The flat wasp family Bethylidae Haliday lacks global scale literature on their alpha taxonomy. The only world revision for the family was by Kieffer in 1914 and is fully out of date and somewhat useless; the only catalog for the family was made by Gordh & Móczár in 1990 and does not include hundreds of changes made since then; and the most recent world genera keys were proposed by Terayama in 2003, but do not reflect the current knowledge we have for the family. Given this scenario, we present a global guide of Bethylidae with diagnoses, taxonomic evaluation, keys, and a checklist of all their extant genera and subfamilies. We visited the main collections around the world, analyzed about 2,000 holotypes, and examined at least 400,000 specimens. To eliminate homonymies, we add the prefix “neo” to the original specific epithet when possible. The family is now composed by 2,920 species allocated in 96 genera distributed in eight subfamilies: Bethylinae, Pristocerinae, Epyrinae, Mesitiinae, Scleroderminae, Lancepyrinae, Holopsenellinae and Protopristocerinae. The latter three are extinct. One new family-group synonym is proposed: Fushunochrysidae Hong syn. nov. of Bethylidae. Two incertae sedis genera are allocated into Bethylinae: Cretobethylellus Rasnytsyn and Omaloderus Walker. One new genus-group synonym is revalidated: Pristepyris Kieffer stat. rev. from Acrepyris Kieffer. Sixteen new genus-group synonyms are proposed: Fushunochrysites Hong syn. nov. and Sinibethylus Hong syn. nov. of Eupsenella Westwood; Messoria Meunier syn. nov. of Goniozus Förster; Acrepyris Kieffer syn. nov. of Pristepyris Kieffer; Apristocera Kieffer syn. nov. and Parapristocera Brues syn. nov. of Pristocera Klug; Usakosia Kieffer syn. nov. of Prosapenesia Kieffer; Isobrachium Förster syn. nov., Leptepyris Kieffer syn. nov., Neodisepyris Kurian syn. nov., Rhabdepyris Kieffer syn. nov. of Epyris Westwood; Codorcas Nagy syn. nov., Hamusmus Argaman syn. nov. and Ukayakos Argaman syn. nov. of Heterocoelia Dahlbom; Domonkos Argaman syn. nov. of Incertosulcus Móczár; Ateleopterus Förster syn. nov. of Sclerodermus Latreille. One new genus-group synonym is revalidated: Topcobius Nagy syn. rev. of Sulcomesitius Móczár. One new genus-group revalidation is proposed: Incertosulcus Móczár stat. rev. from Anaylax Móczár. The following species-group nomenclatural acts are established: 153 new or revalidated combinations, 16 new names to avoid secondary homonyms, 11 species with revalidated status, and one synonym. Keys to the subfamilies and genera are provided. The text is supported by 599 illustrations organized onto 92 plates.

Epyrini (Epyrinae) are the largest and morphologically most diverse group within Bethylidae. Evans' conception of Epyrini induces misclassification of several taxa and their diagnostic features, such as median lobe of clypeus projected, and fovea in lateroposterior angle of propodeal disc, are insufficient to place species in the tribe. No previous comprehensive cladistic study has recovered Epyrini as a clade, and no synapomorphies are reported for the tribe, suggesting that the group is artificial. We infer a phylogeny based on 391 informative morphological characters for 42 terminal taxa (31 ingroup, 11 outgroup). Monophyly of Epyrini is supported by a petiole with fused root and body as an autapomorphy. Anisepyris Kieffer, Laelius Ashmead, Holepyris Kieffer, Formosiepyris Terayama, Disepyris Kieffer, Bakeriella Kieffer and Trachepyris Kieffer were retrieved as monophyletic. Epyris Westwood and Chlorepyris Kieffer are paraphyletic. The addition of new characters was essential to define Epyrini, and the ventral region of the mesosoma, petiole and male genitalia may help to better understand genera and species limits. All resulting trees indicate that Epyrinae are polyphyletic. Tribes Epyrini and Sclerodermini are revalidated to subfamily status.

