Braconid Subfamily Research Articles

First records of Miracinae (Hymenoptera: Braconidae) from Borneo with description of two new species

The braconid subfamily Miracinae is recorded for the first time from Borneo. Two new species are described: Rugosimirax obsolescens sp. nov. and Rugosimirax bimacula sp. nov. based on specimens in the Natural History Museum, UK. The relationships of Miracinae genera are discussed.

Ophiclypeus, a new genus of Cardiochilinae Ashmead (Hymenoptera, Braconidae) from the Oriental region with descriptions of three new species.

A new genus of the braconid subfamily Cardiochilinae, Ophiclypeusgen. nov., is described and illustrated based on three new species: O.chiangmaiensis Kang, sp. nov. type species (type locality: Chiang Mai, Thailand), O.dvaravati Ghafouri Moghaddam, Quicke & Butcher, sp. nov. (type locality: Saraburi, Thailand), and O.junyani Kang, sp. nov. (type locality: Dalin, Taiwan). We provide morphological diagnostic characters to separate the new genus from other cardiochiline genera. A modified key couplet (couplet 5) and a new key couplet (couplet 16) are provided with detailed images for Dangerfield's key to the world cardiochiline genera to facilitate recognition of Ophiclypeusgen. nov.

A new genus in the braconid subfamily Miracinae from the Oriental region, with descriptions of seven new species from India and Sri Lanka.

A new genus in the braconid subfamily Miracinae is described from the Oriental region including a new species from south India. Additionally, six new species of the genus Centistidea Rohwer is described from India and Sri Lanka. Taxonomic keys to separate the new genus from other miracine genera, to the extant species of Rugosimirax gen. nov. and to the Oriental species of Centistidea are provided.

A new fossil euphorine genus and species (Hymenoptera, Braconidae) with the longest known ovipositor from Dominican amber

A new genus and species of the braconid subfamily Euphorinae, Palaeorionis longicaudisgen. et sp. nov., is described and illustrated from Miocene Dominican amber. This new genus is characterised by the long and tube-shaped petiole, presence of both radiomedial (2-SR and r-m) veins on the infuscate fore wing and long ovipositor.

Chelonus inanitus bracovirus encodes lineage-specific proteins and truncated immune IκB-like factors.

Bracoviruses and ichnoviruses are endogenous viruses of parasitic wasps that produce particles containing virulence genes expressed in host tissues and necessary for parasitism success. In the case of bracoviruses the particles are produced by conserved genes of nudiviral origin integrated permanently in the wasp genome, whereas the virulence genes can strikingly differ depending on the wasp lineage. To date most data obtained on bracoviruses concerned species from the braconid subfamily of Microgastrinae. To gain a broader view on the diversity of virulence genes we sequenced the genome packaged in the particles of Chelonus inanitus bracovirus (CiBV) produced by a wasp belonging to a different subfamily: the Cheloninae. These are egg-larval parasitoids, which means that they oviposit into the host egg and the wasp larvae then develop within the larval stages of the host. We found that most of CiBV virulence genes belong to families that are specific to Cheloninae. As other bracoviruses and ichnoviruses however, CiBV encode v-ank genes encoding truncated versions of the immune cactus/IκB factor, which suggests these proteins might play a key role in host-parasite interactions involving domesticated endogenous viruses. We found that the structures of CiBV V-ANKs are different from those previously reported. Phylogenetic analysis supports the hypothesis that they may originate from a cactus/IκB immune gene from the wasp genome acquired by the bracovirus. However, their evolutionary history is different from that shared by other V-ANKs, whose common origin probably reflects horizontal gene transfer events of virus sequences between braconid and ichneumonid wasps.

A new genus and three new species of fossil braconid wasps (Hymenoptera, Ichneumonoidea) from Eocene Baltic and Rovno ambers

AbstractNew parasitoid wasps of the family Braconidae are documented from Eocene Baltic and Rovno ambers. A new fossil genus belonging to the braconid subfamily Exothecinae, Palaeocolastes n. gen., with its type species P. bruesi n. sp., is described and illustrated from Baltic amber. This represents the first reliable fossil record for a member of Exothecinae. Two additional new species from Baltic amber are also described: Ascogaster (Syntaphus) latitibialis n. sp. (Cheloninae) and Meteorus arasnitsyni n. sp. (Euphorinae). Another fossil species, Microtypus eocenus n. sp. (Microtypinae), is described from coeval Rovno amber (Ukraine), representing the first braconid species described from this deposit. A new record of a female of Diospilites brevicornis Brues, 1933 (Diospilitinae) from Baltic amber, together with variation of some diagnostic features of the species and redescription of its subfamily and genus, are also provided.UUID: http://zoobank.org/656cb1a3-b9cf-4696-ae24-0d4df9545101.

