Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae

A new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses. Eulophidae was strongly supported as monophyletic only after exclusion of the enigmatic genus Trisecodes. The subfamilies Eulophinae, Entiinae (=Euderinae) and Tetrastichinae were consistently supported as monophyletic, but Entedoninae was monophyletic only in combined analyses. Six contiguous bases in the 3e′ subregion of the 28S D2 rDNA contributed to placement of nominal subgenus of Closterocerus outside Entedoninae. In all cases, Euderomphalini was excluded from Entiinae, and we suggest that it be retained in Entedoninae. Opheliminae n. stat. is raised from tribe to subfamily status. Trisecodes is removed from Entedoninae but retained as incertae sedis in Eulophidae until its family placement can be determined new placement . The genera Neochrysocharis stat. rev. and Asecodes stat. rev. are removed from synonymy with Closterocerus because strong molecular differences corroborate their morphological differences. Closterocerus (Achrysocharis) germanicus is transferred to the genus Chrysonotomyia n. comb. based on molecular and morphological characters.     

Phylogenetic implications of the mesofurca in Chalcidoidea (Hymenoptera), with emphasis on Aphelinidae

The skeletomusculature of the mesofurcal–mesopostnotal complex is surveyed within the Chalcidoidea. Four internal character systems are assessed for their phylogenetic significance: the mesofurcal bridge, the structure and position of the furcal–laterophragmal muscle, the structure of the lateral arms of the mesofurca, and the supporting structures for the interfurcal muscles. Among Hymenoptera, Chalcidoidea are unique in having the furcal–laterophragmal muscle attached along the entire length of the laterophragmal apodeme. Also the furcal–laterophragmal muscle originates medial to the junction of the mesofurcal bridge and lateral mesofurcal arm in most Chalcidoidea. Mymarommatidae do not share either of these apomorphic states with Chalcidoidea. Within Chalcidoidea, apomorphic character states were found in each of Aphelinidae, Encyrtidae, Eulophidae, Mymaridae, Rotoitidae, Signiphoridae, Tanaostigmatidae and Trichogrammatidae. For taxa classified as Aphelinidae, the plesiomorphic complement of structures and muscle attachments is retained in Eriaphytinae and Eriaporinae. The mesofurcal bridge is considered to have been lost at least twice in each of Aphelininae and Coccophaginae. Similar interfurcal processes, resulting from loss of the mesofurcal bridge, support the monophyly of Aphelininae (Aphelinini, Aphytini and Eutrichosomellini). Azotinae are placed as the sister group of Aphelininae because of a similar lateral origin of the laterophragmal muscle and the shape of the mesofurcal arms. Other than loss of the mesofurcal bridge, no character states were shared by Azotinae and Coccophaginae. Coccophaginae (Coccophagini and Pteroptricini) are regarded as monophyletic based on the loss of the mesofurcal bridge, the peculiar shape of the mesofurca, and a unique modification of the laterophragmal muscle. Euxanthellus is removed from synonomy with Coccophagus and may be best treated as a separate tribe of Coccophaginae based on the shape of the lateral mesofurcal arms and the presence of a mesofurcal bridge. The shape of the mesofurca suggests a monophyletic grouping of Cales, Eretmocerus and Trichogrammatidae that could render Aphelinidae paraphyletic.     

The phylogeny of lower Hymenoptera (Insecta), with a summary of the early evolutionary history of the order

A cladistic analysis of the lower Hymenoptera, including all the ‘symphytan’ families and the apocritan families Stephanidae, Megalyridae, Trigonalyidae, Ibaliidae, Vespidae and Gasteruptiidae, has been undertaken. A total of 98 characters were scored for 21 taxa. Twenty equally parsimonious minimum-length trees were obtained. The phylogenetic status of the Xyelidae is uncertain: they might be monophyletic. or the Xyelinae might be the sistergroup of the rest of the Hymenoptera. The non-xyelid Hymenoptera are probably monophyletic; the phylogeny Tenthredinoidea + (Megalodontoidea + (Cephidae + (Anaxyelidea + (Siricidae + (Xiphydriidae + (Orussidae + Apocrita)))))) is proposed for this clade. The Blasticotomidae are probably the sistergroup of all othe Tenthredinoidea, but tenthredinoid phylogeny is otherwise uncertain. Substantial homoplasy occurs within the ‘siricoid’ families, making the relative positions of the Anaxyelidae and Siricidae uncertain. The Stephanidae might be the sistergroup of the rest of the Apocrita; the phylogeny of the remaining apocritan taxa included is insufficiently elucidated. The phylogeny proposed here supports the hypothesis that the appearance of parasitism in the Hymenoptera took place in the common ancestor of Orussidae + Apocrita, the host of which was probably wood boring insect larvae. The exact larval mode of feeding of the ancestral hymenopteran cannot be determined due ot the diversity of lifestyles in the basal lineages of the order.     

