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Annotated and illustrated world checklist of Microgastrinae parasitoid wasps (Hymenoptera, Braconidae)
expand article infoJose Fernandez-Triana, Mark R. Shaw§, Caroline Boudreault, Melanie Beaudin|, Gavin R. Broad
‡ Canadian National Collection of Insects, Ottawa, Canada
§ National Museums of Scotland, Edinburgh, United Kingdom
| Carleton University, Ottawa, Canada
¶ Natural History Museum, London, United Kingdom
Open Access

Abstract

A checklist of world species of Microgastrinae parasitoid wasps (Hymenoptera: Braconidae) is provided. A total of 81 genera and 2,999 extant species are recognized as valid, including 36 nominal species that are currently considered as species inquirendae. Two genera are synonymized under Apanteles. Nine lectotypes are designated. A total of 318 new combinations, three new replacement names, three species name amendments, and seven species status revised are proposed. Additionally, three species names are treated as nomina dubia, and 52 species names are considered as unavailable names (including 14 as nomina nuda). A total of three extinct genera and 12 extinct species are also listed. Unlike in many previous treatments of the subfamily, tribal concepts are judged to be inadequate, so genera are listed alphabetically. Brief diagnoses of all Microgastrinae genera, as understood in this paper, are presented. Illustrations of all extant genera (at least one species per genus, usually more) are included to showcase morphological diversity. Primary types of Microgastrinae are deposited in 108 institutions worldwide, although 76% are concentrated in 17 collections. Localities of primary types, in 138 countries, are reported. Recorded species distributions are listed by biogeographical region and by country. Microgastrine wasps are recorded from all continents except Antarctica; specimens can be found in all major terrestrial ecosystems, from 82°N to 55°S, and from sea level up to at least 4,500 m a.s.l. The Oriental (46) and Neotropical (43) regions have the largest number of genera recorded, whereas the Palaearctic region (28) is the least diverse. Currently, the highest species richness is in the Palearctic region (827), due to more historical study there, followed by the Neotropical (768) and Oriental (752) regions, which are expected to be the most species rich. Based on ratios of Lepidoptera and Microgastrinae species from several areas, the actual world diversity of Microgastrinae is expected to be between 30,000–50,000 species; although these ratios were mostly based on data from temperate areas and thus must be treated with caution, the single tropical area included had a similar ratio to the temperate ones. Almost 45,000 specimens of Microgastrinae from 67 different genera (83% of microgastrine genera) have complete or partial DNA barcode sequences deposited in the Barcode of Life Data System; the DNA barcodes represent 3,545 putative species or Barcode Index Numbers (BINs), as estimated from the molecular data. Information on the number of sequences and BINs per genus are detailed in the checklist. Microgastrinae hosts are here considered to be restricted to Eulepidoptera, i.e., most of the Lepidoptera except for the four most basal superfamilies (Micropterigoidea, Eriocranioidea, Hepialoidea and Nepticuloidea), with all previous literature records of other insect orders and those primitive Lepidoptera lineages being considered incorrect. The following nomenclatural acts are proposed: 1) Two genera are synonymyzed under Apanteles: Cecidobracon Kieffer & Jörgensen, 1910, new synonym and Holcapanteles Cameron, 1905, new synonym; 2) Nine lectotype designations are made for Alphomelon disputabile (Ashmead, 1900), Alphomelon nigriceps (Ashmead, 1900), Cotesia salebrosa (Marshall, 1885), Diolcogaster xanthaspis (Ashmead, 1900), Dolichogenidea ononidis (Marshall, 1889), Glyptapanteles acraeae (Wilkinson, 1932), Glyptapanteles guyanensis (Cameron, 1911), Glyptapanteles militaris (Walsh, 1861), and Pseudapanteles annulicornis Ashmead, 1900; 3) Three new replacement names are a) Diolcogaster aurangabadensis Fernandez-Triana, replacing Diolcogaster indicus (Rao & Chalikwar, 1970) [nec Diolcogaster indicus (Wilkinson, 1927)], b) Dolichogenidea incystatae Fernandez-Triana, replacing Dolichogenidea lobesia Liu & Chen, 2019 [nec Dolichogenidea lobesia Fagan-Jeffries & Austin, 2019], and c) Microplitis vitobiasi Fernandez-Triana, replacing Microplitis variicolor Tobias, 1964 [nec Microplitis varicolor Viereck, 1917]; 4) Three names amended are Apanteles irenecarrilloae Fernandez-Triana, 2014, Cotesia ayerzai (Brèthes, 1920), and Cotesia riverai (Porter, 1916); 5) Seven species have their status revised: Cotesia arctica (Thomson, 1895), Cotesia okamotoi (Watanabe, 1921), Cotesia ukrainica (Tobias, 1986), Dolichogenidea appellator (Telenga, 1949), Dolichogenidea murinanae (Capek & Zwölfer, 1957), Hypomicrogaster acarnas Nixon, 1965, and Nyereria nigricoxis (Wilkinson, 1932); 6) New combinations are given for 318 species: Alloplitis congensis, Alloplitis detractus, Apanteles asphondyliae, Apanteles braziliensis, Apanteles sulciscutis, Choeras aper, Choeras apollion, Choeras daphne, Choeras fomes, Choeras gerontius, Choeras helle, Choeras irates, Choeras libanius, Choeras longiterebrus, Choeras loretta, Choeras recusans, Choeras sordidus, Choeras stenoterga, Choeras superbus, Choeras sylleptae, Choeras vacillatrix, Choeras vacillatropsis, Choeras venilia, Cotesia asavari, Cotesia bactriana, Cotesia bambeytripla, Cotesia berberidis, Cotesia bhairavi, Cotesia biezankoi, Cotesia bifida, Cotesia caligophagus, Cotesia cheesmanae, Cotesia compressithorax, Cotesia delphinensis, Cotesia effrena, Cotesia euphobetri, Cotesia elaeodes, Cotesia endii, Cotesia euthaliae, Cotesia exelastisae, Cotesia hiberniae, Cotesia hyperion, Cotesia hypopygialis, Cotesia hypsipylae, Cotesia jujubae, Cotesia lesbiae, Cotesia levigaster, Cotesia lizeri, Cotesia malevola, Cotesia malshri, Cotesia menezesi, Cotesia muzaffarensis, Cotesia neptisis, Cotesia nycteus, Cotesia oeceticola, Cotesia oppidicola, Cotesia opsiphanis, Cotesia pachkuriae, Cotesia paludicolae, Cotesia parbhanii, Cotesia parvicornis, Cotesia pratapae, Cotesia prozorovi, Cotesia pterophoriphagus, Cotesia radiarytensis, Cotesia rangii, Cotesia riverai, Cotesia ruficoxis, Cotesia senegalensis, Cotesia seyali, Cotesia sphenarchi, Cotesia sphingivora, Cotesia transuta, Cotesia turkestanica, Diolcogaster abengouroui, Diolcogaster agama, Diolcogaster ambositrensis, Diolcogaster anandra, Diolcogaster annulata, Diolcogaster bambeyi, Diolcogaster bicolorina, Diolcogaster cariniger, Diolcogaster cincticornis, Diolcogaster cingulata, Diolcogaster coronata, Diolcogaster coxalis, Diolcogaster dipika, Diolcogaster earina, Diolcogaster epectina, Diolcogaster epectinopsis, Diolcogaster grangeri, Diolcogaster heterocera, Diolcogaster homocera, Diolcogaster indica, Diolcogaster insularis, Diolcogaster kivuana, Diolcogaster mediosulcata, Diolcogaster megaulax, Diolcogaster neglecta, Diolcogaster nigromacula, Diolcogaster palpicolor, Diolcogaster persimilis, Diolcogaster plecopterae, Diolcogaster plutocongoensis, Diolcogaster psilocnema, Diolcogaster rufithorax, Diolcogaster semirufa, Diolcogaster seyrigi, Diolcogaster subtorquata, Diolcogaster sulcata, Diolcogaster torquatiger, Diolcogaster tristiculus, Diolcogaster turneri, Diolcogaster vulcana, Diolcogaster wittei, Distatrix anthedon, Distatrix cerales, Distatrix cuspidalis, Distatrix euproctidis, Distatrix flava, Distatrix geometrivora, Distatrix maia, Distatrix tookei, Distatrix termina, Distatrix simulissima, Dolichogenidea agamedes, Dolichogenidea aluella, Dolichogenidea argiope, Dolichogenidea atreus, Dolichogenidea bakeri, Dolichogenidea basiflava, Dolichogenidea bersa, Dolichogenidea biplagae, Dolichogenidea bisulcata, Dolichogenidea catonix, Dolichogenidea chrysis, Dolichogenidea coffea, Dolichogenidea coretas, Dolichogenidea cyane, Dolichogenidea diaphantus, Dolichogenidea diparopsidis, Dolichogenidea dryas, Dolichogenidea earterus, Dolichogenidea ensiger, Dolichogenidea eros, Dolichogenidea evadne, Dolichogenidea falcator, Dolichogenidea gelechiidivoris, Dolichogenidea gobica, Dolichogenidea hyalinis, Dolichogenidea iriarte, Dolichogenidea lakhaensis, Dolichogenidea lampe, Dolichogenidea laspeyresiella, Dolichogenidea latistigma, Dolichogenidea lebene, Dolichogenidea lucidinervis, Dolichogenidea malacosomae, Dolichogenidea maro, Dolichogenidea mendosae, Dolichogenidea monticola, Dolichogenidea nigra, Dolichogenidea olivierellae, Dolichogenidea parallelis, Dolichogenidea pelopea, Dolichogenidea pelops, Dolichogenidea phaenna, Dolichogenidea pisenor, Dolichogenidea roepkei, Dolichogenidea scabra, Dolichogenidea statius, Dolichogenidea stenotelas, Dolichogenidea striata, Dolichogenidea wittei, Exoryza asotae, Exoryza belippicola, Exoryza hylas, Exoryza megagaster, Exoryza oryzae, Glyptapanteles aggestus, Glyptapanteles agynus, Glyptapanteles aithos, Glyptapanteles amenophis, Glyptapanteles antarctiae, Glyptapanteles anubis, Glyptapanteles arginae, Glyptapanteles argus, Glyptapanteles atylana, Glyptapanteles badgleyi, Glyptapanteles bataviensis, Glyptapanteles bistonis, Glyptapanteles borocerae, Glyptapanteles cacao, Glyptapanteles cadei, Glyptapanteles cinyras, Glyptapanteles eryphanidis, Glyptapanteles euproctisiphagus, Glyptapanteles eutelus, Glyptapanteles fabiae, Glyptapanteles fulvigaster, Glyptapanteles fuscinervis, Glyptapanteles gahinga, Glyptapanteles globatus, Glyptapanteles glyphodes, Glyptapanteles guierae, Glyptapanteles horus, Glyptapanteles intricatus, Glyptapanteles lamprosemae, Glyptapanteles lefevrei, Glyptapanteles leucotretae, Glyptapanteles lissopleurus, Glyptapanteles madecassus, Glyptapanteles marquesi, Glyptapanteles melanotus, Glyptapanteles melissus, Glyptapanteles merope, Glyptapanteles naromae, Glyptapanteles nepitae, Glyptapanteles nigrescens, Glyptapanteles ninus, Glyptapanteles nkuli, Glyptapanteles parasundanus, Glyptapanteles penelope, Glyptapanteles penthocratus, Glyptapanteles philippinensis, Glyptapanteles philocampus, Glyptapanteles phoebe, Glyptapanteles phytometraduplus, Glyptapanteles propylae, Glyptapanteles puera, Glyptapanteles seydeli, Glyptapanteles siderion, Glyptapanteles simus, Glyptapanteles speciosissimus, Glyptapanteles spilosomae, Glyptapanteles subpunctatus, Glyptapanteles thespis, Glyptapanteles thoseae, Glyptapanteles venustus, Glyptapanteles wilkinsoni, Hypomicrogaster samarshalli, Iconella cajani, Iconella detrectans, Iconella jason, Iconella lynceus, Iconella pyrene, Iconella tedanius, Illidops azamgarhensis, Illidops lamprosemae, Illidops trabea, Keylimepie striatus, Microplitis adisurae, Microplitis mexicanus, Neoclarkinella ariadne, Neoclarkinella curvinervus, Neoclarkinella sundana, Nyereria ituriensis, Nyereria nioro, Nyereria proagynus, Nyereria taoi, Nyereria vallatae, Parapanteles aethiopicus, Parapanteles alternatus, Parapanteles aso, Parapanteles atellae, Parapanteles bagicha, Parapanteles cleo, Parapanteles cyclorhaphus, Parapanteles demades, Parapanteles endymion, Parapanteles epiplemicidus, Parapanteles expulsus, Parapanteles fallax, Parapanteles folia, Parapanteles furax, Parapanteles hemitheae, Parapanteles hyposidrae, Parapanteles indicus, Parapanteles javensis, Parapanteles jhaverii, Parapanteles maculipalpis, Parapanteles maynei, Parapanteles neocajani, Parapanteles neohyblaeae, Parapanteles nydia, Parapanteles prosper, Parapanteles prosymna, Parapanteles punctatissimus, Parapanteles regalis, Parapanteles sarpedon, Parapanteles sartamus, Parapanteles scultena, Parapanteles transvaalensis, Parapanteles turri, Parapanteles xanthopholis, Pholetesor acutus, Pholetesor brevivalvatus, Pholetesor extentus, Pholetesor ingenuoides, Pholetesor kuwayamai, Promicrogaster apidanus, Promicrogaster briareus, Promicrogaster conopiae, Promicrogaster emesa, Promicrogaster grandicula, Promicrogaster orsedice, Promicrogaster repleta, Promicrogaster typhon, Sathon bekilyensis, Sathon flavofacialis, Sathon laurae, Sathon mikeno, Sathon ruandanus, Sathon rufotestaceus, Venanides astydamia, Venanides demeter, Venanides parmula, and Venanides symmysta.

Keywords

Microgastrinae, world fauna, checklist, nomenclature changes, genus diagnosis, genus illustration, distribution, Lepidoptera

Introduction

With almost 3,000 described species and estimates of up to 46,000+ worldwide (Rodriguez et al. 2013), the parasitoid wasp subfamily Microgastrinae (Hymenoptera: Ichneumonoidea, Braconidae) is an important and hyperdiverse group, which has long played a central role in our understanding of insect parasitism in the context of many areas of ecological, agricultural, and basic science (Whitfield et al. 2018). Because of their diversity, prevalence in most terrestrial habitats, and the fact that species are exclusively parasitoids of larval Lepidoptera across nearly the full range of families within the taxon (Eulepidoptera, sensu Aarvik et al. 2017), microgastrine wasps are one of the most important groups in the biological control of agricultural and forestry lepidopterous pests worldwide (Whitfield 1997).

A world checklist of Microgastrinae has never been published, although Shenefelt (1972, 1973) listed the species as part of his monumental work cataloguing the world species of Braconidae. Unfortunately, those papers are outdated, especially since Mason (1981) published a seminal study that changed the generic and tribal classifications. In addition to taxonomic changes (many nominal species had been placed in synonymy), the number of newly described species has increased dramatically since Shenefelt’s catalogue: 1,446 new species of Microgastrinae (48.2%) were described between 1974 and 2019. In the past six years alone (2014–2019), 720 new species have been described (an average of 120 new species/year), which represents, by far, the largest increase in species for any subfamily of Braconidae in that time span (data extracted from this paper and Yu et al. 2016).

The database Taxapad, originally produced as a CD (Yu et al. 2005), and later available as a USB drive (Yu et al. 2012, 2016) or, partially, as a web product (now offline), has been used as the de facto catalogue of Ichneumonoidea (and associated data comprising some 350,000 names) for almost fifteen years. It is important to understand that it is essentially a compilation of all published information, whether correct or not. Nevertheless, Taxapad is an extraordinary product that contains copious information about the taxonomy, distribution, hosts and associated host plants, morphology, etc., of Ichneumonoidea that is easy to collate and analyze. As a result, it is widely consulted by researchers worldwide, and it has been adopted and (unfortunately uncritically) used in many other databases, websites, and publications pertinent to Ichneumonoidea.

However, for Microgastrinae, Taxapad follows a classification based on van Achterberg (2003), which is far from being universally accepted. A different classification, based on an older, more comprehensive paper (Mason 1981), is the one preferred and used by most researchers worldwide (e.g., Papp 1988, Kotenko 2007a, Shaw 2012, Broad et al. 2016 in the Palearctic; Whitfield 1995a, Fernandez-Triana 2010 in the Nearctic; Whitfield 1997, Fernandez-Triana et al. 2014e in the Neotropical region; Rousse and Gupta 2013 in the Afrotropical region; Chen and Song 2004, Liu et al. 2017, 2018 in the Oriental region; Austin and Dangerfield 1992 in Australasia). Thus, the Microgastrinae arrangement in Taxapad conflicts with that used by most taxonomists working on the subfamily, a situation that becomes even more confusing for ecologists, biocontrol researchers and other non-taxonomist users of Taxapad.

To complicate matters further, neither Mason (1981) nor van Achterberg (2003) treated all world species, having left many nominal species without checking their generic placement, especially those described in older literature. As a result, many of those species have remained where they were originally described or as Nixon (1965) interpreted them, usually in one of the three traditional genera historically considered to constitute practically all Microgastrinae: Apanteles Foerster, Microgaster Latreille, and Microplitis Foerster; or they were placed as part of an expanded Apanteles and Protapanteles Ashmead (sensu van Achterberg 2003). Some exceptions fared slightly better, e.g., Papp (1988) assigned many European species to Mason’s (1981) genera, Whitfield (1995a) did the same for North America, and Austin and Dangerfield (1992) for Australasia.

In this paper we a) summarize general information about Microgastrinae, including a historical outline of the internal classification, estimates of specific and generic diversity, distribution at local and world levels, advances in regional taxonomic studies, and general trends in host use; b) characterize all 81 currently accepted genera of extant Microgastrinae, including brief morphological diagnostic features, colour illustrations, available DNA barcodes and general comments on known host families; c) revise, to the best of our knowledge, the generic placement of all described species of Microgastrinae; d) compile an updated checklist of the extant and fossil world species of Microgastrinae, including recorded geographical distribution and taxonomic notes; and e) provide all information as a supplementary Excel file, to facilitate future use of the data. As work on Microgastrinae advances, we hope to provide updates in future versions of this checklist.

Materials and methods

We used the last two versions of Taxapad (Yu et al. 2012, 2016) as the starting point to compile a list of world genera and species of Microgastrinae and their recorded geographical distribution. Because the last version of Taxapad includes only information published up to the end of 2015, with some data from early 2016 (Yu, pers. comm.), we checked Zoological Record and Google Scholar for all papers published after 2015. The information presented in this paper has the cut off date of 31 December 2019.

We also compiled information from some of the world’s largest collections of Microgastrinae. All primary types (representing almost 500 species) of the Canadian National Collection of Insects (Ottawa, Canada) were studied, and unpublished information on the distribution of many species and genera was extracted from that collection, probably the largest depository of world Microgastrinae, with 120,000+ pinned specimens. We examined all primary types (representing almost 500 species of Microgastrinae) in The Natural History Museum (London, United Kingdom). Most of the primary types (representing almost 400 species of Microgastrinae) in the National Museum of Natural History (Washington, United States) were either examined or studied from images (available at http://www.usnmhymtypes.com/). Types and non-type material were extensively studied in the Finnish Museum of Natural History (Helsinki, Finland), the National Museums of Scotland (Edinburgh, United Kingdom), four major Japanese collections (Hokkaido University, Sappporo; Kobe University, Kobe; Meijo University, Nagoya; and the Osaka Museum of Natural History, Osaka), the New Zealand Arthropod Collection (Auckland, New Zealand), Naturalis (Leiden, the Netherlands), the Hungarian Natural History Museum (Budapest, Hungary), and the Austrian Natural History Museum (Vienna, Austria). Extensive non-type material, representing thousands of specimens worldwide, were borrowed for study from several institutions in Canada, Costa Rica, France, Sweden, Thailand, and the United States. Several online databases such as the Barcoding of Life Data Systems (http://v4.boldsystems.org/) and Area de Conservación Guanacaste (ACG), Costa Rica (http://janzen.sas.upenn.edu/caterpillars/database.lasso) were searched as well. The final data were input into an Excel file, which is provided here as a supplementary file to facilitate access to all information for personal use and editing (Suppl. material 1). We also provide an index of all available species names of Microgastrinae in strict alphabetical order; with the valid names in bold and italics, and the synonyms, homonyms, and nomina dubia just in italics (Suppl. material 2).

After the initial list was compiled, all species were assessed as comprehensively as possible, including: a) examination of primary types whenever possible (in a few cases we examined high quality illustrations of the primary types, which were sufficient to establish their generic placement unambiguously; in those cases we clearly indicate the source of the illustrations); b) study of secondary types and/or authenticated specimens (= specimens in collections identified by experts on the group; in those cases we mention the name of the expert identifying the species); and c) checking relevant literature, either the original description (including illustrations whenever available) or subsequent references where the species was treated (e.g., taxonomic revision, regional checklist, etc.). Throughout the checklist, “not examined but original description checked” or “not examined but subsequent treatment of the species checked” means that one of us checked those references. For every species, we detail how we assessed its status, as it is evident that the conclusion will be more reliable if the primary type was examined as opposed to secondary types, authenticated specimens, or the reading of a description. For species where we could neither examine specimens nor check for relevant literature we (explicitly) maintain the original generic combination.

For a few species, mostly in Apanteles and Microgaster, the available information (usually only the original description) was enough to suggest that they belonged to a different genus, but not enough to confidently place them in another genus (usually because several alternatives were possible, or none was clear). In those cases we considered the species as species inquirendae and add a question mark before the genus name it was originally described in (e.g., ? Apanteles) to indicate the questionable generic placement.

In the checklist, at the beginning of each genus we detail its author, year of publication and page (of the original description of the genus), gender of the genus name, type species, genus synonyms, and comments (if needed). As far as we know, the gender of every Microgastrinae genus has not been stated in a single publication before (e.g., Shenefelt (1972, 1973) did not address that; Mason (1981) only discussed the gender of some of the new genera described there; Yu et al. (2016) did not present that information either). For our checklist we follow the original publication (if the gender was stated there), or expert advice from an ICZN commissioner (Doug Yanega, pers. comm.).

For each species in the checklist we provide current name, original combination, synonyms, homonyms, and details of the primary type (including sex, holding institution, and country of the type locality), as well as details of the recorded geographical distribution of the species. Where necessary, additional comments are added at the end of the species’ treatment under “Notes”. We do not include full details on the combination history of the species name or further taxonomic details (other than the ones detailed above). For such details, Taxapad (Yu et al. 2012, 2016) and Shenefelt (1972, 1972) must be consulted.

The spelling of some author’s last names was found to vary in the literature: de Saeger/De Saeger, de Santis/De Santis, Fernandez-Triana/Fernandez-Triana, Foerster/Förster, van Achterberg/Van Achterberg. For the sake of consistency, in this paper we are using the first alternative in each of the above cases. The only exception is María Teresa Oltra Moscardó (Spain), as she has recorded her last name in several publications as either Oltra (referring to species authorship and also as paper authorship for most of her papers) or Oltra-Moscardó (only applying to one paper cited in our checklist: Oltra-Moscardó and Jiménez-Peydró 2005). In this case we use the appropriate alternative according to the corresponding reference cited, but for all eight species that she has described we refer to her as Oltra.

The availability of species names was assessed following the latest version of the International Commission on Zoological Nomenclature (ICZN); throughout the text any reference to ICZN articles follows the online version (https://www.iczn.org/the-code/the-international-code-of-zoological-nomenclature/the-code-online/).

Details on species distribution are first presented by biogeographical regions, and then by countries within biogeographical regions, in both cases arranged in alphabetical order. For biogeographical boundaries we follow the O’Hara et al. (2009) approach of combining the Australasian and Oceanian regions into one, with the name of the former. Throughout the text we use six regions (there are no Microgastrinae recorded from their Antarctic region), abbreviated as follows: NEO Neotropical (sometimes referred to as Neotropics), NEA Nearctic, PAL Palaearctic, OTL Oriental, AFR Afrotropical (sometimes referred to as Afrotropics), and AUS Australasian.

