Terebra steering in chalcidoid wasps.

Various chalcidoid wasps can actively steer their terebra (= ovipositor shaft) in diverse directions, despite the lack of terebral intrinsic musculature. To investigate the mechanisms of these bending and rotational movements, we combined microscopical and microtomographical techniques, together with videography, to analyse the musculoskeletal ovipositor system of the ectoparasitoid pteromalid wasp Lariophagus distinguendus (Förster, 1841) and the employment of its terebra during oviposition. The ovipositor consists of three pairs of valvulae, two pairs of valvifers and the female T9 (9th abdominal tergum). The paired 1st and the 2nd valvulae are interlocked via the olistheter system, which allows the three parts to slide longitudinally relative to each other, and form the terebra. The various ovipositor movements are actuated by a set of nine paired muscles, three of which (i.e. 1st valvifer-genital membrane muscle, ventral 2nd valvifer-venom gland reservoir muscle, T9-genital membrane muscle) are described here for the first time in chalcidoids. The anterior and posterior 2nd valvifer-2nd valvula muscles are adapted in function. (1) In the active probing position, they enable the wasps to pull the base of each of the longitudinally split and asymmetrically overlapping halves of the 2nd valvula that are fused at the apex dorsally, thus enabling lateral bending of the terebra. Concurrently, the 1st valvulae can be pro- and retracted regardless of this bending. (2) These muscles can also rotate the 2nd valvula and therefore the whole terebra at the basal articulation, allowing bending in various directions. The position of the terebra is anchored at the puncture site in hard substrates (in which drilling is extremely energy- and time-consuming). A freely steerable terebra increases the chance of contacting a potential host within a concealed cavity. The evolution of the ability actively to steer the terebra can be considered a key innovation that has putatively contributed to the acquisition of new hosts to a parasitoid's host range. Such shifts in host exploitation, each followed by rapid radiations, have probably aided the evolutionary success of Chalcidoidea (with more than 500,000 species estimated).

Delimiting the cryptic diversity and host preferences of Sycophila parasitoid wasps associated with oak galls using phylogenomic data.

Cryptic species diversity is a major challenge regarding the species-rich community of parasitoids attacking oak gall wasps due to a high degree of sexual dimorphism, morphological plasticity, small size and poorly known biology. As such, we know very little about the number of species present, nor the evolutionary forces responsible for generating this diversity. One hypothesis is that trait diversity in the gall wasps, including the morphology of the galls they induce, has evolved in response to selection imposed by the parasitoid community, with reciprocal selection driving diversification of the parasitoids. Using a rare, continental-scale data set of Sycophila parasitoid wasps reared from 44 species of cynipid galls from 18 species of oak across the USA, we combined mitochondrial DNA barcodes, ultraconserved elements (UCEs), morphological and natural history data to delimit putative species. Using these results, we generate the first large-scale assessment of ecological specialization and host association in this species-rich group, with implications for evolutionary ecology and biocontrol. We find most Sycophila target specific subsets of available cynipid host galls with similar morphologies, and generally attack larger galls. Our results suggest that parasitoid wasps such as Sycophila have adaptations allowing them to exploit particular host trait combinations, while hosts with contrasting traits are resistant to attack. These findings support the tritrophic niche concept for the structuring of plant-herbivore-parasitoid communities.

Barcoding of parasitoid wasps (Braconidae and Chalcidoidea) associated with wild and cultivated olives in the Western Cape of South Africa 1.

Wild and cultivated olives harbor and share a diversity of insects, some of which are considered agricultural pests, such as the olive fruit fly. The assemblage of olive-associated parasitoids and seed wasps is rich and specialized in sub-Saharan Africa, with native species possibly coevolving with their hosts. Although historical entomological surveys reported on the diversity of olive wasp species in the Western Cape Province of South Africa, no comprehensive study has been performed in the region in the molecular era. In this study, a dual approach combining morphological and DNA-based methods was used for the identification of adult specimens reared from olive fruits. Four species of Braconidae and six species of Chalcidoidea were identified, and DNA barcoding methodologies were used to investigate conspecificity among individuals, based on randomly selected representative specimens. Morphological identifications were congruent with DNA data, as NJ and ML trees correctly placed the sequences for each species either at the genus or species level, depending on the available taxa coverage, and genetic distances strongly supported conspecificity. No clear evidence of cryptic diversity was found. Overall seed infestation and parasitism rates were higher in wild olives compared to cultivated olives, and highest for Eupelmus spermophilus and Utetes africanus. These results can be used for early DNA-based detection of wasp larvae in olives and to further investigate the biology and ecology of these species.