Grooming behaviours in the Hymenoptera (Insecta): potential phylogenetic significance

Grooming behaviours from representatives of 36 families of Hymenoptera were video recorded and analysed. Thirty-three distinct types of grooming movements were recognized. The evolutionary pattern of each behaviour is discussed. Some behaviours displayed consistent variation between taxa, and appear to be informative regarding higher level relationships within the order. Putative synapomorphies are reported that support the monophyly of the Apocrita, Cynipoidea, Platygastroidea, Ichneumonoidea, and Chalcidoidea including the Mymaridae.     

A new species of Aprostocetus Westwood (Hymenoptera: Eulophidae), parasitizing mango leaf gall midge (Diptera: Cecidomyiidae), from India

A new species of Aprostocetus doonensis Singh sp. nov. is described from northern India. The new species is a parasitoid of mango leaf gall midge, Procontarinia mangiferae (Felt) (Diptera: Cecidomyiidae). Some observations on biology and parasitization rate are also given.     

Susceptibility to insecticides of Glyptapanteles militaris (Hymenoptera: Braconidae), a Parasitoid of Pseudaletia unipuncta (Lepidoptera: Noctuidae)

The susceptibility of cocoons and adults of Glyptapanteles militaris (Walsh) were studied. One organophosphate insecticide (trichlorfon), one organochlorine insecticide (endosulfan), one pyrethroid (deltamethrin) and a commercial formulation of Bacillus thuringiensis subsp. Kurstaki were selected for testing. All the tests were carried out with fresh solutions of commercial insecticides applied on host larvae at the recommended concentration. One- and 6-day-old cocoons were sprayed with the insecticide solutions by means of a Potter Tower and held for adult emergence. Adults were exposed to residues of insecticides inside plastic vials. The B. thuringiensis formulation had no harmful effect on the cocoons nor on the adults. Trichlorfon and endosulfan were highly toxic to all the tested stages. Deltamethrin was highly toxic to adults but relatively safe when applied on parasitoid cocoons. Based on these results, field applications of deltamethrin would be least disruptive of tested insecticides to populations of G. militaris.     

Laying the foundations for a new classification of Agaonidae (Hymenoptera: Chalcidoidea), a multilocus phylogenetic approach

A phylogeny of the Agaonidae (Chalcidoidea) in their restricted sense, pollinators of Ficus species (Moraceae), is estimated using 4182 nucleotides from six genes, obtained from 101 species representing 19 of the 20 recognized genera, and four outgroups. Data analysed by parsimony and Bayesian inference methods demonstrate that Agaonidae are monophyletic and that the previous classification is not supported. Agaonidae are partitioned into four groups: (i) Tetrapus, (ii) Ceratosolen + Kradibia, (iii) some Blastophaga + Wiebesia species, and (iv) all genera associated with monoecious figs and a few Blastophaga and Wiebesia. The latter group is subdivided into subgroups: (i) Pleistodontes, (ii) Blastophaga psenes and neocaledonian Dolichoris, (iii) some Blastophaga and Wiebesia species, and (iv) Platyscapa, all afrotropical genera and all genera associated with section Conosycea. Eleven genera were recovered as monophyletic, six were para‐ or polyphyletic, and two cannot be tested with our data set. Based on our phylogeny we propose a new classification for the Agaonidae. Two new subfamilies are proposed: Tetrapusiinae for the genus Tetrapus, and Kradibiinae for Ceratosolen + Kradibia. Liporrhopalum is synonymized with Kradibia and the subgenus Valisia of Blastophaga is elevated to generic rank. These changes resulted in 36 new combinations. Finally, we discuss the hypothesis of co‐speciation between the pollinators and their host species by comparing the two phylogenies. © The Willi Hennig Society 2009.     

Long-term monitoring of the introduced emerald ash borer (Coleoptera: Buprestidae) egg parasitoid,Oobius agrili (Hymenoptera: Encyrtidae), in Michigan,USA and evaluation of a newly developed monitoring technique

Emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a serious invasive pest of ash trees (Fraxinus spp.) in North America from China. The egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) was introduced from China as a biological control agent for this pest in Michigan and throughout the infested area of the United States. A critical component of any biological control program is post-release monitoring and evaluation; however, because of the small size and cryptic nature of O. agrili, evaluation of its impact is difficult. We compared two methods for measuring parasitism of emerald ash borer eggs: (1) timed visual searches of bark on standing ash trees and (2) bark collection, sifting, and sorting. Both methods were carried out in paired parasitoid-release and control plots, the visual search method over a six-year period (2008–2013) and the more recently developed bark-collection and sifting method for 2 years (2012–2013). The visual search method found parasitism in release plots remained low (0.7–4.2%) in samples taken from 2008 to 2012 and reached 10.6% in 2013. In comparison, the bark-sifting method found that rates of egg parasitism were considerably higher in release plots, 21.8% and 18.9% for samples taken in 2012 and 2013, respectively. These findings indicate that the population-level impact of O. agrili is increasing and may be an important source of mortality for EAB populations. We recommend the bark-collection and sifting method as the more effective method to recover parasitoids and estimate parasitism rates of O. agrili.     