Occasionally, we use wider terms such as Holarctic (NEA and PAL), New World (NEA and NEO), Old World tropics (AFR, OTL and AUS), and pantropical (NEO, AFR, OTL, AUS). Some of these terms can be vague or hard to define precisely (e.g., some of the Australasian or southern Neotropical taxa are not really “tropical”, and the southern limits of the Holarctic region have a mix of temperate and subtropical taxa). However, they are used throughout the paper as a way to discuss trends in generic distribution and are not meant to be taken as strictly defined boundaries.

The list of countries follows the Standard ISO 3166 (codes for names of countries and their subdivisions: https://www.iso.org/obp/ui/#search). Throughout the text, we abbreviate United States of America as USA. For the six largest countries by area (Russia, Canada, China, USA, Brazil and Australia) we also present finer species distributions by country subdivisions (provinces, republics, states, territories, etc.). For Australian states and territories, we follow http://www.bda-online.org.au/help/bda-conventions/abbreviations-states/. For states of the USA and for Canadian provinces and territories, acronyms consisting of two capital letters are used, following Canada Post (http://www.canadapost.ca/tools/pg/manual/PGaddress-e.asp). We follow Standard ISO 3166 for China provinces (https://www.iso.org/obp/ui/#iso:code:3166:CN) and Brazil states (https://www.iso.org/obp/ui/#iso:code:3166:BR). For Russia subdivisions we mostly follow Standard ISO 3166 (https://www.iso.org/obp/ui/#iso:code:3166:RU), but see next paragraph for explanation on exceptions.

In most cases the information on species distribution per subdivisions was summarized from Yu et al. (2016), with updates from publications after that date. For Brazil we followed Shimbori et al. (2019). For Russia we mostly followed Yu et al. (2016), but we also added information from a recent update from Belokobylskij et al. (2019). However, Belokobylskij et al. (2019) combined several of the Russian subdivisions (according to the Standard ISO 3166, followed by Yu et al. 2016 and also by us in this paper) into broader categories, its “geoscheme for Russia” being different. As a result, some species recorded from Russia have its distribution detailed only to the level of those broader categories, as dealt with by Belokobylskij et al. (2019). The acronyms for those categories are as follow: C Centre, E East, N North, NC North Caucasus, NW North-West and S South, in the “European Part of Russia”; IR Irkutsk Province, in “Eastern Siberia”; UR Ural in the “Ural” (no province or territory detailed); KA Kamchatka Territory and PR Primorskii Territory, in the “Far East” (for more details see Belokobylskij et al. 2019: 9, fig. 1 on page 10).

Some countries have political units located in different biogeographical regions (or, in some cases, islands which are separate from the continent where the country is located), we considered those units as separate entities in our checklist (and the “country” in those cases is recorded as the separate entity and not the actual country it politically belongs to). Those cases are: Chile (Juan Fernández Islands), France (French Guiana, Guadeloupe, Marquesas Islands, Réunion, Society Islands), Japan (Ryukyu Islands), the Netherlands (Netherlands Antilles), Portugal (Azores, Madeira Islands, Selvagens Islands), Spain (Canary Islands), United Kingdom (British Virgin Islands, Saint Helena), and USA (American Samoa, Hawaiian Islands, and the USA Virgin Islands).

For all species historically recorded from the former Czechoslovakia we were able to separate the records that belong to either Czech Republic or Slovakia, based on Capek and Lukas (1989). However, for some species historically recorded from the former Yugoslavia (currently six or seven different countries, depending on the source) and also from the former Sudan (currently two countries: Sudan and South Sudan), the sources of the species records did not contain enough information to determine to which country they currently belong; therefore we annotate those records just as Yugoslavia and Sudan respectively.

Apart from some general comments on Microgastrinae hosts, we have not attempted to add host information for particular species; we intend to publish a critical assessment of Microgastrinae host records at a later date. We do, however, state general trends in host parasitization on a generic level. We follow the arrangement in Aarvik et al. (2017) when referring to families and superfamilies of Lepidoptera. Taxapad (Yu et al. 2016) gives almost complete information on published host records up to the end of 2015, but that source is inevitably very far from a reliable indication of true host associations. A complete and critical analysis of those records would require a huge effort, and in many cases it might be very difficult to determine unambiguously which ones are correct. In this respect, the amount of misinformation in the general literature is far larger than generally realised and can completely mask any real understanding of a parasitoid’s host range; Noyes (1994), Shaw (1994) and Shaw and Aeschlimann (1994) discuss this with examples.

For collection acronyms we mostly follow the website “Insect and Spider Collections of the World” (http://hbs.bishopmuseum.org/codens/codens-r-us.html). In cases where institutions were not listed there, we propose codens based on some abbreviation of the institution name. The complete list of institutions mentioned in this paper is:

AEIC American Entomological Institute, Utah State University, Logan, USA

AMNH American Museum of Natural History, New York, New York, USA

AMUZ Aligarh Muslim University, Zoological Museum, Aligarh, Uttar Pradesh, India

ANIC Australian National Insect Collection, CSIRO, Canberra City, Australia

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA

BAMU Dr. Babasaheb Ambedkar Marathwada University, Aurangabad, India

BGM Beth Gordon Agriculture and Nature Study Institute, Deganya, Israel

BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA

CAS California Academy of Sciences, San Francisco, California, USA

CBGP Centre de Biologie pour la Gestion des Populations, Montpellier, France

CFRB Chinese Academy of Forestry, Forest Research Institute, Beijing, China

CNC Canadian National Collection of Insects, Ottawa, Canada

CUIC Cornell University, Ithaca, New York, USA

DCBU Departamento de Ecologia e Biologia Evolutiva, Universidad Federal de São Carlos, São Carlos, Brazil

DCMP Universidade Federal do Paraná, Curitiba, Paraná, Brazil

DPBA Departamento de Patologia Vegetal, Buenos Aires, Argentina

DPPZ Department of Plant Protection, University of Zabol, Zabol, Iran

DZCU Department of Zoology, University of Calicut, Kerala, India

DZUC University of Ceylon, Department of Zoology, Colombo, Sri Lanka

EBW Deutsches Entomologisches Institut, Eberswalde, Germany

EIHU Hokkaido University, Sapporo, Hokkaido, Japan

ESUW University of Wyoming, Laramie, USA

FAFU Fujian Agriculture and Forestry University, Fuzhou, China

FNIC Fiji National Insect Collection, Suva, Fiji

FSCA Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, USA

GUGC Guizhou University, Guiyang, China

HNHM Hungarian Natural History Museum, Budapest, Hungary

HUNAU Hunan Agricultural University, Changsha, China

IAVH Instituto Alexander von Humboldt, Bogotá, Colombia

IEAS Academia Sinica, Institute of Entomology, Shanghai, Shanghai, China

IEBR Institute of Ecology and Biological Resources, Hanoi, Vietnam

IECA Institute of Entomology, České Budějovice, Czech Republic

IFRI Indian Forest Research Institute, Dehradun, Uttarakhand, India

IIAF Instituto de Investigaciones Agropecuarias y Forestales, Universidad Michoacana San Nicolás de Hidalgo, México

INBio Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica

INHS Illinois Natural History Survey, Champaign, Illinois, USA

INPC National Pusa Collections, Indian Agricultural Research Institute, New Delhi, India

KUEC Kyushu University, Fukuoka, Japan

LNKD Landessammlung für Naturkunde, Karlsruhe, Germany

LSUK The Linnean Society of London, London, United Kingdom

LUNZ Lincoln University, Lincoln, New Zealand

MACN Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, USA

MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland

MIUP Museo de Invertebrados Graham Bell Fairchild, Universidad de Panamá, Panama

MLP Museo de La Plata, La Plata, Argentina

MMBC Moravske Muzeum [Moravian Museum], Brno, Czech Republic

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain

MNHN Muséum National d'Histoire Naturelle, Paris, France

MNNC Museo Nacional de Historia Natural, Santiago, Chile

MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru

MVMMA Museums Victoria, Melbourne Museum, Melbourne, Australia

MZH Finnish Museum of Natural History, Helsinki, Finland

MZLU Lund University, Lund, Sweden

MZUSP Museum of Zoology, University of São Paulo, Brazil

NBAIR National Bureau of Agricultural Insect Resources, Bangalore, India

NHMO Zoological Museum, University of Oslo, Oslo, Norway

NHMUK Natural History Museum, London, United Kingdom

NHMW Naturhistorisches Museum Wien, Vienna, Austria

NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden

NIAES National Institute for Agro-Environmental Sciences, Tsukuba, Japan

NMID National Museum of Ireland, Dublin, Ireland

NMKE National Museum of Kenya, Nairobi, Kenya

NZAC New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand

NZSI National Zoological Collection, Zoological Survey of India, Kolkata, West Bengal, India

OUMNH Museum of Natural History, Oxford University, United Kingdom

PCMAG Plymouth City Museum and Art Gallery, Plymouth, United Kingdom

PPRI Plant Protection Research Institute, Pretoria, Gauteng, South Africa

QM Queensland Museum, South Brisbane, Queensland, Australia

QSBG Queen Sirikit Botanic Garden, Chaing Mai, Thailand

QCAZ Pontificia Universidad Católica del Ecuador, Quito, Ecuador

RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium

RMCA Musée Royal de l'Afrique Centrale, Tervuren, Belgium

RMNH Naturalis Biodiversity Centre, Leiden, Netherlands

RSME National Museums of Scotland, Edinburgh, United Kingdom

SAMA South Australian Museum, Adelaide, South Australia, Australia

SAMC Iziko Museum of Capetown, Cape Town, South Africa

SAUC Shandong Agricultural University, Tai'an, China

SCAC South China Agricultural College, Guangzhou, Guangdong, China

SEMC Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA

SIZK Schmalhausen Institute of Zoology, Kiev, Ukraine

SJCA St. John's College, Agra, Uttar Pradesh, India

SMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt-am-Main, Germany

SUKI Shivaji University, Kolhapur, India

TARI Taiwan Agricultural Research Institute, Taichung, Taiwan, China

TFRI Insect Museum, Tropical Forest Research Institute, Jabalpur, Madhya Pradesh, India

TMAG Tasmanian Museum and Art Gallery, Hobart, Tasmania, Australia

TMSA Ditsong National Museum of Natural History, Pretoria, Gauteng, South Africa

TMUC Department of Entomology, Tarbiat Modares University, Tehran, Iran

TUDTG Technische Universität Dresden, Department of Forest Science, Tharandt, Germany

UCDC R.M. Bohart Museum of Entomology, University of California, Davis, California, USA

UFSM Universidade Federal de Santa Maria, Rio Grande do Sul, Brazil

UFVB Universidade Federal de Viçosa, Museum of Entomology, Viçosa, Minas Gerais, Brazil

UKM Universiti Kebangsaan, Bangi, Selangor, Malaysia

UKZMP Universiti Kebangsaan, Bangi, Selangor, Malaysia

ULQC University of Laval, Quebec City, Canada

USNM National Museum of Natural History, Washington, USA

UUZM Uppsala University, Uppsala, Sweden

UVS University of Valencia, Valencia, Spain

VNMN Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, Hanoi, Vietnam

WAM Western Australian Museum, Perth, Western Australia, Australia

ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia

ZJUH Parasitic Hymenoptera Collection, Zhejiang University, Hangzhou, China

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany

ZMTU Zoological Museum, Trakya University, Turkey

ZMUC Zoological Museum, University of Copenhagen, Copenhagen, Danmark

ZMUK Zoologisches Museum, Universität Kiel, Kiel, Germany

ZSM Zoologische Staatssammlung, Munich, Germany

The concept of DNA barcoding as a tool for species discovery and identification was proposed approximately 15 years ago (Hebert et al. 2003a, 2003b). A short DNA sequence, approximately 650 base pairs (bp) in the mitochondrial gene encoding cytochrome c oxidase subunit 1 (CO1), has been accepted as a practical and standardized DNA barcode for many groups of animals (e.g., Kress et al. 2015). The Barcode Index Number (BIN) System uses DNA barcodes to indicate possible species limits (see more details on the BIN concept in Ratnasingham and Hebert 2013), and it has been used in taxonomic studies of Microgastrinae (e.g., Fernandez-Triana and Boudreault 2018, Fagan-Jeffries et al. 2018b). In the checklist below we provide details of the number of DNA barcode sequences and BINs for every genus of Microgastrinae currently available in the Barcoding of Life Data Systems (BOLD, see also http://v4.boldsystems.org/index.php) as of 31 December 2019. Sequences were considered as “barcode compliant” if they fulfilled the requirements set in Ratnasingham and Hebert (2007), namely: the sequence has at least 500 nucleotides with fewer than 1% ambiguous base calls (Ns); it has a species name (assigned by an expert taxonomist) or a provisional name; it has a unique specimen identifier, information related to the voucher specimen (including the name of the institution storing the voucher), and a collection record (e.g., collector, collection date, collection location, geospatial coordinates); and it has the sequence of PCR primers used to generate the CO1 amplicon and the trace files (Santschi et al. 2013).

We provide brief morphological diagnostic features and colour illustrations for all 81 valid genera of Microgastrinae (at least one species per genus is illustrated, usually more). For morphological terms we follow several published references (Huber and Sharkey 1993, Sharkey and Wharton 1997, Karlsson and Ronquist 2012, Fernandez-Triana et al. 2014e) as well as the Hymenoptera Anatomy Ontology (HAO) website (http://portal.hymao.org/projects/32/public/ontology/). We use the abbreviations T1, T2, and T3 for metasomal mediotergites 1, 2, and 3; and the fore wing second submarginal cell is mentioned throughout the text as areolet for the sake of brevity.

Photographs were taken with either a Keyence VHX-1000 Digital Microscope or with a Leica camera on a Leica M165 C Microscope, using lenses with a range of 10–130 ×. Multiple images were taken of a structure through the focal plane and then combined to produce a single in-focus image using the software associated with the Keyence System or, for the images taken with the Leica camera, the Zerene Stacker program (http://zerenesystems.com/cms/stacker). Images were corrected using Adobe Photoshop CS4 and Gimp 2.10.12; the plates were prepared using Microsoft PowerPoint 2010 and later saved as .tiff files. For seven figures in our paper we used other sources, all of which are acknowledged in the corresponding figure caption and in the Acknowledgements section below.

In the Results section, we discuss several topics concerning Microgastrinae before providing the checklist of world species. These include a detailed explanation of the generic concepts used here, geographical patterns, general overview of host data in the subfamily, extinct taxa, and limitations of both Taxapad and our checklist. It is very important to understand the limitations, as the user must be aware of the areas where Taxapad and/or our list lack strong support, e.g., critical review of host data, and/or missing information, such as examination of primary types. Further, there will undoubtedly be some yet to be recognised synonymy. We hope future versions of our world checklist will address some of the shortcomings of the present one. We also hope to prepare an online version that is continuously updated, probably in the style of a similar effort currently outdated (http://microgastrinae.myspecies.info/).

Results

Overview of the present paper and its limitations

In the checklist below, a total of 81 genera and 2,999 extant species are recognized as valid, including 36 nominal species that are currently considered to be species inquirendae.

Two genera are synonymized under Apanteles: Cecidobracon Kieffer & Jörgensen, 1910 syn. nov., and Holcapanteles Cameron, 1905 syn. nov. Nine lectotypes are designated. A total of 318 new combinations, three new replacement names, three species name amendments, and seven species status revised are proposed. Additionally, three species names are treated as nomina dubia, and 52 species names are considered to be unavailable (including 14 as nomina nuda), listed at the end of the checklist.

Extinct taxa, only known as fossils (three genera and 12 species) are listed in a separate section below (Table 3).

The pace of species description in Microgastrinae has been steadily increasing since the first species was described in 1758 and has shown no signs of slowing down (Fig. 1). The total number of genera has also increased substantially, especially since 1965; the information is summarized in Whitfield et al. (2018), Fernandez-Triana and Boudreault (2018), and below.

Figure 1. 

Microgastrinae species described since 1758 based on data in present paper A Total numbers per decade B Cumulative number (1758–2019).

Primary types of Microgastrinae are deposited in 108 institutions worldwide, although 76% of those types are concentrated in seventeen collections (Table 1), seven of which have more than 100 primary types each. Localities of primary types are reported from 138 different countries.

Microgastrine wasps have been recorded in most countries and all continents except Antarctica. Only 16 countries do not yet have any recorded species of Microgastrinae: Bahrain, Botswana, Bhutan, Cambodia, Djibouti, Equatorial Guinea, Gabon, Gambia, Guinea, Guinea-Bissau, Kuwait, Laos, Liberia, Mauritania, Qatar, and Swaziland. This is of course just an artifact of insufficient collecting and/or lack of studies in those countries; each is expected to harbour many species.

The current data (Table 2) show two countries with 400+ Microgastrinae species each (China with 448 and Costa Rica with 427), another two with 300+ species each (Russia with 388 and Hungary with 327) and five with 200+ species each (USA, Germany, India, United Kingdom, and Canada). Overall, 34 countries have more than 100 described species recorded, although those numbers can be misleading. For example, the reason Hungary ranks so high is because of extensive studies in that country, done over many years by Jenö Papp while working in the Hungarian Natural History Museum. A similar situation applies to both the United Kingdom and Germany, where a long European tradition of experts on the group coupled with extensive collecting have provided figures that are close to the actual diversity in those countries. While the microgastrine fauna of those three countries is relatively well known, the opposite occurs in large and/or mostly tropical countries, where more species are still undescribed. For example, in Costa Rica, DNA barcoding has already identified more than 1,200 species just in ACG (Janzen and Hallwachs 2016). And the figures for China and India (which are considered to be “megadiverse countries”, sensu Myers et al. 2000), are still very far from being complete, as both countries should easily reach more than 1,000 species each. Other megadiverse countries such as Australia, Brazil, Colombia, Democratic Republic of Congo, Indonesia, Madagascar, Mexico, Peru, Malaysia, Papua New Guinea and USA are all likely to have similar (in some cases higher) totals, but studies thus far have been insufficient, leading to most of those countries having “only” a hundred species or fewer recorded at present.

There are three main limitations in our paper that we want to point out. The first relates to the coverage of primary types in our study. We were able to examine primary types for 1,394 species (46.5%), and for another 1,568 species (52.3%) we studied authenticated specimens, checked original descriptions, or read subsequent revisions. However, for 37 species (1.2%) we could not check any source of information, or it was considered inadequate, and they are left in the genus in which they were originally described (or as species inquirendae), with explanatory annotations. In future versions, we aim to increase the number of species for which we have examined primary types, but for the present paper the reader must consider the relatively large number of species still needing to be thoroughly studied. It is especially important to keep in mind that for some of those species for which we could only study descriptions (which may not be detailed or clear enough), the generic placement made in this paper might be incorrect.

A second limitation is the coverage of references concerning Microgastrinae. In the References section we tried to list all papers where original descriptions of Microgastrinae were published (those references in turn are cited under the corresponding treatment of every species in the checklist below). However, our list is not complete and we are aware of some omissions; in that sense, the latest versions of Taxapad (Yu et al. 2012, 2016) have more comprehensive lists of references than our paper. Especially important and comprehensive is Yu et al. (2016), which lists 6,200+ references related to Microgastrinae.

A third limitation of our paper is that we do not treat host records in detail. We expect to present host data for microgastrine species with verified information in a subsequent version of the world checklist, although it is improbable that we will be able to comment with reliability on all published records. The latest versions of Taxapad (Yu et al. 2012, 2016) provide the best coverage of references on hosts of Microgastrinae; however, that is only an uncritical compilation of literature, and many of those references report incorrect data. The reader is strongly advised to double check host references and be very cautious in interpreting information from secondary sources.

World collections with the largest numbers of primary types of Microgastrinae (data from valid species as recognized in the present paper).

Collection code Country Number of primary types
NHMUK UK 491
CNC Canada 476
USNM USA 380
ZJUH China 160
RMCA Belgium 122
ZIN Russia 113
HNHM Hungary 108
MNHN France 84
FAFU China 63
ANIC Australia 52
SIZK Ukraine 44
ZMHB Germany 40
MACN Argentina 36
RMNH The Netherlands 35
AEIC USA 32
EIHU Japan 29
HUNAU China 29

Alphabetic list of countries with described species of Microgastrinae (data based on this paper). Countries with political units located in different biogeographical regions (mostly islands) have species recorded from those entities listed separately below; those species are not added to the total for the country to which the entities belong politically.

Countries No. of Species Countries No. of Species
Afghanistan 20 Lithuania 70
Albania 7 Luxembourg 1
Algeria 7 Macedonia 37
Andorra 2 Madagascar 67
Angola 1 Malawi 11
Argentina 68 Malaysia 70
Armenia 105 Mali 1
Australia 129 Malta 18
Austria 97 Mauritius 12
Azerbaijan 126 Mexico 54
Bahamas 1 Moldova 113
Bangladesh 11 Mongolia 161
Barbados 2 Montenegro 23
Belarus 23 Morocco 14
Belgium 61 Mozambique 7
Belize 7 Myanmar 9
Benin 3 Namibia 1
Bolivia 10 Nepal 6
Bosnia and Herzegovina 6 Netherlands 105
Brazil 120 Netherlands (Netherlands Antilles) 1
Brunei 1 New Zealand 27
Bulgaria 128 Nicaragua 5
Burkina Faso 1 Niger 1
Burundi 1 Nigeria 16
Cape Verde 32 Norway 15
Cameroon 13 Oman 1
Canada 213 Pakistan 20
Central African Republic 2 Panama 22
Chad 1 Papua New Guinea 47
Chile 21 Paraguay 10
Chile (Juan Fernández Islands) 2 Peru 39
China 448 Philippines 90
Colombia 31 Poland 170
Comoros 1 Portugal 7
Democratic Republic of Congo 135 Portugal (Azores) 3
Costa Rica 427 Portugal (Madeira Islands) 14
Croatia 70 Portugal (Selvagens Islands) 2
Cuba 20 Romania 174
Cyprus 11 Russia 388
Czech Republic 90 Rwanda 59
Denmark 20 Saint Kitts & Nevis 2
Dominica 3 Saint Lucia 2
Dominican Republic 5 Saint Vincent 18
Ecuador 101 Saudi Arabia 2
Egypt 12 Senegal 51
El Salvador 1 Serbia 95
Eritrea 3 Sierra Leone 3
Estonia 12 Singapore 11
Ethiopia 11 Slovakia 161
Fiji 29 Slovenia 18
Findland 162 Solomon Islands 5
France 122 Somalia 2
France (French Guiana) 6 South Africa 98
France (Guadeloupe) 2 Spain 103
France (Marquesas Islands) 1 Spain (Canary Islands) 18
France (Réunion) 34 Sri Lanka 37
France (Society Islands) 2 Sudan 8
Gambia 1 Suriname 5
Georgia 73 Sweden 121
Germany 248 Switzerland 166
Ghana 6 Syria 2
Greece 92 Tajikistan 42
Grenada 15 Tanzania 23
Guatemala 6 Thailand 30
Guyana 12 Togo 3
Haiti 2 Tonga 2
Honduras 8 Trinidad & Tobago 19
Hungary 327 Tunisia 40
Iceland 5 Turkey 173
India 245 Turkmenistan 63
Indonesia 63 Uganda 35
Iran 109 Ukraine 154
Iraq 2 United Arab Emirates 3
Ireland 81 United Kingdom 242
Israel 72 United Kingdom (British Virgin Islands) 1
Italy 149 United Kingdom (Saint Helena) 1
Ivory Coast 16 United States 299
Jamaica 6 United States (American Samoa) 3
Japan 96 United States (Hawaiian Islands) 14
Japan (Ryukyu Islands) 7 United States (USA Virgin Islands) 1
Jordan 10 Uruguay 11
Kazakhstan 121 Uzbekistan 72
Kenya 30 Vanuatu 8
Korea 130 Venezuela 21
Kyrgyzstan 18 Vietnam 137
Latvia 37 Western Samoa 10
Lebanon 2 Yemen 17
Lesotho 1 Zambia 3
Libya 2 Zimbabwe 7

Fossil Microgastrinae taxa

Extinct genera and species of Microgastrinae have been found in Eocene and Oligocene deposits, from 37–44 million years ago (MYA). Many specimens from the Miocene (20–30 MYA) are known from Dominican and Chiapas ambers, but most appear to be undescribed representatives of extant genera (Murphy et al. 2008). Belokobylskij (2014) revised the taxonomic status of all previously known taxa of fossil Microgastrinae and described one new genus as well as two new species. The origin of Microgastrinae has been estimated at ~ 54 MYA (Murphy et al. 2008).