Ophelimus mediterraneus sp. n. (Hymenoptera, Eulophidae): a new Eucalyptus gall wasp in the Mediterranean region.

We report here for the first time the presence of Ophelimus mediterraneus sp. n. in Mediterranean Europe. This species appears to be closely related to Ophelimus maskelli, a well-known invasive pest of Eucalyptus. Based on molecular (cytochrome oxidase I, 28S), morphological (multivariate ratio analysis) and bio-ecological investigations, our study gives unambiguous relevant criteria that allow the discrimination between these two species. A full description of O. mediterraneus sp. n. is also provided. The geographic distribution of O. mediterraneus sp. n. as well as its impact on Eucalyptus species needs to be more widely assessed since its presence may have been confused with O. maskelli in their sympatric introduced areas. Further investigations of potential parasitoids in the native area may thus be welcomed to evaluate classical biological control achievability.

Comparative genomics of the miniature wasp and pest control agent Trichogramma pretiosum.

BACKGROUND: Trichogrammatids are minute parasitoid wasps that develop within other insect eggs. They are less than half a millimeter long, smaller than some protozoans. The Trichogrammatidae are one of the earliest branching families of Chalcidoidea: a diverse superfamily of approximately half a million species of parasitoid wasps, proposed to have evolved from a miniaturized ancestor. Trichogramma are frequently used in agriculture, released as biological control agents against major moth and butterfly pests. Additionally, Trichogramma are well known for their symbiotic bacteria that induce asexual reproduction in infected females. Knowledge of the genome sequence of Trichogramma is a major step towards further understanding its biology and potential applications in pest control. RESULTS: We report the 195-Mb genome sequence of Trichogramma pretiosum and uncover signatures of miniaturization and adaptation in Trichogramma and related parasitoids. Comparative analyses reveal relatively rapid evolution of proteins involved in ribosome biogenesis and function, transcriptional regulation, and ploidy regulation. Chalcids also show loss or especially rapid evolution of 285 gene clusters conserved in other Hymenoptera, including many that are involved in signal transduction and embryonic development. Comparisons between sexual and asexual lineages of Trichogramma pretiosum reveal that there is no strong evidence for genome degradation (e.g., gene loss) in the asexual lineage, although it does contain a lower repeat content than the sexual lineage. Trichogramma shows particularly rapid genome evolution compared to other hymenopterans. We speculate these changes reflect adaptations to miniaturization, and to life as a specialized egg parasitoid. CONCLUSIONS: The genomes of Trichogramma and related parasitoids are a valuable resource for future studies of these diverse and economically important insects, including explorations of parasitoid biology, symbiosis, asexuality, biological control, and the evolution of miniaturization. Understanding the molecular determinants of parasitism can also inform mass rearing of Trichogramma and other parasitoids for biological control.

Latitudinal variation in responses of a forest herbivore and its egg parasitoids to experimental warming.

Disrupted biotic interactions are a predicted consequence of anthropogenic climate change when interactants differ in the magnitude or direction of phenological responses. Here, we examined the responses to artificial warming of northern, southern and central populations of the eastern tent caterpillar and its hymenopteran egg parasitoids. We subjected egg masses from each region to the typical conditions they experience in their source locality or to a warmer temperature regime, to quantify the effects of simulated warming on their relative phenology, survival and neonate starvation endurance. In addition, we characterized spring heat accumulation and cloud cover at each collection site using 30 years of hourly weather station data. As predicted, degree-day accumulation rates decreased with latitude; however, the mid-latitude site experienced what we predict to be the harshest spring conditions for tent caterpillars: slow heat accumulation combined with thick cloud cover. Remarkably, caterpillars from this site exhibited the largest phenological plasticity, hatching a month earlier under warmer than under typical conditions and doubling caterpillar survival. Survival of caterpillars from all regions was enhanced at warmer temperatures, whereas parasitoid survival was unaffected. The starvation endurance of hatchlings increased under warmer conditions in the central and southern populations only. We show that phenological responses to warming differed between hosts and parasitoids, resulting in a 5-day reduction in the relative phenology of wasps and caterpillars in the northern population. Our findings caution that responses to global warming are likely to be population or region specific and cannot be readily generalized, particularly for wide-ranging organisms.

