The role of parasitoids in regulation of Polistes wasp population (Hymenoptera, Vespidae: Polistinae)

The study conducted in 2005–2010 analyzes the behavioral response of the parasitoids Latibulus argiolus (Rossi) (Hymenoptera, Ichneumonidae) and Elasmus schmitti Ruschka (Hymenoptera, Eulophidae) to the distribution of their host, Polistes wasps (Hymenoptera, Vespidae). Various conditions of the parasitoid-host system and conditions of regulation of the host abundance are discussed. The parasitoid females are more active in wasp colony clusters and tend to infest larger nests. If the parasitoids are abundant, infestation of host colonies starts earlier, sometimes before the worker emergence; therefore, density-dependent behavioral response of parasitoids is caused primarily by the impact of the aggregation component. Thus, the host population density factor appears to be mediated not only by the non-uniform development rates of colonies and their spatial distribution, but also by the seasonal (temporal) aspect of their development. Low density of the host population, at which the parasitoids regulate the wasp abundance, corresponds to a certain phase of the seasonal colony development, namely to the period before the emergence of workers. On the whole, we are dealing with a host-parasitoid system in which the spatial and temporal factors are closely interrelated.     

Brood size, sex ratio and egg load of Metaphycus lounsburyi (Hymenoptera: Encyrtidae) when parasitizing adult females of black scale Saissetia oleae (Hemiptera: Coccidae) in the field

Black scale Saissetia oleae (Olivier) (Hemiptera: Coccidae) is a widespread olive pest in California and Europe. Metaphycus lounsburyi (Howard) (Hymenoptera: Encyrtidae) is often one of its main parasitoids. Augmentative releases of M. lounsburyi have been proposed in those areas in which biological control is ineffective. In this paper, we study the relationship between black scale and M. lounsburyi in the field. According to our data, M. lounsburyi parasitizes mainly ovipositing females of black scale. Parasitism rates of ovipositing females reach high levels. In those scales, M. lounsburyi develops as a gregarious parasitoid, with an average of 13 and a maximum of 40 parasitoids developing per scale. The secondary sex ratio is female biased (proportion of males = 0.13) and appears to be brood size dependent. Metaphycus lounsburyi eggs are encyrtiform; however, its egg load is much higher than other Metaphycus that also parasitize black scale. The implications of these results on the behavioral ecology of M. lounsburyi and its use in biological control of black scale are discussed.     

Maternal and progeny quality of Habrobracon hebetor Say (Hymenoptera: Braconidae) after cold storage

The ectoparasitoid Habrobracon hebetor Say (Hymenoptera: Braconidae) is an important biological control agent for lepidopterous pests of stored products. We investigated the effects of low temperature storage on the quality of adult parasitoids and their progeny. Newly emerged females were stored for 10, 20, 30, 40, 50, and 70 days at 5 ± 1 °C. Several reproductive and developmental parameters were then assessed to determine the quality of the adult parasitoids and their progeny. After more than 30 days of storage, there was a decrease in parental parasitism, but low temperature storage of parents had no effect on parasitism of the F1 generation. Parental longevity and fecundity decreased after more than 20 days of storage, but there was no effect of storage duration on the fecundity and longevity of the F1 generation until a storage duration of 50 days. Development time varied with storage duration but differences were within 2 days. Storage duration had no effect on the sex ratio of F1 and F2 generations. Our data show that H. hebetor can be cold stored for up to 20 days without adversely affecting the performance of the parasitoid. Therefore, short-term storage of H.hebetor adults could be used for maintaining and accumulating large numbers of parasitoids in mass rearing programs.     