Unlike previous work (Mason 1981, Yu et al. 2005, 2012, 2016), we exclude fossil genera or species from our world checklist. Instead, we tabulate in this section the three genera and 12 species of fossil Microgastrinae currently described (Table 3).

Extinct genera and species of Microgastrinae, compiled from Yu et al. (2012, 2016) and Belokobylskij (2014).

Genera only known from fossils Species only known from fossils
Dacnusites Cockerell, 1921 Apanteles concinnus Statz, 1938
Eocardiochiles Brues, 1933 Apanteles macrophthalmus Statz, 1938
Palaeomicrogaster Belokobylskij, 2014 Dacnusites reductus Cockerell, 1921
Dacnusites sepultus Cockerell, 1921
Eocardiochiles fritschii Brues, 1933
Microplitis elegans Timon-David, 1944
Microplitis primordialis (Brues, 1906)
Microplitis vesperus Brues, 1910
Semionis nixoni Tobias, 1987
Semionis wightensis Belokobylskij, 2014
Snellenius succinalis Brues, 1933
Palaeomicrogaster oculatus Belokobylskij, 2014

Generic limits and taxonomic arrangement of the subfamily Microgastrinae

Microgastrinae was originally described at family rank, as ‘Microgasteroidae’, by Foerster (1863). At that time, it comprised only three genera: Microgaster Latreille, 1804 (the genus that provides the root for the subfamily name, meaning “small abdomen”, in reference to the relatively short length of the metasoma compared to other Braconidae), as well as two genera described by Foerster (1863): Microplitis (which means “small sword” or “small weapon”, referring to the generally short ovipositor in that genus) and Apanteles (meaning “incomplete”, in reference to the fore wing lacking the second intercubitus, leaving the second submarginal cell open or incomplete). Fornicia, although described by Brullé (1846) before Foerster’s work, was at the time considered to belong to other subfamilies in Braconidae (e.g., Dalla Torre (1898) placed the genus in Cheloninae; Ashmead (1900a) placed it in Sigalphinae; Granger (1949) placed it in Triaspidinae), and it was not recognized to be part of Microgastrinae until a century later (Baltazar 1962, Nixon 1965).

The high diversity of Microgastrinae quickly became evident, and so attempts to split the group into further genera started shortly after Foerster’s (1863) paper, e.g., by Reinhard (1880). Many additional genera (15 recognized in this paper) were described between 1882 and 1958, although some were not associated with Microgastrinae at the time, and others were not accepted as valid genera by some authors of the period, e.g., Muesebeck (1921) and Telenga (1955).

This view changed with two seminal works in 1965 and 1981. Nixon (1965) reclassified the subfamily limits and provided some structure to what was being recognized as a huge assemblage of parasitoids of Lepidoptera. He recognized 20 genera, eight of which were new, and reclassified the species within Apanteles sensu lato into a more practical and useful array of 44 species groups to facilitate identification. Mason (1981) fundamentally changed the taxonomy of Microgastrinae by recognizing 50 genera (23 of which he described as new), including numerous taxa that mostly corresponded to particular species groups of Nixon (1965, 1973), and additionally proposing new combinations for some 350 species.

Since Mason (1981) 32 genera have been described. Whitfield et al. (2018: fig. 2) graphically showed the increase in description of new genera during the past 150 years. Nevertheless, there are still many more genera of Microgastrinae that remain to be described, e.g., Fernandez-Triana and Boudreault (2018). Additionally, several genera, as currently understood, are probably polyphyletic and need to be split, e.g., Diolcogaster and Glyptapanteles. A comprehensive phylogenetic analysis of the subfamily is needed before we can achieve a clearer picture. However, just based on the material we have seen in collections, we estimate that the Microgastrinae is likely to comprise close to one hundred genera.

For the past few years the main problem with the generic concepts is that two different classifications of Microgastrinae have been proposed and are widely used: those based on Mason (1981) and on van Achterberg (2003). For a visual depiction of how the two classifications differ (based on the number of species assigned to each of the most speciose genera), see Figure 2.

Figure 2. 

Number of extant species per larger genera of Microgastrinae A Data from Taxapad 2016, which is mostly an update, with slight modifications, of van Achterberg (2003), total number of species: 2,710 B Data from present paper, which is mostly based on Mason (1981) but extensively updated, total number of species: 2,999.

The classification proposed by Mason (1981) had a narrower concept of Apanteles and Protapanteles, which resulted in a larger number of Microgastrinae genera treated as valid. Many of the new combinations resulting from that classification are in Mason (1981), although not all species have been properly transferred to the corresponding genus. Mason’s system has been followed by most researchers (see examples cited in the Introduction) and has remained largely stable for the past 38+ years, with a few exceptions: his genus Teremys was synonymized under Pholetesor (Whitfield 2006); and his arrangement of genera within tribes, largely based on phylogenetic grounds, has not been universally accepted (Austin 1990, Austin and Dangerfield 1992, Whitfield 1995a, Fernandez-Triana 2010; see also Walker et al. 1990 for further criticism of tribes within Microgastrinae). Mason (1981) based his paper on studies of the world fauna; however, a careful examination of the CNC collection (Mason’s base) and other material available to him at the time shows that specimens from the Afrotropical, Oriental, and Australasian regions were much more poorly represented than the remaining regions. Thus, most of the new genera from those regions described by Mason (1981) have later been found to have a wider distribution and greater morphological variation than originally thought, and some of those genera will need redefinition. Another consequence of the limited geographical coverage of the studied specimens is that the keys to tribes and genera in Mason (1981) work reasonably well for the temperate areas, but not as well for the tropical areas, especially the Old World tropics.

The classification proposed by van Achterberg (2003) reduced the number of genera by treating eleven genera recognised by Mason as synonyms or subgenera of Apanteles and Protapanteles. That system was later implemented in Taxapad (Yu et al. 2005, 2012, 2016) and other, mostly European, databases, e.g., Fauna Europaea (https://fauna-eu.org/) and Dyntaxa (https://www.dyntaxa.se/). Shortcomings of this approach have been pointed out by other authors, e.g., Broad et al. (2016) and Whitfield et al. (2018). The main issue with van Achterberg’s approach is that his classification was based mainly on the European species, a region with relatively little diversity in genera and species (see sections below), and is thus clearly insufficient to capture the rich fauna of Microgastrinae worldwide. Second, and more worrisome, van Achterberg’s generic concepts were applied in Taxapad to the entire world fauna, effectively producing numerous (perhaps hundreds) of new name combinations which have never been formally published, let alone critically assessed. The validity of those names may be questionable, but van Achterberg’s classification has been embraced uncritically by some users of Taxapad.

To complicate things further, generic concepts changed slightly in Taxapad from the 2012 to the 2016 version (Table 4). For example, Taxapad 2016 considers some taxa as subgenera that the 2012 version had listed as synonyms of Apanteles (Dolichogenidea, Exoryza, Iconella, Illidops, and Pholetesor) or as synonyms of Protapanteles (Nyereria, Rasivalva, Sathon, and Venanides). Other genera were treated differently, e.g., Distatrix is treated as a valid genus in the 2012 version but as a subgenus of Protapanteles in 2016, and Glyptapanteles is a synonym of Protapanteles in 2012 but a valid genus in 2016. Some of those decisions may have merit, but three are highly questionable:

a) Rasivalva should never have been considered to be part of Protapanteles as it has a complete areolet in the fore wing (a character not present in any Protapanteles or related genera);

b) Ectadiophatnus is listed as a genus of Microgastrinae in both the 2012 and 2016 versions, following Shenefelt (1973), despite having been published as belonging to the subfamily Blacinae since at least 1935 (Ferrière 1935, Mani 1938, Varshney 1976, Mason 1981) [van Achterberg (pers. comm.) has examined the type species and found that it is a new synonym of Eubazus Nees, in Brachistinae-Brachistini];

c) the species listed under Lissogaster have since 1988 been transferred back to Microgaster (see more details about that in Mason (1986) and in the checklist below, in the introductory comments to the genus Microgaster).

The rationale for the changes between versions of Taxapad is not always evident and, as far as we are aware, has never been explained in a published paper. As a result, it is difficult to follow the different arrangements of genera and subgenera, a problem which is further compounded by the use of tribes in the 2012 version, while the 2016 version added sub-tribes (Table 4).

We believe that the classification proposed by Mason (1981), although not entirely free from problems and shortcomings, provides the best framework currently available to deal with the world diversity of Microgastrinae and provides a solid and clear foundation from which to work towards future improvements. In this paper we largely follow that system, except for dividing the subfamily into tribes, as we do not think the tribes proposed by Mason properly reflect the phylogenetic relationships within the subfamily. We here classify the world species in 81 genera of Microgastrinae (Table 4 and checklist below).

Microgastrinae arrangement (genera, subgenera, subtribes, and tribes) used in the 2012 and 2016 versions of Taxapad (Yu et al. 2012, 2016) and the present paper. Each column is independent of the others, so the lists must be read vertically only, as they are not comparable horizontally.

Taxapad 2012 Taxapad 2016 Present paper
MICROGASTRINAE Foerster, 1862 MICROGASTRINAE Foerster, 1863 MICROGASTRINAE Foerster, 1863
MICROGASTRINI Foerster, 1863 (No tribes)
APANTELINI Viereck, 1918 APANTELINA Viereck, 1918 (No subtribes)
Alphomelon Mason, 1981 Alphomelon Mason, 1981 Agupta Fernandez-Triana, 2018
Apanteles (Apanteles) Foerster, 1862 Apanteles (Apanteles) Foerster, 1863 Alloplitis Nixon, 1965
Dolichogenidea Viereck, 1911 Napamus Papp, 1993 Alphomelon Mason, 1981
Iconella Mason, 1981 Apanteles (Choeras) Mason, 1981 Apanteles Foerster, 1863
Illidops Mason, 1981 Apanteles (Dolichogenidea) Viereck, 1911 Austinicotesia Fernandez-Triana, 2018
Napamus Papp, 1993 Apanteles (Exoryza) Mason, 1981 Austrocotesia Austin & Dangerfield, 1992
Apanteles (Choeras) Mason, 1981 Apanteles (Iconella) Mason, 1981 Beyarslania Koçak & Kemal, 2009
Apanteles (Exoryza) Mason, 1981 Apanteles (Illidops) Mason, 1981 Billmasonius Fernandez-Triana, 2018
Austrocotesia Austin & Dangerfield, 1992 Apanteles (Pholetesor) Mason, 1981 Buluka de Saeger, 1948
Exulonyx Mason, 1981 Austrocotesia Austin & Dangerfield, 1992 Carlmuesebeckius Fernandez-Triana, 2018
Miropotes Nixon, 1965 Dasylagon Muesebeck, 1958 Chaoa Luo & You, 2004
Papanteles Mason, 1981 Exulonyx Mason, 1981 Choeras Mason, 1981
Parapanteles Ashmead, 1900 Miropotes Nixon, 1965 Clarkinella Mason, 1981
Pelicope Mason, 1981 Papanteles Mason, 1981 Cotesia Cameron, 1891
Pholetesor Mason, 1981 Parapanteles Ashmead, 1900 Cuneogaster Choi & Whitfield, 2006
Promicrogaster Brues & Richardson, 1913 Promicrogaster Brues & Richardson, 1913 Dasylagon Muesebeck, 1958
Sendaphne Nixon, 1965 Sendaphne Nixon, 1965 Deuterixys Mason, 1981
Xanthapanteles Whitfield, 1995 Xanthapanteles Whitfield, 1995 Diolcogaster Ashmead, 1900
COTESIINI Mason, 1981 COTESIINA Mason, 1981 Distatrix Mason, 1981
Buluka de Saeger, 1948 Buluka de Saeger, 1948 Dodogaster Rousse, 2013
Chaoa Luo & You, 2004 Chaoa Luo & You, 2004 Dolichogenidea Viereck, 1911
Cotesia Cameron, 1891 Cotesia Cameron, 1891 Eripnopelta Xiong, van Achterberg & Chen, 2017
Cuneogaster Choi & Whitfield, 2006 Cuneogaster Choi & Whitfield, 2006 Exix Mason, 1981
Deuterixys Mason, 1981 Deuterixys Mason, 1981 Exoryza Mason, 1981
Diolcogaster Ashmead, 1900 Diolcogaster Ashmead, 1900 Exulonyx Mason, 1981
Distatrix Mason, 1981 Exix Mason, 1981 Fornicia Brullé, 1846
Exix Mason, 1981 Glyptapanteles Ashmead, 1904 Gilbertnixonius Fernandez-Triana, 2018
Larissimus Nixon, 1965 Larissimus Nixon, 1965 Glyptapanteles Ashmead, 1904
Lathrapanteles Williams, 1985 Lathrapanteles Williams, 1985 Hygroplitis Thomson, 1895
Parenion Nixon, 1965 Nyereria Mason, 1981 Hypomicrogaster Ashmead, 1898
Protapanteles (Protapanteles) Ashmead, 1898 Parenion Nixon, 1965 Iconella Mason, 1981
Glyptapanteles Ashmead, 1904 Protapanteles (Protapanteles) Ashmead, 1898 Illidops Mason, 1981
Protapanteles (Nyereria) Mason, 1981 Protapanteles (Distatrix) Mason, 1981 Janhalacaste Fernandez-Triana, 2018
Protapanteles (Rasivalva) Mason, 1981 Protapanteles (Rasivalva) Mason, 1981 Jenopappius Fernandez-Triana, 2018
Protapanteles (Sathon) Mason, 1981 Protapanteles (Sathon) Mason, 1981 Jimwhitfieldius Fernandez-Triana, 2018
Protapanteles (Venanides) Mason, 1981 Protapanteles (Venanides) Mason, 1981 Keylimepie Fernandez-Triana, 2016
Protomicroplitis Ashmead, 1898 Protomicroplitis Ashmead, 1898 Kiwigaster Fernandez-Triana, Ward & Whitfield, 2011
Pseudovenanides Xiao & You, 2002 Pseudovenanides Xiao & You, 2002 Kotenkosius Fernandez-Triana, 2018
Venanus Mason, 1981 Venanus Mason, 1981 Larissimus Nixon, 1965
Wilkinsonellus Mason, 1981 Wilkinsonellus Mason, 1981 Lathrapanteles Williams, 1985
MICROGASTRINI Foerster, 1862 MICROGASTRINA Foerster, 1863 Mariapanteles Whitfield & Fernandez-Triana, 2012
Beyarslania Koçak & Kemal, 2009 Beyarslania Koçak & Kemal, 2009 Markshawius Fernandez-Triana, 2018
Cecidobracon Kieffer & Jörgensen, 1910 Cecidobracon Kieffer & Jörgensen, 1910 Microgaster Latreille, 1804
Clarkinella Mason, 1981 Clarkinella Mason, 1981 Microplitis Foerster, 1863
Dasylagon Muesebeck, 1958 Ectadiophatnus Cameron, 1913 Miropotes Nixon, 1965
Ectadiophatnus Cameron, 1913 Holcapanteles Cameron, 1905 Napamus Papp, 1993
Holcapanteles Cameron, 1905 Hygroplitis Thomson, 1895 Neoclarkinella Rema & Narendran, 1996
Hygroplitis Thomson, 1895 Hypomicrogaster Ashmead, 1898 Nyereria Mason, 1981
Hypomicrogaster Ashmead, 1898 Lissogaster Bengtsson, 1926 Ohenri Fernandez-Triana, 2018
Lissogaster Bengtsson, 1926 Mariapanteles Whitfield & Fernandez-Triana, 2012 Papanteles Mason, 1981
Microgaster Latreille, 1804 Microgaster Latreille, 1804 Parapanteles Ashmead, 1900
Neoclarkinella Rema & Narendran, 1996 Neoclarkinella Rema & Narendran, 1996 Parenion Nixon, 1965
Paroplitis Mason, 1981 Paroplitis Mason, 1981 Paroplitis Mason, 1981
Prasmodon Nixon, 1965 Prasmodon Nixon, 1965 Pelicope Mason, 1981
Pseudapanteles Ashmead, 1898 Pseudapanteles Ashmead, 1898 Philoplitis Nixon, 1965
Rhygoplitis Mason, 1981 Rhygoplitis Mason, 1981 Pholetesor Mason, 1981
Xanthomicrogaster Cameron, 1911 Shireplitis Fernandez-Triana & Ward, 2013 Prasmodon Nixon, 1965
MICROPLITINI Mason, 1981 Xanthomicrogaster Cameron, 1911 Promicrogaster Brues & Richardson, 1913
Alloplitis Nixon, 1965 MICROPLITINI Mason, 1981 Protapanteles Ashmead, 1898
Microplitis Foerster, 1862 Alloplitis Nixon, 1965 Protomicroplitis Ashmead, 1898
Philoplitis Nixon, 1965 Microplitis Foerster, 1863 Pseudapanteles Ashmead, 1898
Snellenius Westwood, 1882 Philoplitis Nixon, 1965 Pseudofornicia van Achterberg, 2015
FORNICIINI Mason, 1981 Snellenius Westwood, 1882 Pseudovenanides Xiao & You, 2002
Fornicia Brullé, 1846 FORNICIINI Mason, 1981 Qrocodiledundee Fernandez-Triana, 2018
SEMIONINI Tobias, 1987 Fornicia Brullé, 1846 Rasivalva Mason, 1981
Semionis Nixon, 1965 Pseudofornicia van Achterberg, 2015 Rhygoplitis Mason, 1981
Kiwigaster Fernandez-Triana, Whitfield & Ward, 2011 SEMIONINI Tobias, 1987 Sathon Mason, 1981
Pelicope Mason, 1981 Semionis Nixon, 1965
Semionis Nixon, 1965 Sendaphne Nixon, 1965
Dodogaster Rousse, 2013 Shireplitis Fernandez-Triana & Ward, 2013
Keylimepie Fernandez-Triana, 2016 Snellenius Westwood, 1882
Kiwigaster Fernandez-Triana, Whitfield & Ward, 2011 Silvaspinosus Fernandez-Triana, 2018
Tobleronius Fernandez-Triana, 2018
Ungunicus Fernandez-Triana, 2018
Venanides Mason, 1981
Venanus Mason, 1981
Wilkinsonellus Mason, 1981
Xanthapanteles Whitfield, 1995
Xanthomicrogaster Cameron, 1911
Ypsilonigaster Fernandez-Triana, 2018
Zachterbergius Fernandez-Triana, 2018

Brief diagnosis of all Microgastrinae genera as they are understood in this paper

The last two published keys to world genera of Microgastrinae were in Nixon (1965) and Mason (1981). Nixon (1965) recognized 19 genera in his key, whereas Mason (1981) included 50 genera (although Mason’s paper started with a key to tribes, and then genera within each tribe are keyed out and treated separately). Some regional generic keys have been published since, e.g., Tobias (1986) for the former Soviet Union, Austin and Dangerfield (1992) for the Australasian region, Whitfield (1997) for the New World, Chen and Song (2004) for China, and Kotenko (2007a) for the Russian Far East. However, with 81 genera considered in this paper, the information to recognize them in the aforementioned references is clearly outdated, and an updated key to world genera is badly needed.

Unfortunately, we still lack a robust phylogeny for the subfamily, which would be needed to provide a useful and comprehensive key. The limits of some genera at present are not well defined, and at times are contradictory; moreover, it is likely that future work will change many groups as currently understood. We anticipate that a few genera will end up as synonyms while several others, which are paraphyletic or polyphyletic as currently defined, will be split. This should likely result in an overall increase in the total number of genera as compared to present (e.g., see Fernandez-Triana and Boudreault 2018).

We divide the 81 genera recognized in this paper into four groups and characterize each group and singular genus with brief morphological diagnoses. We emphasize that these groups are not to be considered as monophyletic, and we caution that the discussion below is not to be taken as a new phylogeny for the subfamily, which is beyond the scope of the present paper. We do not present the information below as a surrogate key either; to key out Microgastrinae genera the reader is advised to initially consider the works mentioned at the beginning of this section. Our only intention here is to provide the reader with some basic information on the concepts we have followed when making decisions about generic placement of species, especially in the new combinations we propose in the checklist below. Besides comments on morphological diagnoses, we also provide illustrations for every Microgastrinae genus (at least one species per genus, usually more), the first time that has been done for the entire subfamily.

We separate Microgastrinae into four broadly defined groups:

a) unplaced genera, all of which have unique morphological characters that make them very distinctive, although they do not share any character in common per se, comprising 18 genera: Austinicotesia, Austrocotesia, Beyarslania, Billmasonius, Clarkinella, Exulonyx, Fornicia, Janhalacaste, Kiwigaster, Mariapanteles, Miropotes, Neoclarkinella, Pelicope, Prasmodon, Qrocodiledundee, Semionis, Xanthomicrogaster, and Zachterbergius;

b) Microplitis group, which includes the Microplitini (sensu Mason 1981) and four additional genera described by Fernandez-Triana and Boudreault (2018), for a total of eight genera: Alloplitis, Gilbertnixonius, Jenopappius, Microplitis, Philoplitis, Silvaspinosus, Snellenius, and Tobleronius;

c) Cotesia group, which includes most but not all of the Cotesiini (sensu Mason 1981), with 29 genera: Buluka, Carlmuesebeckius, Chaoa, Cotesia, Cuneogaster, Deuterixys, Diolcogaster, Distatrix, Eripnopelta, Exix, Glyptapanteles, Jimwhitfieldius, Keylimepie, Larissimus, Lathrapanteles, Markshawius, Nyereria, Ohenri, Parenion, Protapanteles, Protomicroplitis, Pseudofornicia, Pseudovenanides, Rasivalva, Sathon, Ungunicus, Venanides, Venanus, and Wilkinsonellus;

d) Apanteles group, which includes most but not all of the Apantelini + Microgastrini (sensu Mason 1981) with 26 genera: Agupta, Alphomelon, Apanteles, Choeras, Dasylagon, Dodogaster, Dolichogenidea, Exoryza, Hygroplitis, Hypomicrogaster, Iconella, Illidops, Kotenkosius, Microgaster, Napamus, Papanteles, Parapanteles, Paroplitis, Pholetesor, Promicrogaster, Pseudapanteles, Rhygoplitis, Sendaphne, Shireplitis, Xanthapanteles, and Ypsilonigaster.

a) Unplaced genera

Kiwigaster (Figs 136137) is the only genus of Microgastrinae with sexual dimorphism in the number of antennal segments; females have 17 flagellomeres and males have 18 (Fernandez-Triana et al. 2011). All other known microgastrines have 16 flagellomeres in both sexes.

Only five genera of Microgastrinae, Austinicotesia, Austrocotesia, Miropotes, Pelicope, and Semionis, have hind wings without vein 2r-m (all other known Microgastrinae have that vein present, although often weakly pigmented).

Pelicope and Semionis can be recognized within this group because both have the fore wing areolet very large (while the other three genera are without an areolet or have a very small areolet). Pelicope (Fig. 181) has the propodeum unsculptured, notauli at least partially marked, and eyes in frontal view slightly divergent ventrally. Semionis (Figs 221, 222) has the propodeum with a partial transverse carina and many fine striations radiating from the nucha, the notauli not marked, and the eyes in frontal view are not divergent ventrally (Nixon 1965, Mason 1981).

Miropotes (Figs 157159) differs from the other genera by the ovipositor sheaths and ovipositor with a unique shape, in most species strongly bent; eyes enlarged and strongly convergent with malar space totally or almost totally obliterated; metacoxa small and metatibial spurs very short (Fernandez-Triana et al. 2014d).