Four new species of parasitoid wasp (Hymenoptera: Braconidae) described through a citizen science partnership with schools in regional South Australia.

Involving the community in taxonomic research has the potential to increase the awareness, appreciation and value of taxonomy in the public sphere. We report here on a trial citizen science project, Insect Investigators, which partners taxonomists with school students to monitor Malaise traps and prioritise the description of new species collected. In this initial trial, four schools in regional South Australia participated in the program and all collected new species of the braconid subfamily Microgastrinae (Hymenoptera: Braconidae). These four species are here described as new, with the names being chosen in collaboration with the participating school students: Choeras ramcomarmorata Fagan-Jeffries Austin sp. nov., Glyptapanteles drioplanetus Fagan-Jeffries Austin sp. nov., Dolichogenidea franklinharbourensis Fagan-Jeffries Austin sp. nov. and Miropotes waikerieyeties Fagan-Jeffries Austin sp. nov. All four species are diagnosed against the known members of the genera from Australia, New Zealand, Fiji, Samoa and Papua New Guinea, and images and COI DNA barcodes are provided of the holotypes. Students had positive feedback about their experiences of the program, and there is significant potential for it to be expanded and used as a means to connect communities with taxonomic science.

Discovery of the ichneutine genus Paroligoneurus Muesebeck (Hymenoptera: Braconidae) from the Indian subcontinent with the description of a new species from Northwest India.

The small braconid subfamily, Ichneutinae is one of the least studied groups within the Braconidae. Old world species of the widely distributed ichneutine genus, Paroligoneurus (unknown from the Neotropical region) is revised along with description of a new species, P. harishi Ranjith van Achterberg sp. nov. We reinstate the genera Anaprixia Mason and Muesebeckia Mason based on the following characters; exodont mandibles and absence of spiracles in the alternate tergites (in Anaprixia) and presence of carinae on propodeum and anterior subalar depression with carina (in Muesebeckia). Five species, Paroligoneurus cosmopterygivorus (He, 2000), P. crassicornis (He, 2000), P. flavifacialis (He, 2000), P. sinensis (He, 2000) and P. songyangensis (He, 2000) were described from the Oriental part of China are transferred from Oligoneurus to Paroligoneurus. A taxonomic key to the Old world species Paroligoneurus and a provisional checklist of extant species are provided.

Phylogenomics and mitochondrial genome evolution of the gall-associated doryctine wasp genera (Hymenoptera: Braconidae)

The braconid subfamily Doryctinae is a cosmopolitan, highly diverse group of wasps mainly represented by parasitoid species. In this subfamily, however, there is a group of mainly Neotropical genera whose species are associated with galls of several vascular plant families. Previous molecular phylogenetic studies based on a few markers consistently recovered the monophyly of this group, though various relationships among genera and the monophyly of its most speciose genus, Allorhogas Gahan, remained unresolved. We characterized the mitogenomes of 13 representative species of the gall-associated doryctine clade and performed a phylogenomic analysis with both mitogenome and ultraconserved element (UCE) DNA sequence data to assess the classification of Doryctinae and to examine the evolution of phytophagy within the group. We found different patterns of tRNA gene rearrangements, two of which are present in most of the ingroup taxa. All phylogenetic analyses yielded highly similar, well-supported topologies that are congruent with relationships reported in previous studies. The resulting phylogenies confirmed Percnobracon as sister to the remaining genera and supported two separate clades whose genera are exclusively associated with Ficus (Moraceae). Allorhogas is confirmed to be polyphyletic. The combined evidence also supported a main Allorhogas clade composed of multiple phytophagous species that feed on at least five different host plant families. Further rearing records and phylogenetic studies are needed to examine host plant shifts and to broaden our understanding of a poorly studied radiation of gall-associated wasps.