Distribution and Abundance of Mymarid Parasitoids (Hymenoptera: Mymaridae) of Sophonia rufofascia Kuoh and Kuoh (Homoptera: Cicadellidae) in Hawaii

The abundance of mymarid parasitoids attacking the two-spotted leafhopper (Sophonia rufofascia [Kuoh and Kuoh]), a polyphagous pest recently adventive to Hawaii, was monitored using yellow sticky cards deployed in several areas on the islands of Kauai and Hawaii. The yellow cards captured Chaetomymar sp. nr bagicha Narayanan, Subba Rao, & Kaur and Schizophragma bicolor (Dozier), both adventive species, and Polynema sp. Haliday, which is endemic to Hawaii (Hymenoptera: Mymaridae). The former two species were most abundant at all sites. On Kauai, there was a negative correlation between the captures of C. sp. nr bagicha and those of Polynema sp. Throughout the season, the increase in parasitoid numbers generally followed the increase in leafhopper numbers. C. sp. nr. bagicha and S. bicolor showed distinct habitat preferences. Removal of Myrica faya Aiton, an invasive weed that is a highly preferred two-spotted leafhopper host, decreased the overall numbers of captured parasitoids, but led to a twofold increase in the ratio of trapped parasitoids/hosts in weed-free areas.     

Host discrimination and interspecific competition of Trissolcus nigripedius and Telenomus gifuensis (Hymenoptera: Scelionidae), sympatric parasitoids of Dolycoris baccarum (Heteroptera: Pentatomidae)

Trissolcus nigripedius Nakagawa and Telenomus gifuensis Ashmead (Hymenoptera: Scelionidae) are solitary egg parasitoids of Dolycoris baccarum L. (Heteroptera: Pentatomidae), a polyphagous seed-sucking bug of agricultural crops. Field examinations revealed that the two parasitoids are the most common species, but only a single species emerge from a host egg mass. To explain this observation we tested two hypotheses of interspecific host discrimination and asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis. Trissolcus nigripedius and Te. gifuensis could discriminate host egg pre-parasitized by either self or conspecific like other scelionid parasitoids that use external mark on host eggs. When provided with host eggs pre-parasitized by each other, both Tr. nigripedius and Te. gifuensis multiparasitized 94% and 100% of the host eggs without interspecific host discrimination, respectively. Interestingly, from the multiparasitized eggs irrespective of oviposition sequence, progeny of Te. gifuensis always survive better than Tr. nigripedius. Telenomus gifuensis is superior in immature competition probably due to shorter egg incubation period after oviposition, hence progeny of Te. gifuensis become first instar faster than that of Tr. nigripedius. However, adult Tr. nigripedius is always superior competitor in possessing and guarding the host eggs even after oviposition against Te. gifuensis. Therefore, the asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis may explain the emergence of a single species from a host egg mass in the field in spite of no interspecific host discrimination.     

Female reproductive biology of Platygaster diplosisae (Hymenoptera: Platygastridae) and Aprostocetus procerae (Hymenoptera: Eulophidae), two parasitoids associated with the African Rice Gall Midge, Orseolia oryzivora (Diptera: Cecidomyiidae)

We investigated the female reproductive system of Platygaster diplosisae (Hymenoptera: Platygastridae) and Aprostocetus procerae (= Tetrastichus pachydiplosisae) (Hymenoptera: Eulophidae), two parasitoids associated with the African rice gall midge, Orseolia oryzivora (Diptera: Cecidomyiidae). Both optical and electron microscopy were used. The female reproductive system of P. diplosisae includes two large ovaries of the meristic polytrophic‐type, each composed of several tens of ovarioles. The system includes also a venomous gland that extends to a common oviduct. This gland had a filiform secretory portion, in which the epithelium was thin and surrounded a common evacuation canal. The secretory cells secrete into a large reservoir. Parasitism due to P. diplosisae is gregarious. The female reproductive system of A. procerae includes two ovaries of the meristic polytrophic‐type, and each ovary has a few ovarioles. Each ovariole includes one or two oocytes, which can be seen in the vitellarium. Two accessory glands, which extend to the oviduct, are also visible. The secretory epithelium of the accessory gland is made up of a dense network of secretory cells surrounded by muscle fibers. Females of A. procerae pierce the tissues of the gall and probably deposit one egg on or close to the pupa of the midge. Aprostocetus procerae is a solitary parasitoid of the midge. The two parasitoids exploit the same host at different developmental stages. These findings improve our knowledge of the reproductive biology of these two parasitoids associated with the African rice gall midge, an important pest in Africa.     