Austinicotesia (Figs 27, 28) and Austrocotesia (Figs 2932) are similar to each other in several features (Austin and Dangerfield 1992, Fernandez-Triana and Boudreault 2018) but differ as follows: Austinicotesia has the fore wing without areolet (with areolet in Austrocotesia); fore wing with pterostigma relatively thin and long, 3.5 × as long as wide (pterostigma much less than 3.0 × as long as wide in Austrocotesia); fore wing vein 2RS much longer, ca. 1.5 ×, than vein r (fore wing vein 2RS much shorter, ca. 0.5 ×, than vein r in Austrocotesia); metafemur relatively thick and stout (of more normal proportions in Austrocotesia); T1 widening towards posterior margin and with strong hump followed by deeply excavated area and strong carinae (T1 more or less parallel-sided or narrowing towards posterior margin and without hump or excavate area in Austrocotesia); and T2 mostly smooth (usually mostly sculptured in Austrocotesia).

Only six genera of Microgastrinae have the propodeum mostly smooth except for complete longitudinal and transverse carinae: Beyarslania, Clarkinella, Janhalacaste, Neoclarkinella, Mariapanteles, and Prasmodon. We place them together because of the diagnostic value of that unique carination pattern, but it is clear that these genera do not constitute a monophyletic group.

Prasmodon (Figs 191193) is the only genus in this subgroup with notauli strongly marked and fore wing areolet relatively large (Fernandez-Triana et al. 2014f).

Clarkinella and Janhalacaste also have a fore wing areolet (although very small, almost obliterated) and can be distinguished from each other as follows. Clarkinella (Figs 46, 47) has the scutellar disc with a smooth posteromedian band, T1 without a median longitudinal carina, and hypopygium mostly inflexible with only a sharp fold posteriorly (Mason 1981), whereas Janhalacaste (Figs 128, 129) has the scutellar disc with a coarse posteromedian band, T1 with a longitudinal sulcus on the anterior 0.6–0.7 of its length and posterior 0.3 with two short carinae centrally delimiting a slightly raised area, and hypopygium folded medially and with several pleats (Fernandez-Triana and Boudreault 2018).

Neoclarkinella (Figs 161165), Mariapanteles, and Beyarslania all lack a fore wing areolet. Neoclarkinella can be recognized because it has a very distinctive T1 which sharply narrows towards posterior margin and has a wide depression on the anterior half, and a hypopygium with multiple pleats (Chen and Song 2004, Veena et al. 2014).

Mariapanteles and Beyarslania have the hypopygium mostly inflexible, with a posteromedian translucent fold where only a few or no pleats are visible; and T1 has a sharply defined median, longitudinal sulcus, at least on the anterior half. Mariapanteles (Figs 143, 144) has the ovipositor sheaths much longer (0.7 × as long as the metatibia length), and ovipositor mostly straight to slightly curved (Whitfield et al. 2012), whereas Beyarslania (Fig. 33) has the ovipositor sheaths relatively very short (less than 0.3 × metatibial length), and the ovipositor strongly downcurved (Mason 1981, at the time referring to the genus as Xenogaster). Mariapanteles is also the only genus in this group with the propodeum having some additional, small and short transverse carinae that radiate from the median carina (but, nevertheless, the propodeum still appears as if it is crossed by the median and transverse carinae, the defining trait of this group).

The remaining six genera in this group cannot easily be associated with any other genus and are discussed below in alphabetical order.

Billmasonius (Fig. 34) is recognized by T1 with a unique shape and desclerotization, with a relatively wide anterior 0.6 and very narrow posterior 0.4, so that widest part of tergite, near anterior margin, is around 4.0 × the narrowest width, along posterior 0.4, and with anterior 0.6 mostly desclerotized, only with lateral margins and narrow central strip sclerotized; T2 is also diagnostic, with a partially sclerotized area surrounding each spiracle on laterotergite 2 the same colour as T2, giving the impression of T2 having three peaks, the largest and central one being the actual T2, the two smaller lateral ones being the area surrounding the spiracles on each laterotergite (Fernandez-Triana and Boudreault 2018).

Exulonyx (Fig. 95) has a unique combination of features within Microgastrinae: propodeum with a partial median, longitudinal carina on anterior 0.6 and complete areola on posterior 0.4, hypopygium inflexible, ovipositor curving downwards on posterior 0.3, and T1 and T2 coarsely sculptured (Mason 1981).

Fornicia (Figs 9698) is the only Microgastrinae genus with the epicnemial carina complete and the fore wing areolet absent; also, the head in lateral view is relatively small (compared to the mesosoma) (Austin and Dangerfield 1992), and T1–T3 form a carapace covering the entire dorsal surface of the metasoma. Only a few species in the Microplitis group (see below) have a partial to complete epicnemial carina, but all those genera have the fore wing with an areolet (usually relatively large), and the head of normal proportions.

Qrocodiledundee (Fig. 212) can easily be recognized by its propodeal apophysis, unique among Microgastrinae, as well as the flattened mesosoma, metafemur short and stout, pronotum dorsally enlarged, and the propodeum with a median carina and a partially defined areola (Fernandez-Triana and Boudreault 2018).

Xanthomicrogaster (Figs 246249) is unique because of the following combination of features: hind wing with vein 1cu-a strongly sinuous and first submarginal cell tall (height at least 2 × its width), fore wing with a very small areolet, metacoxa very large (almost as large as the metasoma length), propodeum mostly smooth but with a strong and sharp median longitudinal carina, T1 very wide and with a strong median longitudinal sulcus, T2 rectangular and usually sculptured, hypopygium inflexible, and ovipositor sheaths relatively long (more than 0.5 × metatibia length) and with numerous setae. Some of these morphological features would suggest this genus could be placed within the Cotesia group, contrary to Mason’s (1981) opinion when he grouped it within his Microgastrini. However, Xanthomicrogaster has many other features that are so different to both our Cotesia group and Mason’s Microgastrini that we prefer to maintain it as an unplaced genus.

Zachterbergius (Figs 253, 254) has the longest and thinnest T2 among all known Microgastrinae, with T2 length 4.0 × its width at base and apex, 0.7–0.8 × as long as T1 length, and around 1.5× as long as T3 length. Also, the propodeum has a clearly defined median carina, partially defined transverse carina, and the posterior part of an areola; the antennal scape is very transverse, and the labial palpi are very long, extending to the mesopleuron (Fernandez-Triana and Boudreault 2018).

b) Microplitis group

This is one of the best-defined groups of genera within Microgastrinae (see Mason 1918), and most likely to be monophyletic. It is characterized by: tentorial pits relatively large, head mostly coarsely sculptured, stemmaticum usually very well defined and slightly to strongly raised from the surrounding areas, anteromesoscutum and scutellar disc usually coarsely sculptured, notauli almost always defined (often very clearly), propodeum always sculptured and with several strongly defined carinae, fore wing with areolet usually large, metacoxa relatively small, metatibial spurs short, T1 with median longitudinal sulcus, hypopygium inflexible and almost always relatively short, ovipositor sheaths with few setae that are mostly limited to the apex, and ovipositor almost always very short (much shorter than 0.5 × metatibia length).

Philoplitis (Figs 182, 183) has a unique combination of features including an enormous scutellum conically prolonged posteriorly over the propodeum (Mason 1981, Fernandez-Triana and Goulet 2009, Ranjith et al. 2019). It also has an occipital carina, and ocelli forming a very low triangle, to the point that the anterior ocellus seems almost on the same line as the posterior ones.

Silvaspinosus (Fig. 227) has the clypeus extremely long and thin, the malar line extremely short (almost absent), the mandible base separated from the rest of the head by a desclerotized area that looks almost like an opening, and mandibles relatively stout and large. The shape of the clypeus, and the separation of the mandible from the rest of the head by a desclerotized area are unique among Microgastrinae (Fernandez-Triana and Boudreault 2018). It also has the fore tarsus with a spine-like seta, and the scutellar disc with the posteromedian band smooth; both of which are unique and distinctive among the Microplitis group.

Gilbertnixonius (Fig. 99), is the only genus in this group that has the propodeum with both longitudinal and transverse carinae but without an areola (Alloplitis and Tobleronius have those carinae, although sometimes incomplete, but they also have a complete areola on the propodeum). Gilbertnixonius also has an epicnemial carina (otherwise only present in some species of Snellenius and in all species of the unrelated genus Fornicia) and an incomplete occipital carina (otherwise only present in Alloplitis, Philoplitis, and Tobleronius) (Fernandez-Triana and Boudreault 2018).

Alloplitis and Tobleronius are somewhat similar morphologically and distinguished from the other six genera in this group by the propodeum with a complete areola (in addition to partial longitudinal and transverse carinae). Alloplitis (Figs 7, 8) has T1 more or less parallel-sided or slightly widening towards the posterior margin, and T2 more or less rectangular; whereas Tobleronius (Fig. 233) has T1 strongly narrowing towards the posterior margin (width at posterior margin 0.3 × or less of width at anterior margin) and T2 very long and thin (although slightly widening towards the posterior margin) and with the area surrounding the spiracles on laterotergite 2 partially sclerotized and the same colour as T2 giving the impression of T2 having three peaks, the largest and central one being the actual T2, the two smaller lateral ones being the area surrounding the spiracle on each laterotergite (Fernandez-Triana and Boudreault 2018).

Microplitis (Figs 151156) and Snellenius (Figs 228232) are very similar and form one of the most morphologically distinct groups of Microgastrinae (Nixon 1965, Mason 1981, Walker et al. 1990, Shaw and Huddleston 1991, Austin and Dangerfield 1993, Fernandez-Triana et al. 2015b) with the following shared diagnostic features: propodeum with coarse sculpturing and a strong median carina and T2 and T3 with a poorly defined separation between them. Most species of Snellenius are easily distinguished by having the notauli and the scutellar disc strongly excavated and sculptured, and by having the scutoscutellar sulcus very wide and deep; both cases represent the most extreme examples within Microgastrinae. Additionally, the propodeum is divided into two distinct areas (faces) clearly marked by a strong angulation (observed in lateral view) and a transverse carina (observed in dorsal view). The main difficulty when trying to distinguish both genera is that those features appear to grade, from strongly excavated and sculptured notauli and scutellar disc (most Snellenius) to less excavated and less sculptured (a few Snellenius, most Microplitis), to basically smooth and unexcavated (some Microplitis). The only reliable feature to separate the two genera is the presence of an epicnemial carina in Snellenius, which is absent in Microplitis (Mason 1981, Austin and Dangerfield 1992, 1993, Fernandez-Triana et al. 2015b), although in practice it may be difficult to distinguish the epicnemial carina due to setae and/or sculpture on the epicnemium and mesopleuron.

Jenopappius (Figs 130131) resembles Microplitis but with T2 strongly sculptured and rectangular, and T1 mostly sculptured and with a median depression anteriorly. Some Alloplitis may also have a somewhat similar sculpture on either T1 or T2 but the shape of those tergites is very different, and Alloplitis always has the propodeum with a complete areola, defined by strongly raised carinae. The combination of the sculptured propodeum without an areola, T1 with an anteromedian depression, and T1 and T2 with strong sculpture are very unusual and will separate Jenopappius from any other genus of Microgastrinae (Fernandez-Triana and Boudreault 2018).

c) Cotesia group

We place here genera with a completely inflexible hypopygium, ovipositor sheaths relatively short (less than 0.5 × metatibial length, usually much less) and mostly without setae (except apically in some cases). Most of the 29 genera considered here also have the propodeum without a complete areola (although some have it, and others have a complex arrangement of carinae and sculpture where a partial to complete areola can sometimes be defined). Although these features work well to recognize most members of the group, a few species of Sathon, Lathrapanteles, Glyptapanteles, and Ohenri have relatively long ovipositor sheaths, but in these cases the hypopygium is still always inflexible. Most or perhaps all the species within the Cotesia group posses a suite of characters indicative of parasitism of “macrolepidoptera” (sensu Mason 1981: 25), but the group is probably not monophyletic. From the Cotesiini (sensu Mason 1981) we exclude here Parapanteles and instead transfer it to the Apanteles group (see details under that group); the main reason being that this genus, as it had been understood, apparently includes two different sets of taxa: one that seems to be Cotesia species misidentified as Parapanteles (Valerio et al. 2009, Parks 2018, Freitas et al. 2019), and another (representing the majority of the genus, as currently understood, including the type species) that are more related to Dolichogenidea and Apanteles than to any genus in the Cotesia group. We also add here Sathon, which we consider to be closer to Glyptapanteles and related genera, unlike Mason (1981), who considered it to be part of his Microgastrini group.

The Cotesia group can be broadly split into two subgroups, based on whether the fore wing has an areolet (Buluka, Cuneogaster, Diolcogaster, Eripnopelta, Exix, Jimwhitfieldius, Keylimepie, Larissimus, Markshawius, Parenion, Protomicroplitis, Rasivalva, Ungunicus, Venanus) or does not have an areolet (Carlmuesebeckius, Chaoa, Cotesia, Deuterixys, Distatrix, Glyptapanteles, Lathrapanteles, Nyereria, Ohenri, Protapanteles, Pseudofornicia, Pseudovenanides, Sathon, Venanides, Wilkinsonellus).

Among the genera with a fore wing areolet, Jimwhitfieldius (Figs 132, 133) has the metatrochantellus with a unique shape (Fig. 133), the head with a strong depression behind the occiput, the metatibia with a very long and thick inner spur, and the ovipositor and ovipositor sheaths extremely short, probably the shortest in the entire subfamily (Fernandez-Triana and Boudreault 2018).

Venanus (Figs 237240) is quite distinctive, and comprises small species, often with the body slightly depressed, face with a triangular flange between the antennal sockets, fore wing with a relatively large areolet, T2 with strongly defined lateral sulci, and ovipositor sheaths with very few and minute setae (Mason 1981).

The remaining genera in the subgroup seem to share one or several morphological features with Diolcogaster (whether those features are homoplastic or not). Diolcogaster (Figs 6677), as currently understood, is most likely a polyphyletic genus that will need to be split into several genera. Until then, it is difficult to define unequivocally. Instead, we discuss the remaining genera in this subgroup in alphabetical order, with the features that distinguish them from Diolcogaster.

Buluka (Figs 3537) has T1–T3 forming a carapace and occupying the entire dorsal surface of the metasoma, the fore wing has a complete areolet, and females have part of the ventral surface of the distal six or seven flagellomeres without longitudinal placodes, instead having an oblique groove bounded on one side by a row of bent-tipped sensilla (Austin 1989). The carapace is shared with Fornicia and very few species of other genera, e.g., Deuterixys, Pholetesor, none of which have a fore wing areolet. The basimacula species group of Diolcogaster (sensu Saeed et al. 1999) have both the carapace and areolet, but the antenna does not have the special groove and sensilla.

Cuneogaster (Figs 61, 62) resembles Diolcogaster but it has the glossa long and apically bilobed, T1 wedge-shaped, and the scutellar disc with the medioposterior band smooth (Choi and Whitfield 2006) whereas in Diolcogaster the glossa is not elongated, T1 is usually not wedge-shaped, and the scutellar disc has a medioposterior band of rugosity in most species.

Eripnopelta (Figs 87, 88) could be considered an atypical Diolcogaster, but the pronotal lateral surface does not have distinct furrows, the scutellar disc has a smooth and protruding medioposterior band, T1 does not have a distinct median groove on the basal half, and the fore wing areolet is very small, almost obliterated (Xiong et al. 2017).

Exix (Figs 89, 90) also seems morphologically related to Diolcogaster, but it is defined by T2 large and smooth, without submedian grooves, the hind wing has the vannal lobe concave and lacking setae, and the hind wing nervellus is externally concave (Mason 1981).

Keylimepie (Figs 134, 135) can be recognized by the reduced wings in females, relatively small eyes and long malar space. The shape and sculpture of the head, mesosoma sculpture, shape and sculpture of T2, and ovipositor are all similar to some Diolcogaster, but Keylimepie has a T1 without a median sulcus and instead it has the anterior 0.5 rather depressed and concave, and the posterior 0.5 with strong transversal striations (Fernandez-Triana and Boudreault 2016).

Larissimus (Figs 139, 140) is another genus related to Diolcogaster but it can be recognized by the greatly reduced vannal lobe in the hind wing with, almost entirely smooth body, and the only described species is the largest known species of Microgastrinae, with a body and fore wing length of 7–8 mm (Nixon 1965, Mason 1981).

Markshawius (Figs 145, 146) has a unique set of features (Fernandez-Triana and Boudreault 2018) which together are very distinctive (although some, but not all, are shared with other genera). The female head is elongated and strongly concave posteriorly, modified to be tightly appressed to the anterior margin of the pronotum (following its contour); the face has its upper margin produced dorsally between the antennal insertions into a triangular flange; the frons is very elongated, with ocelli clearly much higher than normal; the antenna is very short (much shorter than body length, usually shorter than the combined length of the head and mesosoma), with all flagellomeres except the first having a single row of placodes; the propodeum has a median carina (defined posteriorly) and transverse rugosity which includes a poorly and partially defined transverse carina; and T1 is either extremely long and thin, with length at least 6.0× its width centrally, or very thin on the anterior 0.3–0.4, then strongly widening posteriorly, its width at the posterior margin around 3.0 × its width centrally.

Parenion (Figs 176, 177) can only be confused with some Diolcogaster, but is distinguished by having T2 and T3 smooth and barely or not separated, scutellar disc with the medioposterior band smooth and very small lunules on its lateral surface (Mason 1981).

Protomicroplitis (Figs 201, 202) is closely related to Diolcogaster, both morphologically and molecularly, and some of the criteria used to define it may need revision. The genus is defined by some flagellomeres having three rows of placodes, relatively large fore wing areolet, and T1 very long and narrow (Mason 1981, Fernandez-Triana 2015), although the last two features are also present in a few Diolcogaster species.

Rasivalva (Figs 213, 214) is characterized by the ovipositor sheaths lacking setae, or with very few and minute setae (Mason 1981, Chen and Song 2004, Kotenko 2007b). This separates it from Diolcogaster, which has relatively long setae on the ovipositor sheaths, including a few strong and thickened setae in many species. Other distinguishing features that appear in some species are the scutellar disc with the medioposterior band smooth, body sculpture smoother overall than in Diolcogaster, and propodeum with a median, longitudinal carina that is sometimes reduced or absent.

Ungunicus (Fig. 234) has remarkable and very distinctive tarsal claws, with a very large basal tooth longer than the apex of the tarsal claw, and a median lobe with setae arising from its margin, which seems slightly bilobate. These claws are unique within Microgastrinae (Fernandez-Triana and Boudreault 2018).

Among the genera without the fore wing areolet, Chaoa (Fig. 39) was described from a single specimen (Luo et al. 2004), with little information provided. Based on the original description and illustrations of the holotype, this genus might just represent a species of Glyptapanteles, or perhaps Nyereria but without examining the type we cannot conclude and therefore retain it as a valid genus for the time being.

Carlmuesebeckius (Fig. 38) has the ovipositor and ovipositor sheaths relatively long, and the propodeum with a complete areola, unlike most other genera in this subgroup. Other unique features are T1 with a strong and raised median carina for most of its length, and the ovipositor bulging near apex and with two subapical serrate teeth on the lower (first) valvulae (Fernandez-Triana and Boudreault 2018).

Cotesia (Figs 4860) is a relatively uniform genus morphologically, long considered the easiest group to recognize among all segregates from Apanteles sensu lato (Mason 1981: 113). Defining characters are: fore wing without areolet; T1 and T2 usually mostly to entirely sculptured, T3 also often at least partially sculptured or, more rarely, completely sculptured; T1 either widening towards its posterior margin (very often), more or less parallel-sided or barrel-shaped (often), slightly widening towards the posterior 0.7–0.8 of the tergite length and from that point slightly narrowing towards the posterior margin which is more or less rounded (rarely), or medially constricted (extremely rare), but never completely narrowing towards the posterior margin; ovipositor and ovipositor sheaths are very short to short, very rarely moderately long. The propodeum varies considerably but has a well defined median longitudinal carina (very often), although the median carina may be difficult to distinguish on its own in species with the propodeum strongly sculptured with an irregular pattern of carinae (often), or the median carina may be partially absent (rarely), or the median carina may be combined with a partial to complete areola partially defined by a transverse carina (rarely), or the median carina is absent and/or the propodeal surface is shiny overall and almost without any sculpture (rarely). The only other genus that could be confused here would be Protapanteles, which may eventually be considered as just a species group within Cotesia, with smoother propodeum and T1–T3.

Protapanteles (Figs 198200) usually has T1 either slightly widening towards the posterior 0.7–0.8 of the tergite length and then slightly narrowing towards the posterior margin which is more or less rounded (often), more or less parallel-sided or barrel-shaped (rarely), or slightly widening towards the posterior margin (rarely). The propodeum is variously sculptured, usually having a median longitudinal carina that may be partially or completely defined, and rarely lacking the median carina. A character commonly used to define this genus, a modified spine on the fore tarsus (Nixon 1965, 1972, 1973, 1976, Mason 1981), is present in some species of many related genera, e.g., Cotesia, Glyptapanteles, Distatrix, Nyereria, and even in some non-related genera such as Silvaspinosus, and thus does not have the same diagnostic value as expressed by Mason (1981). Some species may be considered as borderline between Cotesia and Protapanteles, and others may be considered as borderline between Glyptapanteles and Protapanteles; thus, it is difficult to clearly define these three genera. Differences between Protapanteles and Cotesia were given in the previous paragraph. Differences with Glyptapanteles are mostly related to the shape of T1. In Glyptapanteles, T1 is either parallel-sided anteriorly and then strongly narrowing posteriorly, or its sides are gradually to strongly converging posteriorly when compared to Protapanteles which has T1 parallel-sided throughout, except for a strongly rounded apex, and propodeum sculpture that is usually, but not always, more rugose and carinated than in Glyptapanteles. Additionally, Protapanteles larvae have mandibles with a row of 12 or fewer large teeth concentrated distally on the blade, and its species distribution is almost completely confined to the Holarctic region (Mason 1981). However, the morphological features mentioned above vary considerably among different species (Arias-Penna et al. 2019).

Glyptapanteles (Figs 100110) is most likely a polyphyletic assemblage, and may eventually be split into several genera. As a result, it is difficult to define (Arias-Penna et al. 2019). Some of its species may be confused with Protapanteles, Sathon, Lathrapanteles and, to a lesser extent, also Distatrix, Venanides, and Nyereria. The main features defining Glyptapanteles are: fore wing without an areolet; propodeum that is either completely smooth (often) to more or less rugose (more rarely), with a median longitudinal carina that is entirely absent (often), partially defined posteriorly (often) to complete and strong (rarely), or no median carina but instead a series of very short carinae radiating from the nucha (rarely); T1 narrows towards the posterior margin, usually strongly (almost always), or more parallel-sided, or rounded at apex, as in some species of Protapanteles (rarely); T2 is almost always subtriangular or trapezoidal (rarely shaped differently); ovipositor and ovipositor sheaths are relatively short (usually) to moderately long (rarely); setae at apex of ovipositor sheaths relatively long (as long or longer than setae on hypopygium). The differences from Protapanteles were given in the previous paragraph. Sathon has the ovipositor sheaths longer and male specimens have enlarged external genitalia; however, a few Glyptapanteles species have females with longer ovipositor sheaths, and a very few other species have males with external genitalia similarly enlarged; whether those species should be transferred to Sathon requires further study. Lathrapanteles has similar characters to Sathon (see more about those two genera below) and can be separated in the same manner from Glyptapanteles. Distatrix has the pronotum with only one furrow laterally, eyes enlarged and ovipositor sheaths without setae or with very few minute setae, whereas Glyptapanteles has the pronotum with two furrows, eyes that are almost never enlarged (but see Fernandez-Triana 2018, for one exception) and the ovipositor sheaths have much longer setae. Venanides can in turn be separated from Glyptapanteles based on having similar ovipositor sheaths to Distatrix (Mason 1981).

Distatrix (Figs 78, 79) is similar to Venanides, but it has two rows of placodes in the flagellomeres in females, and T2 has a characteristic shape, with the lateral margins widely diverging (Mason 1981, Grinter et al. 2009).

Venanides (Figs 235, 236) can be differentiated from Distatrix because it has only a single row of placodes in the flagellomeres in females, and T2 has less diverging lateral margins (Mason 1981). Additionally, Venanides specimens tend to be smaller and have a dorsoventrally compressed body that is also generally mostly smooth and shiny.

