The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps.

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the "Angiosperm Terrestrial Revolution". Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3-180.5 Ma) and a crown age of 162.2 Ma (153.9-169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history.

Mischochalcis, an enigmatic new genus of Chalcididae (Hymenoptera: Chalcidoidea) from the Old World tropics with the description of a new species.

A new chalcidid genus, Mischochalcis (Hymenoptera: Chalcididae), is erected with the description M. enigmatus sp. nov. from the Western Ghats as the type species. An Afrotropical species, initially described in Haltichella Spinola, is transferred into the new genus as Mischochalcis inermis (Schmitz) comb. nov. The new genus belongs to the very diverse subfamily Haltichellinae and differs from other genera of the subfamily in having an obliquely truncate metatibia in which the truncation ends in a spine formed by the continuation of an additional carina. The systematic placement of Mischochalcis and its presence in two distant biogeographic realms showing disrupted distribution are discussed.

Ultra-Conserved Elements and morphology reciprocally illuminate conflicting phylogenetic hypotheses in Chalcididae (Hymenoptera, Chalcidoidea).

Recent technical advances combined with novel computational approaches have promised the acceleration of our understanding of the tree of life. However, when it comes to hyperdiverse and poorly known groups of invertebrates, studies are still scarce. As published phylogenies will be rarely challenged by future taxonomists, careful attention must be paid to potential analytical bias. We present the first molecular phylogenetic hypothesis for the family Chalcididae, a group of parasitoid wasps, with a representative sampling (144 ingroups and seven outgroups) that covers all described subfamilies and tribes, and 82% of the known genera. Analyses of 538 Ultra-Conserved Elements (UCEs) with supermatrix (RAxML and IQTREE) and gene tree reconciliation approaches (ASTRAL, ASTRID) resulted in highly supported topologies in overall agreement with morphology but reveal conflicting topologies for some of the deepest nodes. To resolve these conflicts, we explored the phylogenetic tree space with clustering and gene genealogy interrogation methods, analyzed marker and taxon properties that could bias inferences and performed a thorough morphological analysis (130 characters encoded for 40 taxa representative of the diversity). This joint analysis reveals that UCEs enable attainment of resolution between ancestry and convergent/divergent evolution when morphology is not informative enough, but also shows that a systematic exploration of bias with different analytical methods and a careful analysis of morphological features is required to prevent publication of artifactual results. We highlight a GC content bias for maximum-likelihood approaches, an artifactual mid-point rooting of the ASTRAL tree and a deleterious effect of high percentage of missing data (>85% missing UCEs) on gene tree reconciliation methods. Based on the results we propose a new classification of the family into eight subfamilies and ten tribes that lay the foundation for future studies on the evolutionary history of Chalcididae.

Description of Eniacomorpha hermetiae Delvare sp. n. (Hymenoptera, Chalcidoidea, Chalcididae) a pupal parasitoid of Hermetia illucens (L.) (Diptera, Stratiomyidae), and a potential threat to mass production of the fly as a feed supplement for domestic animals.

Eniacomorpha hermetiae Delvare sp. n. (Hymenoptera, Chalcididae, Dirhininae), reared from pupae of black soldier fly (BSF), Hermetia illucens (Linnaeus, 1758) (Diptera, Stratiomyidae), is described and illustrated from Africa and compared with other similar species newly considered as forming the ehrhorni species-group within Eniacomorpha Girault, 1915. The newly described parasitic wasp may have a negative impact on efforts to mass produce BSF in Africa as a feed supplement for domestic animals. Eniacomorpha is removed from synonymy under Dirhinus Dalman, 1818, revised status, for the Afrotropical species of Dirhininae previously placed in Dirhinus subgenus Pareniaca Crawford, 1913. A checklist of the 10 recognized species of Afrotropical Eniacomorpha is given, of which 9 are new generic combinations.