Host specialization is ancestral in Torymus (Hymenoptera,Chalcidoidea) cynipid gall parasitoids

The host range of parasitoids varies greatly among species: Some only parasitize one to a few hosts (specialists), while others parasitize multiple species or a variety of host types (generalists). The direction of most host range shifts in parasitoid groups, that is from generalist to specialist or, alternatively, from specialist to generalist, is unknown. To explore the origin of host range shifts, we studied a clade within the genus Torymus (Hymenoptera, Chalcidoidea) that includes both generalist and specialist parasitoids of Cynipidae (Hymenoptera). We analysed the phylogenetic relationships of the species of Torymus on the basis of two gene fragments (cox1 and ITS2) of 246 specimens and performed an ancestral state reconstruction of the specialist/generalist trait. Our results revealed the following: (a) The ancestral state of this group of Torymus is specialist, with the generalist state evolving through a loss of specialization. (b) The species Torymus cyaneus and Torymus flavipes both have a strong genetic structure, suggesting the existence of different biological identities. (c) There has been a host plant shift in the lineage(s) leading to Torymus rubi and Torymus bedeguaris from galls on Quercus to those on Rosaceae. (d) The alien species Torymus sinensis and the native European species Torymus notatus are phylogenetically closely related. (e) Speciation within Torymus was likely associated with the diversification of their cynipid hosts, which itself was driven by the dramatic changes in climate and vegetation that occurred during the Miocene.     

Cold storage effects on maternal and progeny quality of Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae)

This study determined the effects of cold storage on the survival, development and reproduction of the mymarid wasp, Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae). Following storage of the immature parasitoids within host eggs of the glassy-winged sharpshooter, Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae) under a daily fluctuating temperature for up to 50 d, the quality of the parental and F₁ generations was assessed by examining several reproductive and developmental parameters indicating fitness. Immature wasps were stored for 20 d within the host without reducing their subsequent survival, development or progeny fitness parameters. After 30 d of storage, survival declined, post-storage developmental time was extended, and the fecundity of the adult females decreased. Storage for 40 d severely damaged G. ashmeadi, because it not only yielded a 12% survival rate, 44% female sterility and increased the proportion of progeny males by 155%, but it also reduced parasitism and fecundity by 70% and 73%, respectively. No wasps emerged after 50 d of storage. Cold storage affected the emergence pattern of the parental but not the F₁ and F₂ generations. Parental emergence was extended and the pattern displayed two additional peaks after the initial onset. Analysis of several demographic parameters for the parental and F₁ generations further confirmed that the quality of the adult parents declined after they had been stored as immatures for 30 d. The detrimental effects caused by cold storage of the parental generation do not extend to the F₁ generation. Our results indicate that short-term cold storage of G. ashmeadi within its host could be used for maintaining and accumulating these parasitoids during mass propagation for release in a control program.     

Single-step PCR differentiation of Cotesia sesamiae (Cameron) and Cotesia flavipes Cameron (Hymenoptera: Braconidae) using polydnavirus markers

Cotesia sesamiae (Cameron) and Cotesia flavipes Cameron (Hymenoptera: Braconidae) are the main larval parasitoids of cereal stemborers in sub-Saharan Africa. Cotesia sesamiae is endemic to eastern and southern Africa, while C. flavipes was introduced into the region for biological control against the exotic lepidopteran Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). The two are sibling parasitoids, difficult to distinguish morphologically. The introduced insect could potentially lead its African biotype to extinction because of their similar ecological niche. In order to distinguish the two species, multiplex primer-specific and PCR-RFLP tests were developed. Rapid identification of the two species was possible using primer-specific tests on DNA extracts as well as on pieces of tissue in a single PCR step followed by gel electrophoresis. The CRV1 gene of the polydnavirus, a symbiont to the wasps, was used as the marker. The results show that the morphological identifications, validated by molecular tests, are accurate in 93% of cases.     