Sathon (Figs 218220) is distinguished mainly by the enlarged external genitalia in males and relatively long ovipositor sheaths in females; some species probably have the longest sheaths among the entire Cotesiini (sensu Mason 1981). However, these features are not unique: a few Glyptapanteles species have similarly enlarged male genitalia, and all described Lathrapanteles species (Figs 141, 142) are also very similar to Sathon (e.g., Williams 1985, 1988). The limits of Lathrapanteles and Sathon need revision and it is possible that one will eventually be placed in synonymy with the other.

Deuterixys (Figs 64, 65) is a very distinctive genus on account of its T1–T3 sculpture and shape (there appears to be a second constriction between T2 and T3), the propodeum being smooth and shiny and with a complete and strong median, longitudinal carina, and the relatively small body length (Mason 1981, Whitfield 1985, Zeng et al. 2011).

Nyereria (Figs 166169) has T2 divided into three sections by two deep, usually crenulated, longitudinal grooves delimiting a raised, median area that is not wider than long (Mason 1981). This genus can only be confused with a few species of Cotesia and Glyptapanteles that have their T2 with a similar raised, median area, although in those cases T2 is never as strongly defined by grooves.

Pseudovenanides (Fig. 211) has very scarce information available, but from the original description (Xiao and You 2002) it is clear that it is related to Glyptapanteles and, to a lesser extent, to Venanides. Apparently, T1 with a strongly marked longitudinal sulcus on most of the tergite is the defining feature of this genus.

Ohenri (Fig. 170) has many unique features and is only tentatively considered to be part of this subgroup lacking the fore wing areolet. The pronotum is considerably enlarged dorsally, the ovipositor has its lower valvulae with four subapical teeth, the tarsal claws have large teeth, and the propodeum has a median carina with a partially defined areola (Fernandez-Triana and Boudreault 2018).

Pseudofornicia (Figs 208210) superficially resembles the (probably) unrelated Fornicia because its metasoma mostly forms a dorsal carapace, but it differs in lacking the epicnemial carina, the fore wing does not have an areolet, and T1 is movably joined to T2, whereas Fornicia has an epicnemial carina, fore wing with an areolet, and T1 and T2 are immovably joined (van Achterberg et al. 2015).

Wilkinsonellus (Figs 241244) is a very recognizable genus, with T1 very long and thin, propodeum with distinctive sculpture and carination pattern, and fore wing with veins r and 2RS strongly angled (Mason 1981, Long & van Achterberg 2011, Arias-Penna et al. 2013, 2014).

d) Apanteles group

Mason (1981) proposed the tribes Apantelini and Microgastrini to accommodate species with ovipositor sheaths mostly setose and relatively long (at least 0.5 × metatibial length), hypopygium with ventral margin usually flexible and either with one (rarely) or several (commonly) pleats. The latter is the most diagnostic feature for this group; however, there are exceptions (all Alphomelon, most Hygroplitis, and a few species of Apanteles and Microgaster) where the hypopygium is mostly to entirely inflexible. In this paper we combine most of the genera included in the two tribes into a single Apanteles group composed of 26 genera. The group is clearly not monophyletic. Most, if not all, of the species included here have the “microlepidoptera suite of characters” sensu Mason (see further discussion in Mason 1981, Walker et al. 1990). Here we separate the group into several subgroups that can be recognized on simple morphological features, although the genera included in each subgroup are not necessarily related.

The largest subgroup includes 13 genera that lack a fore wing areolet: Alphomelon, Apanteles, Dolichogenidea, Exoryza, Iconella, Illidops, Napamus, Parapanteles, Pholetesor, Pseudapanteles, Rhygoplitis, Shireplitis, and Xanthapanteles. Another two genera could be placed here, at least partially: some species of Choeras lack a fore wing areolet; however, most of the species have a complete or partial areolet so we consider Choeras to be better placed with the subgroup of genera with a complete or partial fore wing areolet; and a similar situation occurs with Promicrogaster, where smaller species tend to lack the areolet whereas the larger species have a complete areolet, and we similarly place that genus in the subgroup with an areolet. These two genera exemplify the challenges of delimiting precise groups in Microgastrinae (a frustration also shared by Mason 1981: 77).

Among the genera without a fore wing areolet, four have the propodeum either with a median longitudinal carina (Iconella, Pseudapanteles, Rhygoplitis) or with a complex pattern that includes full sculpturing and a series of short carinae radiating medially on the posterior 0.2–0.3 near the nucha (Illidops). A fifth genus, Napamus, could also be included in this subgroup, as one of its two described species has the propodeum with a median, longitudinal carina; however, the other species does not (Papp 1993: 170). Nevertheless, Napamus (Fig. 160) can be characterized by its mouth parts elongate, fore wing vein R1 very short (shorter than pterostigma length), inner metatibial spur much longer (1.3 ×) than the outer spur, body and legs black, and wings strongly infumate.

Iconella (Figs 122124) was described by Mason (1981) as a new genus based on the hind wing with a sinuous vein cu-a as a plesiomorphic character that suggests its unique status among similar genera. Fernandez-Triana et al. (2013a, 2014e) also considered the presence of a median longitudinal carina on the propodeum as strong support for its generic status. However, some Oriental species (with large body size and large and bilobate glossae) currently assigned to Iconella may eventually be placed in a different genus.

Illidops (Figs 125127) includes species that have the scutellar disc with a medioposterior band of rugosity, fore wing vein R1 shortened, and the propodeum with a series of short carinae medially on its posterior 0.2–0.3, near the nucha (Fernandez-Triana et al. 2014e). In some, but not all species the lower margins of the eyes converge, and T3–T7 are weakly sclerotized (Mason 1981).

Pseudapanteles (Figs 203207) is characterized by the glossa elongate and strongly bilobed apically, propodeum with a strongly defined median longitudinal carina but no transverse carina (traces of a transverse carina are very rarely present in a few Neotropical species), and T1 with a sharp median sulcus (Mason 1981, Whitfield 1997, Fernandez-Triana et al. 2014a).

Rhygoplitis (Figs 215217) is the only genus in this subgroup with notauli relatively well defined. It also has the propodeum coarsely sculptured (in addition to a median, longitudinal carina), and fore wing with very short vein R1 (Mason 1981, Whitfield 1997).

The other eight genera without a fore wing areolet have the propodeum with a complete to partial areola, although in large genera such as Apanteles, Dolichogenidea, and Pholetesor, some species have lost all carinae and the propodeum is mostly smooth.

Shireplitis (Figs 225, 226) has the propodeum entirely sculptured, without median or transverse carina, but with the areola defined on the posterior 0.5 by two lateral carinae, ovipositor sheaths relatively short (0.4–0.5 × metatibia length), and legs short and robust – with the metafemur usually less than 3.0 × as long as wide (Fernandez-Triana et al. 2013b).

Alphomelon (Figs 911) has the gena with a white/pale spot that is relatively large and very distinctive (Mason 1981, Deans et al. 2003). A few other Microgastrinae genera have some species with a similar pale spot, but it is usually much smaller. Alphomelon is distinguished from the other Microgastrinae with white/pale spot on gena by its ovipositor sheaths being relatively long (much shorter in Cotesia, Glyptapanteles, Protapanteles), mesoscutum anteriorly without strong notauli (strong notauli in Prasmodon), propodeum without a median, longitudinal carina (strong median, longitudinal carina in Pseudapanteles), and the hypopygium inflexible and unpleated (almost always flexible and with several pleats in Apanteles).

Apanteles (Figs 1226) is currently the most speciose genus in Microgastrinae and has some morphological variability. It usually has the propodeum fully to partially areolated, rarely smooth and never with a median longitudinal carina; fore wing without an areolet; hind wing with the vannal lobe usually strongly concave or straight (see next paragraph for more details on that); ovipositor sheaths relatively long; and the hypopygium almost always flexible and pleated. This genus could only be confused with Pholetesor or Dolichogenidea (which seem to be related to one another, see below) and Parapanteles. Most Apanteles species can be distinguished from both Pholetesor and Parapanteles by the flexible, pleated hypopygium and relatively long ovipositor sheaths (usually at least 0.5 × length of metatibia). In contrast, Parapanteles and Pholetesor have the hypopygium either entirely inflexible or at most with a small, translucent area near the posterior margin (which may look like a pleat in a few species); and the ovipositor sheaths are relatively short (less than half the metatibia length, usually much less). However, a few species of Apanteles have relatively short ovipositor sheaths, and very few species may even have an inflexible hypopygium (e.g., Fernandez-Triana et al. 2014e); the generic placement of those species may be revisited, but at present those exceptions make for a more difficult separation of these three genera.

Dolichogenidea (Figs 8186) is even more difficult to distinguish from Apanteles, as there is some overlap in some species groups of both genera (e.g., Mason 1981: 53, 54). The differences are frequently subtle and, at times it is very difficult to assign a species to one or other genus depending on the interpretation of morphological features alone. Apanteles has the hind wing with the vannal lobe usually strongly concave or, more rarely, straight to very slightly convex; the central part of the vannal lobe lacks any setae or has few, sparse setae that are often minute and not continuous. In contrast, Dolichogenidea has the vannal lobe convex to slightly straight; the central part of the vannal lobe is more or less entirely setose so that a continuous fringe of setae is almost always visible (although setae may be small in a few species). The fringe of setae (or lack of them) is the only morphological character that almost always seems to work in distinguishing these genera from each other; we are aware of very few species currently assigned to Dolichogenidea where the fringe is not complete and could lead to the species being placed within Apanteles, despite molecular data strongly suggesting the best generic placement is Dolichogenidea. Other features function only partially and seem to represent trends that are far from being universally present in one genus or the other. For example, the anteromesoscutum punctures (when present) tend to be partially or completely fused near the scutoscutellar sulcus in Apanteles, whereas in Dolichogenidea, which usually does not have punctures on the anteromesoscutum anteriorly and very rarely has them near scutoscutellar sulcus, the punctures never fuse. The scutoscutellar sulcus in many Dolichogenidea species tends to be very narrow and sometimes looks almost obliterated, whereas the sulcus in Apanteles is usually wider. Despite the rather subtle morphological differences, DNA barcodes tend to cluster both genera clearly apart (Smith et al. 2013, Fernandez-Triana et al. 2014e).

Dolichogenidea tends to cluster near Pholetesor (Figs 184190) and these genera seem to be closer to each other than either is to Apanteles. Dolichogenidea has a flexible, pleated hypopygium and relatively long ovipositor sheaths (usually at least 0.5 × metatibia length) whereas Pholetesor has the hypopygium entirely inflexible or with a small, translucent area near the posterior margin that could look like a pleat in a few species, and the ovipositor sheaths are relatively short, less than half the metatibia length (Mason 1981, Whitfield 2006).

The status of Exoryza (Figs 9294) as a valid genus has been questioned by many authors (Valerio et al. 2004, Rousse and Gupta 2013, Fernandez-Triana et al. 2014e, 2016c). Mason (1981) characterized it as having T1 and T2 heavily sculptured, and the propodeum coarsely rugose, with an areola present but obscured by heavy sculpture. However, the distinction between Exoryza and Dolichogenidea may be particularly difficult because many species of the latter genus have the propodeum sculptured, with or without an areola, and T1 is occasionally sculptured, although not as strongly as in Exoryza (Fernandez-Triana et al. 2014e, 2016c).

Parapanteles (Figs 172175) is a very difficult genus to understand at present. Parks (2018) found it to be paraphyletic. Some species of “Parapanteles” with available DNA barcodes cluster within Dolichogenidea and could just be considered as species within that genus, with short ovipositor sheaths and an inflexible hypopygium (similar to Pholetesor and the few borderline species of Apanteles mentioned above). Another group of Parapanteles seems to represent misidentifications of Cotesia (e.g., Valerio et al. 2009, Freitas et al. 2019). Whether a group of species that could be considered true Parapanteles actually exists remains to be seen. For the present, the genus can be defined as having the propodeum completely to mostly areolated (usually with well defined carinae), ovipositor sheaths short, and an inflexible hypopygium.

Xanthapanteles (Fig. 245) is a very distinctive genus, on the basis of the propodeum fully areolated with strongly defined and raised carinae, T1 very large and wide, T1–T3 sculpture like a finely pebble-grained surface (unlike any other Microgastrinae), flagellomere placodes arranged irregularly and fore wing relatively slender and much longer than body length (Whitfield 1995b).

Another subgroup within the Apanteles group includes six genera, Agupta, Dasylagon, Hypomicrogaster, Papanteles, Promicrogaster, and Sendaphne, that can be recognized by the fore wing with a very small areolet, sometimes almost obliterated. They also share (except for Agupta, see below) having the scutellum with lunules relatively high, more than 0.5 × the height of its lateral face. These genera are separated from each other based on different propodeal carination patterns, and T1 and T2 shapes and sculptures. Some described species of Choeras, almost exclusively from the Oriental region, have a very small areolet and thus could be included in this group. However, these are exceptions and are very likely to be transferred elsewhere or classified separately. For now, we place Choeras (see below) within the subgroup with a large fore wing areolet.

Agupta (Figs 5, 6) does not have enlarged lunules; however, it can be recognized by several unusual features: in males (and sometimes in females) the antenna has the first few flagellomeres with placodes irregularly distributed in three rows or no row can be clearly defined; the propodeum has a strongly raised median carina with small radiating carinae across its length; T1 shape (narrowing for first half, then parallel-sided) and T1 sculpture (anterior half mostly smooth, strongly concave and with central sulcus, posterior half punctured and with a polished area on posterior margin) are distinctive; and the body length is among the largest in Microgastrinae (second only to the unrelated genus Larissimus) (Fernandez-Triana and Boudreault 2018). Some large specimens of Choeras in the Oriental region (see previous paragraph) might end being placed within Agupta when more studies are done in the future.

Promicrogaster and Sendaphne can be recognized by the following combination of features: glossa elongate and bilobate, metacoxa very long (0.8–1.0 × metafemur length and 0.6–0.8 × metatibia length), and ovipositor and ovipositor sheaths very long – among the longest in Microgastrinae usually 2.0 × as long as the metatibia or even longer. Most species have the body length longer than the fore wing length, usually by 0.2–0.4 mm (the majority of Microgastrinae species have the fore wing slightly longer than the body length). These two genera are very closely related and may eventually be treated as a single genus. Promicrogaster (Figs 194197) has the ovipositor apically sinuate; propodeum sculptured and usually with some carination (which may include a complete or partial median longitudinal carina, or an indication of a partial areola posteriorly); T1 parallel-sided to slightly narrowing towards the posterior margin; and T2 transverse, its width at the posterior margin 3.0–4.5 × (rarely 2.0 ×) its length medially (Fernandez-Triana et al. 2016b, Fernandez-Triana 2019). Sendaphne (Figs 223, 224) has the ovipositor straight apically, propodeum mostly smooth and without carina (with the rare exception of having sparse punctures and a few rugae near the nucha), T1 strongly narrowing towards the posterior margin, and T2 subtriangular (Fernandez-Triana et al. 2014h).

Dasylagon (Fig. 63) has the propodeum fully areolated (defined by strong carinae), T1 comparatively very wide and large (in dorsal view more than 0.3 of entire metasoma), T2 very transverse, metasomal terga entirely smooth, hind wing with a sinuous vein cu-a, and ovipositor and ovipositor sheaths relatively very long (more than 1.5 × metatibia length) (Mason 1981).

Hypomicrogaster (Figs 113121) has the propodeum with a complex carination pattern, which includes a median carina and a more or less complete areola, although some species have all carinae reduced, but still the propodeum would be mostly sculptured. The head is relatively transverse, i.e., wider than in most other genera of Microgastrinae, and T1 and T2 are mostly to entirely smooth (Mason 1981, Valerio and Whitfield 2015).

Papanteles (Fig. 171) has the propodeum fully areolated, T1 relatively long (ca. 2.0 × its width at posterior margin), T1 and T2 strongly sculptured, T2 and T3 comparatively narrow and not occupying the entire dorsal surface of the segment (dorsal width of T2 and T3 half the width of T5 and following terga), and the ovipositor sheaths are approximately the same length as the metatibia length (Mason 1981).

The remaining eight genera in the Apanteles group all have the fore wing areolet relatively large; even when some species may have a relatively smaller areolet, it never appears almost obliterated.

Ypsilonigaster (Figs 250252) has a very characteristic T1, with a median sulcus shaped like an inverted Y, a unique feature to recognize the genus (Fernandez-Triana and Boudreault 2018).

Hygroplitis and Microgaster have the propodeum with a median carina, fore wing areolet relatively large, anteromesoscutum anteriorly mostly smooth, T1 and T2 heavily sculptured (also T3, partially or entirely), T1 relatively large and wide (width at posterior margin greater than width at anterior margin), and T2 mostly rectangular. The two genera are very closely related and DNA barcodes suggest Hygroplitis may eventually be synonymized under Microgaster. Hygroplitis (Figs 111, 112) has the body somewhat depressed dorsoventrally, notauli more strongly impressed, flagellomeres with three rows of placodes, and the hypopygium usually inflexible although in some cases it is weakly but distinctly pleated (Mason 1981); whereas Microgaster (Figs 147150) does not have the body dorsoventrally depressed, the notauli are barely visible, flagellomeres are usually (but not always) with two rows of placodes, and the hypopygium is usually (but not always) flexible and pleated (Mason 1981).

Paroplitis (Figs 178180) species are relatively small, with a body length of 2.5 mm or less; legs, especially the metafemur, short and robust; antenna short, with flagellomeres in females having only a single row of placodes; hypopygium almost entirely sclerotized but with a sharp fold medially; propodeum rarely entirely sculptured but almost always with a median longitudinal carina, at least on the anterior 0.5, and sometimes also with a complete or partial transverse carina; and T2 usually smooth, rarely sculptured (Mason 1981, Fernandez-Triana et al. 2013b).

Kotenkosius (Fig. 138) has a unique propodeal carination pattern that includes three complete longitudinal carinae, one medially, the other two sublaterally, and a complete transverse carina near posterior 0.6, with additional small striae radiating from the median and sublateral longitudinal carinae, and most carinae strongly defined and raised (Fernandez-Triana and Boudreault 2018).

Choeras (Figs 4045), as presently understood, is clearly a polyphyletic assemblage of species, some of which may eventually be placed in different genera. It is one of the few Microgastrinae genera that has some species without a fore wing areolet (although the shape of the remaining veins r, 2RS, and 2M usually indicate a partially defined areolet), and other species with a complete areolet that can vary from very small in some species to large in others (van Achterberg 2002, Fagan-Jeffries and Austin 2018b). The propodeum also varies, from having a complete longitudinal median carina to having a partial one, to not having any visible median carina, or having just minute carinae radiating from the nucha. T1 is mostly rectangular (slightly narrowing towards the posterior margin in some species), but never much wider on the posterior margin than on the anterior margin, and T2 is mostly transverse. Many Oriental species of “Choeras” most likely represent different lineages from the temperate species and may warrant placement in different genera, e.g., some of the species may be better placed in Agupta.

Dodogaster (Fig. 80) has a unique set of features in the Apanteles group. The propodeum has a more or less complete areola and a partial median carina, the fore wing has a relatively large areolet, and T1–T3 are heavily sculptured and almost form a carapace (Rousse and Gupta 2013).

Diversity and distribution of Microgastrinae genera at world and regional scales

Microgastrinae are present in all continents except Antarctica. Specimens can be found in all major terrestrial ecosystems, from 82°30'N (Canada, Nunavut, Ellesmere Island, Alert) to 55°S (Argentina and Chile, Tierra del Fuego) in the New World and 50°S (New Zealand, Auckland Islands) in the Old World, and from sea level up to at least 4,500 m (Fernandez-Triana 2018). The information currently available allows us to make preliminary comments on species diversity and distribution at the generic level (Table 5 and Fig. 2).

World genera of Microgastrinae, based on the present paper. The column Species richness details the current number of described species and estimated total, for each genus, the two figures separate by a slash. The estimated total is very conservative and is based on specimens we have seen in collections. For many genera, more species are to be expected. World region keys: NEO Neotropical, NEA Nearctic, PAL Palaearctic, OTL Oriental, AFR Afrotropical, AUS Australasian (including Oceanian). X Genus present in specific region. X* New record for that region (based on undescribed species seen in collections). X- Introduced into that region, not native. X? Questionable record for a region. The column Host data tallies the genera that have at least one lepidopteran host recorded (although no critical assessment of how accurate those host records was made). The column DNA barcodes records all genera for which there is at least one DNA barcode available; Yes- denotes a genus with only partial sequence(s) available, without fulfilling the criteria for DNA-barcode compliant sequences (see Materials and methods for definition of a barcode-compliant sequence).

Genera Species richness NEO NEA PAL OTL AFR AUS Host data DNA bar-codes
Agupta 4/30+ X X No Yes
Alloplitis 8/30+ X X* No Yes
Alphomelon 19/50+ X X Yes Yes
Apanteles 633/3,000+ X X X X X X Yes Yes
Austinicotesia 2/5 X No Yes
Austrocotesia 5/10 X? X No Yes-
Beyarslania 1/2 X No Yes
Billmasonius 1/1 X No Yes
Buluka 11/20 X X X Yes Yes
Carlmuesebeckius 1/1 X No No
Chaoa 1/1 X No No
Choeras 80/100+ X* X X X X X Yes Yes
Clarkinella 2/5+ X X No Yes
Cotesia 328/1500+ X X X X X X Yes Yes
Cuneogaster 1/5 X No No
Dasylagon 2/5 X Yes No
Deuterixys 18/20+ X X X X X Yes Yes
Diolcogaster 141/1,000+ X X X X X X Yes Yes
Distatrix 32/40+ X X X X X Yes Yes
Dodogaster 1/1 X No No
Dolichogenidea 366/700+ X X X X X X Yes Yes
Eripnopelta 1/1 X No No
Exix 7/10 X X No Yes-
Exoryza 15/20+ X X X X X Yes Yes
Exulonyx 1/1 X No No
Fornicia 32/50+ X X X X Yes Yes
Gilbertnixonius 1/1 X No Yes
Glyptapanteles 307/3,000+ X X X X X X Yes Yes
Hygroplitis 9/10+ X X X Yes Yes
Hypomicrogaster 48/200+ X X Yes Yes
Iconella 38/50+ X X X X X Yes Yes
Illidops 37/50+ X X X X X X- Yes Yes
Janhalacaste 3/5 X Yes Yes
Jenopappius 3/5+ X No Yes
Jimwhitfieldius 2/5+ X No Yes
Keylimepie 4/10 X* X X No Yes-
Kiwigaster 1/1 X No Yes
Kotenkosius 1/2+ X No Yes
Larissimus 1/5+ X Yes Yes
Lathrapanteles 4/10+ X X Yes Yes
Mariapanteles 2/10+ X No Yes
Markshawius 3/5 X No Yes
Microgaster 104/200+ X X X X X X Yes Yes
Microplitis 192/500+ X X X X X X Yes Yes
Miropotes 15/20 X X* X Yes Yes
Napamus 2/2 X Yes No
Neoclarkinella 7/50+ X* X X* No Yes
Nyereria 29/50+ X X X Yes Yes
Ohenri 1/1 X No No
Papanteles 2/5 X Yes Yes
Parapanteles 62/100+? X X X* X X X Yes Yes
Parenion 3/5+ X X No Yes
Paroplitis 5/10 X X X Yes Yes
Pelicope 1/1 X Yes Yes
Philoplitis 9/10+ X* X X No Yes
Pholetesor 57/100+ X X X X X* X Yes Yes
Prasmodon 18/30+ X Yes Yes
Promicrogaster 46/100+ X X* X* X* X* X* Yes Yes
Protapanteles 25/30+ X X X Yes Yes
Protomicroplitis 3/5 X X Yes Yes
Pseudapanteles 36/100+ X X Yes Yes
Pseudofornicia 4/5+ X X No No
Pseudovenanides 1/5+ X* X Yes No
Qrocodiledundee 1/1 X No No
Rasivalva 12/20+ X* X X X X X* Yes Yes
Rhygoplitis 4/10+ X X Yes Yes
Sathon 23/30+ X X X X X* X Yes Yes
Semionis 1/1 X No No
Sendaphne 11/20 X No Yes
Shireplitis 6/6 X No Yes
Silvaspinosus 1/2+ X No Yes
Snellenius 41/50+ X X X X* X Yes Yes
Tobleronius 1/2+ X No Yes
Ungunicus 1/1 X No Yes
Venanides 14/20+ X X X* X X X Yes Yes
Venanus 11/15+ X X Yes Yes
Wilkinsonellus 23/50+ X X X X Yes Yes
Xanthapanteles 1/1 X No No
Xanthomicrogaster 6/30+ X Yes Yes
Ypsilonigaster 6/10+ X No Yes
Zachterbergius 1/1 X No Yes

The most species-rich genera are Apanteles (in its restricted sense) and Glyptapanteles. The latter is probably the largest, but it may eventually be split into several genera. In contrast, Apanteles, although also likely to have some species reclassified into other genera, is a much more cohesive group and might end up being the larger group if many species are removed from the current Glyptapanteles. Regardless, the diversity of both genera will likely comprise a few thousand species each.

