological control. African farmers often introduce essential oils into granaries at harvest time. In Togo, essential oils were extracted from two Gramineae, Cymbopogon nardus (L.) and Cymbopogon schoenanthus (L.) and from a Lamiaceae, Ocimum basilicum (L.). The major components of these essential oils were citronellal in C. nardus, carene-2 and piperitone in C. schoenanthus and estragol in O. basilicum. Cymbopogon schoenanthus was the most toxic oil for C. maculatus adults. D. basalis adults were more susceptible to the three essential oils than the adults of their hosts C. maculatus. In the presence of cowpea seeds, the LC50s of the three essential oils were lower than in their absence, suggesting that the seeds may absorb a part of the volatiles. High doses of three essential oils slightly affected the survival of the fourth instar or the pupae of C. maculatus. This high survival was due to protection of larvae from volatiles by the surrounding seeds. The D. basalis were more affected by the oil volatiles than their hosts. Sub-lethal doses of essential oils reduced the duration of the adult life of both insect species and fecundity of the females. The differences in sensitivity of the host and its parasitoid could influence their population dynamics. The introduction of the essential oils into storage systems potentially could reduce density of parasitoid populations and increase seed losses.     

Species-diagnostic single-nucleotide polymorphism and sequence-tagged site markers for the parasitic wasp genus Nasonia (Hymenoptera: Pteromalidae)

Wasps of the genus Nasonia are important biological control agents of house flies and related filth flies, which are major vectors of human pathogens. Species of Nasonia (Hymenoptera: Pteromalidae) are not easily differentiated from one another by morphological characters, and molecular markers for their reliable identification have been missing so far. Here, we report eight single-nucleotide polymorphism and three sequence-tagged site markers derived from expressed sequenced tag libraries for the two closely related and regionally sympatric species N. giraulti and N. vitripennis. We studied variation of these markers in natural populations of the two species, and we mapped them in the Nasonia genome. The markers are species-diagnostic and evenly spread over all five chromosomes. They are ideal for rapid species identification and hybrid recognition, and they can be used to map economically relevant quantitative trait loci in the Nasonia genome.     

Males mate guard in absentia through extended effects of postcopulatory courtship in the parasitoid wasp Spalangia endius (Hymenoptera: Pteromalidae)

The proximal mechanisms leading to monandry have been little studied in most insect orders, including Hymenoptera. In the parasitoid wasp Spalangia endius, mated females are less attractive (less often mounted) than virgins and are unreceptive (unlikely to allow copulation). Which aspects of mating are responsible was tested by observing male responses toward females whose mating had been interrupted at various stages. All females were allowed to receive precopulatory courtship and to open their genital aperture to copulate. Then some were interrupted before copulation, some after copulation but before postcopulatory courtship, and some were allowed to complete postcopulatory courtship. Females that had copulated were not less attractive than females that had not. In contrast, females that had received postcopulatory courtship were clearly both less attractive and less receptive. Thus, postcopulatory courtship functions as extended mate guarding, by making the female less attractive and less receptive to subsequent males even after the original male is no longer present. The effect of postcopulatory courtship on female attractiveness was persistent but imperfect: when males were presented sequentially to mated females, most but not all males retreated without mounting, and a female could repulse more than twenty males in succession.     

How host larval age, and nutrition and density of the parasitoid Dinarmus basalis (Hymenoptera: Pteromalidae) influence control of Acanthoscelides obtectus (Coleoptera: Bruchidae)

Choice of the targeted host developmental stage, regulation of parasitoid numbers released and introduction of food supplements are operational factors with a potential to influence the level of biological control. In a closed laboratory storage system maintained over two generations of the host, the impact of these three parameters on the control potential of the parasitoid Dinarmus basalis Rondani was investigated for high populations of larvae of Acanthoscelides obtectus (Say) feeding inside dry common bean seeds Phaseolus vulgaris. The beans were already infested with immature bruchids at the beginning of the storage period to simulate harvest conditions, characterized in a previous study. Treatments resulted in a reduction of 48-75% of the bruchid population within 16 weeks of storage. The best timing of parasitoid release was at the simulated harvest, as later releases reduced the bruchid population only by about half this percentage. Host feeding is postulated to be the key factor involved in the observed difference. The effect of increasing the number of parasitoids strongly depended on host age and food supplement. Addition of vials with honey had no direct effect on the bruchid population or on the parasitoid progeny. The ecological significance of these findings and implications for biological control are discussed.     

Disruption of pupariation and eclosion behavior in the flesh fly, Sarcophaga bullata Parker (Diptera: Sarcophagidae), by venom from the ectoparasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae)

The action of venom from the ectoparasitic wasp, Nasonia vitripennis, was monitored by examining alterations in patterned muscular movements characteristic of pupariation and eclosion behavior in the flesh fly, Sarcophaga bullata. Venom injected into larvae prior to pupariation caused a dose-dependent delay in pupariation. Eventually, such larvae did pupariate, but puparia were abnormally formed. Barographic records revealed that all elements of pupariation behavior were present in venom-injected larvae, but pupariation behavior was not well synchronized with tanning, thus implying that the venom caused disruption in the temporal organization of central motor programs. When larvae were ligated and injected with venom posterior to the ligature, no response was evident in the posterior region, suggesting that the venom does not directly stimulate muscles or neuromuscular junctions. Injection of exogenous ecdysteroid into venom-injected larvae restored some elements of pupariation behavior, consistent with ecdysone's role in stimulating the release of anterior retraction factor and puparium tanning factor, two factors that are released from the CNS to regulate pupariation. When the venom was injected into newly emerged imagoes, the duration of extrication behavior was shortened, whereas all phases of post-eclosion behavior were lengthened. These observations imply that the venom affects CNS centers that regulate the muscular systems engaged in extrication and post-eclosion behavior.     

Delayed inhibition of host development by the nonparalyzing venoms of parasitic wasps

The nonparalyzing venoms of two unrelated parasitic wasps, Eulophus larvarum and Clinocentrus gracilipes, in separate ways cause delayed yet well-defined arrests in their hosts' development, apparently by disrupting events normally under endocrine control. The venom of E. larvarum (for which species host acceptance and oviposition behavior is described) prevents apolysis in its host after a depressed feeding period. The venom employed by C. gracilipes switches the host to a pharate pupal stage irrespective of its larval instar.