Fitness of Encarsia sophia (Hymenoptera: Aphelinidae) parasitizing Trialeurodes vaporariorum and Bemisia tabaci (Hemiptera: Aleyrodidae)

Abstract Fitness and efficacy of Encarsia sophia (Girault & Dodd) (Hymenoptera: Aphelinidae) as a biological control agent was compared on two species of whitefly (Hemiptera: Aleyrodidae) hosts, the relatively smaller sweetpotato whitefly, Bemisia tabaci (Gennadius) biotype ‘B’, and the larger greenhouse whitefly, Trialeurodes vaporariorum (Westwood). Significant differences were observed on green bean (Phaseolus vulgaris L.) in the laboratory at 27 ± 2°C, 55%± 5% RH, and a photoperiod of 14: 10 h (L: D). Adult parasitoids emerging from T. vaporariorum were larger than those emerging from B. tabaci, and almost all biological parameters of E. sophia parasitizing the larger host species were superior except for the developmental times of the parasitoids that were similar when parasitizing the two host species. Furthermore, parasitoids emerging from T. vaporariorum parasitized more of these hosts than did parasitoids emerging from B. tabaci. We conclude that E. sophia reared from larger hosts had better fitness than from smaller hosts. Those from either host also preferred the larger host for oviposition but were just as effective on smaller hosts. Therefore, larger hosts tended to produce better parasitoids than smaller hosts.     

Taxonomic review and key to European Ichneumon flies (Hymenoptera, Ichneumonidae), parasitoids of gall-forming sawflies of the genera Pontania Costa, Phyllocolpa Benson, and Euura Newman (Hymenoptera, Tenthredinidae) on willows: Part I

Extensive material on parasitoids of the family Ichneumonidae reared from gall-making sawflies on Salix is identified. The general host—parasite list includes 75 species of hosts—Pontania (40 spp.), Phyllocolpa (21 spp.), Euura (14 spp.), their food plants (33 species of Salix), and their parasitoids of the family Ichneumonidae; many data on hosts and parasitoids are recorded for the first time. The complex of ichneumonids on gallforming sawflies is composed of about 45 species belonging to 15 genera of 6 subfamilies. The annotated list of about 30 species of ichneumonids (except for species of Saotis) reared from gall-making sawflies is given with data on the literature, studied material, distribution, and hosts. Two new species Anoncus gallicola sp. n. and Rhinotorus brachycerus sp. n. are described. A key to the species of the Ichneumonidae of this complex is given.     

Parasitism of the Wheat Stem Sawfly (Hymenoptera: Cephidae) in Montana

The wheat stem sawfly,Cephus cinctusNorton (Hymenoptera: Cephidae) was first found in wild grasses and soon became an economically important pest of wheat after cultivation began in the northern Great Plains. Of the species of parasitoids that attackC. cinctusin wild grasses, onlyBracon cephi(Gahan) andB. lissogasterMuesebeck (Hymenoptera: Braconidae) have been found in wheat. Levels of parasitism vary between wheat-producing regions in Montana. Parasitism levels were increased by releasing individuals from a region with high levels of parasitism into a region where parasitism levels were low. This suggests that there has been unequal rates of parasitoid adaptation from wild grasses to wheat in different regions of Montana.     

Host suitability and preference studies of Trichogramma cordubensis (Hymenoptera: Trichogrammatidae)

Studies of host suitability and preferences of Trichogramma cordubensis Vargas and Cabello (Hymenoptera: Trichogrammatidae) were performed with eggs of six Lepidoptera (Noctuidae) species: Thysanoplusia orichalcea Fabricius, Peridroma saucia (Hübner), Xestia c-nigrum L., Phlogophora meticulosa (L.), Noctua pronuba (L.), and N. atlantica (Warren). Host suitability was studied by analysing separately the effects of the attacked host species and the influence of the rearing host species on different biological parameters of T. cordubensis. Host preference was analysed by offering eggs of two host species simultaneously to a single female wasp without previous oviposition experience (dual-choice tests). Results show that P. saucia, followed by P. meticulosa were the least suitable hosts for T. cordubensis, since on these species the preimaginal development of the parasitoids was significantly longer and, the number of parasitized eggs as well the number of offspring per female were significantly lower. Contrarily, T. cordubensis parasitized at a higher rate the eggs of the endemic non-target species, N. atlantica. Dual choice tests showed that the option of the first host to be accepted by the wasp was random; however, the mean number of parasitized eggs differed significantly when two host species were offered simultaneously to T. cordubensis, always being the host species with heavier eggs the most parasitized.     

