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.

The morphological diversity of Mymaridae (Hymenoptera): an atlas of scanning electron micrographs. Part 1. General overview and structure of the head.

This is the first in a series of studies that aim to provide a comprehensive overview of the morphological diversity of Mymaridae (Hymenoptera), a monophyletic family of small parasitic wasps that are postulated as the sister group of other Chalcidoidea. The external cranial morphology of 65-75 genera and subgenera of Mymaridae (fairyflies) is described and illustrated with almost 430 scanning electron micrographs, including 73 micrographs of the anterior, 68 of the posterior, 75 of the dorsal, 75 of the lateral, and 67 of the ventral views of the head, plus 71 micrographs of the ventral view of the mouthparts. Twenty-one annotated figures illustrate the terms used for morphological structures. Two appendices list the 64 morphological terms and 5 measurements that are defined and illustrated, and the 116 currently recognized valid genera and subgenera of Mymaridae, including collection localities for those that are illustrated. Discussion of head morphology characteristic of Mymaridae is preceded by an overview that includes discussion of best practices for taxonomic descriptions and why these and accurate identifications require well preserved and imaged specimens. Aspects of intraspecific variation, colour, secondary sexual dimorphism, setation (chaetotaxy), surface sculpture and morphometrics are also treated as all of these are often important for describing and distinguishing species. Many of the features illustrated have not previously been used in Mymaridae systematics but may prove to be useful for helping to identify and describe genera and species.

The morphological diversity of Mymaridae (Hymenoptera): an atlas of scanning electron micrographs. Part 2. Structure of the mesosoma.

This is the second in a series of studies that aim to provide a comprehensive overview of the morphological diversity of Mymaridae (Hymenoptera) or fairyflies, a monophyletic family of small parasitic wasps postulated to be the sister group of all other Chalcidoidea. The external morphology of the mesosoma of about 6575 taxa, representing 5565% of the 115 currently valid described genera and subgenera, is described and illustrated with almost 269 scanning electron micrographs, including 77 micrographs of the dorsal, 71 micrographs of the lateral, 59 micrographs of the ventral, 53 micrographs of the anterior, and 9 micrographs of the posterior views of the mesosoma. Twenty annotated figures of the external and major internal structures are given. Two appendices list the morphological terms used, and names of the 75 genera and subgenera of Mymaridae illustrated. The variety of characters and their features that could be used to help define morphologically the genera, and possibly also the species, of Mymaridae is discussed.

A new, remarkable species of Platystethynium (Platypatasson) (Hymenoptera: Mymaridae) from New Zealand.

Platystethynium (Platypatasson) earlyi Huber, sp. n. (Hymenoptera: Mymaridae), is described from both sexes reared from an egg of Rhaphidophoridae (Orthoptera) found in an old mining tunnel in New Zealand. The male is micropterous and has lateral ocelli but no median ocellus, a unique feature in Mymaridae found so far only in males of Platystethynium Ogloblin species. The remaining described Eastern Hemisphere species of Platystethynium are discussed and Platystethynium glabrum Jin Li, syn. n., is placed in synonymy under P. onomarchicidum Ogloblin. A key to females, and males where known, of the described Eastern Hemisphere species of Platystethynium is given.

Illustrated key to the genera and catalogue of Mymaridae (Hymenoptera) in the Afrotropical region.

Separate identification keys for females, and for males where known, of the 40 genera of Mymaridae (Hymenoptera: Chalcidoidea) in the Afrotropical region are given. The subgenera of four genera are also included in the key to females. The genera are illustrated with over 300 photographs. The 122 named, valid species reported from the region are catalogued. Reliable host records are reported for 6 genera and 11 species from rearings undertaken in the region. An appendix lists the 27 species from northern Africa and offshore islands (Canary Islands, Madeira) but not recorded in the Afrotropical region as defined in this publication. Camptoptera (Zemicamptoptera) Ogloblin Annecke, syn. n. is synonymized with Camptoptera (Camptoptera) Foerster. Eofoersteria Mathot, syn. n. is synonymized with Camptoptera Foerster and is treated as subgenus C. (Eofoersteria) Mathot, stat. n.; its type species Eofoersteria camptopteroides Mathot is transferred to Camptoptera as C. (Eofoersteria) camptopteroides (Mathot), comb. n. Polynema megacephala (Risbec) is transferred to Lymaenon as L. megacephala (Risbec), comb. n. Limacis opuntiae Risbec is transferred to Encarsia Foerster (Hymenoptera: Aphelinidae) as E. opuntiae (Risbec), comb. n. A few corrections to Huber et al. (2020) are given.

