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 camobiensis (Hym.: Eulophidae), a new hyperparasitoid of Cotesia invirae (Hym.: Braconidae) in Opsiphanes invirae (Lep.: Nymphalidae) pupae

Abstract Horismenus camobiensis sp. nov. (Hymenoptera: Eulophidae), is described based on morphological, molecular and ecological data; this new species of chalcid wasp acts as hyperparasitoid of Opsiphanis invirae (Hübner, 1818) (Lepidoptera: Nymphalidae) in its parasitoid Cotesia invirae Salgado-Neto and Whitfield, 2019 (Hymenoptera: Braconidae). Diagnoses with morphological and molecular characters and illustrations are provided.

Resumo Horismenus camobiensis sp. nov. (Hymenoptera: Eulophidae) é descrita com base em dados morfológicos, moleculares e ecológicos; esta nova espécie Chalcididae atua como hiperparasitoide de Opsiphanis invirae (Hübner, 1818) (Lepidoptera: Nymphalidae) em pupas de seu parasitoide Cotesia invirae Salgado-Neto and Whitfield, 2019 (Hymenoptera: Braconidae). Caracteres diagnósticos morfológicos e moleculares e ilustrações de H. camobiensis são fornecidos.

Horismenus camobiensis (Hym Eulophidae), a new hyperparasitoid of Cotesia invirae (Hym Braconidae) in Opsiphanes invirae (Lep Nymphalidae) pupae / Horismenus camobiensis (Hym Eulophidae), um novo hiperparasitóide de Cotesia invirae (Hym Braconidae) em pupas de Opsiphanes invirae (Lep Nymphalidae)

Abstract Horismenus camobiensis sp. nov. (Hymenoptera: Eulophidae), is described based on morphological, molecular and ecological data; this new species of chalcid wasp acts as hyperparasitoid of Opsiphanis invirae (Hübner, 1818) (Lepidoptera: Nymphalidae) in its parasitoid Cotesia invirae Salgado-Neto and Whitfield, 2019 (Hymenoptera: Braconidae). Diagnoses with morphological and molecular characters and illustrations are provided.

Resumo Horismenus camobiensis sp. nov. (Hymenoptera: Eulophidae) é descrita com base em dados morfológicos, moleculares e ecológicos; esta nova espécie Chalcididae atua como hiperparasitoide de Opsiphanis invirae (Hübner, 1818) (Lepidoptera: Nymphalidae) em pupas de seu parasitoide Cotesia invirae Salgado-Neto and Whitfield, 2019 (Hymenoptera: Braconidae). Caracteres diagnósticos morfológicos e moleculares e ilustrações de H. camobiensis são fornecidos.

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.

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).

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.

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.

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.

A Compositional Heterogeneity Analysis of Mitochondrial Phylogenomics in Chalcidoidea Involving Two Newly Sequenced Mitogenomes of Eupelminae (Hymenoptera: Chalcidoidea).

As next-generation sequencing technology becomes more mature and the cost of sequencing continues to fall, researchers are increasingly using mitochondrial genomes to explore phylogenetic relationships among different groups. In this study, we sequenced and analyzed the complete mitochondrial genomes of Eupelmus anpingensis and Merostenus sp. We predicted the secondary-structure tRNA genes of these two species and found that 21 of the 22 tRNA genes in Merostenus sp. exhibited typical clover-leaf structures, with trnS1 being the lone exception. In E. anpingensis, we found that, in addition to trnS1, the secondary structure of trnE was also incomplete, with only DHU arms and anticodon loop remaining. In addition, we found that compositional heterogeneity and variable rates of evolution are prevalent in Chalcidoidea. Under the homogeneity model, a Eupelmidae + Encyrtidae sister group relationship was proposed. Different datasets based on the heterogeneity model produced different tree topologies, but all tree topologies contained Chalcididae and Trichogrammatidae in the basal position of the tree. This is the first study to consider the phylogenetic relationships of Chalcidoidea by comparing a heterogeneity model with a homogeneity model.

Hiding in plain sight: Cryptic enemies are found on cochineal (Hemiptera: Dactylopiidae), a scale insect of economic and cultural significance.

Cochineal is the common name for cactus-feeding scale insects in the Dactylopiidae. These ruby-red insects include the domesticated dye insect Dactylopius coccus. Dactylopius coccus and congeners have been introduced around the world, some accidentally, to become pests of prickly pear cactus species (Opuntia), and some intentionally, for dye production or biological control of pest Opuntia. In the northern Sonoran Desert (Tucson, AZ, USA), we studied the enemy complex of D. opuntiae and D. confusus on Opuntia and characterized two cryptic enemies, a coccinellid beetle predator and a parasitoid wasp. (1) Hyperaspis sp. The coccinellid predator Hyperaspis trifurcata shares a niche with a similar, typically all-black beetle. Morphological data, crossing tests, and phylogenetic results showed the black beetle to be a distinct, undescribed species in the genus Hyperaspis, with a rare spotted phenotype that is similar in appearance to H. trifurcata. Crossing tests among black and spotted forms showed the spotted morph is inherited as a single-locus dominant allele. (2) Formicencyrtus thoreauini. Rearing of this ant-like parasitoid wasp (Encyrtidae) in pure culture of D. opuntiae showed it to be a semi-gregarious primary parasitoid of cochineal. To our knowledge, this is the first confirmed instance of a cochineal parasitoid. Observations of development show early instar larvae keep their posterior end within the egg chorion, attached to an aeroscopic plate with a connection to the cochineal body wall. Late instar larvae are eventually surrounded by a membrane, likely of larval origin. Wasps then pupate in a dry air-filled chamber within the desiccated scale remains before chewing out as an adult. Both Hyperaspis sp. and F. thoreauini may have restricted distributions. Hyperaspis sp. does not appear to be a member of the cochineal community in Mexico or Texas, and scant records suggest F. thoreauini may also be restricted to the Southwestern USA.