Deciphering host-parasitoid interactions and parasitism rates of crop pests using DNA metabarcoding.

An accurate estimation of parasitism rates and diversity of parasitoids of crop insect pests is a prerequisite for exploring processes leading to efficient natural biocontrol. Traditional methods such as rearing have been often limited by taxonomic identification, insect mortality and intensive work, but the advent of high-throughput sequencing (HTS) techniques, such as DNA metabarcoding, is increasingly seen as a reliable and powerful alternative approach. Little has been done to explore the benefits of such an approach for estimating parasitism rates and parasitoid diversity in an agricultural context. In this study, we compared the composition of parasitoid species and parasitism rates between rearing and DNA metabarcoding of host eggs and larvae of the millet head miner, Heliocheilus albipunctella De Joannis (Lepidoptera, Noctuidae), collected from millet fields in Senegal. We first assessed the detection threshold for the main ten endoparasitoids, by sequencing PCR products obtained from artificial dilution gradients of the parasitoid DNAs in the host moth. We then assessed the potential of DNA metabarcoding for diagnosing parasitism rates in samples collected from the field. Under controlled conditions, our results showed that relatively small quantities of parasitoid DNA (0.07 ng) were successfully detected within an eight-fold larger quantity of host DNA. Parasitoid diversity and parasitism rate estimates were always higher for DNA metabarcoding than for host rearing. Furthermore, metabarcoding detected multi-parasitism, cryptic parasitoid species and differences in parasitism rates between two different sampling sites. Metabarcoding shows promise for gaining a clearer understanding of the importance and complexity of host-parasitoid interactions in agro-ecosystems, with a view to improving pest biocontrol strategies.

Four-horned wasps, description of some remarkable Dirhinus (Hymenoptera, Chalcididae) from Kenya, with a discussion of their taxonomic placement.

Five new species belonging to Dirhinus Dalman are described: D. quadrhinus Delvare sp. nov., D. gigasetosus Delvare sp. nov., D. kambae Delvare sp. nov., D. maasaii Delvare sp. nov. and D. leakeyorum Delvare sp. nov. These species belong to the newly defined and characterized quadrhinus species group, included in the subgenus Dirhinus of Dirhinus Dalman. Evidence for this taxonomic placement is provided despite the distinctive morphology exhibited by the relevant species. The himalayanus, claviger, anthracia and hesperidum groups, also included in subgenus Dirhinus, are similarly defined and characterized and lists of the relevant described species are provided. A key to the females of the quadrhinus group is given. The phylogeny of the genus Dirhinus, especially the other subgenera Pareniaca and Hontalia is discussed as well as the morphological characters the genus Dirhinus in relation to the potential hosts, and the restricted distribution of the members of the quadrhinus group in Kenya.

The subfamily Cratocentrinae (Hymenoptera: Chalcididae): reappraisal of their morphological characters and review of the West Palaearctic species, with the description of two new species.

The Cratocentrinae (Hymenoptera: Chalcididae) of the West Palaearctic region are reviewed. A reappraisal of the morphological characters used for their recognition and phylogeny is provided as is a key to separate the genera and species of the region. Their distribution in the West Palaearctic is updated and the subfamily is reported for the first time in Europe. Cratocentrus inermus Delvare sp. nov. and Philocentrus papillus Abul-Sood Gadallah sp. nov. are described. A neotype is designated for Philocentrus argenteopilosus (Cameron), which is revalidated and transferred to Philocentrus comb. nov. stat rev.

Review of Iranian Chalcididae (Hymenoptera, Chalcidoidea) with nomenclatural notes.