Conserved community structure and simultaneous divergence events in the fig wasps associated with Ficus benjamina in Australia and China.

BACKGROUND: Localised patterns of species diversity can be influenced by many factors, including regional species pools, biogeographic features and interspecific interactions. Despite recognition of these issues, we still know surprisingly little about how invertebrate biodiversity is structured across geographic scales. In particular, there have been few studies of how insect communities vary geographically while using the same plant host. We compared the composition (species, genera) and functional structure (guilds) of the chalcid wasp communities associated with the widespread fig tree, Ficus benjamina, towards the northern (Hainan province, China) and southern (Queensland, Australia) edges of its natural range. Sequence data were generated for nuclear and mtDNA markers and used to delimit species, and Bayesian divergence analyses were used to test patterns of community cohesion through evolutionary time. RESULTS: Both communities host at least 14 fig wasp species, but no species are shared across continents. Community composition is similar at the genus level, with six genera shared although some differ in species diversity between China and Australia; a further three genera occur in only China or Australia. Community functional structure remains very similar in terms of numbers of species in each ecological guild despite community composition differing a little (genera) or a lot (species), depending on taxonomic level. Bayesian clustering analyses favour a single community divergence event across continents over multiple events for different ecological guilds. Molecular dating estimates of lineage splits between nearest inter-continental species pairs are broadly consistent with a scenario of synchronous community divergence from a shared "ancestral community". CONCLUSIONS: Fig wasp community structure and genus-level composition are largely conserved in a wide geographic comparison between China and Australia. Moreover, dating analyses suggest that the functional community structure has remained stable for long periods during historic range expansions. This suggests that ecological interactions between species may play a persistent role in shaping these communities, in contrast to findings in some comparable temperate systems.

Transcriptome analysis provides insight into venom evolution in a seed-parasitic wasp, Megastigmus spermotrophus.

One of the most striking host range transitions is the evolution of plant parasitism from animal parasitism. Parasitoid wasps that have secondarily evolved to attack plants (ie gall wasps and seed-feeders) demonstrate intimate associations with their hosts, yet the mechanism of plant-host manipulation is currently not known. There is, however, emerging evidence suggesting that ovipositional secretions play a role in plant manipulation. To investigate whether parasites have modified pre-existing adaptations to facilitate dramatic host shifts we aimed to characterize the expression of venom proteins in a plant parasite using a collection of parasitoid venom sequences as a guide. The transcriptome of a seed-feeding wasp, Megastigmus spermotrophus, was assembled de novo and three putative venoms were found to be highly expressed in adult females. One of these putative venoms, aspartylglucosaminidase, has been previously identified as a major venom component in two distantly related parasitoid wasps (Asobara tabida and Leptopilina heterotoma) and may have originated via gene duplication within the Hymenoptera. Our study shows that M. spermotrophus, a specialized plant parasite, expresses putative venom transcripts that share homology to venoms identified in Nasonia vitripennis (both superfamily Chalcidoidea), which suggests that M. spermotrophus may have co-opted pre-existing machinery to develop as a plant parasite.

Diversity and evolution of Ty1-copia retroelements within Chalcidoidea by reverse transcriptase domain analysis.

Ty1-copia retrotransposons are widespread and diverse in insects. Some features of their hosts, such as mating and genetic systems, are predicted to influence the spread of selfish genetic elements like Ty1-copia. Using part of the reverse transcriptase gene as a reference, we experimentally surveyed Ty1-copia elements in eight species of fig wasps (Hymenoptera: Chalcidoidea), and performed an in silico analysis of six available genomes of chalcid wasps. Contrary to initial expectations that selfish elements such as Ty1-copia would be purged from the genomes of these species because of inbreeding and haplodiploidy, almost all of these wasps harbour an abundance of diverse Ty1-copia elements. Phylogenetic analyses suggest that the families of Ty1-copia elements found in these species have had a long association with their chalcid hosts. These results suggest an evolutionary scenario in which there was ancestral polymorphism followed by some taxa-specific events including stochastic loss and further diversification. Furthermore, estimating natural selection within the internal and terminal portions of the Ty1-copia phylogenies demonstrated that the elements are under strong evolutionary constraints for their long-term survival, but evolve like pseudogenes in the short term, accompanied by the rise and fall of parasitic elements in the history of wasp lineage.