Apanteles already contains more than 630 described species (see checklist below); just in ACG, Costa Rica, 186 new species were recently described (Fernandez-Triana et al. 2014e). The world fauna of Apanteles could number many more than 3,000 species. The genus is notably absent from New Zealand (although a few species have been introduced there), where it is replaced by Dolichogenidea and an undescribed genus. It also has not been found in the high Arctic (Fernandez-Triana et al. 2017b).

Glyptapantes contains more than 300 species, with hundreds of undescribed species from all biogeographical regions seen in collections; we estimate that the world total could be more than 3,000 species. However, the generic limits are controversial (see previous section) and it may eventually be restricted to a slightly smaller, although still substantial, number of species. Regardless, its status as one of the two largest genera of Microgastrinae is certain.

The following genera are also very speciose: Cotesia, Diolcogaster, Dolichogenidea, Hypomicrogaster, and Microplitis. Among these, Diolcogaster is clearly the largest, and it could attain more than 1,000 species. But it will almost certainly be split into several genera and thus it could potentially end up having just a few hundred species. Cotesia, already with more than 320 described species, will also attain more than 1,000 species (Mason (1981) estimated between 1,500–2,000 species), and is a more cohesive group, unlikely to be severely split. The other three genera will certainly surpass 500 species each, probably substantially (e.g., Dolichogenidea already has more than 360 described species). Diolcogaster and Hypomicrogaster are more speciose in tropical areas, whereas Cotesia, Dolichogenidea and Microplitis tend to be richer in temperate areas.

Other relatively large genera are Microgaster, Choeras, and Pholetesor in temperate areas, and Parapanteles and Pseudapanteles in tropical areas. All of them are likely to have more than one hundred (in most cases several hundred) species. A few other genera might be equally large, but the material in collections is not comprehensive enough to provide estimates.

In regional composition, the tropical areas have a larger representation than temperate areas (as expected) with the Oriental (46 genera) and Neotropical (43 genera) regions being of comparable diversity, and the Afrotropical (36 genera) and Australasian (28 genera) regions following. Furthermore, we have seen in collections several putative additional (undescribed) genera from all tropical regions. In temperate areas, the Nearctic region (33 genera, including several Neotropical genera having a few species entering North America) has the highest generic diversity and the Palaearctic region (28 genera, including some Oriental genera that have a few species entering the southernmost areas of the Palearctic) has the lowest diversity. Considered as a whole, the entire Holarctic region would have a relatively high diversity of 39 genera.

The distribution of individual genera worldwide (Fig. 3) shows that 20 genera (24.7%) are cosmopolitan or almost so: 15 are present in all biogeographical regions (Apanteles, Choeras, Cotesia, Diolcogaster, Dolichogenidea, Glyptapanteles, Illidops, Microgaster, Microplitis, Parapanteles, Pholetesor, Promicrogaster, Rasivalva, Sathon, and Venanides) while another five are present in five out of the six biogeographical regions (Deuterixys, Distatrix, Exoryza, Iconella, and Snellenius). A few additional genera may eventually be found to be cosmopolitan.

Figure 3. 

Biogeographical distribution of the 81 Microgastrinae genera currently known worldwide. Data from the present paper.

Eleven genera (13.6%) are restricted to the New World tropics (Neotropical region): Cuneogaster, Dasylagon, Janhalacaste, Larissimus, Mariapanteles, Papanteles, Prasmodon, Sendaphne, Venanus, Xanthapanteles, and Xanthomicrogaster. Another nine genera (Alphomelon, Clarkinella, Exix, Hypomicrogaster, Lathrapanteles, Protomicroplitis, Pseudapanteles, Rhygoplitis, and Venanus) are almost exclusively found in the Neotropics, with few species reaching the Nearctic. The only genus that can be considered as a Nearctic endemic is Pelicope.

Ten genera (12.3%) are relatively widespread in, but restricted to, the Old World tropics: Agupta, Alloplitis, Buluka, Miropotes, Parenion, and Pseudofornicia. We also consider here Neoclarkinella, Nyereria, Philoplitis, and Pseudovenanides as almost exclusively present in the Old World tropics, as only a few species reach the southernmost areas of the Palaearctic.

Only two genera (2.5%) (Fornicia and Wilkinsonellus) seem to be pantropical, and completely absent in the Holarctic. Because almost all undescribed genera of Microgastrinae in collections are from tropical areas, this proportion could increase. No genus has a strictly Holarctic distribution, but three genera almost fulfill that criterion, as just a few species of each reach the northern limits of either the Oriental region (Hygroplitis and Paroplitis) or the Neotropical region (Rhygoplitis).

A total of 35 genera (43.2%) are presently known only from a single biogeographical region, with the Neotropical and Oriental regions each having ten endemic genera, respectively, and the Afrotropical having eight (Table 5). However, some of those genera will almost certainly be found to have a wider distribution.

DNA barcoding and Microgastrinae

During the past 12+ years, an extensive library of DNA barcodes for Microgastrinae has been assembled (Smith et al. 2013), resulting in the subfamily comprising 37% of all DNA sequences of Braconidae and almost 5% of all Hymenoptera sequences currently available in BOLD. At present, 44,739 specimens of Microgastrinae have sequences deposited in BOLD; 40,812 of those specimens have DNA barcodes representing 3,545 public BINs (http://v4.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxid=2099). The number of BINs will certainly increase, as most of the Microgastrinae specimens currently in BOLD come from just two countries: Canada and Costa Rica (Fig. 4).

Figure 4. 

Overview of Microgastrinae data in the Barcoding of Life Data System (BOLD) as of 31 December 2019.

BINs usually match well with putative species (as identified by an expert taxonomist), and thus could be used as a surrogate for analyses of species diversity, like other Operational Taxonomic Units (e.g., Ratnasingham and Hebert 2013, Fagan-Jeffries et al. 2018b). Based on our unpublished data, the correspondence between BINs and putative species in Microgastrinae may exceed 90%. For example, the number of Microgastrinae public BINs from Canada and Alaska (combined) currently found in BOLD is 551, very similar to the 550 species estimated for that area by Fernandez-Triana (2010; see also Fagan-Jeffries et al. 2018b). Even with the limited geographical coverage presently available, the total number of worldwide Microgastrinae BINs already surpasses the total of described species in our checklist by almost 200.

At the genus level, a significant proportion (67 genera or 83%) have some DNA data (Table 5). In most cases (64 genera or 79%) that includes at least one barcode compliant sequence, usually many more. Many of the 14 genera without molecular data in BOLD include taxa that are very rare in collections, i.e., only known from one or very few specimens, and/or the available specimens are very old (collected many decades ago) and did not yield any DNA. However, for at least a few of those genera it is expected that it will soon be possible to have DNA data.

Estimating species richness in Microgastrinae

With 2,999 valid species of Microgastrinae recognized here, an interesting question is how many species remain undescribed, whether or not known from collections. The actual species richness of Microgastrinae worldwide has been variously estimated during the past 35 years. At the lower end, Dolphin and Quicke (2001) extrapolated species richness of Braconidae based on data from butterflies and (primarily) mammals, arriving at an estimated 3,617–4,178 species of Microgastrinae. Jones et al. (2009) obtained similar results by comparing taxonomic revision data, with their estimates ranging from 3,900–5,500 species. Mason (1981) thought that 5,000–10,000 species would be a reasonable estimate, based on museum specimens he had seen. At the higher end of the spectrum, Rodriguez et al. (2013) compared the number of Lepidoptera and Microgastrinae species in several areas to arrive at estimates ranging from 17,000 to 46,000+ species.

Obviously, these estimates vary considerably: if the lowest one (3,617) were accurate, then we would already know 82% of the Microgastrinae species; if the highest one (46,620) were accurate, then the described species would represent only 6% of the actual diversity worldwide. Which estimate is more likely to be correct?

While a definite answer cannot be provided, some refinement of the current estimates is possible. The lowest range (3,000–5,000 species) is clearly too low based on what is currently known (2,999 described, valid species are recognized in this paper). As mentioned in the previous section, and despite its limited geographical coverage, Microgastrinae public BINs already represent 3,545 putative species. But, even if DNA data is disregarded, we have certainly seen in collections a few thousand undescribed species, which are clearly distinct based on morphological features alone. In that sense, Mason’s estimate of 10,000 species seems very reasonable.

But could the figures from Rodriguez et al. (2013) also be considered reasonable, or are they way off the mark? Although this might be seen just as a numbers game, the implications are significant. If indeed there were 30-, 40- or even 50,000 species of Microgastrinae worldwide, that could extrapolate to the entire family Braconidae having at least 150–200,000 species, and the entire Hymenoptera having much more than one million species. Those values are an order of magnitude higher than the values presently known for subfamily, family, and order, although they agree with estimates of the entire Hymenoptera suggested by other authors (e.g., LaSalle and Gauld 1991, Hanson and Gauld 1995, Foottit and Adler 2017).

Rodriguez et al. (2013) based their estimates on what Fernandez-Triana (2010) had referred to as the Lepidoptera/Microgastrinae ratio (L/M). Briefly explained, the ratio between lepidopteran and microgastrine species (where sufficient data are available) seems to be surprisingly similar in different regions, regardless of the area and diversity of such regions. The initial calculations were limited and only included three separate areas in Canada (Table 2 in Fernandez-Triana 2010). Based on the average ratio calculated from those three areas (L/M = 12/1) it was concluded that the richness of Microgastrinae in Canada and Alaska would be approximately 550 species. Rodriguez et al. (2013: Table 1) expanded the dataset to eleven different regions, mostly from North America and Europe, but also including New Zealand and ACG in Costa Rica; the resulting L/M ratios were still remarkably close, mostly ranging from 10/1 to 20/1, with an average of 16.4/1.

But just a few years later, some of the numbers used by Fernandez-Triana (2010) and Rodriguez et al. (2013) are already outdated. For Microgastrinae, the species richness in Ottawa, based on Fernandez-Triana et al. (2016a) and subsequent unpublished data, is now approaching 180 species, which represents a 20% increase compared to the total published in 2010; ACG in Costa Rica has surpassed 1,200 species, a 50% increase (based on Janzen and Hallwachs 2016); the Canadian High Arctic now has 26 recorded species or 30% more than initially reported (based on Fernandez-Triana et al. 2017b); the New Zealand fauna will increase by more than 25% compared to previous estimates (Fernandez-Triana & Ward, unpublished); even for the UK, arguably the most thoroughly studied region, the microgastrine count increased by at least 15% (based on Broad et al. 2016). Those revised figures all share one element in common: the species richness of Microgastrinae in those areas was underestimated by both Fernandez-Triana (2010) and Rodriguez et al. (2013).

Thus, the updated L/M ratios calculated for the above regions decreased, from an average of 16/1 in Rodriguez et al. (2013) to around 10/1 at present (also including Finland, where comprehensive data have become available since the Rodriguez et al. paper was published). But the lower the L/M ratio the higher the actual species richness of Microgastrinae. For example, assuming an estimated world number of Lepidoptera between 300,000 (Kristensen et al. 2007) and 500,000 species (Foottit and Adler 2017), and a world average L/M ratio of 10/1, the estimated number of Microgastrinae would then range from 30,000–50,000 species. If anything, the current data still seem to support higher, rather than lower, estimates for the subfamily.

As far as we know, there is only one major caveat in using L/M ratios to extrapolate and calculate the world fauna of Microgastrinae: at present all known figures come from temperate areas, with the sole exception of ACG. There is no other tropical area in the world with sufficient data to allow for meaningful L/M ratios to be calculated. Thus, it may be argued that if different ratios were prevalent in temperate areas compared to the tropics, which harbour, by far, the highest richness of Microgastrinae, then the overall world estimates could not be as high as Rodriguez et al. (2013) suggested. Only more data will allow this to be answered in a definite way; however, for the present it is worth noting that the L/M ratio in ACG (10/1) is actually very similar to those of temperate areas.

Overview of regional taxonomic studies on Microgastrinae

As with many insect groups, knowledge of Microgastrinae has been historically concentrated on the Northern Hemisphere temperate fauna. However, numerous recent studies are starting to shift focus to the tropics, with most new species in the past few years being described from the hitherto poorly worked Neotropical and Oriental regions, chiefly Costa Rica, China, and India.

In the Western Palearctic subregion, papers from the 1960s–1990s from Nixon and Papp treated most of the Microgastrinae species known up to that time, following careful work by Wilkinson from the 1920s–1940s aimed largely at interpreting poorly understood names (see papers of these three authors cited in the References section). Recent works have described a relatively small number of new species, although their papers sometimes included detailed accounts of species biology, and there is an ongoing concomitant deposition of DNA barcodes, etc. (Oltra and Michelena 1989, Oltra et al. 1995, 1996, Oltra-Moscardó & Jiménez-Peydró 2005, Shaw 1992, 2004, 2007, 2009, 2012b, van Achterberg 2002, Fernandez-Triana et al. 2014c). The Eastern Palearctic subregion is less well known, although progress has also been made (Tobias 1986, Kotenko 1981, 1986, 1992, 1993, 2004, 2007a, 2007b), and most of the new Palaearctic species to be discovered will probably come from the Eastern Palearctic. Some southern areas of the Palearctic, e.g., Iran, Turkey, and the Palearctic area of the Arabian Peninsula have also seen an increase in the number of publications in the last few years (Inanç 1992, 2002a, 2002b, Inanç and Çetin Erdogan 2004, Gadallah et al. 2015, Farahani et al. 2016, Ghahari and van Achterberg 2016, Fernandez-Triana and van Achterberg 2017, Ghafouri Moghaddam et al. 2018, 2019, Samin et al. 2018, Abdoli et al. 2019a, 2019b, Zargar et al. 2019); however, there have been few taxonomic revisions, with most of the work being biodiversity estimates, local checklists, or isolated species descriptions. With 827 described species of Microgastrinae, the Palearctic is currently the most speciose region, although it will almost certainly become the least when more studies in the other regions are undertaken.

In the Nearctic region progress has been slower than in the Palearctic. After two seminal papers from Muesebeck (1921, 1922), most of the new taxa have been described in isolated papers, mostly treating species of biocontrol relevance (Marsh 1975, 1978, 1979b, 1979c, Wharton 1983, Whitfield et al. 1999, Fernandez-Triana 2010, 2018; cf. other papers from Muesebeck cited in the References section), with some taxonomic revisions also produced (Whitfield 1985, 2006, Whitfield et al. 2011, Grinter et al. 2009, Valerio et al. 2009, Valerio and Whitfield 2015, Fernandez-Triana 2015, 2019 Fernandez-Triana and Boudreault 2016, Fernandez-Triana et al. 2013a). Hundreds of additional species from this region have been revealed by DNA barcoding, but the southernmost areas and west coast, which also happen to be the most species rich, have barely been studied (Fernandez-Triana 2018). It is expected that the actual numbers in the Nearctic will be several times higher than the current 350 described species.

The Neotropical region has been the focus of recent efforts, including the description of more than 400 new species and revision of many genera. However, most of those papers deal almost exclusively with the fauna of ACG, Costa Rica (Janzen et al. 2003, Valerio and Whitfield 2003, Valerio et al. 2005a, Fernandez-Triana 2015, Fernandez-Triana et al. 2013a, 2014a, 2014e, 2014f, 2014g, 2014h, 2015b, 2016b, 2016c), with only some marginal coverage of other countries (Austin and Dangerfield 1989, Penteado-Dias 1995, Penteado-Dias et al. 2000, 2002, 2011, Valerio and Whitfield 2005, 2015, Valerio et al. 2004, 2009, Choi and Whitfield 2006, Grinter et al. 2009, Arias-Penna et al. 2014, 2019, Salgado-Neto et al. 2018, 2019). Large collections have been amassed in South America, e.g., French Guiana, Colombia, Brazil, Ecuador, and Peru, but an impediment to assessing that material is the difficulty in exchanging specimens with colleagues from other countries. In general, most of the Neotropics are extremely understudied, with several thousand species awaiting description but only 768 species described so far. For Microgastrinae, this is likely to be the most speciose region of the world.

The Oriental region, with 752 described species, currently ranks third after the Palearctic and Neotropical regions. It also contains thousands of undescribed species and may rival the Neotropical region as the most speciose. Recent advances have mostly been made in China and India, but we are also aware of large collections of specimens from other countries such as Indonesia, Malaysia, Thailand and Vietnam, which have already resulted in several publications (Austin 1987, 1989, Chen et al. 1994, Long and van Achterberg 2003, 2008, 2011, 2013, 2014, 2015, Chen and Song 2004, Long 2007, 2010, 2015, Fernandez-Triana and Goulet 2009, Fernandez-Triana et al. 2014d, Zeng et al. 2011a, 2011b, Gupta 2013a, 2013b, Gupta and Kalesh 2012, Gupta and Fernandez-Triana 2014, 2015, Gupta et al. 2011, 2013a, 2013b, 2014a, 2014b, 2016a, 2016b, Liu et al. 2014, 205, 2016, 2018, Veena et al. 2014, van Achterberg et al. 2015, Xiong et al. 2017, Zhang et al. 2017, Ranjith et al. 2015a, 2015b, 2019; cf. papers from authors Chen, Sathe, Song, Xu, and You cited in the References section). The main problem (other than difficulties in exchanging material) is the lack of revisions covering the entire region; the available taxonomic keys and papers tend to cover single countries, with few efforts to coordinate work at a larger (regional) scale. There is also a number of species described from India in publications that do not comply with ICZN Article 16, and thus those names are unavailable (see section Unavailable names below).

No significant progress has been made in the Afrotropical region for the past half a century. The very few exceptions include recent papers on the fauna of Réunion (Rousse and Gupta 2013), the Afrotropical area of the Arabian Peninsula (Fernandez-Triana and van Achterberg 2017), and some new species of importance in biocontrol (Kaiser et al. 2017, Fiaboe et al. 2017), or more general papers not specifically devoted to the Afrotropics (Walker 1994, Valerio et al. 2009, Fernandez-Triana and Goulet 2009, Fernandez-Triana and Boudreault 2018). However, relatively large collections from Kenya, Madagascar, Republic of Congo, and South Africa have been amassed during the past few years (Fernandez-Triana and Boudreault 2018), and there is potential to add hundreds, if not thousands of new species. Although the current total of described species is just 429 it is estimated that this will be the third most species-rich region of the planet for Microgastrinae.

Since the 1990s, several papers have treated the Australasian species (Austin 1990, Austin and Dangerfield 1992, 1993, Walker 1996, Saeed et al. 1999, Fernandez-Triana et al. 2011, 2013b, Fagan‐Jeffries and Austin 2018, Fagan‐Jeffries et al. 2018a, 2018b, 2019), but progress has been comparatively slow. At present 222 species are described from this region. Work on Pacific islands is basically non-existent but, when done, may reveal many more new and interesting taxa.

Hosts of Microgastrinae

The host range of a parasitoid is one of its most important features, linking its evolutionary past with its present autecology (Shaw 1994, Shaw and Aeschlimann 1994). Through knowledge of the host range it is possible to understand and to predict a parasitoid’s behaviour within current ecosystems (Shaw 2017b), and also gain some understanding of the speciation processes that brought them into existence (Shaw 2003).

Microgastrinae are the single most important group of parasitoids of Lepidoptera in the world, both in economic terms and in species richness (Whitfield 1995a, 1997). They are all koinobiont endoparasitoids and parasitize almost the entire taxonomic and biological spectrum of Lepidoptera (Shaw and Huddleston 1991, Whitfield 1997, Whitfield et al. 2018), with the probable exception of the four most basal superfamilies.

Adult female wasps typically oviposit into early instar larvae (with a few species known to oviposit into host eggs), within which the wasp eggs hatch and larval development takes place with the aid of venom and polydnavirus (PDV) effects on the host’s immune and endocrine system (summarized in Whitfield et al. 2018). All microgastrines fully depend on mutualistic PDVs to successfully parasitize hosts, the relationship between wasps and PDVs being the most remarkable known example of the evolution of a mutualistic endosymbiotic association between eukaryotes and viruses (Strand and Burke 2012, 2014).

Numerous literature records of non-Lepidoptera as hosts of Microgastrinae exist (Table 6), comprising at least 29 families within five orders of Insecta (data compiled from Yu et al. 2012). However, these records are wrong or at the very least highly questionable.

Historical account of Microgastrinae hosts that are not Lepidoptera, based on the compilation of Yu et al. (2012).

Order Families
Coleoptera Anobiidae, Anthomyiidae, Attelabidae, Bostrichidae, Buprestidae, Cerambycidae, Chrysomelidae, Coccinellidae, Curculionidae, Melandryidae, Phalacridae, Scirtidae
Diptera Agromyzidae, Cecidomyiidae, Chloropidae, Muscidae, Syrphidae, Tephritidae
Hymenoptera Apidae, Argidae, Cimbicidae, Cynipidae, Diprionidae, Eurytomidae, Pteromalidae, Tenthredinidae, Vespidae
Mantodea Mantidae
Trichoptera Limnephilidae

For example, the record of Apidae (Bombus sp.) as “host” of Microgastrinae can be easily rejected. Bombus nests have associated case-bearing moth caterpillars (Tineidae) feeding within the nest and the three known species of Microgastrinae that emerge from those nests actually parasitize the caterpillars, not the bees (Whitfield and Cameron 1993, Whitfield et al. 2001).

Two other recent examples are equally illustrative. The record of Enoicyla pusilla (Burmeister) (Trichoptera: Limnophilidae) as a host of the microgastrine Choeras gielisi (van Achterberg 2002) was at times considered to be a reliable example of a non-Lepidoptera host record; however, subsequent examination of the situation has called that record into doubt as it was reared from a substrate from which the host remains were not recovered (Shaw 2017a). Similarly, Kopelke (2011) reported two different species of Dolichogenidea (each from a single specimen), as part of his extensive rearing of inhabitants of 34,210 galls of nematine sawflies (Hymenoptera: Tenthredinidae) in Europe; he asserted those two cases to be accidental, but genuine (Kopelke 2011: 9). Unfortunately, it is not clear from the publication if host remains were available (in those two specific cases) to confirm host identity, and under such circumstances we consider it appropriate to regard those records as highly dubious. Sawfly galls are nutritious and frequently fed on by caterpillars. It is relatively easy for a small parasitized lepidopteran larva to enter such a structure to die and become practically entirely consumed by the parasitoid, leaving almost only the head capsule. This happens with most Dolichogenidea species, which have a final external feeding period that leaves the host remains easily overlooked or misinterpreted. Many similar deductions concerning other recorded supposed non-lepidopteran hosts are easily made.

Even if examples of parasitization of other insect orders by Microgastrinae are well founded, we consider such cases would be highly abnormal. Shaw (1994) provided a conceptual definition for the host range of a particular parasitoid species, which should include only those species of potential hosts that the parasitoid is usually able to attack successfully, following a pattern of searching behaviour enabling it to encounter them regularly. That rather loose definition implies that some perfectly correct rearing records should be excluded from the host range if they represent only freak events of no importance to the autecology of the parasitoid or the host, and lack phylogenetic significance. It also implies that some hosts within the host range may be intrinsically more important than others that are encountered less frequently, or attacked less enthusiastically, or with a less successful outcome.