The current paper presents new data on the nomenclature and distribution of Chalcididae (Hymenoptera, Chalcidoidea) collected in various regions of Iran. The following new synonymies are proposed: Brachymeria argenteopilosa (Radoszkowski, 1876) = Chalcis persica Masi, 1824 syn. nov., Brachymeria albicrus (Klug, 1834) = Chalcis rufigaster Masi, 1924 syn. nov. and Bucekia differens (Boucek, 1949) = B. Bouceki Nikol'skaya, 1960 syn. nov. The status of Lasiochalcidia [described under Euchalcis] igiliensis (Masi, 1916) is resurrected and is removed from synonymy with L. pubescens (Klug). Lectotypes are designated for Chalcis kassalensis Kirby, 1886, Halticella subelongata Kohl, 1906 and Chalcis pubescens Klug, 1834. Two genera, Bucekia Steffan, 1951 and Euchalcis Dufour, 1861, as well as eight species, Brachymeria kassalensis (Kirby, 1886), Dirhinus anthracia Walker, 1846, Antrocephalus subelongatus (Kohl, 1906), Bucekia differens (Boucek, 1949), Euchalcis unicolor (Lucas, 1871), L. rubripes (Kieffer, 1899), Psilochalcis dentata (Steffan, 1951) and P. subdola (Nikol'skaya, 1960), are new records for the Iranian insect fauna. A new host is recorded for E. unicolor. A list of 68 species in 18 genera of Chalcididae known from Iran is also included and some previous erroneous records are discussed.

Torymidae (Hymenoptera, Chalcidoidea) revised: molecular phylogeny, circumscription and reclassification of the family with discussion of its biogeography and evolution of life-history traits.

A phylogeny of the Torymidae (Chalcidoidea) is estimated using 4734 nucleotides from five genes. Twelve outgroups and 235 ingroup taxa are used, representing about 70% of the recognized genera. Our analyses do not recover Torymidae as monophyletic and we recognize instead two families: Megastigmidae (stat. rev.) and Torymidae s.s. (stat. rev.). Within Torymidae s.s., we recognize six subfamilies and six tribes, including Chalcimerinae, Glyphomerinae and Microdontomerinae (subf. nov.), and two new tribes: Boucekinini and Propalachiini (trib. nov.). Seven unclassified genera (i.e. Cryptopristus, Echthrodape, Exopristoides, Exopristus, part of Glyphomerus, Thaumatorymus, Zaglyptonotus) are assigned to tribes within our new classification. Five genera are restored from synonymy-Ameromicrus and Didactyliocerus from under Torymoides (stat. rev.), Iridophaga and Iridophagoides from under Podagrionella (stat. rev.) and Nannocerus from under Torymus (stat. rev.)-and three genera are synonymized-Allotorymus under Torymussyn. nov., Ditropinotus under Eridontomerussyn. nov. and Pseuderimerus under Erimerussyn. nov. A Palaearctic or Eurasian origin for Torymidae is proposed. The ancestral area of Megastigmidae is indicated as the Australian region. The most probable ancestral life strategy for Torymidae s.s. is ectoparasitism on gall-forming Cynipidae. The life strategy and putative hosts of the common ancestor of Megastigmidae remain uncertain.

Description of Brachymeria philornisae sp. n. (Hymenoptera: Chalcididae), a parasitoid of the bird parasite Philornis trinitensis (Diptera: Muscidae) in Tobago, with a review of the sibling species.

In this paper, we describe and illustrate a new species of parasitoid wasp as Brachymeria philornisae Delvare, sp. nov. The new species was reared from Philornis trinitensis Dodge & Aitken (Diptera: Muscidae) puparia that were found in the nests of the bird species Mimus gilvus (Vieillot) (Mimidae) and Tiaris bicolor (L.) (Thraupidae) in Tobago. The new species is of particular interest as it may be considered a potential biological control agent in locations where Philornis species are invasive, such as the Galapagos Islands. Closely related Brachymeria species had taxonomically ambiguous relationship in the past and are compared and reviewed. The species have been classified in the subgenus Pseudobrachymeria, but are here treated within the newly defined subconica species-group of Brachymeria as part of a sibling species complex designated as the subrugosa complex. Species assigned to the subconica species-group are listed and five, one unnamed, are assigned to the subrugosa complex and their females keyed. Species are separated by qualitative characters and morphometry using distance measurements. Trigonura annulipes Costa Lima is renamed as Brachymeria costalimai Delvare nom. nov. because of secondary homonymy.