Parasitoid complex associated with the overwintering generation of Swammerdamia pyrella (Lepidoptera: Yponomeutidae) in Poland.

The study was conducted on fruit trees where bands of corrugated cardboard were attached around the trunks of the trees, which were used to catch the larvae of overwintering generation of the rufous-tipped swammerdamia moth, Swammerdamia pyrella (Villers) (Lepidoptera: Yponomeutidae). Twenty-five species of parasitic Hymenoptera have been described from S. pyrella in Poland including the report in this article of seven species belonging to the family of Ichneumonidae (three species) and superfamily Chalcidoidea (four species). The parasitoids Gelis agilis F. (Ichneumonidae), Chrysocharis aquilegiae (Erdös) (Eulophidae), Catolaccus ater (Ratzeburg) (Pteromalidae), and Eupelmus urozonus (Dalman) (Eupelmidae) had not been reported from the host before. Triclistus pallipes Holmgren (Ichneumonidae), Dibrachys cavus Walker (Pteromalidae) had the greatest effect on the natural regulation of S. pyrella population. Parasitization for the wintering cocoons of S. pyrella changed each year, but it was high throughout the study. The contribution of secondary parasitoids was much higher than primary parasitoids.

A taxonomic study of Cheiloneurus Westwood (Hymenoptera, Encyrtidae) from China.

Fourteen species of Cheiloneurus from China are studied. Cheiloneurusguangxiensis Zu, sp. nov., is described as new to science, and C.boldyrevi Trjapitzin & Agekyan, 1978, C.bouceki Anis & Hayat, 2002, C.gonatopodis Perkins, 1906, and C.hadrodorys Anis & Hayat, 2002 are newly recorded from China. A key to Chinese species based on females is also presented.

Horismenus saturnus n. sp. (Hymenoptera, Eulophidae) a New Parasitoid Wasp from Eggs of Saturniidae (Lepidoptera) from the Atlantic Forest, Minas Gerais, Brazil.

A new species of Horismenus Walker, H. saturnus Schoeninger & Hansson (Hymenoptera: Eulophidae), is described from material reared from eggs of an unidentified species of Saturniidae (Lepidoptera). The new species is compared to H. cupreus (Ashmead, 1894), a species it is very similar to, and to H. ancillus (Brèthes), a species with the same type of host as H. saturnus. A total of 30 adult specimens developed from the eggs of Saturniidae. This is the second record of a Horismenus species parasitizing eggs of Saturniidae and the first record of this host from Brazil. Here, we provide a diagnosis and description of the new species including morphological and molecular characters, and multiple illustrations.

Three new species of Quadrastichus Girault (Hymenoptera, Eulophidae) from China with a key to Chinese species.

Six species of Quadrastichus Girault (Eulophidae: Tetrastichinae) from China are reviewed, including three new species: Q.longisetasp. nov., Q.flavomaculatussp. nov., Q.longiscapussp. nov. and one new country record, Q.vacuna (Walker, 1839). New distributional data for Q.anysis (Walker, 1839) and Q.sajoi (Szelényi, 1941), and a key to the Chinese species of Quadrastichus based on females, are included.

Description of three new species and new distributional data for three species of Homalotylus (Hymenoptera, Encyrtidae) from China.

Homalotylustianjinensis Zu, sp. nov., H.bicolor Zu, sp. nov. and H.guangxiensis Zu, sp. nov. are described as new to science; H.agarwali Anis & Hayat, 1998, H.hemipterinus (De Stefani, 1898) and H.varicolorus Krishnachaitanya & Manickavasagam, 2016 are newly recorded from China. A key to Chinese species based on females is also presented.

A new species of Entedon Dalman (Hymenoptera, Eulophidae) and three newly recorded species from China.