The flat wasp Apenesia Westwood is one of the largest genera in Pristocerinae with 190 species worldwide. The lack of a single diagnostic morphological character has resulted in many distinct ground plans within Apenesia. Some authors have suggested Apenesia as polyphyletic. Here we aimed to investigate and delimit the genus based on morphological and molecular data. We analysed 163 morphological characters on tnt using ‘traditional search’, whereas Bayesian (BI) and maximum likelihood (ML) inferences were performed on 1553 concatenated nucleotide base pairs from COI and 28S sequences. Apenesia was recovered as polyphyletic in all analyses with up to 10 different lineages spread throughout the tree. We mapped female morphological characters onto the molecular tree to enlighten morphological evolution in the apterous females. Based on our morphological and molecular analyses, and on the taxonomic revision of holotypes and specimens, we propose six new genera in the Pristocerinae and revalidate the generic status of two: Acrenesia gen.n., Austranesia gen.n., Cleistepyris stat. rev., Dracunesia gen.n., Eleganesia gen.n., Epynesia gen.n., Pristonesia gen.n., and Propristocera stat. rev. We also propose four new generic synonymies: Neoapenesia syn.n. under Apenesia, Dipristocera syn.n. under Cleistepyris, Afrocera syn.n. and Neopristocera syn.n. under Propristocera. At the species level, we indicate Apenesia minor syn.n. under Propristocera tagala. In all, 162 new combinations, eight reinstated combinations and one new name to avoid homonymy are proposed. Apenesia is now defined as flat wasps having males with the mesoscutum gibbous, the genitalia with the paramere narrow and densely hairy, and the aedeagus with the ventral apical lobe elliptical and covered with lumps. Females of Apenesia can be distinguished from other Pristocerinae by having the head wider than the mesosoma, the antennae short, the mandible long, and the clypeus surpassing the toruli in the frons. This study reinforces the difficulties in defining Apenesia and other Pristocerinae genera. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:65FB3087-0F30-4851-B1BA-ED6E4518B958.

Melanepyris Kieffer, 1913 was proposed to accommodate a single species of Epyris Westwood, 1832, E. imicola Kieffer, 1913, mainly based on the absence of the posterior propodeal carina. Today, Melanepyris includes only two nominal species. The type-material of these species has been considered lost since their original description. In this study, the single known adult male (holotype) of Melanepyris asiaticus Kieffer, 1922 from the Philippines has been rediscovered, redescribed and illustrated. Melanepyris asiaticus is transferred to Epyris Westwood due to the following features: scutellar groove absent, well separated scutellar pits and lower mesopleural fovea large and with undefined upper margin. We checked the original description of M. imicola and concluded that it also fits the definition of Epyris perfectly, except for the (described) absence of a posterior carina. However, the thickness of the posterior carina of the propodeal disc varies within species of different Epyrinae genera. The diagnostic characters used by Kieffer to create Melanepyris and other genera from Epyris are briefly discussed. Melanepyris is proposed as a new junior synonym of Epyris, with the transfer of M. imicola to Epyris

ABSTRACT. It is difficult to make reliable sex associations in several species of Hymenoptera due to sexual dimorphism. Only a few species of the flat wasp genus Dissomphalus Ashmead, 1893 have had their sexes associated, since females are rarely collected and differ morphologically from their conspecific males. Collecting couples during their phoretic copula is difficult, but it is the most reliable way to associate the sexes in Dissomphalus. Herein, we associate the sexes and discuss the mating briefly discuss the genital structures used during phoretic copulation, analyzing the male and female genitalia in copulation.During copulation, the mating female is attached to the male only by the genitalia and is apparently grasped by muscles attached to the male's metasoma.