Investigating Biological Control Agents for Controlling Invasive Populations of the Mealybug Pseudococcus comstocki in France.

Pseudococcus comstocki (Hemiptera: Pseudococcidae) is a mealybug species native to Eastern Asia and present as an invasive pest in northern Italy and southern France since the start of the century. It infests apple and pear trees, grapevines and some ornamental trees. Biocontrol programmes against this pest proved successful in central Asia and North America in the second half of the 20th century. In this study, we investigated possible biocontrol agents against P. comstocki, with the aim of developing a biocontrol programme in France. We carried out systematic DNA-barcoding at each step in the search for a specialist parasitoid. First we characterised the French target populations of P. comstocki. We then identified the parasitoids attacking P. comstocki in France. Finally, we searched for foreign mealybug populations identified a priori as P. comstocki and surveyed their hymenopteran parasitoids. Three mealybug species (P. comstocki, P. viburni and P. cryptus) were identified during the survey, together with at least 16 different parasitoid taxa. We selected candidate biological control agent populations for use against P. comstocki in France, from the species Allotropa burrelli (Hymenoptera: Platygastridae) and Acerophagus malinus (Hymenoptera: Encyrtidae). The coupling of molecular and morphological characterisation for both pests and natural enemies facilitated the programme development and the rejection of unsuitable or generalist parasitoids.

Availability of eleven species names of Eupelmus (Hymenoptera, Eupelmidae) proposed in Al khatib et al. (2014).

This paper is an addendum for the availability of the names of 11 new species proposed in Al khatib et al. (2014).

A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera).

Chalcidoidea (Hymenoptera) is extremely diverse with an estimated 500 000 species. We present the first phylogenetic analysis of the superfamily based on both morphological and molecular data. A web-based, systematics workbench mx was used to score 945 character states illustrated by 648 figures for 233 morphological characters for a total of 66 645 observations for 300 taxa. The matrix covers 22 chalcidoid families recognized herein and includes 268 genera within 78 of 83 subfamilies. Morphological data were analysed alone and in combination with molecular data from ribosomal 18S (2105 bp) and 28S D2-D5 expansion regions (1812 bp). Analyses were analysed alone and in combined datasets using implied-weights parsimony and likelihood. Proposed changes in higher classification resulting from the analyses include: (i) recognition of Eriaporidae, revised status; (ii) recognition of Cynipencyrtidae, revised status; (iii) recognition of Azotidae, revised status; (iv) inclusion of Sycophaginae in Agaonidae, revised status; (v) reclassification of Aphelinidae to include Aphelininae, Calesinae, Coccophaginae, Eretmocerinae and Eriaphytinae; (vi) inclusion of Cratominae and Panstenoninae within Pteromalinae (Pteromalidae), new synonymy; (vii) inclusion of Epichrysomallinae in Pteromalidae, revised status. At a higher level, Chalcidoidea was monophyletic, with Mymaridae the sister group of Rotoitidae plus the remaining Chalcidoidea. A eulophid lineage was recovered that included Aphelinidae, Azotidae, Eulophidae, Signiphoridae, Tetracampidae and Trichogrammatidae. Eucharitidae and Perilampidae were monophyletic if Eutrichosomatinae (Pteromalidae) was included, and Eupelmidae was monophyletic if Oodera (Pteromalidae: Cleonyminae) was included. Likelihood recovered a clade of Eupelmidae + (Tanaostigmatidae + (Cynipencyrtus + Encyrtidae). Support for other lineages and their impact on the classification of Chalcidoidea is discussed. Several life-history traits are mapped onto the new phylogeny.