Qualitative aspects of sperm stock in males and females from Eupelmus orientalis and Dinarmus basalis (Hymenoptera: Chalcidoidea) as revealed by dual fluorescence

Abstract The quality of a sperm population can be characterized physiologically and its fecundity predicted by its viable : non-viable sperm ratio. To improve the knowledge of reproductive strategies in two ectoparasitoid hymenopteran species, Eupelmus orientalis Crawford (Hymenoptera: Eupelmidae) and Dinarmus basalis Rondani (Hymenoptera: Pteromalidae) , the assessment of sperm viability using the dual fluorescence staining procedure SYBR-14 : propidium iodide was developed. The aim of the study was to provide a comparative test in vitro applicable to both sexes to study the evolution of sperm quality at various stages of the reproductive processes. The reliability of propidium iodide to detect non-viable sperm (stained in red) was confirmed in both species on the basis of two stress tests (ethanol and Triton X-100) but our study also revealed that propidium iodide concentrations must be adequately adjusted for each single species. This experiment also demonstrated the physiological heterogeneity of sperm populations in E. orientalis and D. basalis males and females. In both species, 40% of the sperm in the seminal vesicles was found to be non-viable. By contrast with E. orientalis, the populations of non-viable sperm estimated from the seminal vesicles of D. basalis were found to be strongly different from those observed in the spermatheca. From the present results, the population of viable sperm detected in the spermatheca of females from both species proved a reliable predictor of fertilization achieved in ovipositing females.     

Re-curation of the Linnaean Hymenoptera (Insecta), with a reassessment of the taxonomic importance of the collection

The history of acquisition and maintenance of the Linnaean insect collections is briefly summarized. Curatorial details (such as kinds of label) are discussed, chiefly with reference to Hymenoptera. Some misleading opinions concerning the taxonomic value of the collections and the nature of Linnaean type-material are examined and a practical approach is recommended. The rationale governing re-curation of the collections is detailed, and the present contents of the collection of Hymenoptera are listed in order to inform interested specialists.     

Phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with a reclassification of Eulophinae and the recognition that Elasmidae are derived eulophids

Eulophidae is a large and biologically varied family of parasitoid wasps, traditionally split into four subfamilies; Elasmidae is a uniform (single genus) and morphologically distinct family of wasps that are thought to be related to Eulophidae. The D2 region of the 28S rDNA gene (≈ 560 bp) of eighty‐seven species of eulophid, three species of elasmid and sixteen outgroup species in five families was sequenced. Cladograms were constructed, and the results compared with conclusions drawn from morphological studies. The gene was most informative at the level of subfamily and tribe. The monophyly of both Eulophinae and Tetrastichinae is supported; that of Entedoninae and Euderinae is less clear. Results indicate that Eulophinae is a derived group within Eulophidae, rather than an ancestral group as previously thought, and that Elasmus, the sole genus of Elasmidae, belongs within this subfamily. The tribes of Eulophinae are reassessed and only three accepted: Eulophini (including Euplectrini and Elachertini), Elasmini and Cirrospilini LaSalle trib.n. for Bou?ek's Ophelimini with Ophelimus and Australsecodes excluded. Three small Australian tribes, Anselmellini, Ophelimini and Platytetracampini, are removed from Eulophinae and Entedoninae, respectively, but their exact relationships and subfamily status cannot as yet be decided. Another tribe, Keryini, known from a single Australian genus, is excluded from both Eulophinae and Eulophidae.     

Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters

In a previous study of the phylogeny of basal Hymenoptera, Vilhelmsen (2001; Zool. J. Linn. Soc . 131 : 393–442) compiled an extensive morphological data matrix for a phylogenetic analysis of basal Hymenoptera, comprising 38 hymenopteran genera. In this study, his characters are revised. This results in a cladogram whose relationships largely agree with those proposed by Vilhelmsen, except that the relationships at the base of the Hymenoptera are unresolved. The revised data matrix is expanded by 17 sawfly and three apocritan taxa. Moreover, 112 new morphological characters from different parts of the larval and adult morphology are also added to the data matrix, including 82 from a recent study of the terminal abdominal segments of male Hymenoptera. The addition of the new characters leads to Xyelidae, again, being the sister-group of all other Hymenoptera. The relationships among the sawfly families as proposed by Vilhelmsen are confirmed, except that the relationships among Syntexis , Siricidae and Xiphydriidae + Vespina are unresolved and that the monophyly of Apocrita is not convincingly supported. A separate analysis is performed which includes all extant genera of Xyelidae. The internal phylogeny of Xyelidae is determined as (( Macroxyela Megaxyela ) Xyelecia ( Xyela Pleroneura )).  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 209–243.     

Morphology and evolution of the tarsal plantulae in Hymenoptera (Insecta), focussing on the basal lineages

The morphology of the plantulae (= tarsal pulvilli = plantar lobes), structures attached to the underside of the tarsus in Hymenoptera, was examined in 55 genera from all 14 families of the basal lineages of Hymenoptera ('Symphyta') and a few species of Apocrita, using scanning electron microscopy. Two distinct types of plantula were found: (1) integrated , an unsclerotized patch positioned ventro-distally on each tarsomere, and (2) distal , a membranous vesicle attached to the apical end of each tarsomere. The evolution of these two types is discussed in the light of current phylogenetic hypotheses. The plantulae exhibit an astonishing variety of form and structural details; their potential phylogenetic and taxonomic value is discussed.     

Biology of Coeloides sordidator (Hymenoptera: Braconidae), a Possible Candidate for Introduction Against Pissodes strobi (Coleoptera: Curculionidae) in North America

Various aspects of the biology of Coeloides sordidator, an ectoparasitoid of pine weevils of the genus Pissodes, were studied. When reared in the laboratory on P. castaneus, females preferred to oviposit on third and fourth instar larvae in galleries, but ovipositions were also observed on second instars and prepupae. Pupae were not accepted as hosts. At 23°C and a 16 h day length, pre-imaginal development lasted 15-20 days. For each parasitoid population tested, between 8 and 28% of the individuals diapaused as prepupae in cocoons. The rate of diapause was not influenced by the photoperiod during larval development. In contrast, diapause was induced by a short day photoperiod on parents before and during oviposition and by low temperatures during the larval stage. Diapause termination was achieved by cold storage at 2°C for at least 8 weeks. Rearing of C. sordidator on P. castaneus under natural conditions showed that in the Swiss Jura it is mainly univoltine, with less than 10% of the parasitoids emerging for a second generation. When C. sordidator females were offered healthy P. castaneus larvae and larvae parasitized by the braconid endoparasitoid Eubazus semirugosus, there was no evidence that they were able to discriminate between healthy and parasitized hosts. The potential of C. sordidator as a biological control agent against the North American species P. strobi is discussed.     

安徽稻田小蜂分科检索（膜翅目：小蜂总科）

对中国安徽芜湖稻田中的小蜂总科进行了分类研究,共发现9科12属,编制了分科检索表,提供了各科主要的分类特征图。     

Discovery of a couple of Foenobethylus (Hymenoptera,Bethylidae) in copulation suggests a synonym under Parascleroderma

Foenobethylus Kieffer 1913 is a rare and small genus of Pristocerinae in the flat wasp family Bethylidae, with 10 described species worldwide, all of which are known only from males. In this study, we describe a new species, F. syndesis Chen & Azevedo, sp. nov., based on a couple in copulation and one additional male from China. An updated key to species of the genus is provided. In light of the first discovery of the female of Foenobethylus, the differences between Foenobethylus and Parascleroderma are discussed.     

Suitability of three aphid species for Aphidius gifuensis (Hymenoptera: Braconidae): Parasitoid performance varies with hosts of origin

Oviposition behavior and offspring fitness of the parasitoid Aphidius gifuensis (Ashmead) were compared on three aphid species, Sitobion avenae F., Myzus persicae (Sulzer), and Aphis gossypii Glover using wasps collected from both S. avenae and M. persicae. A. gifuensis produced more mummies and adults on S. avenae and M. persicae than on A. gossypii regardless of the host of origin. Mummy production was influenced by attack rate and percentage of aphids superparasitized. The F₁ generations from S. avenae and M. persicae were more female-biased and wasps were larger than those from A. gossypii. Although there were significant differences in development time of A. gifuensis in the three aphid species, the difference was generally shorter than one day. Fewer mummies were produced when A. gifuensis was transferred between S. avenae and M. persicae, but no significant difference was observed in emergence rate, percentage of female offspring, or body size. The effects of host species on A. gifuensis female performance and offspring fitness are discussed, along with the potential for using A. gifuensis to control M. persicae and A. gossypii.