New species of Australian microgastrine parasitoid wasps (Hymenoptera: Braconidae: Microgastrinae) documented through the 'Bush Blitz' surveys of national reserves.

The braconid subfamily Microgastrinae are ecologically important parasitoids of larval lepidopterans, but are poorly studied in many regions of the world. In this study, we focus on describing new species of microgastrine wasps, in part from specimens collected on six different 'Bush Blitz' surveys of regional reserves in South Australia and Tasmania. Ten species of Microgastrinae are described as new and DNA barcodes of the genes COI and wingless are provided: three species in the genus Choeras Mason: C. bushblitz Fagan-Jeffries Austin sp. nov., C. parvoculus Fagan-Jeffries Austin sp. nov., and C. zygon Fagan-Jeffries Austin sp. nov.; six species in the genus Dolichogenidea Viereck: D. bonbonensis Fagan-Jeffries Austin sp. nov., D. brabyi Fagan-Jeffries Austin sp. nov., D. forrestae Fagan-Jeffries Austin sp. nov., D. garytaylori Fagan-Jeffries Austin sp. nov., D. kelleri Fagan-Jeffries Austin sp. nov., and D. lobesiae Fagan-Jeffries Austin sp. nov.; and one species from the genus Sathon Mason: S. oreo Fagan-Jeffries Austin sp. nov. These new species represent just a small fraction of the potential of 'Bush Blitz' surveys in regional Australia, which provide DNA-quality material allowing an integrative taxonomic approach and offer a window into the biodiversity of some of the least studied areas of the continent.

New insights on the biology and larval morphology of Neotropical Mesostoinae (Hymenoptera, Braconidae): Hydrangeocola llaollin sp. nov. an endoparasitoid of Centrodiplosis crassipes (Diptera, Cecidomyiidae)

The Southern Hemisphere subfamily Mesostoinae (Hymenoptera: Braconidae) includes, in most cases, species associated with plant galls, although the details regarding their biological traits are largely unknown. In this contribution, Hydrangeocola llaollin sp. nov. is described based on specimens that develop as larval endoparasitoids of Centrodiplosis crassipes (Diptera: Cecidomyiidae) in stem galls on Lycium chilense (Solanaceae). The mature larva of a Neotropical mesostoine wasp is described here for the first time and is compared with the only known larva from Australia. Larval morphology is entirely congruent with features typical of endoparasitoid species, which together with male genital features clearly relates Hydrangeocola with the South American genus Andesipolis, but separates it from the Australian phytophagous genus Mesostoa. These findings reveal the relevance of larval and male genital characters for our comprehension of the relationships among representatives of this poorly known braconid subfamily.

Systematics of the Neotropical braconid wasps of the Pedinotus genus group (Doryctinae)

AbstractThe braconid subfamily Doryctinae (Hymenoptera) is a cosmopolitan, species‐rich group of parasitoid wasps whose known species richness and morphological diversity is mainly concentrated in the Neotropical region. Among the doryctine taxa that are endemic to this region, there is a group of six genera (the Pedinotus genus group) that are mainly characterized by having lateral, posteriorly converging or subparallel grooves on the second metasomal tergite (including Gymnobracon Szépligeti, Hybodoryctes Szépligeti, Lamquetia Braet et van Achterberg, Osmophila Szépligeti, Pedinotus Szépligeti and Trigonophasmus Enderlein). Most of these genera were described more than a century ago, and thus their limits and diagnostic morphological features are not well defined. We present an analysis of the phylogenetic relationships among various representative species of five of the six genera belonging to the Pedinotus genus group based on one mitochondrial (cytochrome oxidase I) and two nuclear (28S, Wingless) gene markers. We also estimated the timing of origin and diversification within the group and investigated the external morphology of the genera involved in order to determine reliable identification characters. The group as a whole, whilst being recovered as monophyletic with the inclusion of Acanthorhogas Szépligeti, was not statistically supported. However, the respective monophyly of all of the morphologically supported genera are confirmed, as are the intergeneric relationships (Gymnobracon, Trigonophasmus) ((Acanthorhogas, Osmophila) (Lamquetia, Pedinotus)). We propose that the pattern of sublateral grooves or depressions appears to be a useful systematic character for recognition of an endemic Neotropical radiation that could have originated c. 41–27 Ma. We redescribe six of the seven genera that we recognize for the Pedinotus genus group and provide a key to enable their identification.

