Inter and intra-guild interactions in egg parasitoid species of the soybean stink bug complex

The objective of this research was to evaluate the parasitism behavior of Telenomus podisi Ashmead, Trissolcus basalis (Wollaston) e Trissolcus urichi Crawford (Hymenoptera: Scelionidae) on eggs of Nezara viridula L., Euschistus heros F., Piezodorus guildinii Westwood and Acrosternum aseadum Rolston (Heteroptera: Pentatomidae), in no choice and multiple choice experiments. For all parasitoid species, the results demonstrated the existence of a main host species that maximizes the reproductive success. The competitive interactions among the parasitoid species were investigated in experiments of sequential and simultaneous release of different combinations of parasitoid pairs on the hosts N. viridula, E. heros and A. aseadum. Exploitative competition was observed for egg batches at the genus level (Telenomus vs. Trissolcus) and interference competition at the species level (T. basalis vs. T. urichi). Trissolcus urichi was the most aggressive species, interfering with the parasitism of T. basalis. Generally, T. basalis showed an opportunistic behavior trying to parasitise eggs after T. urichi had abandoned the egg batch. The selection of parasitoid species for use in augmentative biological control programs should take into account the diversity of pentatomids present in soybean in addition to the interactions among the different species of parasitoids.

Sensory response of the egg parasitoid Telenomus podisi to stimuli from the bug Euschistus heros

The objective of this work was to study the foraging behavior of Telenomus podisi Ashmead (Hymenoptera: Scelionidae) in the presence of stimuli from its host, Euschistus heros (Heteroptera: Pentatomidae). The stimuli selected were: egg mass; virgin males and females; volatile extracts of sexually mature males and females; components of male sex pheromone; a component of the alarm pheromone, hexane and an empty cage as control. In a closed arena, the parasitoids were given the choice between single and combined stimuli presented to them simultaneously. To find the host egg, T. podisi primarily uses the sensory cues released from the male insects. The orientation toward odors of male chemical extract indicates that a source of kairomone was detected. Gas chromatographic analyses of this substance showed peak of methyl 2,6,10-trimethyltridecanoate, the main component of male sexual pheromone. The sensory response to methyl 2,6,10-trimethyltridecanoate confirms that this compound may act as a kairomone to find host eggs. Females and egg mass stimuli were weakly attractive to the parasitoid.

Egg parasitoid wasps as natural enemies of the neotropical stink bug Dichelops melacanthus

The objective of this work was to determine the potential of five species of Scelionidae wasps - Telenomus podisi, Trissolcus basalis, Trissolcus urichi, Trissolcus teretis and Trissolcus brochymenae - as natural enemies of the neotropical stink bug Dichelops melacanthus, and to determine if the presence of eggs of other stink bug species influences the parasitism and development of the parasitoids. Two kinds of experiments were done in laboratory: without choice of hosts (eggs of D. melacanthus) and with choice (eggs of D. melacanthus and of Euschistus heros). Biological parameters, including proportion of parasitism, immature survivorship, progeny sex ratio, immature stage development period, and host preference were recorded. All the evaluated parasitoids can parasitize and develop on D. melacanthus eggs. The first choice of eggs did not influence the proportion of D. melacanthus eggs parasitized by Tr. basalis, Tr. teretis or Tr. brochymenae. However, D. melacanthus eggs as the first choice of Te. podisi and Tr. urichi increased, respectively, 9 and 14 times the chance for parasitism on eggs of this species. Behavioral and ecological aspects of parasitoids should be considered prior to their use in biological control programs.

Identification of parasitoid species using PCR amplification and restriction enzyme digestion

Spodoptera frugiperda is a pest of great economic importance in the Americas. It is attacked by several species of parasitoids, which act as biological control agents. Parasitoids are morphologically identifiable as adults, but not as larvae. Laboratory rearing conditions are not always optimal to rear out parasitic wasps from S. frugiperda larvae collected from wild populations, and it frequently happens that parasitoids do not complete their life cycle and stop developing at the larval stage. Therefore, we explored ways to identify parasitoid larvae using molecular techniques. Sequencing is one possible technique, yet it is expensive. Here we present an alternate, cheaper way of identifying seven species of parasitoids (Cotesia marginiventris, Campoletis sonorensis, Pristomerus spinator, Chelonus insularis, Chelonus cautus, Eiphosoma vitticolle and Meteorus laphygmae) using PCR amplification of COI gene followed by a digestion with a combination of four restriction endonucleases. Each species was found to exhibit a specific pattern when the amplification product was run on an agarose gel. Identifying larvae revealed that conclusions on species composition of a population of parasitic wasps can be biased if only the emerging adults are taken into account.