Post-release survey to assess impact and potential host range expansion by Amitus hesperidum and Encarsia perplexa, two parasitoids introduced for the biological control of the citrus blackfly, Aleurocanthus woglumi in Dominica

Four years after the release of two exotic parasitoids, Amitus hesperidum Silvestri (Hymenoptera: Platygasteridae) and Encarsia perplexa Huang and Polaszek (Hymenoptera: Aphelinidae) for the classical biological control of the citrus blackfly (CBF), Aleurocanthus woglumi Ashby (Hemiptera: Aleyrodidae) in Dominica, a survey was conducted to assess establishment as well as potential nontarget effects especially on Aleyrodidae and other related taxa. CBF populations were low to non-existent in 50 of 51 field sites examined. At the site where CBF was encountered, both E. perplexa and A. hesperidum were present and CBF populations were declining. The two parasitoids were not among the several species collected on nontarget Aleryodidae and Hemiptera. It is concluded that E. perplexa and A. hesperidum have kept CBF populations under effective biological control in Dominica and there is no evidence of any nontarget effects on other Aleyrodidae or their natural enemies. Handling Editor: Dirk Babendreier.     

Effect of insecticides on the mortalities of three whitefly parasitoid species, Eretmocerusmundus, Eretmoceruseremicus and Encarsiaformosa (Hymenoptera: Aphelinidae)

The toxicities of 24 insecticides for the biological control of whiteflies were evaluated for Eretmocerus mundus (Mercet), Eretmocerus eremicus Rose and Zolnerowich and Encarsia formosa Gahan using the residual film method (for adults) and the dipping method (for pupae). Mortalities from insect growth regulators (IGRs) (flufenoxuron and lufenuron), Bacillus thuringiensis (Bt), pymetrozine and sulfur were <30% for both pupae and adults of all three species, indicating that the parasitoids were not seriously affected by these insecticides. Neonicotinoids (acetamiprid, clothianidin, dinotefuran, imidacloprid and nitenpyram), synthetic pyrethroids (etofenprox and permethrin), organophosphates (acephate and fenitrothion), chlorphenapyr, emamectin benzoate, spinosad and tolfenpyrad were seriously harmful (100% mortality) and acaricides (chinomethionat, milbemectin and pyridaben) were moderately harmful or seriously harmful to adult parasitoids (leading to mortalities of >92%). For each insecticide, the mortality of pupae was generally lower than that of adults, even though the toxicity classification for the two groups was similar. The results indicate that IGRs, Bt, pymetrozine and sulfur are relatively harmless, and are compatible with the use of parasitoids to help control whiteflies for integrated pest management in greenhouses.     

Larvicidal activity of lectins onLucilia cuprina: mechanism of action

Foraging behaviour and host-instar preference of young and old females of the solitary aphid parasitoid,Lysiphlebus cardui Marshall (Hymenoptera: Aphidiidae), were studied in the laboratory. The analysis of interactions between parasitoids and different stages ofAphis fabae cirsiiacanthoidis Scop. (Homoptera: Aphididae) revealed that encounter rates between aphids and parasitoid females and defence reactions of the aphids influenced the degree to which a particular aphid age class is parasitized. Encounter rates between hosts and parasitoid females depended on the foraging pattern of the parasitoid, which varied with age. In mixed aphid colonies patch residence time increased with parasitoid age. Furthermore, younger parasitoids (≦1 day old) laid more eggs into second and third instars, while older parasitoids (≧4 days old) did not show distinct host instar preferences. It is suggested that the oviposition behaviour ofL. cardui is influenced by the physiological state, i.e. the age of the wasp.     