We also consider that there is no convincing evidence that the four most basal superfamilies of Lepidoptera (sensu Aarvik et al. 2017) are parasitized by Microgastrinae. There is no published record of Microgastrinae parasitizing Micropterigoidea and Eriocranioidea, and the few literature records of hosts in Hepialoidea and Nepticuloidea are highly questionable; we discuss and reject them below.

Sathon falcatus (Nees, 1834) was recorded in two broods (of 45 and 37 individuals) parasitizing Hepialus humulis (Linnaeus, 1758) (Hepialidae) in the United Kingdom (Hammond and Smith 1957). We have located those specimens in the NHMUK but, although the relevant cocoon masses are present, there are no host remains. Sathon falcatus is a known parasitoid of the noctuid moth Apamea monoglypha (Hufnagel, 1766), whose larvae are superficially very similar to those of Hepialus humuli. Thus, we distrust the record strongly enough to refute it. It should also be noted that the rearings were not done by Hammond, who was the expert on Lepidoptera larvae.

The other known record is for Cotesia spuria (Wesmael, 1837) parasitizing Triodia sylvina (Linnaeus, 1761) (Hepialidae), published by Telenga (1955) with no details whatsoever, i.e., no information was provided on who identified the host or the parasitoid, or where and when the sample was collected, nor the depository of specimens. Cotesia spuria does have a wide host range, but confirmed hosts are all folivorous macrolepidoptera. Under these circumstances it is best to simply refute the record; although of course, if a rearing is repeated with appropriate credentials the refuted record could be recalled to stand as a possible previous instance.

The two published records of Nepticuloidea as hosts are also highly suspicious. Nixon’s (1976) record of Fomoria weaveri (Stainton, 1855) (Nepticulidae) as a host of Apanteles contaminatus (Haliday, 1834) has recently been refuted by Shaw (2012b), who commented on the rearing. The inflated mines of F. weaveri are superficially similar to those Epinotia nemorivaga (Tengström, 1848) (Tortricidae) from which A. contaminatus has been reliably reared; thus, in this case an error in host identification was almost certainly involved. Unfortunately, the specimen could not be found in the cited depository.

Gates et al. (2002: 221) recorded Stigmella ? variella (Nepticulidae) being parasitized by Dolichogenidea tischeriae Viereck (1912b) from a leaf mine on oak (Quercus agrifolia Née). However, that record is quoted as “parasitoids lot-reared from more than one leafmines from a single plant” (see caption of Table 2 on page 230 of Gates et al. 2002), and in that same Table other Lepidoptera families were recorded from that host plant, including several species of Gracillariidae and Tischeriidae, both of which had been reported as hosts of D. tischeriae in other papers and most likely represent the actual host(s). In that case, it is clear that the sample (leaves with mines) contained several lepidopteran species, and that Stigmella was wrongly assigned as a host of D. tischeriae.

Adeloidea and Tischerioidea are the most basal superfamilies of Lepidoptera (and the only non-Ditrysia groups) for which there is reasonably solid evidence supporting them as being hosts of Microgastrinae. There is reliable data showing that a few Microgastrinae indeed parasitize species of Adelidae (Shaw 2012b), Incurvariidae (Fernandez-Triana 2010), Prodoxidae (Nixon 1972, Shaw 2012b, Whitfield et al. 2005), Tischeriidae (Shaw 2012b) and even Heliozelidae (Fernandez-Triana et al., unpublished data).

Ditrysia (sensu Kristensen and Skalski 1999, Roe et al. 2009) constitutes the most derived clade of Lepidoptera, comprising more than 98% of all lepidopteran species, and representing by far the group most commonly parasitized by Microgastrinae. Eulepidoptera (sensu Aarvik et al. 2017) consists of Adeloidea + Tischerioidea + Ditrysia, which are the three groups for which we have solid evidence of parasitism by Microgastrinae. Thus, in this paper we propose that Microgastrinae hosts are restricted to Eulepidoptera, i.e., most of the Lepidoptera except for the four most basal superfamilies: Micropterigoidea, Eriocranioidea, Hepialoidea and Nepticuloidea. We consider all previous literature records of other insect orders and of the four early branching lineages of Lepidoptera as incorrect. Claims for hosts other than Eulepidoptera, which are made with conviction from time to time, are in our experience never supported by the recovery and preservation of associated host remains for careful assessment.

The published sources we compiled so far include Lepidoptera host data for 44 genera (54%) and around 1,250 species (42%) of Microgastrinae. Although the coverage is insufficient, those records include 3,200+ species of Lepidoptera and represent 5,500+ supposed host/parasitoid associations. In addition, there is a large amount of unpublished but databased host information (e.g., http://janzen.sas.upenn.edu/caterpillars/database.lasso; http://www.caterpillars.org/), with hundreds of additional host/parasitoid records from currently undescribed microgastrine species (e.g., Whitfield et al. 2009, 2018, Hrcek et al. 2013). Still, more than half of the described species of microgastrines lack any information about their hosts. Even worse, an unknown but probably very large proportion of the published associations are also almost certainly wrong. Clearly, there is much to be learned, and for the existing information to be a good basis for understanding host records there needs to be a critically examination of the data to (try to) prune out wrong host/parasitoid associations, an effort that would require years of work, and even then would leave much uncertainty. A better approach to secure real knowledge may be to ensure that higher standards of data collection and specimen deposition take place for the future: in fact, without that we cannot expect much improvement in our understanding.

From the data presently available, the top ten families of Lepidoptera (as per number of species recorded as host) which are parasitized by Microgastrinae are Noctuidae, Tortricidae, Pyralidae, Crambidae, Geometridae, Gracillariidae, Depressariidae, Hesperiidae, Gelechiidae, and Nymphalidae. Altogether those families account for two-thirds of all known host/microgastrine parasitoid associations, which is not surprising given that they are also among the most species- rich Lepidoptera families. That probably also reflects a bias in collecting effort: these families provide most of the economically important crop and forestry pests, which are accordingly the most intensely sampled taxa for their parasitoids. Further, in some of these families there are large and/or spectacular caterpillars that are the most often seen and reared by hobbyists. Other groups such as stem borers, leaf litter, and canopy caterpillars tend to be less commonly reared.

Earlier compilations for species within particular microgastrine genera are dominated by records from the northern temperate region which are unlikely to reflect the complete spectrum of host associations when the ongoing (but currently mostly unpublished) massive number of rearings from tropical surveys are taken into account, e.g., Whitfield et al. (2018). In addition, there is a need to recognise phenological aspects of host range, especially in temperate climates: many parasitoid species are plurivoltine yet use univoltine hosts, each available to only one generation of the parasitoid; sometimes it happens that the parasitoid is, at least locally, entirely dependent on a single host species at one time of year but able to use another host or a wider range of hosts at another (see Shaw and Aeshlimann 1994). Last but not least, a parasitoid’s realized host range may not be constant in either space or time unless, of course, the parasitoid is strictly monophagous, and thus the relative abundance of co-occurring hosts will also vary (Shaw 2006). Recognition of the realized host range at a point in space and time is often of more practical significance for population dynamics, conservation biology, or biological control (Shaw 1994, 2003, 2017).

Despite the constraints mentioned above and the relatively poor state of knowledge, some general comments can be made for some of the most speciose Microgastrinae genera. For example, most Microgaster, Choeras, Apanteles, and Dolichogenidea species parasitize more or less concealed host larvae, allowing the final instar larvae of these parasitoids to carry out their external feeding phase in a sheltered environment, and host Lepidoptera with this amenable larval biology overwhelmingly belong to the families of the so-called microlepidoptera. Other genera such as Pholetesor and Deuterixys specialize on leaf-miners and parasitize hosts that feed in at least moderate concealment, as is required by the final external tissue-feeding phase of their parasitoid larvae. This is correlated with their use of hosts primarily from microlepidopteran families, which tend to be small, resulting in most of the parasitoids of microlepidoptera being solitary. In contrast, genera such as Microplitis, Cotesia, Distatrix, Diolcogaster, Protapanteles, and Glyptapanteles are fully endophagous and well-suited to parasitize exposed Lepidoptera larvae, such as those of many macrolepidoptera, which tend to be large and are thus more suited to support gregariousness, which is much more expressed in these microgastrine genera. There are exceptions, but they can often be understood in autecological terms, e.g., the few Microgaster that parasitize macrolepidopterans have hosts that feed or rest in concealed sites (Shaw 2004); the few Cotesia that parasitize microlepidopterans are usually associated with semi-exposed hosts in webs which feed partly exposed (see Nixon 1974); the flavipes group of Cotesia parasitizes stem borers in the families Pyralidae and Crambidae (e.g., Fujie et al. 2018).

Whenever comprehensive data are available, be it in temperate (e.g., in Europe and especially the United Kingdom), or tropical areas (e.g., ACG), patterns emerge. Often, they show that many species within most genera of Microgastrinae appear to have a high host specificity, often having been recorded from only a single or very few taxonomically closely related species. An alternative is having ecologically similar hosts (Shaw 2003, Fernandez-Triana 2018). Earlier studies often did not differentiate these levels of host specificity clearly, partly due to the presence of many morphologically cryptic species in large genera of Microgastrinae but also because it is very much harder to discover all or most of the hosts of a particular parasitoid species than it is to discover all or most of the parasitoid species using a given host. Only recently have they been detected through integrative taxonomy that incorporates DNA barcoding and other molecular methods, as well as much greater levels of field ecological data (Shaw 2017b, Whitfield et al. 2018).

However, some species of Microgastrinae seem to be much less restricted. Examples include Glyptapanteles vitripennis (Curtis, 1830), an incredibly polyphagous species with an immense host range of mainly (but not entirely) exposed macrolepidoptera found on trees and bushes in Europe (Nixon 1973, Shaw unpublished data), or Glyptapanteles pseudotsugae Fernandez-Triana, 2018, which parasitizes several lepidopteran species (Geometridae and Erebidae) feeding on Douglas fir across a range of 2,500 km in western North America (Fernandez-Triana 2018).

A few Microgastrinae genera seem to be restricted to only one host Lepidoptera family, e.g., Alphomelon (only reared from Hesperiidae), Fornicia (Limacodidae), Janhalacaste (Depressariidae), Papanteles and Xanthomicrogaster (Crambidae), and Pelicope (Prodoxidae). However, these microgastrine genera are not very species rich and it is difficult to know whether more data would extend their apparent associations.

For more speciose genera, the patterns are less clear or consistent, as the number of host families increases, in some cases dramatically. This may in part be a consequence of some Microgastrinae genera not being well defined, comprising at present an arrangement of different lineages that may be separated into different genera in future, e.g., Choeras, Diolcogaster, Glyptapanteles, and Hypomicrogaster. But some large and relatively well-defined genera, e.g., Apanteles, Cotesia, Dolichogenidea, Microplitis, and Microgaster, have large host ranges, including both early and more recently branching lepidopteran families, and ecological factors in their radiations have clearly been of importance.

There is no comprehensive account of the impact of Microgastrinae in biological control. Whitfield (1997) estimated that more than one hundred species had been studied and used in biocontrol programs against caterpillar pests worldwide, but he did not provide details or references to support that number. We have compiled the available information and have found that 800+ species of Lepidoptera considered as pests of some sort in agriculture and forestry are parasitized by Microgastrinae (Fernandez-Triana et al. unpublished data; host data for individual species of Microgastrinae is not presented in this paper, see next paragraph). That includes 110+ major pests, highlighting the importance of this group of parasitoid wasps in biological control programs anywhere.

In summary, Microgastrinae are the most abundant and diverse taxon of hymenopteran parasitoids reared from lepidopteran caterpillars worldwide. However, our current level of knowledge is still poor, as more than half of the wasp species have no host association records, and of the records that do exist, many of them are doubtful or plainly wrong. Considerable effort will be needed before we have a better and more accurate picture of the host/parasitoid associations of most species of Microgastrinae. Thus, in this paper we only provide general comments; details on individual host/parasitoid associations are intentionally omitted to avoid repeating and perpetuating inaccurate information.

Checklist of world genera and species of Microgastrinae

[Genera, and species within each genus, are arranged in alphabetical order. At the end of the list we place the species we consider as species inquirendae, nomina dubia, and nomina nuda, also in alphabetical order. For a complete list of all Microgastrinae available names in strict alphabetical order see also Suppl. material 1, 2]

Genus Agupta Fernandez-Triana, 2018

Agupta Fernandez-Triana, 2018: 28. Gender: neuter. Type species: Agupta jeanphilippei Fernandez-Triana & Boudreault, 2018, by original designation.

Four species are described from the Oriental region (Fernandez-Triana and Boudreault 2018); those authors stated that there are dozens of undescribed species, based on collection holdings and specimens with available DNA barcodes, from the Australasian and Oriental regions. No revision of the genus has yet been produced. No host data are currently available for this genus. There are dozens of DNA-barcode compliant sequences of Agupta in BOLD, representing more than 25 different BINs (but none of those sequences have been identified in BOLD as belonging to Agupta, see Fernandez-Triana and Boudreault (2018) for more details on that).

Agupta danyi Fernandez-Triana & Boudreault, 2018

Agupta danyi Fernandez-Triana & Boudreault, 2018.

Type information. Holotype female, RMNH (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Agupta jeanphilippei Fernandez-Triana & Boudreault, 2018

Agupta jeanphilippei Fernandez-Triana & Boudreault, 2018.

Type information. Holotype female, RMNH (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Agupta raymondi Fernandez-Triana & Boudreault, 2018

Agupta raymondi Fernandez-Triana & Boudreault, 2018.

Type information. Holotype female, RMNH (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Agupta solangeae Fernandez-Triana & Boudreault, 2018

Agupta solangeae Fernandez-Triana & Boudreault, 2018.

Type information. Holotype female, RMNH (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Genus Alloplitis Nixon, 1965

Alloplitis Nixon, 1965: 268. Gender: masculine. Type species: Alloplitis guapo Nixon, 1965, by original designation.

Eight species are currently described from the Oriental and Afrotropical regions, but we have seen in collections (CNC, RMNH) numerous additional species from those regions. No revision of the genus has been produced, although a key to all four species known from Vietnam (Long & van Achterberg 2008) covers half of the described species. No host data are currently available for the genus. There are 20 DNA-barcode compliant sequences of Alloplitis in BOLD representing eight different BINs, most of them undescribed species from Thailand.

Alloplitis albiventris Long & van Achterberg, 2008

Alloplitis albiventris Long & van Achterberg, 2008.

Type information. Holotype female, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Alloplitis completus Mason, 1981

Alloplitis completus Mason, 1981.

Type information. Holotype female, CNC (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Alloplitis congensis (de Saeger, 1944), new combination

Microplitis congensis de Saeger, 1944.

Type information. Holotype male, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo.

Notes. Even in the original description (de Saeger 1944), this species was considered not likely to belong to Microplitis. Without examining the holotype (and only known specimen), the best generic placement at present would be Alloplitis based on the propodeal areola, T1 with an impression on the basal third and striae on lateral margins, T2 rectangular in shape, and T3 shorter than T2.

Alloplitis detractus (Walker, 1860), new combination

Microgaster detractus Walker, 1860.

Type information. Holotype male, NHMUK (examined). Country of type locality: Sri Lanka.

Geographical distribution. OTL.

OTL: Sri Lanka.

Notes. From the original description and subsequent treatment of the species (Wilkinson 1927, 1929), it is clear that this species does not belong to Microgaster. After examining the holotype, we here transfer detractus to Alloplitis based on its short metatibial spurs, propodeum with a complete areola defined by strong carinae, T1 with a broad impression on anterior half, T2 broadly rectangular, and anteromesoscutum, scutellar disc, T1 and T2 heavily sculptured.

Alloplitis guapo Nixon, 1965

Alloplitis guapo Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines, Vietnam.

Alloplitis laevigaster Long & van Achterberg, 2008

Alloplitis laevigaster Long & van Achterberg, 2008.

Type information. Holotype male, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Alloplitis typhon Nixon, 1965

Alloplitis typhon Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Alloplitis vietnamicus Long & van Achterberg, 2008

Alloplitis vietnamicus Long & van Achterberg, 2008.

Type information. Holotype female, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Genus Alphomelon Mason, 1981

Alphomelon Mason, 1981: 54. Gender: neuter. Type species: Urogaster nigriceps Ashmead, 1900, by original designation.

Known from 19 described species from the New World (mostly Neotropical, with a few extending north into the Nearctic). The revision by Deans et al. (2003) is outdated; we have seen in collections (CNC) dozens of additional species, and the genus will easily surpass 50 species with additional study of the Neotropical fauna. All data currently available suggest that Alphomelon species may exclusively be parasitoids of Hesperiidae. There are 1,200+ DNA-barcode compliant sequences of this genus in BOLD, representing 32 BINs, most of them undescribed species from Costa Rica.

Alphomelon arecaphile Deans, 2003

Alphomelon arecaphile Deans, 2003.

Type information. Holotype female, USNM (not examined but paratype examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Brazil (PA), Costa Rica.

Alphomelon brachymacher Deans, 2003

Alphomelon brachymacher Deans, 2003.

Type information. Holotype female, USNM (not examined but authoritatively identified specimens examined). Country of type locality: Colombia.

Geographical distribution. NEO.

NEO: Brazil (ES, MT, PA, SC), Colombia, Costa Rica, Ecuador, Peru.

Notes. The specimens we studied were identified by the author of the species.

Alphomelon brasiliensis Shimabukuro & Penteado-Dias, 2003

Alphomelon brasiliensis Shimabukuro & Penteado-Dias, 2003.

Type information. Holotype female, DCBU (not examined but original description checked). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (MG, SP, RS).

Alphomelon bromeliphile Deans, 2003

Alphomelon bromeliphile Deans, 2003.

Type information. Holotype female, USNM (not examined but paratype examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica, Mexico.

Alphomelon citroloma Deans, 2003

Alphomelon citroloma Deans, 2003.

Type information. Holotype female, USNM (not examined but paratype examined). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina, Bolivia, Brazil (PE, RJ, RO), Costa Rica, Ecuador, Panama, Paraguay, Trinidad & Tobago, Venezuela.

Alphomelon conforme (Muesebeck, 1958)

Apanteles conformis Muesebeck, 1958.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: Venezuela.

Geographical distribution. NEO.

NEO: Brazil (RJ), Costa Rica, Venezuela.

Notes. This species was transferred from Apanteles to Alphomelon by Deans et al. (2003), although the new combination was not clearly formalized (but is implicit, see pages 1 and 18 of that paper). Deans et al. (2003) did not change the ending of the species name to agree in gender with the generic name (Article 34.2 of the ICZN). The genus Alphomelon was described by Mason (1981) as neuter, but conformis is a masculine adjective, and thus it must be changed to the neuter form conforme. Until now, no published paper had ever referred to this species as Alphomelon conforme although Taxapad (Yu et al. 2012, 2016) correctly did so.

Alphomelon crocostethus Deans, 2003

Alphomelon crocostethus Deans, 2003.

Type information. Holotype female, USNM (not examined but paratype examined). Country of type locality: Jamaica.

Geographical distribution. NEO.

NEO: Bolivia, Brazil (ES, MG, RJ), Colombia, Jamaica, Puerto Rico.

Alphomelon disputabile (Ashmead, 1900), lectotype designation

Urogaster disputabilis Ashmead, 1900.

Type information. Lectotype male, NHMUK (examined). Country of type locality: Grenada.

Geographical distribution. NEA, NEO.

NEA: USA (KS, TX); NEO: Argentina, Belize, Bolivia, Brazil (ES, MT, PA, RJ, SC), Costa Rica, Cuba, Dominica, Ecuador, Grenada, Guatemala, Mexico, Panama, Paraguay, Puerto Rico, Saint Vincent, Trinidad & Tobago, Venezuela.

Notes. Ashmead (1900c: 286) did not designate a type in the original description of the species, which was based on 'several specimens'. Subsequent references to the species (e.g., Muesebeck 1921, Shenefelt 1972, Marsh et al. 1979) did not address that either. In the most complete nomenclatural account of the species (Shenefelt 1972: 494), it is implied that the type series, including both male and female specimens, was deposited in London (NHMUK), and could be from either Grenada or Saint Vincent. Much later Deans et al. (2003) mentioned that they had examined the holotype of the species, which they wrote was a male and was deposited in the USNM (with USNM type #6446). However, there cannot be a 'holotype' when Ashmead’s paper makes it clear that the species description was based on a series of specimens. From the Introduction section of the original paper (Ashmead 1900c: 207) it is also clear that the specimens studied were loaned to him from London (NHMUK). Thus, what likely happened was that, after studying the loaned material, Ashmead retained one specimen in Washington from the original type series and returned the rest to London. That means that the male specimen examined by Deans et al. (2003) in Washington is a syntype. The Washington specimen cannot be considered as the lectotype either, following ICZN Article 74.7 “Lectotype designation after 1999”, which clearly states that “To be valid, a lectotype designation made after 1999 must, 74.7.1. employ the term “lectotype” or an exact translation (e.g., “lectotypus” but not “the type”), 74.7.2. contain information sufficient to ensure recognition of the specimen designated, and 74.7.3. contain an express statement of deliberate designation (merely citing a specimen as “lectotype” is insufficient)”. For the sake of clarity, here we designate a male specimen as the lectotype [NHMUK, type number 3c.2395, specimen number 010636228, ‘St’ Vincent, | W.I. | H.H. Smith’, ‘W. Indies | 99-331.’]. There are an additional four paralectotype males in NHMUK, three from Grenada and one from St. Vincent, that from St. Vincent labelled by Ashmead as ‘Type male’ and with a yellow ‘co-type’ label. The specimen designated lectotype here is in better condition, albeit lacking its antennae.

Alphomelon melanoscelis Deans, 2003

Alphomelon melanoscelis Deans, 2003.

Type information. Holotype female, ESUW (not examined but paratype examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Belize, Brazil (AL, MT), Costa Rica, Mexico, Venezuela.

Alphomelon nanosoma Deans, 2003

Alphomelon nanosoma Deans, 2003.

Type information. Holotype female, USNM (not examined but authoritatively identified specimens examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Brazil (MT), Costa Rica, Ecuador, Mexico, Panama, Trinidad & Tobago.

Notes. The specimens we studied were identified by the author of the species.

Alphomelon nigriceps (Ashmead, 1900), lectotype designation

Urogaster nigriceps Ashmead, 1900.

Type information. Lectotype female, NHMUK (examined). Country of type locality: Saint Vincent.

Geographical distribution. NEA, NEO.

NEA: USA (FL, NC, TX); NEO: Argentina, Belize, Brazil (RO), Colombia, Cuba, Dominica, Grenada, Netherlands Antilles, Peru, Saint Lucia, Saint Vincent, Trinidad & Tobago, Venezuela.

Notes. Ashmead (1900c: 284) did not designate a type in the original description of the species, which was based on eight female specimens. Subsequent references to the species (e.g., Szépligeti 1904, Muesebeck 1921, Shenefelt 1972, Marsh et al. 1979) did not address that either. The most complete nomenclatural account of the species (Shenefelt 1972: 580) mentioned that the type series was in London (NHMUK), and a female specimen, with code 3c.1125 is referred to as the type. Much later, Deans et al. (2003) mentioned that they had examined the holotype of the species, which they wrote was a female and was deposited in the USNM (with USNM type #6443). Deans et al. (2003) probably overlooked Shenefelt’s account, but in any case, there cannot be a holotype when the original paper makes clear that it was a series of specimens. From the Introduction section of the original paper (Ashmead 1900c: 207) it is clear that the specimens studied were loaned to him from London (NHMUK). Thus, what likely happened was that, after studying the loaned material, Ashmead retained one specimen in Washington from the original type series and returned the rest to London. That means that the female specimen that Deans et al. (2003) saw in Washington is a syntype. We have seen in London the specimen referred to by Shenefelt (1972) with code 3c.1125. It is a female in good condition and, in addition to the standard type label from the NHMUK, it also has an additional, handwritten label that reads “Urogaster nigriceps, ♀ type, Ash.” For the sake of clarity, here we designate that female specimen as the lectotype; the female specimen examined by Deans et al. (2003) and deposited in the USNM, as well as the rest of the female specimens deposited in NHMUK are thus to be considered as paralectotypes.

Alphomelon paurogenum Deans, 2003

Alphomelon paurogenum Deans, 2003.