First survey of fruit fly (Diptera: Tephritidae) and parasitoid diversity among myrtaceae fruit across the state of Bahia, Brazil

The objective of this study was to evaluate the diversity of fruit fly (Diptera: Tephritidae) species that use myrtaceous fruit, particularly guava, as hosts in several localities in the state of Bahia and to determine the infestation rates, pupal viability rates, and fruit fly-parasitoid associations. Sampling of myrtaceous fruit was carried out in 24 municipalities in different regions in the state of Bahia. Four fruit fly species, Anastrepha fraterculus, Anastrepha zenildae, Anastrepha sororcula, and Ceratitis capitata were obtained from the collected fruit. Three parasitoid species (Hymenoptera: Braconidae) emerged from Anastrepha larvae/pupae, Doryctobracon areolatus, Utetes anastrephae, and Asobara anastrephae. Doryctobracon areolatus emerged from A. fraterculus, A. sororcula and A. zenildae; Utetes anastrephae emerged from A. fraterculus and A. zenildae; and Asobara anastrephae emerged from A. fraterculus. Fruit fly and myrtaceous fruit associations are reported for the first time in several municipalities in the state of Bahia. A. zenildae was found infesting Syzygium malaccense for the first time in Brazil.

Diploid males support a two-step mechanism of endosymbiont-induced thelytoky in a parasitoid wasp.

BACKGROUND: Haplodiploidy, where females develop from diploid, fertilized eggs and males from haploid, unfertilized eggs, is abundant in some insect lineages. Some species in these lineages reproduce by thelytoky that is caused by infection with endosymbionts: infected females lay haploid eggs that undergo diploidization and develop into females, while males are very rare or absent. It is generally assumed that in thelytokous wasps, endosymbionts merely diploidize the unfertilized eggs, which would then trigger female development. RESULTS: We found that females in the parasitoid wasp Asobara japonica infected with thelytoky-inducing Wolbachia produce 0.7-1.2 % male offspring. Seven to 39 % of these males are diploid, indicating that diploidization and female development can be uncoupled in A. japonica. Wolbachia titer in adults was correlated with their ploidy and sex: diploids carried much higher Wolbachia titers than haploids, and diploid females carried more Wolbachia than diploid males. Data from introgression lines indicated that the development of diploid individuals into males instead of females is not caused by malfunction-mutations in the host genome but that diploid males are most likely produced when the endosymbiont fails to activate the female sex determination pathway. Our data therefore support a two-step mechanism by which endosymbionts induce thelytoky in A. japonica: diploidization of the unfertilized egg is followed by feminization, whereby each step correlates with a threshold of endosymbiont titer during wasp development. CONCLUSIONS: Our new model of endosymbiont-induced thelytoky overthrows the view that certain sex determination mechanisms constrain the evolution of endosymbiont-induced thelytoky in hymenopteran insects. Endosymbionts can cause parthenogenesis through feminization, even in groups in which endosymbiont-diploidized eggs would develop into males following the hosts' sex determination mechanism. In addition, our model broadens our understanding of the mechanisms by which endosymbionts induce thelytoky to enhance their transmission to the next generation. Importantly, it also provides a novel window to study the yet-poorly known haplodiploid sex determination mechanisms in haplodiploid insects.

Absence of complementary sex determination in the parasitoid wasp genus Asobara (Hymenoptera: Braconidae).

An attractive way to improve our understanding of sex determination evolution is to study the underlying mechanisms in closely related species and in a phylogenetic perspective. Hymenopterans are well suited owing to the diverse sex determination mechanisms, including different types of Complementary Sex Determination (CSD) and maternal control sex determination. We investigated different types of CSD in four species within the braconid wasp genus Asobara that exhibit diverse life-history traits. Nine to thirteen generations of inbreeding were monitored for diploid male production, brood size, offspring sex ratio, and pupal mortality as indicators for CSD. In addition, simulation models were developed to compare these observations to predicted patterns for multilocus CSD with up to ten loci. The inbreeding regime did not result in diploid male production, decreased brood sizes, substantially increased offspring sex ratios nor in increased pupal mortality. The simulations further allowed us to reject CSD with up to ten loci, which is a strong refutation of the multilocus CSD model. We discuss how the absence of CSD can be reconciled with the variation in life-history traits among Asobara species, and the ramifications for the phylogenetic distribution of sex determination mechanisms in the Hymenoptera.