-The generic richness and abundance of Bethylidae collected in four different hillside areas of Atlantic rain forest from Espírito Santo, Brazil were studied. The sites are Santa Maria de Jetibá (SMJ), Domingos Martins (DM), Pancas (P) and Atílio Vivacqua (AV). A total of 2,840 specimens of 12 genera were collected. Lepidosternopsis Ogloblin and Bakeriella Kieffer are fi rst recorded from the State. Richness of taxa was calculated using fi rst-order Jackknife richness with EstimateS program. Genera accumulation curves were ran to evaluate the samples. Abundance data were adjusted to the geometric distribution. Parameter k was used to compare areas. The generic profi le was not equal for the sites we studied. The areas were considered disturbed. SMJ and DM presented genera richness bigger than in P and AV. The differences in the sites refl ect the different preservation of each environment. Pseudisobrachium Kieffer and Dissomphalus Ashmead are most dominant genera in SMJ, DM and P, and Anisepyris Kieffer in AV. This study emphasizes the fact of Dissomphalus as the most abundant genus in rain forests. The generic profi le found in AV is similar to that of some areas of Brazilian savannah.KEY WORDS: Alpha diversity, fi rst-order Jackknife, Neotropical region, new record, parasitoid wasp RESUMO -Estudou-se a riqueza e abundância de gêneros de Bethylidae coletados em quatro áreas de mata de encosta da Mata Atlântica do Espírito Santo, com estados de preservação diferentes: Santa Maria de Jetibá (SMJ), Domingos Martins (DM), Pancas (P) e Atílio Vivácqua (AV). Foram coletados 2.840 espécimes alocados em 12 gêneros, sendo Lepidosternopsis Ogloblin e Bakeriella Kieffer citados pela primeira vez para esse estado. A riqueza dos táxons foi obtida através do procedimento Jackknife com auxílio do programa EstimateS. Curvas de acumulação de gêneros foram construídas para avaliar o esforço amostral. Os dados se ajustaram à distribuição geométrica e calculou-se o parâmetro k para comparar as localidades. O perfi l genérico não foi equivalente em todas as localidades, e todas foram consideradas perturbadas. SMJ e DM apresentaram riqueza de gêneros maior em comparação com P e AV. As diferenças relatadas neste estudo para as áreas amostradas refl etem o grau diferente de preservação das matas. Pseudisobrachium Kieffer e Dissomphalus Ashmead foram os gêneros mais abundantes em SMJ, DM e P e Anisepyris Kieffer em AV. Este estudo reforça o fato de Dissomphalus ser mais abundante em fl orestas tropicais úmidas e que o perfi l genérico encontrado em AV assemelhase a dados publicados para o cerrado. PALAVRAS-CHAVE: Diversidade alfa, Jackknife de primeira ordem, Região Neotropical, citação nova, vespa parasitóide Bethylidae são vespas ectófagas quando adultas e ectoparasitóides idiobiontes primárias quando imaturas. Seus hospedeiros são larvas de Lepidoptera e Coleoptera que vivem em situações crípticas, tais como solo, serrapilheira, madeira e sementes (Evans 1964). Corpo achatado, cabeça prognata, fêmures anteriores largos e au...

Abstract. The genus Neurepyris Kieffer, 1905 is revised. The adult male holotypes N. rufiventer Kieffer, 1913 from Eritrea and N. tagala (Ashmead, 1905) from the Philippines are redescribed and illustrated. Both species are transferred from the subfamily Epyrinae to Pristocerinae because they have the metanotum well developed medially. Neurepyris rufiventer is transferred to Pristocera Klug, 1808 because the hypopygium is deeply divided into two apical lobes, the subdiscoidal and cubital veins do not reach the border of the forewing, the pronotal disc has the anterior region slightly elevate medially, and the stigma is elongate. Neurepyris tagala is transferred to Apenesia Westwood, 1874 because the basal tooth of mandible is not curved inward, the median lobe of clypeus is not depressed near the antennal insertions, and the aedeagus consists of one lamina. The genus Neurepyris is considered a junior synonym of Pristocera because its type species is N. rufiventer. Key words. Epyrinae, Pristocerinae, Apenesia, Afrotropical region, Oriental region. Kieffer (1905) created Neurepyris Kieffer, 1905 to accommodate a species of Epyrini, which has a longitudinal carina on the propodeal disc absent, unidentate tarsal claw and lanceolate stigma. Kieffer & Marshall's key (1904-1906 indicated that there was one species from Eritrea known from one female, but they did not described in that time. This species was formally described years later as N. rufiventer Kieffer, 1913by Kieffer (1913. Kieffer (1914a) transferred a doubtful Philippine species Rhabdepyris tagala (Ashmead, 1905) to Neurepyris. Both species are known only from their types. However our initial analyses indicate that both species are males of Pristocerinae because both of them have the metanotal well developed medially, propodeum with convex sides, metasoma depressed and genitalia with complex aedeagus. IntroductionIn this study, we aimed to reconsider the genus, presenting a new view about both species here redescribed and illustrated.

The Epyrinae genus Pristepyris Kieffer is revised. The genus has four species. Pristepyris rugicollis Kieffer from Malaysia and P. levicollis Kieffer from Madagascar are rediscovered. Their adult male holotypes are redescribed and illustrated. Specimens of Pristepyris agraensis Kurian and P. flavicornis Kieffer were unavailable and reassigned based on their original descriptions. All species are considered to be Pristocerinae, because they have the metanotum well developed medially. Pristepyris rugicollis is transferred to Acrepyris Westwood because it has the aedeagus divided in three valvae, P. agraensis is transferred to Acrepyris, because it has an undivided hypopygium; and P. levicollis and P. flavicornis are transferred to Pristocera Klug, because they have a hypopygium that is deeply divided into two lobes. Pristepyris is considered junior synonym of Acrepyris, because Pristepyris rugicollis is its type-species.