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.

A revision of the Palaearctic species of Reikosiella (Hirticauda) (Hymenoptera, Eupelmidae).

The Palaearctic species of Reikosiella Yoshimoto, subgenus R. (Hirticauda Boucek), are revised. Illustrated keys are given to identify females of the ten recognized species and all known males. In addition to R. (Hirticauda) hungarica (Erd6s), previously the only formally recognized Palaearctic species, two species are newly transferred to the genus and subgenus, R. (Hirticauda) bolivari (Kalina) comb. nov. and R. (Hirticauda) rostrata (Ruschka) comb. nov., both from Eupelmus Dalman. Seven species are described as new: R. (Hirticauda) andriescui sp. nov. from Canary Islands, R. (Hirticauda) gordoni sp. nov. and R. (Hirticauda) graeca sp. nov. from Greece, R. (Hirticauda) vanharteni sp. nov. from United Arab Emirates, and R. (Hirticauda) cornuta sp. nov., R. (Hirticauda) koreana sp. nov., and R. (Hirticauda) tripotinorum. sp. nov. from Korea. A lectotype is designated for Eupelmus rostratus Ruschka. Host records are critically discussed for several species in the light of their new generic placement.

Leptoomidae, a new family of Eocene fossil Chalcidoidea (Hymenoptera), and family classification of Eocene fossil genera originally described in Neanastatinae (Eupelmidae).

Leptoomidae Gibson fam. nov. (Chalcidoidea) is described for the Eocene Baltic amber fossil genera Leptoomus Gibson, type genus, reassigned from Tanaostigmatidae, and Neanaperiallus Gibson, reassigned from Neanastatinae (Chalcidoidea: Eupelmidae) sensu Gibson (2009). One new species of Neanaperiallus, N. defunctus Fusu sp. nov., is described. The new family is differentiated from other families of Chalcidoidea that are partly characterized by a greatly enlarged acropleuron. In species of Leptoomidae the prepectus is anteriorly rounded to angulate and extends to or slightly over the posterolateral margin of the pronotum, with the dorsal prepectal margin intersecting the base of the tegula distinctly anterior to and forming an almost right-angle with the posterior margin of prepectus, and the posterior margin truncate along the anterior margin of the acropleuron. This prepectal structure is similar to that in Tanaostigmatidae and Cynipencyrtidae, except the prepectus is elongated anteriorly exterior to the pronotum in Tanaostigmatidae and interior to the lateral surface of the pronotum in Cynipencyrtidae. A difference in prepectal structure also indicates that an anteriorly elongated mesoscutal process internal to the pronotum in Encyrtidae is convergent to that of Cynipencyrtidae, and similarity in shape of the prepectus among Encyrtidae, Eopelma Gibson and Neanastatus Girault might be functionally correlated with an anterior elongation of the mesoscutal process. New or corrected morphological data are provided for the two included genera. Of other Eocene fossil genera originally classified in Neanastatinae, Brevivulva Gibson and Propelma Trjapitzin, are assigned to Neanastatidae sensu Burks et al. (2022) based on similar mesoscutellar structures. Possible relationships of Aspidopleura Gibson, a taxon with a puzzling combination of features, are discussed. Because Aspidopleura cannot be placed with confidence in any extinct or extant higher taxon it is treated as incertae sedis at family level within Chalcidoidea.

Ooencyrtus pitosina (Hymenoptera: Encyrtidae)-A natural enemy of Samoan swallowtail butterfly Papilio godeffroyi (Lepidoptera: Papilionidae).

A new species of encyrtid wasp, Ooencyrtus pitosina Polaszek, Noyes & Fusu sp. n., (Hymenoptera: Encyrtidae: Encyrtinae) is described as a gregarious parasitoid in the eggs of the endemic Samoan swallowtail butterfly Papilio godeffroyi (Lepidoptera: Papilionidae) in the Samoan archipelago. It is described here because it is an important natural enemy of this butterfly, and to facilitate identification for future work with this parasitoid and its host.

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.

Revision of the Oriental and East Palaearctic genus Coryptilus Gibson (Hymenoptera, Chalcidoidea, Eupelmidae).

The previously monotypic genus Coryptilus Gibson (Hymenoptera: Eupelmidae) is revised and two new species are described based on females: Coryptilus circalatus Fusu sp. nov. from the Oriental region and Coryptilus longicervix Kawano Fusu sp. nov. from both the Oriental and far eastern Palaearctic regions. Coryptilus is known only from females and previously only from the Oriental region. An illustrated key to the three known species is provided.

Optimized DNA extraction and library preparation for minute arthropods: Application to target enrichment in chalcid wasps used for biocontrol.