Phylogeny of the parasitic wasp subfamily Euphorinae (Braconidae) and evolution of its host preferences

AbstractThe braconid subfamily Euphorinae is a large, cosmopolitan group of endoparasitoid wasps. The majority of species attack adult hosts, a strategy that is rare among parasitic wasps, but there are also many species that attack nymphs and larval stages. Euphorine hosts may belong to a variety of insect orders (Coleoptera, Hemiptera, Hymenoptera, Neuroptera, Psocoptera, Orthoptera and Lepidoptera) although most euphorine tribes are confined to Coleoptera. Here we investigate the phylogenetic relationships of the Euphorinae based on molecular data (3 kb of nucleotide data from four markers:18S,28S,CADandCOI) and propose a higher‐level classification based upon the resulting phylogeny. We also infer the evolution of host associations and discuss the diversification of the Euphorinae. Results from both Bayesian inference and maximum‐likelihood analysis show that the subfamily, as previously circumscribed, is paraphyletic. We propose that the subfamily be expanded to include the tribes Meteorini and Planitorini (Mannokeraia + Planitorus), so that it corresponds to a clade that is strongly supported as monophyletic in our analyses. Based on our results, a revised higher classification of the Euphorinae is proposed, in which 52 extant genera and 14 tribes are recognized. We reinstate the genusMicroctonusbelonging to the tribe Perilitini, and synonymizeUssurarideluswithHoldawayella, SinuatophoruswithEucosmophorus. Furthermore, we propose the following tribal rearrangements:SpathicopisandStenothremmaare transferred to Perilitini;Tuberidelus, EucosmophorusandPlynopsto Cosmophorini;Eccliturato Dinocampini;Chrysopophthorus, HoldawayellaandWesmaeliato Helorimorphini;ProclithroporusandHeiato Townesilitini. The monotypic tribe Cryptoxilonini is synonymized with Cosmophorini. The generaPygostolusandLitostolusare placed in a separate tribe, Pygostolini, previously recognized as a subtribe among the Centistini. Parsimony‐based ancestral state reconstructions suggest that the ancestor of Euphorinae was a parasitoid of lepidopteran larvae, and that a host shift to larval Coleoptera occurred only in one clade of the Meteorini, some members of which secondarily shifted back to larval lepidopteran hosts. In the remainder of the subfamily, there was an initial shift from larval to adult coleopterans, followed by subsequent shifts to adults or larvae of Hemiptera, Hymenoptera, Neuroptera, Orthoptera and Psocoptera.

DNA barcoding species inventory of Microgastrinae wasps (Hymenoptera, Braconidae) from a Mexican tropical dry forest

The cosmopolitan Microgastrinae is probably the most diverse braconid subfamily of parasitoid wasps, yet its species diversity is far from being known. As part of a global initiative for DNA barcoding Microgastrinae species, here we show the results of a study that assessed the species richness of this subfamily in a Mexican tropical dry forest located in the Chamela region, near the Pacific coast of Jalisco. Barcoding sequences of a total of 551 microgastrine specimens were generated, corresponding to 238 haplotypes. Performance of two species delineation approaches, a 2% corrected pairwise distance criterion and the general mixed Yule-coalescent (GMYC) method, yielded 100 and 112 putative species, respectively, which belong to 13 genera. The species delimited by the above two approaches were mostly congruent with our morphospecies identification. Ten molecular operational taxonomic units (MOTUs) were split into twenty-two species by the GMYC approach. We found morphological differences between the GMYC species corresponding to three of these MOTUs. Thus, a total of 103 microgastrine species were confirmed for the region of study. Thirty-three species were only represented by males, and therefore, their generic assignment is only tentatively proposed. Fornicia, Dolichogenoidea, Distatrix, Glyptapanteles and Pholetesor represent new generic records for the Mexican territory. A new record for the country is also provided for the Diolcogaster-basimacula species group. Based on a comparison of nearly 20000 barcoding sequences released for Microgastrinae from 75 countries, only five microgastrine species from Chamela were found to occur in other countries, four in Costa Rica and one in Canada and the United States.