Two new species of Camptopteroides (Camptopteroides) (Hymenoptera: Mymaridae) from the Oriental region with a key to Old World species.

An overview of the Oriental species of the nominate subgenus of Camptopteroides Viggiani (Hymenoptera: Mymaridae) is given. Two new species, C. (Camptopteroides) formosa Manickavasagam Sankararaman sp. n. from India and C. (Camptopteroides) reducta Triapitsyn sp. n. from Thailand and Malaysia, are described, and C. formosa additionally compared to two unnamed species. The holotype male of the type species of this genus, C. armata Viggiani from Sri Lanka, is diagnosed and illustrated. A key to Old World species of Camptopteroides is provided.

Illustrated key to genera and catalogue of Mymaridae (Hymenoptera) in America north of Mexico.

Identification keys for females, and for males where known, to the 39 genera and 11 subgenera of Mymaridae in America north of Mexico are given. The genera are illustrated with over 1010 photographs and/or scanning electron micrographs. The 202 currently named and valid species reported from America north of Mexico are catalogued. Reliable host records are reported for 16 genera and almost 90 species from rearings in the region. The five genera represented by more than one subgenus in the region are: Anagrus Haliday, Anaphes Haliday, Erythmelus Enock, Kalopolynema Ogloblin, and Polynema Haliday. Five other genera are represented only by unnamed and possibly undescribed species. About 1000 literature references, mainly the North American ones, are listed. Three new species are described: Callodicopus floridanus Huber, sp. n., Neostethynium americanum Huber, sp. n., and Stephanocampta xanthogaster Huber, sp. n. Cosmocomoidea marilandica (Girault), stat. rev., is placed in synonymy under C. dolichocerus (Ashmead). Platypatasson Ogloblin is given subgeneric status as Platystethynium (Platypatasson Ogloblin), stat. rev. Corrections and additions to the mymarid fauna of Mexico by Guzmán-Larralde et al. (2017) are also given.

Illustrated key to European genera, subgenera and species groups of Mymaridae (Hymenoptera), with new records for the Czech Republic.

Illustrated keys to females and males of the European genera and subgenera of Mymaridae (Chalcidoidea) are presented, including twenty-four genera plus two subgenera in both of Erythmelus Enock and Polynema Haliday. The key to females also includes three subgenera recognized in Anagrus Haliday, two subgenera in Anaphes Haliday, and two species groups in Anagrus s.s.; males of these either cannot be distinguished or are difficult to key. The species of Mymaridae recorded from the Czech Republic are listed and distribution records are provided based on material in three collections. Seven genera (Anagrus, Caraphractus Walker, Dicopus Enock, Macrocamptoptera Girault, Omyomymar Schauff, Pseudocleruchus Donev Huber, Stethynium Enock) and 12 species are newly recorded for the country.

Description of the male of Tanyxiphium harriet (Hymenoptera: Mymaridae), with new distribution records and synonymy.

Tanyxiphium longissimum Huber, syn. n. (Hymenoptera: Mymaridae) is synonymized under T. harriet (Zeya) based on examination of specimens from Hainan Island and Yunnan Province in the Oriental part of China, Sulawesi Island in Indonesia, Peninsular Malaysia, and Thailand. These are new country distribution records except for Thailand. The previously unknown male of T. harriet is described from Sulawesi Island, and the female is redescribed and illustrated based on non-type material examined.

Two new species of Anaphes Haliday (Hymenoptera: Mymaridae) from India and Indonesia.

Two species of Anaphes Haliday with distinctive females are described from India (A. triapitsyni Anwar Zeya, sp. nov.) or from India and Indonesia (A. kailashchandrai Anwar Zeya, sp. nov.). The only other Anaphes species known to occur in India is A. quinquearticulatus Huber Triapitsyn because previous records of A. pallipes (Ashmead) and A. pullicrurus (Girault) are based on misidentifications. These latter two names are therefore removed from the faunal list of India. An illustrated key to females of the three described species of Anaphes known from India is given. The described species of Anaphes known from Australia are also briefly reviewed, with type material of four previously unillustrated species being newly imaged and compared to the Indian species.