Target enrichment is increasingly used for genotyping of plant and animal species or to better understand the evolutionary history of important lineages through the inference of statistically robust phylogenies. Limitations to routine target enrichment are both the complexity of current protocols and low input DNA quantity. Thus, working with tiny organisms such as microarthropods can be challenging. Here, we propose easy to set up optimizations for DNA extraction and library preparation prior to target enrichment. Prepared libraries were used to capture 1,432 ultraconserved elements (UCEs) from microhymenoptera (Chalcidoidea), which are among the tiniest insects on Earth and the most commercialized worldwide for biological control purposes. Results show no correlation between input DNA quantities (1.8-250 ng, 0.4 ng with an extra whole genome amplification step) and the number of sequenced UCEs on an Illumina MiSeq. Phylogenetic inferences highlight the potential of UCEs to solve relationships within the families of chalcid wasps, which has not been achieved so far. The protocol (library preparation + target enrichment) allows processing 96 specimens in five working days, by a single person, without requiring the use of expensive robotic molecular biology platforms, which could help to generalize the use of target enrichment for minute specimens.

Rediscovery of Calymmochilus russoi Gibson, 1995 (Hymenoptera, Chalcidoidea, Eupelmidae), and revision of European Calymmochilus Masi, 1919.

The European species of Calymmochilus Masi (Hymenoptera, Eupelmidae) are revised. Calymmochilus atratus Masi stat. rev. is removed from synonymy under C. subnubilus (Walker) and treated as a valid species. A lectotype is designated for Calymmochilus atratus. The single extant type specimen of Eupelmus subnubilus Walker is considered as lectotype. Calymmochilus bini Fusu sp. n. is described from a single female collected in Sardinia. A female of Calymmochilus russoi Gibson is reported from Spain as a parasitoid in galls of Parapodia sinaica (Frauenfeld) (Lepidoptera, Gelechiidae) on Tamarix (Tamaricaceae), a new national and host record. The species is redescribed and illustrated, this being the first record of the species after its original description. An illustrated key to females and, when known, males of the now six recognized European species of Calymmochilus is given and available biological and distributional data are reviewed.

Description, DNA barcoding and phylogenetic placement of a remarkable new species of Eopelma (Hymenoptera: Eupelmidae) from Borneo.

Eopelma gibsoni sp. nov. is described based on a male recently collected in Borneo. It is the second species of the genus to be described, and the first species of chalcid wasp in which a pattern of dark stripes on the compound eye is described. The presence of similar dark stripes on the eyes of other chalcid wasps is discussed, highlighting the importance of citizen science. The description is accompanied by a DNA barcode sequence to assist future identification and association of the sexes. The phylogenetic position of E. gibsoni based on 28S DNA sequences is discussed.

Revision of the Palaearctic species of Eupelmus (Eupelmus) Dalman (Hymenoptera: Chalcidoidea: Eupelmidae).