Predation on the woodwasp Tremex longicollis Konow (Hymenoptera: Siricidae) and its parasitoid Megarhyssa jezoensis (Matsumura) (Hymenoptera: Ichneumonidae) adults during oviposition

Adults of the woodwasp Tremex longicollis Konow (Hymenoptera: Siricidae) oviposit in the wood of dying Celtis sinensis trees using their ovipositors, and adults of the parasitoid Megarhyssa jezoensis (Matsumura) (Hymenoptera: Ichneumonidae) deposit eggs into the woodwasp larvae that live in the wood, also using their long ovipositors. Many T. longicollis ovipositors, both with and without abdominal segments, and a M. jezoensis ovipositor were found stuck in C. sinensis wood at two parks in Osaka Prefecture, central Japan. Field observations showed that ovipositing woodwasps and parasitoids were attacked by predators during the daytime, and their ovipositors were left behind in the wood. Woodwasps and their parasitoids, which have long ovipositors, may be vulnerable to predation during oviposition.     

Host-plant mediated effects of transgenic maize on the insect parasitoid Campoletis sonorensis (Hymenoptera: Ichneumonidae)

Determining the impact of genetically modified (GM) crops on beneficial organisms is an important aspect of the environmental risk assessment of GM crops. In the present study, the impact of Bt maize expressing Cry1Ab on the development and behaviour of the parasitoid Campoletis sonorensis was compared to individuals reared on hosts fed conventionally bred plants partially resistant to the European corn borer (Ostrinia nubilalis Hübner) and on susceptible maize hybrids. Adult parasitoids reared on Bt maize-fed Spodoptera frugiperda larvae were significantly smaller (15–30%) than those reared in hosts fed either of the conventional maize hybrids. The magnitude of this effect was dependent on the size of the host at oviposition and its subsequent growth rate. The development time of C. sonorensis was not affected by the maize treatment. In choice tests, female parasitoids displayed no preference for hosts fed a specific maize hybrid. No Cry1Ab was detected within adult parasitoids.     

Determination of Homalodisca coagulata (Hemiptera: Cicadellidae) egg ages suitable for oviposition by Gonatocerus ashmeadi, Gonatocerus triguttatus, and Gonatocerus fasciatus (Hymenoptera: Mymaridae)

Homalodisca coagulata (Say) (Hemiptera: Cicadellidae) eggs 1–10 days of age were exposed to Gonatocerus ashmeadi Girault, Gonatocerus triguttatus Girault, and Gonatocerus fasciatus Girault (all Hymenoptera: Mymaridae) in no choice laboratory trials to investigate egg age utilization and to determine which egg ages are vulnerable to attack by these three parasitoids. The H. coagulata egg ages that were most suitable for oviposition by G. ashmeadi, G. triguttatus, and G. fasciatus were eggs 3, 4, and 2 days of age, respectively. Egg ages least suitable for parasitoid development were 6–10 days for G. ashmeadi (resulting in <50% parasitism), 1–2 and 7–10 days for G. triguttatus (resulting in <25% parasitism), and 3–10 days for G. fasciatus (resulting in <11% parasitism). Pooling parasitism data across all egg ages showed that parasitism by G. ashmeadi was 12.9 and 28.5% higher compared with G. triguttatus and G. fasciatus, respectively, and G. triguttatus resulted in 15.6% higher percentage parasitism compared with G. fasciatus. Egg age had a significant effect on the percentage of female G. ashmeadi offspring produced, but this was not significant for G. triguttatus, and low G. fasciatus parasitism prevented statistical analyses for comparisons. Results from tests where females were offered a choice for oviposition between eggs 1, 3, and 5 days of age demonstrated that G. ashmeadi and G. triguttatus showed no significant oviposition preference, while percentage parasitism by G. fasciatus was 29.4 and 7.4% higher when females were presented eggs 1 and 3 days of age, respectively, compared with eggs 5 days of age. Choice tests indicated that an overlap in egg age suitability for oviposition exists between G. ashmeadi, G. triguttatus, and G. fasciatus, and that interspecific competition for eggs 1, 2, and 3 days of age may occur in the field environment.