Phylogenetic relationships and description of Bolivar, a new genus of Neotropical doryctine wasps (Hymenoptera : Braconidae)

Metasomal elongation is a common feature in species of various parasitoid Hymenoptera, probably due to adaptive morphological convergence to similar parasitoid strategies. The braconid subfamily Doryctinae is perhaps where this feature has evolved the most times independently. Here we recognise a new Neotropical doryctine wasp genus with a petiolate first metasomal tergum, based on molecular and morphological analysis. The phylogenetic affinities of the new genus within Doryctinae and the relationships among six of its described and three potentially cryptic, undescribed species were reconstructed using sequence data from three genes, wingless, 28SrDNA and COI. The new genus is resolved in a clade together with Semirhytus Szépligeti, Johnsonius Marsh and Parallorhogas Marsh. These four genera share vein m-cu of the hind wing slightly curved distally and the propodeum with a distinct lateral and median longitudinal carina and an apical areola. The relationships recovered among the examined species suggest a South American origin for the new genus and its subsequent diversification into Central America and Mexico. Described as Bolivar, gen. nov., this new taxon comprises eight species, two species previously placed within Notiospathius Matthews & Marsh, B. ornaticornis (Cameron), comb. nov., and B. bribri (Marsh), comb. nov., and six new species (B. ecuadorensis, sp. nov., B. helmuthi, sp. nov., B. pittieri, sp. nov., B. risaraldae, sp. nov., B. teres, sp. nov. and B. tuxtlae, sp. nov.).

Diversity, distribution and taxonomy of the Australian agathidine genera Camptothlipsis Enderlein, Lytopylus Foerster and Therophilus Wesmael (Hymenoptera: Braconidae: Agathidinae)

The braconid subfamily Agathidinae is a large group of koinobiont endoparasitic wasps of lepidopteran larvae. Until recently, three of the 10 agathidine genera that occur in Australia, Camptothlipsis Enderlein, Lytopylus Foerster and Therophilus Wesmael, were treated as synonyms of Bassus F. s.l. Of these three genera, Therophilus is the most speciose and widely distributed in Australia, and is one of only two agathidine genera whose members are associated with a putative mimicry complex of braconid wasps and other insects comprising species that have a distinctive black, red-orange and white colour pattern. Australian species, previously considered under Bassus s.l., have received little attention since their original description nearly 90 years ago and, not surprisingly, this earlier work is insufficient for reliable species identification. The present study updates the taxonomy of the described species, presents a more thorough assessment of intraspecific variation, and provides a key for species of Therophilus. Four new species are described that support morphological and molecular phylogenetic studies on the Australian fauna: Camptothlipsis oliveri Stevens n. sp., representing the first described species for this genus in Australia, and Therophilus aalvikorum Stevens n. sp., T. mishae Stevens n. sp., and T. stephensae Stevens n. sp., whose descriptions also extend the morphological limits of Therophilus in Australia. In addition, the introduced Lytopylus rufipes (Nees von Esenbeck) is redescribed, this species representing the only member of the genus known from Australia. Significantly, two species of Therophilus, T. unimaculatus (Turner) and T. rugosus (Turner), are important parasitoids of the native Australian lepidopterans Etiella behrii Zeller (Pyralidae) and Epiphyas postvittana (Walker) (Tortricidae) that have become significant pests in southern and eastern Australia, as well as in several other countries.

Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics.

BackgroundThe braconid subfamily Rogadinae is a large, cosmopolitan group of endoparasitoid wasps characterised by 'mummifying' their lepidopteran host larvae, from which the adult subsequently emerges. Rogadines attack a variety of both macro- and microlepidopteran taxa, although the speciose genus Aleiodes almost exclusively attacks macrolepidopterans. Here, we investigate the phylogenetic history of the Rogadinae, revise their higher-level classification and assess the evolution of their host ranges and mummy types. We also assess the divergence times within the subfamily and discuss the reasons for the extraordinary evolutionary diversification of Aleiodes.ResultsOur Bayesian analyses weakly support the monophyly of the subfamily. A clade comprising all Aleiodes species and some other taxa is not nested within the tribe Rogadini as previously supposed, but instead is recovered as sister to the Yeliconini, with the remaining Rogadini genera being recovered as sister to the Stiropiini. The Rogadinae is estimated to have originated during the mid to late Eocene, 36.1–51.62 MYA. Molecular dating gives a more recent origin for the Aleiodes clade (17.98–41.76 MYA) compared to the origins proposed for two of its principal lepidopteran host groups (Noctuidae: 60.7–113.4 MYA; Geometridae 48–62 MYA). The Bayesian ancestral reconstruction of the emergence habits from the mummified hosts weakly recovered an anterior emergence as the ancestral condition for the subfamily. Producing a hard mummy has evolved at various times independently, though most of the species with this biology belong to the Aleiodes clade.ConclusionBased on our results, we erect the tribe Aleiodini nov. to include Aleiodes and Heterogamus stat. rev. Cordylorhogas, Pholichora and Hemigyroneuron are synonymised with Aleiodes. The molecular dating of clades and the ancestral reconstruction of host ranges support the hypothesis that radiation within Aleiodes s. s. was due to host recruitment leading to host range expansion followed by speciation, and not to parasitoid-host coevolution. Within the Rogadinae, variation in the site of emergence from the mummified host probably evolved as a consequence of the mummy's site and mode of formation, and the extent of mummy tanning/hardness to the degree of protection needed in relation to the cost of providing it.