One hundred-four extant species of Eupelmus Dalman (Hymenoptera: Eupelmidae: Eupelminae) are recognized from the Palaearctic region, of which 76 species of E. (Eupelmus) are recognized following a revision of the Palaearctic fauna of the subgenus. The following 25 species are described as new: E. (Eupelmus) adustus Gibson & Fusu n. sp., E. (Eupelmus) angustifrons Gibson & Fusu n. sp., E. (Eupelmus) bicolor Gibson & Fusu n. sp., E. (Eupelmus) brachypterus Fusu & Gibson n. sp., E. (Eupelmus) brachystylus Gibson & Fusu n. sp., E. (Eupelmus) brachyurus Fusu & Gibson n. sp., E. (Eupelmus) fasciatus Gibson & Fusu n. sp., E. (Eupelmus) gelechiphagus Gibson & Fusu n. sp., E. (Eupelmus) hayei Gibson & Fusu n. sp., E. (Eupelmus) infimbriatus Gibson & Fusu n. sp., E. (Eupelmus) iris Fusu & Gibson n. sp., E. (Eupelmus) kamijoi Gibson & Fusu n. sp., E. (Eupelmus) lanceolatus Gibson & Fusu n. sp., E. (Eupelmus) luteipes Fusu & Gibson n. sp., E. (Eupelmus) magdalenae Fusu & Gibson n. sp., E. (Eupelmus) mehrnejadi Gibson & Fusu n. sp., E. (Eupelmus) melanostylus Gibson & Fusu n. sp., E. (Eupelmus) punctatifrons Fusu & Gibson n. sp., E. (Eupelmus) setosus Fusu & Gibson n. sp., E. (Eupelmus) tanystylus Gibson & Fusu n. sp., E. (Eupelmus) tetrazostus Gibson & Fusu n. sp., E. (Eupelmus) vanharteni Fusu & Gibson n. sp., E. (Eupelmus) weilli Fusu & Gibson n. sp., E. (Eupelmus) xenium Fusu & Gibson n. sp., and E. (Eupelmus) zebra Fusu & Gibson n. sp. Of previously described species of Eupelmus, 17 are newly assigned to E. (Eupelmus), 10 to E. (Episolindelia Girault), and 8 to E. (Macroneura Walker). Formally transferred to E. (Macroneura) from Macroneura are E. (M.) algiricus (Kalina 1981), E. (M.) coleophorae (Kalina 1981), E. (M.) impennis (Nikol'skaya 1952), E. (M.) longicornis (Kalina 1981), E. (M.) pleuratus (Kalina 1981) and E. (M.) sugonyaevi (Kalina 1981) n. combs. Eupelmus (Eupelmus) kalinai Gibson & Fusu n. name is given to replace E. (Eupelmus) algiricus Kalina 1988, a secondary homonym of E. (M.) algiricus (Kalina 1981). New synonyms proposed are Eupelmus scolyti Liao 1987 n. syn. under E. (Eupelmus) formosae Ashmead 1904, and Eupelmus nigricauda Nikol'skaya 1952 n. syn. under E. (Eupelmus) microzonus Förster 1860. Eupelmus gueneei Giraud 1870 and Eupelmus xambeui Giard 1900 are transferred to Arachnophaga (Parasolindenia Brues) as A. (P.) gueneei (Giraud) and A. (P.) xambeui (Giard) n. combs., and Eupelmus kim Nikol'skaya 1952 is transferred to Brasema Cameron as B. kim (Nikol'skaya) n. comb. Eupelmus puparum Newport 1840 is transferred to Pteromalus Swederus (Pteromalidae) as P. puparum (Newport) n. comb., a secondary homonym of P. puparum (Linnaeus 1758), and Ceraphron brachynterae Schwägrichen 1835 is removed from Eupelmus and Eupelmidae, and the name treated as incertae sedis. Lectotypes are designated for Eupelmus azureus Ratzeburg 1844, Pteromalus cordairii Ratzeburg 1844, Eupelmus hostilis Förster 1860, and Eupelmus splendens Giraud 1872. Neotypes are designated for Pteromalus audouinii Ratzeburg 1844 and Eupelmus bedeguaris Ratzeburg 1852. Newly recorded from the Palaearctic are E. (Eupelmus) orthopterae (Risbec 1951) and E. (Eupelmus) peculiaris Narendran (2011). Excluded from the Palaearctic are E. (Eupelmus) afer Silvestri 1914 (Afrotropical) and E. (Eupelmus) longicorpus (Girault 1915) (Australasian), the former being compared to E. confusus Al khatib 2015 and the latter to E. iranicus Kalina 1988 and E. kalinai. Seven informal species groups are recognized for the purpose of species comparisons, the fulgens-, fulvipes-, iranicus-, orientalis-, splendens-, stramineipes-, and urozonus-groups. The latter group is restricted to E. urozonus Dalman and five other species that were differentiated initially using molecular evidence. Females of all 76 species of E. (Eupelmus) recognized from the Palaearctic are keyed, described and illustrated. Males are recognized for 44 of the species, and keyed and illustrated, though not all males of the fulvipes- and urozonus-groups are distinguished from each other.

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).

Fauna Europaea: Hymenoptera - Apocrita (excl. Ichneumonoidea).

Fauna Europaea provides a public web-service with an index of scientific names (including important synonyms) of all living European land and freshwater animals, their geographical distribution at country level (up to the Urals, excluding the Caucasus region), and some additional information. The Fauna Europaea project covers about 230,000 taxonomic names, including 130,000 accepted species and 14,000 accepted subspecies. This represents a huge effort by more than 400 contributing specialists throughout Europe and is a unique (standard) reference suitable for many users in science, government, industry, nature conservation and education. Hymenoptera is one of the four largest orders of insects, with about 130,000 described species. In the Fauna Europaea database, 'Hymenoptera - Apocrita (excluding Ichneumonoidea)' comprises 13 superfamilies, 52 families, 91 subfamilies, 38 tribes and 13,211 species. The paper includes a complete list of taxa dealt with, the number of species in each and the name of the specialist responsible for data acquisition. As a general conclusion about the European fauna of Hymenoptera, the best known countries in terms of recorded species are those from northwestern Europe, with the least known fauna probably in the more eastern and southeastern parts of Europe.