Review of Anaphes Haliday (Hymenoptera: Mymaridae) with key to species in Europe and a world catalogue.

The Walter Soyka species of Anaphes Haliday are reviewed and placed in context among the remaining world species. An analysis of his descriptions and identification keys is presented, and discussed in light of a broader interpretation of intraspecific variation in Anaphes species than Soyka accepted. An illustrated diagnosis of Anaphes is given and the species in Europe considered to be valid are keyed-14 species in Anaphes (Anaphes) and 15 species in Anaphes (Patasson). The 244 names of world Anaphes are tabulated in various ways and their type localities, if known, are mapped. All available valid names and their synonyms, mostly from among the 155 nominal species described by Soyka, are catalogued, with details of type material and collecting information. A total of 167 synonyms are listed, 17 of which are synonyms proposed by earlier authors and 149 of which are proposed as new synonyms. Lectotypes are designated for Mymar ferreirei Soyka and Yungaburra acutiventris Soyka. Nine nomina dubia, 2 names of fossil species that most likely do not belong to Anaphes, 3 unavailable names and 20 species transferred to other genera are listed separately; among the latter, Anagrus foersteri (Ratzeburg) is transferred to Erythmelus Enock as E. foersteri (Ratzeburg), comb. n. Host records published from 1985-2015 are tabulated; most are from the literature but a few are from unpublished records found on slide mounted Anaphes specimens in various collections, mostly in Europe. The merits of classifying the available names of nominal species into an earlier classification with subgenera Anaphes (Anaphes) and Anaphes (Patasson) instead a later classification consisting of species groups within Anaphes (Anaphes) and Anaphes (Yungaburra) are discussed; the earlier classification is preferred.

Introduction to the Mymaridae (Hymenoptera) of Bangladesh.

An identification key to the 15 genera of Mymaridae found so far in Bangladesh is given, based on about 520 specimens collected using yellow pan traps placed in agricultural habitats and at the edge of ponds, mainly at Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur. Species already reported from Bangladesh are listed and three more are added: Acmopolynema orientale (Narayanan, Subba Rao & Kaur), Himopolynema hishimonus Taguchi, and Mymar pulchellum Curtis.

Generic key and catalogue of Mymaridae (Hymenoptera) of Mexico.

The Mexican genera of Mymaridae (Hymenoptera: Chalcidoidea) are keyed in English and Spanish, and a catalogue of species occurring in Mexico is presented. Thirty-six genera, including 79 named species in 20 of the genera, are reported. These are mentioned in about 100 publications either as original species descriptions or as publications that specifically mention species and/or specimens from Mexico. In the catalogue, species distributions by state are given based on literature records, and collection data are compiled from about 3630 specimens examined in eight collections in Canada, Mexico and USA. Host are listed for specimens reared mainly in Mexico. A few extralimital host records are also given.

Two genera of Mymaridae (Hymenoptera) new to Africa, a remarkable new species of Anaphes and new generic synonymy.

Bakkendorfia Mathot, syn. n. is placed in synonymy under Cleruchus Enock and its only described species transferred as Cleruchus musangae (Mathot), comb. n.Anaphes quinquearticulatus Huber & Triapitsyn, sp. n. is described; it is the only known Anaphes Haliday species with a 5-segmented funicle in females. Two genera are reported for the first time from the Afrotropical region and two species are described: Paranaphoidea (Idiocentrus) africana Huber & Triapitsyn, sp. n., and Allanagrus occidentalis Huber & Triapitsyn, sp. n.

Nomenclatural changes in Mymaridae (Hymenoptera: Chalcidoidea).