In this paper, a new species of Entedon Dalman, E.flavifemursp. nov. is described from Tibet and three species, E.albifemur Kamijo, E.crassiscapus Erdös, and E.nomizonis Kamijo are reported from China for the first time. A detailed description and illustrations of the new species are provided, as well as diagnoses and illustrations of the three newly recorded species.

Evolutionary timescale of chalcidoid wasps inferred from over one hundred mitochondrial genomes.

Chalcidoidea is one of the most biologically diverse groups among Hymenoptera. Members are characterized by extraordinary parasitic lifestyles and extensive host ranges, among which several species attack plants or serve as pollinators. However, higher-level chalcidoid relationships remain controversial. Here, we performed mitochondrial phylogenomic analyses for major clades (18 out of 25 families) of Chalcidoidea based on 139 mitochondrial genomes. The compositional heterogeneity and conflicting backbone relationships in Chalcidoidea were assessed using various datasets and tree inferences. Our phylogenetic results supported the monophyly of 16 families and polyphyly of Aphelinidae and Pteromalidae. Our preferred topology recovered the relationship (Mymaridae+(Signiphoridae+Leucospidae)+(Chalcididae+((Perilampidae+Eucharitidae)+ remaining Chalcidoidea)))). The monophyly of Agaonidae and Sycophaginae was rejected, while the gall-associated ((Megastigmidae+Ormyridae)+(Ormocerinae+Eurytomidae)) relationship was supported in most results. A six-gene inversion may be a synapomorphy for most families, whereas other derived gene orders may introduce confusion in phylogenetic signals at deeper nodes. Dating estimates suggested that Chalcidoidea arose near the Jurassic/Cretaceous boundary and that two dynamic shifts in diversification occurred during the evolution of Chalcidoidea. We hypothesized that the potential codiversification between chalcidoids and their hosts may be crucial for accelerating the diversification of Chalcidoidea. Ancestral state reconstruction analyses supported the hypothesis that gall-inducers were mainly derived from parasitoids of gall-inducers, while other gall-inducers were derived from phytophagous groups. Taken together, these findings advance our understanding of mitochondrial genome evolution in the major interfamilial phylogeny of Chalcidoidea.

A new species and three newly recorded species of Tetrastichinae (Hymenoptera, Eulophidae) from China.

Five species of five genera in Tetrastichinae (Hymenoptera, Eulophidae) from China are reviewed, including one new species, Mestocharellaqingdaoensis sp. nov., and three new country record species: Nesolynxthymus (Girault, 1916), Holcotetrastichusrhosaces (Walker, 1839), and Peckelachertusdiprioni Yoshimoto, 1970. New distributional data for Ceratoneuraindi Girault, 1917 are provided.

Two new species of Hemiptarsenus Westwood (Hymenoptera, Eulophidae) from China, with a key to Chinese species.

Two new species of Hemiptarsenus Westwood, H.tianshuiensis sp. nov. and H.longjiangensis sp. nov., are described from China. New distributional data for H.jilinus Tao, 2021 are provided, and a key to Chinese species of the genus is given based on females.

Four new species of Diglyphomorphomyia Girault (Hymenoptera, Eulophidae) from China, with a key to Chinese species.

Four new species of Diglyphomorphomyia Girault, D.depressa sp. nov., D.fossa sp. nov., D.hainana sp. nov., and D.octoseta sp. nov., are described from China. A key to the eight species of the genus Diglyphomorphomyia occurring in China is provided.

Two new species and a new record of the Encarsialongifasciata-group (Hymenoptera, Aphelinidae) from Malaysia and China.

Background: The genus Encarsia Förster, 1878, which is the largest genus of the family Aphelinidae, contains 453 valid species worldwide. Most species of Encarsia with known biology are primary endoparasitoids of Aleyrodidae and Diaspididae. New information: Species of the Encarsialongifasciata-group from Malaysia and China are reviewed. This is the first record of this group from Malaysia. Two new species, E.borneensis Geng & Li sp. n. and E.pauroseta Geng & Li sp. n., are described and illustrated. Encarsialongifasciata is newly recorded from Malaysia (Borneo). An updated key to the longifasciata-group species (females) worldwide is provided.