Uncovering Cryptic Parasitoid Diversity in Horismenus (Chalcidoidea, Eulophidae).

Horismenus parasitoids are an abundant and understudied group of eulophid wasps found mainly in the New World. Recent surveys based on morphological analyses in Costa Rica have quadrupled the number of named taxa, with more than 400 species described so far. This recent revision suggests that there is still a vast number of unknown species to be identified. As Horismenus wasps have been widely described as parasitoids of insect pests associated with crop plants, it is of high importance to properly establish the extant diversity of the genus, in order to provide biological control practitioners with an exhaustive catalog of putative control agents. In this study, we first collected Horismenus wasps from wild Phaseolus bean seeds in Central Mexico and Arizona to assess the genetic relatedness of three morphologically distinct species with overlapping host and geographical ranges. Sequence data from two nuclear and two mitochondrial gene regions uncovered three cryptic species within each of the three focal species (i.e., H. missouriensis, H. depressus and H. butcheri). The monophyly of each cryptic group is statistically supported (except in two of them represented by one single tip in which monophyly cannot be tested). The phylogenetic reconstruction is discussed with respect to differences between gene regions as well as likely reasons for the differences in variability between species.

Unique features of odorant-binding proteins of the parasitoid wasp Nasonia vitripennis revealed by genome annotation and comparative analyses

Insects are the most diverse group of animals on the planet, comprising over 90% of all metazoan life forms, and have adapted to a wide diversity of ecosystems in nearly all environments. They have evolved highly sensitive chemical senses that are central to their interaction with their environment and to communication between individuals. Understanding the molecular bases of insect olfaction is therefore of great importance from both a basic and applied perspective. Odorant binding proteins (OBPs) are some of most abundant proteins found in insect olfactory organs, where they are the first component of the olfactory transduction cascade, carrying odorant molecules to the olfactory receptors. We carried out a search for OBPs in the genome of the parasitoid wasp Nasonia vitripennis and identified 90 sequences encoding putative OBPs. This is the largest OBP family so far reported in insects. We report unique features of the N. vitripennis OBPs, including the presence and evolutionary origin of a new subfamily of double-domain OBPs (consisting of two concatenated OBP domains), the loss of conserved cysteine residues and the expression of pseudogenes. This study also demonstrates the extremely dynamic evolution of the insect OBP family: (i) the number of different OBPs can vary greatly between species; (ii) the sequences are highly diverse, sometimes as a result of positive selection pressure with even the canonical cysteines being lost; (iii) new lineage specific domain arrangements can arise, such as the double domain OBP subfamily of wasps and mosquitoes.

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra-Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in large colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV2 of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV2>1 for A. funebris between plant patches, while for T. centaureae CV2>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50-200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Differential susceptibilities of peach-potato aphid, Myzus persicae (Homoptera: Aphididae) and its parasitoid, Aphidius matricariae (Hymenoptera: Aphidiidae) to foliar insecticides on partially resistant and susceptible potato cultivars

In a field experiment the effects of Sumicidin (super) 5EC (fenitrothion), Metasystox EC25 (oxydemeton-methyl) and Tamaron SL600 (methamidophos), applied at different dosages, were evaluated against peach-potato aphid, Myzus persicae (Sulzer) and its parasitoid Aphidius matricariae Haliday on Cardinal and Desiree (respectively partially resistant and susceptible potato cultivars to M. persicae). Sumicidin (super) 5EC was found about 30% more effective in reducing aphid populations than the other insecticides tested. The highest doses of each insecticide caused maximum aphid mortality; in general aphid mortality appeared dose dependent. Almost all the higher and lower doses of the tested insecticides were about 19% more effective on Cardinal than on Desiree. The most significant result was the synergistic interaction at the lower doses with plant resistance, so that the same level of control was recorded with second highest dose on Cardinal as with the highest dose on Desiree. Also the same control level was achieved at the lowest dosage rate on Cardinal compared with the next higher dose on the Desiree. Sumicidin (super) 5EC was found least toxic to the parasitoid, A. matricariae in terms of percent parasitism, emergence of parasitoids and number of mature eggs in the emerging female parasitoids; increase of about 22, 67 and 47% respectively were found in parasitoid performance with Tamaron SL600 which was found comparatively highly toxic. The highest doses of all insecticides were found clearly toxic to the parasitoid. In general, effects on the parasitoid were dose dependent. Maximum yield was obtained from the second highest dose of Sumicidin (super) 5EC.