Three New World species are transferred from Polynema Haliday to other genera of Mymaridae and redescribed based on examination of their type specimens: Polynema albicoxa Ashmead to Acmopolynema Ogloblin as Acmopolynema albicoxa (Ashmead), comb. n., Polynema porteri (Brèthes) to Agalmopolynema Ogloblin as Agalmopolynema porteri (Brèthes), comb. n., and Polynema giraulti Perkins, for which a lectotype is designated, to Stephanodes Enock as Stephanodes giraulti (Perkins), comb. n. Agalmopolynema shajovskoii Fidalgo, syn. n. is synonymized under Agalmopolynema porteri; Polynema (Dorypolynema) gaucho Triapitsyn & Aquino, syn. n. and P. grenadensis Ashmead, syn. n., are synonymized under Polynema (Dorypolynema) magniceps Ashmead, for which taxonomic notes are given. Caenomymar Yoshimoto, syn. n. is synonymized under Omyomymar Schauff, and its only species Caenomymar howdeni Yoshimoto is transferred to Omyomymar as Omyomymar howdeni (Yoshimoto), comb. n. Gonatocerus chula Triapitsyn & Bernal is transferred to Cosmocomoidea Howard as Cosmocomoidea chula (Triapitsyn & Bernal), comb. n. Erythmelus mirus Girault is transferred to Cleruchoides Lin & Huber as Cleruchoides mirus (Girault), comb. n. and compared with Cleruchoides noackae Lin & Huber, a species introduced from Australia into South America as a biological control agent of Thaumastocoris peregrinus Carpintero & Dellapé (Hemiptera: Thaumastocoridae), an introduced pest of Eucalyptus. Four spelling corrections are made to species names in Cosmocomoidea and Lymaenon Walker.

Review of the Mexican species of Erythmelus (Hymenoptera: Mymaridae), with description of two new species.

The Mexican species of Erythmelus Enock (Hymenoptera: Mymaridae) are revised. A key to females of 13 species is provided in both English and Spanish. Two new taxa are described--E. maya Guzmán-Larralde & Triapitsyn, sp. n. and E. tigres Guzmán-Larralde & Triapitsyn, sp. n. Six species are newly recorded from Mexico--E. angustatus Ogloblin, E. cingulatus Ogloblin, E. clavatus Ogloblin, E. gracilis (Howard), E. nanus Dozier, and E. noeli (Dozier), besides new geographic records for E. miridiphagus Dozier, E. picinus (Girault), E. psallidis Gahan, and E. rex (Girault) which were previously known from the country.

World reclassification of the Gonatocerus group of genera (Hymenoptera: Mymaridae).

The 400+ nominal species of the worldwide genus Gonatocerus Nees are reclassified into 14 genera that are placed in Gonatocerini, which is defined by three putative autapomorphies. A key to the 13 extant genera of Gonatocerini is given, based on females. Five previously described genus-group taxa are recognized: Cosmocomoidea Howard stat. rev. (= ater group, of authors), Gahanopsis Ogloblin stat. rev. (= deficiens group, of authors), Gastrogonatocerus Ogloblin stat. n. (= membraciphagus group, of authors), Gonatocerus (= sulphuripes group, of authors), and Lymaenon Walker stat. rev. (= litoralis group, of authors). One new fossil genus, Archigonatocerus Huber gen. n., with two fossil species, A. balticus Huber sp. n., and A. longivena Huber sp. n. and one fossil species in Gonatocerus, G. janzeni Huber sp. n., are described, all from Baltic amber from the Eocene epoch. Eight new extant genera and 16 new extant species are described and their species keyed: Cosmocomopsis Huber gen. n., with C. flopsis Huber sp. n. and C. mopsis Huber sp. n.; Heptagonatocerus Huber gen. n., with H. madagascarensis Huber sp. n., H. magnificus Huber sp. n., H. parvus Huber sp. n., and H. pulchellus Huber sp. n.; Krateriske Huber gen. n., with K. ecuadorensis Huber sp. n., K. guianensis Huber sp. n., and K. peruensis Huber sp. n.; Octomicromeris Huber gen. n., with O. compacta Huber sp. n. and O. brevis Huber sp. n.; Pro-gonatocerus Huber gen. n., with P. albiclava Huber sp. n. and P. brunneiclava Huber sp. n; Tanyxiphium Huber gen. n., with T. breviovipositor Huber sp. n., T. longissimum Huber sp. n., and T. seychellense Huber sp. n. Yoshimotoana Huber gen. n. (= masneri group, of authors) with one included species and Zeyanus Huber, gen. n. (= asulcifrons group, of authors) with 9 included species. Keys to the species of seven genera: Archigonatocerus, Cosmocomopsis, Heptagonatocerus, Krateriske, Octomicromeris, Progonatocerus, and Tanyxiphium are provided. Information for each nominal species catalogued includes the original reference, kind, sex and depository of primary type, and subsequent references that include relevant previous generic combinations, if applicable. The type locality is given, based on original descriptions or, where necessary, subsequent publications that provide clarification on the collection locality. Two new synonyms are proposed: Gonatocerus similis Gupta & Poorani, 2008, syn. n. under G. bialbifuniculatus Subba Rao, 1989; and Gonatocerus hispaniolus Triapitsyn & Huber, 2010, syn. n. under G. masneri Yoshimoto, 1990. Among the species, 245 new combinations are proposed: 82 in Cosmocomoidea, 1 in Cosmocomopsis, 4 in Gahanopsis, 8 in Gastrogonatocerus, 3 in Gonatocerus, 135 in Lymaenon, 2 in Tanyxiphium, 1 in Yoshimotoana, and 9 in Zeyanus. Revived combinations are proposed for Twelve species: 1 in Cosmocomoidea, 1 in Gahanopsis, 2 in Gonatocerus, and 8 in Lymaenon. The 410 nominal species group names are catalogued under their currently accepted genus and also listed alphabetically in an appendix. A tentative generic phylogeny is proposed.