Mexican cardiochiline genera (Hymenoptera: Braconidae), including a preliminary assessment of species-groups in Toxoneuron Say and Retusigaster Dangerfield, Austin and Whitfield

The Mexican species of the braconid subfamily Cardiochilinae are reviewed. Forty-five species are recognized from Mexico, though many of these remain undescribed. Seven species are described as new and one, Toxoneuron levis (Mao), is redescribed. Species-groups are delineated for the New World species of Toxoneuron Say and Retusigaster Dangerfield, Austin and Whitfield and the genera are re-defined with respect to one another. Two new species are described in both Retusigaster and Toxoneuron. The atypical nature of the two included in Toxoneuron is discussed. Hansonia, Heteropteron and Schoenlandella are newly recorded for Mexico with one new species described for each. The placement of all New World species previously included in Schoenlandella is discussed, and several of these species are returned to Cardiochiles s. s. Distribution patterns of the Mexican species are discussed and an annotated key is presented for the 28 described species recorded from Mexico.

The biology and early stages of Acampsis alternipes (Nees), with comments on the relationships of the Sigalphinae (Hymenoptera: Braconidae)

Details of the biology of the braconid subfamily Sigalphinae, based on rearings of Acampsis alternipes (Nees), are provided for the first time. A. alternipes is a solitary, koinobiont larval endoparasitoid of the geometrid moth Alsophila Hubner, though there is an external feeding phase prior to spinning a cocoon and pupation. Two features are of potential phylogenetic significance indicating a close relationship with the subfamily Agathidinae: (1) eggs are deposited in the host's nerve ganglia, and (2) the first instar larva is polypodiform. A. alternipes completes development in the autumn and overwinters as a fully eclosed adult within its cocoon, enabling it to emerge and search for hosts early in spring. The posterior of the eclosed adult within the cocoon is immersed in a viscous liquid that also surrounds the meconium. Chemical analysis of this liquid revealed that it comprises at least 25% glycerol and a further 25% consists of various fatty acid derivatives. Oviposition and developmental stages are illustrated. The overwintering behaviour of the agathidine braconid Earinus elator (Fabricius) is described and compared.

A Phylogeny of Aphid Parasitoids (Hymenoptera: Braconidae: Aphidiinae) Inferred From Mitochondrial NADH 1 Dehydrogenase Gene Sequence

Wasps of the braconid subfamily Aphidiinae are solitary endoparasitoids of aphids. Several aspects of their biology have been the focus of intuitive evolutionary hypotheses which could be tested with a robust phylogeny. Phylogenetic hypotheses have been proposed previously for aphidiines based on morphology, embryology, and DNA sequences. However, many of them are based on a limited number of characters and/or taxa and lack congruence. In addition, many of the inferred phylogenies have not been based upon cladistic analysis. Therefore, a phylogenetic study of Aphidiinae was undertaken, utilizing 465 bp of DNA sequence of the mitochondrial NADH1 dehydrogenase gene. DNA sequences were obtained from 40 taxa, including 14 genera and three outgroups. It is suggested that in agreement with most of the previously proposed phylogenies, the aphidiines, each of the three recognized tribes (Praini, Ephedrini, Aphidiini), and most genera are monophyletic. In contrast to previously proposed phylogenies, the clade ofPraon+Dyscritulus(=Praini), rather than Ephedrini, is basal among the aphidiines.