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.

Revision of the World Species of Megaphragma Timberlake (Hymenoptera: Trichogrammatidae).

Megaphragma species are important models for basic organismal research, and many are potential biological control agents. We present the first extensive revision of species of the genus Megaphragma based on morphological and molecular data. Our revision includes all previously described species, 6 of which are synonymized, and 22 of which are described here as new. We also provide the first key to all species of the genus and reconstruct their phylogeny based on 28S and CO1 molecular markers. The following species are synonymized with M. longiciliatum Subba Rao: M. aligarhensis Yousuf and Shafee syn. nov.; M. amalphitanum Viggiani syn. nov.; M. decochaetum Lin syn. nov.; M. magniclava Yousuf and Shafee syn. nov.; M. shimalianum Hayat syn. nov.M. anomalifuniculi Yuan and Lou syn. nov. is synonymized with M. polychaetum Lin. The following species are described as new: M. antecessor Polaszek and Fusu sp. nov.; M. breviclavum Polaszek and Fusu sp. nov.; M. chienleei Polaszek and Fusu sp. nov.; M. cockerilli Polaszek and Fusu sp. nov.; M. digitatum Polaszek and Fusu sp. nov.; M. fanenitrakely Polaszek and Fusu sp. nov.; M. funiculatum Fusu, Polaszek, and Viggiani sp. nov.; M. giraulti Viggiani, Fusu, and Polaszek sp. nov.; M. hansoni Polaszek, Fusu, and Viggiani sp. nov.; M. kinuthiae Polaszek, Fusu, and Viggiani sp. nov.; M. liui Polaszek and Fusu sp. nov.; M. momookherjeeae Polaszek and Fusu sp. nov.; M. nowickii Polaszek, Fusu, and Viggiani sp. nov.; M. noyesi Polaszek and Fusu sp. nov.; M. pintoi Viggiani sp. nov.; M. polilovi Polaszek, Fusu, and Viggiani sp. nov.; M. rivelloi Viggiani sp. nov.; M. tamoi Polaszek, Fusu, and Viggiani sp. nov.; M. tridens Fusu, and Polaszek sp. nov.; M. uniclavum Polaszek and Fusu sp. nov.; M. vanlentereni Polaszek and Fusu sp. nov.; M. viggianii Fusu, Polaszek, and Polilov sp. nov.

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.

Notes on the Genus Aceratoneuromyia Girault (Hymenoptera: Eulophidae).

Fruit flies in the family Tephritidae are well known as economically important pests of edible fruits and can often cause serious damage and losses to both agriculture and the economy. One of the common parasitoids of fruit flies, Aceratoneuromyia indica (Silvestri), has been used in biological programs. However, the biocontrol utilities of parasitoids are impeded by the difficulties of proper identification. Species of the genus Aceratoneuromyia Girault (Hymenoptera: Eulophidae), usually developed as parasitoids of fruit flies, are studied here. Trjapitzinichus Kostjukov and Kosheleva is proposed as a new synonym under Aceratoneuromyia. Three new species of Aceratoneuromyia, A. bilinis Huangfu and Cao sp. nov., A. carinata Cao and Zhu sp. nov., and A. trilinus Cao and Zhu sp. nov., are described and illustrated from China. Aceratoneuromyia indica is also treated here with diagnosis and illustrations. DNA barcodes of A. bilinis and A. indica and a key to the world species of Aceratoneuromyia are provided. This study provided important identification information of parasitoids with morphology and molecular evidence, which is useful for imperative needs regarding the identity of parasitoids attacking fruit flies.

On the genus Coccophagus Westwood (Hymenoptera, Aphelinidae) from Xishuangbanna Rainforest. Contribution I: Two new species of the Coccophagusvarius group, with an identification key and phylogenetic analysis.

Two new species belonging to the varius group of Coccophagus, C.breviclavulus sp. nov. and C.perlucidus sp. nov., are described from Xishuangbanna Rainforest (China, Yunnan). Coccophagusanchoroides (Huang) and C.yunnana Wang, Huang & Polaszek are recorded. A tentative key to world species of this group is provided. Partial nuclear ribosomal 28S-D2 of these four species and other six species were sequenced and subjected to a phylogenetic analysis. Phylogeny of C.varius group is discussed.

Chinese Species of the Genus Pseudanaphes Noyes & Valentine (Hymenoptera: Mymaridae) with Description of a New Species.

The fairyfly Mymaridae (Hymenoptera: Chalcidoidea) are widely distributed worldwide, but species of this family have rarely been collected and recorded from the Qinghai-Tibet Plateau. In this study, mymarids collected in Tibet, China, are identified based on morphology and molecular data. Two species of the genus Pseudanaphes Noyes & Valentine are treated and illustrated here, including a known species, P. zhaoi Lin, and a new species, P. yadongicus Aishan & Cao sp. nov. In addition, a key to the world species of Pseudanaphes (females) and DNA barcodes for P. yadongicus and P. zhaoi are provided.

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 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.

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

Two new species of Deutereulophus Schulz, D.felix sp. nov. and D.daguisiensis sp. nov., are described from China. A key to species of Deutereulophus known from China is 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.