Redescription of Chrysoctonus and description of Chrysoctonoides (Hymenoptera, Mymaridae), a new genus from the Australian Region.

Chrysoctonoideslongisetosa Huber & Triapitsyn (Hymenoptera: Mymaridae), gen. n. and sp. n., is described from Australia. It is compared with the related genus Chrysoctonus, known from Africa and the New World. Myrmecomymar Yoshimoto, syn. n., is synonymized under Chrysoctonus Mathot and its type species is transferred to Chrysoctonus as Chrysoctonusmasneri (Yoshimoto), comb. n.

Redescription of Mymarilla Westwood, new synonymies under Cremnomymar Ogloblin (Hymenoptera, Mymaridae) and discussion of unusual wings.

The monotypic genus Mymarilla Westwood is known only from St. Helena, a remote island in the South Atlantic Ocean. The peculiar species M. wollastoni Westwood (Mymaridae) is redescribed and illustrated from non-type material. Mymarilla is compared with Cremnomymar Ogloblinspp. from the Juan Fernández Islands in the South Pacific Ocean. Stephanodes Enock is shown to be the most likely sister genus to Mymarilla. Nesopolynema Ogloblin, syn. n., Oncomymar Ogloblin, syn. n., Scolopsopteron Ogloblin, syn. n., are placed in synonymy under Cremnomymar and their species transferred as Cremnomymar caudatum (Ogloblin 1952), comb. n., C. dipteron (Ogloblin 1957), comb. n., and C. kuscheli (Ogloblin 1952), comb. n. Wing shape and wing reductions in Mymaridae are discussed in relation to biogeography, particularly with respect island faunas and to four genera, Cremnomymar, Mymarilla, Parapolynema Fidalgo, and Richteria Girault, some or all of whose species have more or less convex fore wings.

DNA barcoding reveals diversity of Hymenoptera and the dominance of parasitoids in a sub-arctic environment.

BACKGROUND: Insect diversity typically declines with increasing latitude, but previous studies have shown conflicting latitude-richness gradients for some hymenopteran parasitoids. However, historical estimates of insect diversity and species richness can be difficult to confirm or compare, because they may be based upon dissimilar methods. As a proxy for species identification, we used DNA barcoding to identify molecular operational taxonomic units (MOTUs) for 7870 Hymenoptera specimens collected near Churchill, Manitoba, from 2004 through 2010. RESULTS: We resolved 1630 MOTUs for this collection, of which 75% (1228) were ichneumonoids (Ichneumonidae + Braconidae) and 91% (1484) were parasitoids. We estimate the total number of Hymenoptera MOTUs in this region at 2624-2840. CONCLUSIONS: The diversity of parasitoids in this sub-Arctic environment implies a high diversity of potential host species throughout the same range. We discuss these results in the contexts of resolving interspecific interactions that may include cryptic species, and developing reproducible methods to estimate and compare species richness across sites and between surveys, especially when morphological specialists are not available to identify every specimen.