Selective toxicity of different granular insecticides to the peach-potato aphid, Myzus persicae (Homoptera: Aphididae) and its parasitoid, Aphidius matricariae (Hymenoptera: Aphidiidae) in two differentially resistant potato cultivars

Field studies were conducted in Pakistan to examine the effects and the interaction of two differentially resistant potato cultivars i.e. Cardinal and Desiree (one partially resistant and one susceptible to Myzus persicae (Sulzer), respectively) with different dosage rates of granular insecticides, at different time intervals after application in relation to percent kill of M. persicae and effects on the parasitoid Aphidius matricariae Haliday (i.e. the third trophic level) within the aphid mummies, percent parasitism and Thimet 10G (phorate) was found about 30% more effective in reducing aphid population than the Furadan 3G (carbofuran). The highest doses of each insecticide caused maximum aphid mortality, in general aphid mortality appeared dose dependent. Mostly all the higher and lower doses of the tested insecticides were about 10% more effective on Cardinal than on Desiree. The most significant result was the synergistic interaction at the lower doses with plant resistance, so that the same level of control was recorded with the second highest dose on Cardinal as with the highest dose on Desiree. Also the same level of control was observed at the lowest dose on Cardinal as with the second last lowest dose on Desiree. Furadan 3G was found least toxic to the A. matricariae in terms of percent parasitism, emergence of parasitoids and number of mature eggs in the emerging females. Furadan 3G gave 13, 15 and 6% higher figures, respectively from the parasitoid characteristics than Thimet 10G. The highest doses of both insecticides were clearly toxic to the parasitoid. In general, the effects on the parasitoid were dose dependent. The second highest dose of Thimet 10G, gave the maximum yield

Host niches and defensive extended phenotypes structure parasitoid wasp communities

Oak galls are spectacular extended phenotypes of gallwasp genes in host oak tissues and have evolved complex morphologies that serve, in part, to exclude parasitoid natural enemies. Parasitoids and their insect herbivore hosts have coevolved to produce diverse communities comprising about a third of all animal species. The factors structuring these communities, however, remain poorly understood. An emerging theme in community ecology is the need to consider the effects of host traits, shaped by both natural selection and phylogenetic history, on associated communities of natural enemies. Here we examine the impact of host traits and phylogenetic relatedness on 48 ecologically closed and species-rich communities of parasitoids attacking gall-inducing wasps on oaks. Gallwasps induce the development of spectacular and structurally complex galls whose species- and generation-specific morphologies are the extended phenotypes of gallwasp genes. All the associated natural enemies attack their concealed hosts through gall tissues, and several structural gall traits have been shown to enhance defence against parasitoid attack. Here we explore the significance of these and other host traits in predicting variation in parasitoid community structure across gallwasp species. In particular, we test the "Enemy Hypothesis,'' which predicts that galls with similar morphology will exclude similar sets of parasitoids and therefore have similar parasitoid communities. Having controlled for phylogenetic patterning in host traits and communities, we found significant correlations between parasitoid community structure and several gall structural traits (toughness, hairiness, stickiness), supporting the Enemy Hypothesis. Parasitoid community structure was also consistently predicted by components of the hosts' spatiotemporal niche, particularly host oak taxonomy and gall location (e.g., leaf versus bud versus seed). The combined explanatory power of structural and spatiotemporal traits on community structure can be high, reaching 62% in one analysis. The observed patterns derive mainly from partial niche specialisation of highly generalist parasitoids with broad host ranges (>20 hosts), rather than strict separation of enemies with narrower host ranges, and so may contribute to maintenance of the richness of generalist parasitoids in gallwasp communities. Though evolutionary escape from parasitoids might most effectively be achieved via changes in host oak taxon, extreme conservatism in this trait for gallwasps suggests that selection is more likely to have acted on gall morphology and location. Any escape from parasitoids associated with evolutionary shifts in these traits has probably only been transient, however, due to subsequent recruitment of parasitoid species already attacking other host galls with similar trait combinations.