Type information. Holotype female, MCZ (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina, Chile.

Alphomelon pyrrhogluteum Deans, 2003

Alphomelon pyrrhogluteum Deans, 2003.

Type information. Holotype female, MCZ (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Alphomelon rhyssocercus Deans, 2003

Alphomelon rhyssocercus Deans, 2003.

Type information. Holotype female, CNC (examined). Country of type locality: Ecuador.

Geographical distribution. NEO.

NEO: Argentina, Costa Rica, Ecuador, Panama, Peru, Trinidad & Tobago, Venezuela.

Alphomelon rugosum Shimabukuro & Penteado-Dias, 2003

Alphomelon rugosum Shimabukuro & Penteado-Dias, 2003.

Type information. Holotype female, DCBU (not examined but original description checked). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (DF, SP).

Alphomelon simpsonorum Deans, 2003

Alphomelon simpsonorum Deans, 2003.

Type information. Holotype female, CNC (examined). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (PR, SC), Costa Rica, Paraguay.

Alphomelon talidicida (Wilkinson, 1931)

Apanteles talidicida Wilkinson, 1931.

Type information. Holotype female, NHMUK (examined). Country of type locality: Guyana.

Geographical distribution. NEO.

NEO: Belize, Brazil, Colombia, Costa Rica, Ecuador, Guyana, Mexico, Panama, Peru, Trinidad & Tobago, Venezuela.

Alphomelon winniewertzae Deans, 2003

Alphomelon winniewertzae Deans, 2003.

Type information. Holotype female, USNM (not examined but authoritatively identified specimens examined). Country of type locality: USA.

Geographical distribution. NEA, NEO.

NEA: Canada (ON, QC), USA (AR, DC, FL, KS, MA, MI, NC, OH, TN, TX, VA); NEO: Costa Rica, Mexico.

Notes. The specimens we studied were identified by the author of the species.

Alphomelon xestopyga Deans, 2003

Alphomelon xestopyga Deans, 2003.

Type information. Holotype female, USNM (not examined but paratype examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Genus Apanteles Foerster, 1863

Apanteles Foerster, 1863: 245. Gender: masculine. Type species: Microgaster obscurus Nees, 1834, by original designation and monotypy.

Urogaster Ashmead, 1898: 166. Type species: Urogaster vulgaris Ashmead, 1898, by subsequent designation (Viereck 1914).

Holcapanteles Cameron, 1905: 44. Type species: Holcapanteles sulciscutis Cameron, 1905, by monotypy. New synonymy.

Xestapanteles Cameron, 1910: 447. Type species: Xestapanteles latiannulatus Cameron, 1910, by monotypy.

Cecidobracon Kieffer & Jörgensen, 1910: 436. Type species: Cecidobracon asphondyliae Kieffer & Jörgensen, 1910, by monotypy. New synonymy.

Allapanteles Brèthes, 1915: 404. Type species: Allapanteles cecidiptae Brèthes, 1915, by monotypy.

The year of publication of Foerster’s paper, with the original description of Apanteles, was until recently almost universally cited as 1862 (e.g., Dalla Torre 1898, Szépligeti 1904, Shenefelt 1972, Marsh 1979a, Yu et al. 2012); however, it has been shown that the actual year of publication was 1863 (Foley et al. 2003), which has been followed by Yu et al. (2016) and it is also accepted here.

The type species of Holcapanteles is H. sulciscutis Cameron, 1905, from Indonesia. The holotype is apparently lost (Shenefelt 1973, van Achterberg 1980, Mason 1981). The type species of Cecidobracon is C. asphondyliae Kieffer & Jörgensen, 1910, from Argentina. Unfortunately, the type depository was never stated in the original description, and the specimen has not been located subsequently (Shenefelt 1973, Mason 1981). A second species, Cecidobracon braziliensis Kieffer & Tavares, 1925, was described from Brazil a few years later, but the type depository is also unknown. Without seeing the type specimens it may never be possible to establish with certainty the validity of Holcapanteles and Cecidobracon as Microgastrinae genera; however, based on the original descriptions, Mason (1981: 26, 27) considered that both genera were likely to be synonyms of Apanteles, although he did not formally synonymize the names. After reading the three original descriptions (Cameron 1905a: 44, Kieffer and Jörgensen 1910: 436–437, Kieffer and Tavares 1925: 48), including the associated illustrations of the wings of the two Cecidobracon species, we concur with Mason’s opinion and thus formally synonymize both genera under Apanteles for the sake of clarity and stability. The three species are also formally transferred below.

Currently Apanteles is the largest genus of Microgastrinae with 633 described species from all biogeographical regions (although, interestingly, there are no native species in New Zealand and the genus has not been recorded from the high Arctic). Several regional revisions are available, but some are very outdated and the taxonomic coverage of world species is far from complete. We have seen a large number of undescribed species in collections, mostly from tropical areas, and the actual species richness may well attain several thousand species. The name Apanteles was traditionally applied to all species with the fore wing areolet open: subsequently Apanteles auctt. has been split into numerous genera starting as early as 1880 and resulting in more than two dozen new genera being proposed since (see Mason 1981, Whitfield et al. 2002b, and Fernandez-Triana et al. 2014e for summaries of the history of Apanteles and its different concepts). van Achterberg (2003) synonymised several of these genera under Apanteles, thus potentially increasing the number of described species to 1,290 (Fig. 2A; see also Yu et al. 2016); however, we do not follow that arrangement here (Fig. 2B; also, see above, under the section Brief diagnosis of all Microgastrinae genera as they are understood in this paper, a more detailed discussion on the generic limits of the subfamily). Even with the restricted generic concept that we use in this paper, Apanteles is still a huge and varied assemblage of species. Nixon (1965) proposed 44 species groups for the world fauna (although that was before Mason (1981) split the genus, meaning some of those groups are not currently in Apanteles); and Fernandez-Triana et al. (2014e) proposed 30 new species groups just for Mesoamerica. Many of the Apanteles species groups represent monophyletic or at least morphologically cohesive groups, but others are poorly defined, and some are just containers for species that do not fit into any other group. Many families of Lepidoptera have been recorded as hosts for Apanteles, but many records are likely to be incorrect and/or need further verification. In Costa Rica most of the known hosts belong to three families: Crambidae, Depresariidae, and Hesperiidae (Fernandez-Triana et al. 2014e; in that paper depressarids were treated as elachistids). There are 7,800+ DNA-barcode compliant sequences of Apanteles in BOLD representing almost 600 different BINs, mostly from Costa Rica and North America.

Apanteles abdera Nixon, 1965

Apanteles abdera Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: Cape Verde, South Africa.

Apanteles abditus Muesebeck, 1957

Apanteles abditus Muesebeck, 1957.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (SP), Uruguay, Venezuela.

Apanteles acoris Nixon, 1965

Apanteles acoris Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles acutissimus Granger, 1949

Apanteles acutissimus Granger, 1949.

Type information. Syntypes female and male, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Notes. The original description mentions 15 female and 16 male specimens but does not explicitly designate a holotype, thus all are here considered to be syntypes.

Apanteles adelinamoralesae Fernandez-Triana, 2014

Apanteles adelinamoralesae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles adoxophyesi Minamikawa, 1954

Apanteles adoxophyesi Minamikawa, 1954.

Type information. Holotype female, depository unknown (not examined but authoritatively identified specimens examined). Country of type locality: Japan.

Geographical distribution. OTL, PAL.

OTL: China (ZJ); PAL: China (AH, SD), Japan.

Notes. Our concept of Apanteles adoxophyesi is based on two female specimens we examined (EIHU), presumably identified by Watanabe. The digital collection of TARI also contains images of this species, although we could not confirm the accuracy of that identification (https://digiins.tari.gov.tw/tarie/treelist003E.php?id=Brac11122001&lev1=3&lev2=0/1/7/&lev3=01&page=5).

Apanteles adreus Nixon, 1965

Apanteles adreus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles adrianachavarriae Fernandez-Triana, 2014

Apanteles adrianachavarriae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles adrianaguilarae Fernandez-Triana, 2014

Apanteles adrianaguilarae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles adrianguadamuzi Fernandez-Triana, 2014

Apanteles adrianguadamuzi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles afer Wilkinson, 1932

Apanteles afer Wilkinson, 1932.

Type information. Holotype female, NHMUK (examined). Country of type locality: Uganda.

Geographical distribution. AFR.

AFR: Uganda.

Apanteles agatillus Nixon, 1965

Apanteles agatillus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles aglaope Nixon, 1965

Apanteles aglaope Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: Indonesia.

Apanteles aglaus Nixon, 1965

Apanteles aglaus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Fiji.

Geographical distribution. AUS.

AUS: Fiji.

Apanteles agrus Nixon, 1965

Apanteles agrus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles aichagirardae Fernandez-Triana, 2014

Apanteles aichagirardae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles aidalopezae Fernandez-Triana, 2014

Apanteles aidalopezae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles alaspharus Nixon, 1965

Apanteles alaspharus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles alastor de Saeger, 1944

Apanteles alastor de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo.

Apanteles alazoni Lozan, 2008

Apanteles alazoni Lozan, 2008.

Type information. Holotype female, IECA (not examined but original description checked). Country of type locality: Canary Islands.

Geographical distribution. PAL.

PAL: Canary Islands.

Apanteles albanjimenezi Fernandez-Triana, 2014

Apanteles albanjimenezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles albinervis (Cameron, 1904)

Urogaster albinervis Cameron, 1904.

Apanteles albinervicam Shenefelt, 1972.

Type information. Holotype male, NHMUK (examined). Country of type locality: Mexico.

Geographical distribution. NEO.

NEO: Mexico.

Apanteles alejandromasisi Fernandez-Triana, 2014

Apanteles alejandromasisi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles alejandromorai Fernandez-Triana, 2014

Apanteles alejandromorai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles alexanderi Brèthes, 1922

Apanteles alexanderi Brèthes, 1922.

Type information. Lectotype female, MACN (not examined but subsequent treatment of the species checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina, Uruguay.

Notes. Our concept of Apanteles alexanderi is based on Martinez et al. (2012), who examined and designated the lectotype, and provided images and DNA barcodes of the species.

Apanteles allofulvigaster Long, 2007

Apanteles allofulvigaster Long, 2007.

Type information. Holotype female, VNMN (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Notes. The holotype depository was not stated in the English version of the original description (Long 2007). That paper was written in two languages, the first part in Vietnamese, followed by a second part in English; based on the extent of both versions, we suspect that the English part is just a translation from the Vietnamese. However, we do not know if it is a literal translation or just a summarized (= shorter) version; thus, we do not know if the holotype depository is mentioned in the Vietnamese part of the paper. If the holotype was not stated in the Vietnamese version, then this species name would be unavailable (a subsequent paper (Long and Achterberg 2014) records the holotype depository; however, that alone does not comply with the ICZN requirements and would not make the name available). Although we have not been able to establish with certainty what is stated in the Vietnamese part of Long (2007), we provisionally consider here the species name as available.

Apanteles alvarougaldei Fernandez-Triana, 2014

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles anabellecordobae Fernandez-Triana, 2014

Apanteles anabellecordobae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles anamarencoae Fernandez-Triana, 2014

Apanteles anamarencoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles anamartinezae Fernandez-Triana, 2014

Apanteles anamartinezae Fernandez-Triana, 2014.

Apanteles anamartinesae Fernandez-Triana, 2014 [incorrect original spelling].

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Notes. In the paper where this species was originally described, the name was spelled in two different ways: as anamartinezae (in the species list of Table 3, species description, references to ZooBank and caption of Figure 227) or as anamartinesae (in the Abstract, key to species, and caption to Figure 25). The correct spelling is obviously anamartinezae, as the species was named after Ana Martínez, and it is the one to be preserved, following Article 32 of the ICZN.

Apanteles anariasae Fernandez-Triana, 2014

Apanteles anariasae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles anatole Nixon, 1965

Apanteles anatole Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Notes. The holotype specimen has the vannal lobe with very few, very sparse setae across lobe length.

Apanteles andreacalvoae Fernandez-Triana, 2014

Apanteles andreacalvoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles angaleti Muesebeck, 1956

Apanteles angaleti Muesebeck, 1956.

Type information. Holotype female, USNM (examined). Country of type locality: India.

Geographical distribution. AFR, OTL, PAL.

AFR: Kenya; OTL: China (SN, ZJ), India, Indonesia, Pakistan, Vietnam; PAL: Iraq.

Notes. Introduced into Mexico and the USA (e.g., Mcgough and Noble 1957, Bartlett et al. 1978). In total more than 150,000 specimens were released but the species was never recovered in any of the USA states where it was released (Mcgough and Noble 1957), and a subsequent citation of the species from Mexico (Coronado-Blanco et al. 2004) is merely a repetition of the information cited in older references, not a confirmation of the species’ presence in the country. Thus, in this paper we do not consider A. angaleti as an established species in the Nearctic or Neotropical regions.

Apanteles angelsolisi Fernandez-Triana, 2014

Apanteles angelsolisi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles angulatus Granger, 1949

Apanteles angulatus Granger, 1949.

Type information. Syntypes female and male, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Apanteles angustibasis Gahan, 1925

Apanteles angustibasis Gahan, 1925.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: China (HN), India, Malaysia, Pakistan, Philippines, Vietnam.

Notes. This species was transferred to Cotesia by Gupta and Pawar (1992), a non-taxonomic paper, in which it could be argued that those authors did not study the holotype. We have studied the holotype as well as illustrations of specimens from Malaysia identified by C. Watanabe that are deposited in EIHU. Both the holotype and the Malaysian specimens are clearly not Cotesia but Apanteles, and thus we restore the combination of this species here.

Apanteles anodaphus Nixon, 1965

Apanteles anodaphus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Papua New Guinea.

Geographical distribution. AUS.

AUS: Papua New Guinea.

Apanteles ansata Song & Chen, 2004

Apanteles ansata Song & Chen, 2004.

Type information. Holotype female, FAFU (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ).

Apanteles anthozelae de Saeger, 1941

Apanteles anthozelae de Saeger, 1941.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo.

Apanteles anticlea Nixon, 1965

Apanteles anticlea Nixon, 1965

Type information. Holotype female, USNM (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia.

Apanteles antilla Nixon, 1965

Apanteles antilla Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles arachidis Risbec, 1951

Apanteles arachidis Risbec, 1951.

Type information. Holotype male, MNHN (not examined but original description checked). Country of type locality: Senegal.

Geographical distribution. AFR.

AFR: Senegal.

Notes. The original description is not clear enough to determine the correct generic placement of the species, thus is best kept in the genus it was originally described. Future study of the type specimen may change its current generic status.

Apanteles araeceri Wilkinson, 1928

Apanteles araeceri Wilkinson, 1928.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: India, Indonesia, Malaysia.

Apanteles aragatzi Tobias, 1976

Apanteles aragatzi Tobias, 1976.

Type information. Holotype female, depository unknown (not examined but subsequent treatment of the species checked). Country of type locality: Armenia.

Geographical distribution. PAL.

PAL: Armenia, Russia (KDA), Sweden, Turkey.

Notes. Our concept of the species is based on the descriptions provided by Papp (1984a) and Tobias (1986).

Apanteles arielopezi Fernandez-Triana, 2014

Apanteles arielopezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles arion Nixon, 1965

Apanteles arion Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles ariovistus Nixon, 1965

Apanteles ariovistus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: Indonesia.

Apanteles aristaeus Nixon, 1965

Apanteles aristaeus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: China (TW), India, Indonesia.

Apanteles aristoteliae Viereck, 1912

Apanteles aristoteliae Viereck, 1912.

Apanteles gelechiae Viereck, 1912.

Type information. Holotype male, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (NB, ON, QC), USA (AZ, CA, CO, CT, KS, LA, MI, NJ, NY, NC, OH, OR, PA, TX, UT, VT, WA).

Apanteles arsanes Nixon, 1965

Apanteles arsanes Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Kenya.

Geographical distribution. AFR.

AFR: Kenya.

Notes. Despite its relatively short ovipositor sheaths, we are retaining this species in Apanteles because of its pleated hypopygium, strongly concave vannal lobe lacking setae, and anteromesoscutum punctures which are fusing near scutoscutellar disc.

Apanteles articas Nixon, 1965

Apanteles articas Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Senegal.

Geographical distribution. AFR, PAL.

AFR: Senegal; PAL: Israel, Tunisia, Turkey.

Apanteles artustigma Liu & Chen, 2015

Apanteles artustigma Liu & Chen, 2015.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (GD, ZJ).

Apanteles arundinariae de Saeger, 1944

Apanteles arundinariae de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Rwanda.

Apanteles asphondyliae (Kieffer & Jörgensen, 1910), new combination

Cecidobracon asphondyliae Kieffer & Jörgensen, 1910.

Type information. Holotype male, lost (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Notes. The type depository was not stated in the original description, and the specimen has never been located (Shenefelt 1973, Mason 1981). See comments at the beginning of Apanteles for details on the decision to transfer this species to Apanteles.

Apanteles assis Nixon, 1965

Apanteles assis Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines, Vietnam.

Apanteles atrocephalus Granger, 1949

Apanteles atrocephalus Granger, 1949.

Type information. Holotype female, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Notes. Based on some morphological features described by Granger (1949), e.g., the areolated propodeum, shape and sculpture of T1–T3, acute hypopygium, ovipositor sheaths half the metatibia length, we think that this species could potentially be placed in one of the following genera: Apanteles, Parapanteles, or Cotesia. Because the original description (the only source available, apart from the single known specimen, which we could not examine), is not sufficient to determine the correct generic placement, we maintain atrocephalus within the genus in which it was originally described.

Apanteles attevae Yousuf, Hassan & Singh, 2008

Apanteles attevae Yousuf, Hassan & Singh, 2008.

Type information. Holotype female, TFRI (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles audens Kotenko, 1986

Apanteles audens Kotenko, 1986.

Type information. Holotype female?, ZIN (not examined but original description checked). Country of type locality: Georgia.

Geographical distribution. PAL.

PAL: Georgia, Russia (NC).

Notes. The paper in which the original description is included does not clarify the sex of the type material, nor is it specified if there is a holotype (or syntypes) on which the species description was based (Tobias 1986: 805). Without examining the actual specimen(s) is impossible to determine its sex or type status; however, in the Foreword section of the paper (Tobias 1986: page numbered as ix) it is stated that, to comply with nomenclature rules, the type material is specified for all species. The author then explicitly says that the paper includes lectotype and paralectotype designations for species described from the USSR in the past. Such a statement allows the assumption that all new species descriptions must have been based on holotypes – and not a type series (syntypes) as was presumably done in the past. Thus, we are assuming that there is a holotype for Apanteles audens Kotenko, 1986. Regarding the sex of the type, again only assumptions can be made until the specimen is examined, but the key is based on female specimens, including a brief original description that mentions the ovipositor sheaths. Thus, we consider here as very likely that the holotype is a female but add a question mark to clarify that it is only an educated guess.

Apanteles aurangabadensis Rao & Chalikwar, 1970

Apanteles aurangabadensis Rao & Chalikwar, 1970.

Type information. Holotype male, NZSI (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles azollae Sumodan & Sevichan, 1989

Apanteles azollae Sumodan & Sevichan, 1989.

Type information. Holotype female, RMNH (not examined but subsequent treatment of the species checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Notes. See van Achterberg and Narendran (1997) for details about the type, and for the generic placement of the species. Apanteles azollae has been misspelled twice, as azolae and azolla, as previously noted by Yu et al. (2016).

Apanteles bajariae Papp, 1975

Apanteles bajariae Papp, 1975.

Type information. Holotype female, HNHM (not examined but original description checked). Country of type locality: Hungary.

Geographical distribution. PAL.

PAL: Bulgaria, Canary Islands, Greece, Hungary, Montenegro, Turkey.

Notes. Based on the position this species occupies in the key of Papp (1984a), it is possible that bajariae would actually belong to Dolichogenidea. However, the details in both the original description and Papp (1984a) are not definite to conclude with certainty, thus it is here kept in the genus it was originally described.

Apanteles baldufi Muesebeck, 1968

Apanteles baldufi Muesebeck, 1968.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (ON), USA (MI, MN).

Apanteles balteatae Lal, 1942

Apanteles balteatae Lal, 1942.

Type information. Holotype male, INPC (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles balthazari (Ashmead, 1900)

Urogaster balthazari Ashmead, 1900.

Urogaster meridionalis Ashmead, 1900.

Apanteles meridionalis Ashmead, 1900.

Type information. Holotype female, NHMUK (examined). Country of type locality: Saint Vincent.

Geographical distribution. NEO.

NEO: Brazil (CE, PA, PB, PE, RN, SP), Cuba, Grenada, Saint Vincent.

Notes. The original description (Ashmead 1900c) does not match the holotype, as his description of the T1 shape, T2 sculpture and colouration of meso- and metafemora are completely different from the actual specimen examined (see Fernandez-Triana et al. 2014e).

Apanteles bannaensis Song, Chen & Yang, 2001

Apanteles bannaensis Song, Chen & Yang, 2001.

Type information. Holotype female, FAFU (not examined but subsequent treatment of the species checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (YN).

Notes. Our species concept is based on Chen and Song (2004).

Apanteles baoli Risbec, 1951

Apanteles baoli Risbec, 1951.

Type information. Holotype male, depository unknown (not examined but original description checked). Country of type locality: Senegal.

Geographical distribution. AFR.

AFR: Senegal.

Apanteles basicavus Liu & Chen, 2015

Apanteles basicavus Liu & Chen, 2015.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. PAL.

PAL: China (JL, LN).

Apanteles bellatulus de Saeger, 1944

Apanteles bellatulus de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo.

Apanteles bernardoespinozai Fernandez-Triana, 2014

Apanteles bernardoespinozai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles bernyapui Fernandez-Triana, 2014

Apanteles bernyapui Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles bettymarchenae Fernandez-Triana, 2014

Apanteles bettymarchenae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles bienvenidachavarriae Fernandez-Triana, 2014

Apanteles bienvenidachavarriae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles biroicus Papp, 1973

Apanteles biroicus Papp, 1973.

Type information. Holotype female, HNHM (not examined but paratype examined). Country of type locality: Hungary.

Geographical distribution. PAL.

PAL: Hungary, Romania, Tunisia.

Notes. This species was transferred from Apanteles to Illidops by Papp (1988), but examination of two paratype specimens in the CNC revealed that those specimens do not have a median band of rugosity posteriorly on the scutellum, and the propodeum sculpture is also different from that found in Illidops (sensu Fernandez-Triana et al. 2014e). Thus, here we transfer the species back to Apanteles.

Apanteles bitalensis de Saeger, 1944

Apanteles bitalensis de Saeger, 1944.

Type information. Syntypes female, RMCA (not examined but original description checked). Country of type locality: Rwanda.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Rwanda.

Apanteles bordagei Giard, 1898

Apanteles bordagei Giard, 1898.

Type information. Type lost (not examined but original description checked). Country of type locality: Réunion.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Kenya, Réunion, Tanzania.

Notes. The year of description for this species has been incorrectly cited as 1902 by most authors (e.g., Granger 1949, Shenefelt 1972, Rousse and Gupta 2013, Yu et al. 2016), in all cases based on Giard (1902: 22). Having read that paper, it is clear that it only refers to the species as being described by the author in a previous work (Giard 1898: 202, which we have also read). This was correctly mentioned by de Saeger (1944: 316) and Wilkinson (1934: 150). Wilkinson comprehensively redescribed the species, based on specimens from Kenya and Tanzania, and he considered the type(s) to be lost based on his enquiry to a curator of the MNHN at the time, who could not find the specimen(s). Subsequent authors have provided shorter redescriptions, based on specimens from Democratic Republic of Congo (de Saeger 1944), Madagascar (Granger 1949), or Réunion (Rousse and Gupta 2013). Our species concept is based on Wilkinson (1934). We accept the following comments from Madl and van Achterberg (2014): “Known from the Afrotropical region. The record from Madagascar mentioned in Risbec (1960: 629) is doubtful. Brénière (1965b: 347) mentions Apanteles bordagei from Madagascar, citing Granger (1949: 359) as reference, but Granger recorded this species only from Réunion and Africa. The record from Madagascar in Appert et al. (1969: 568) is based on Brénière (1965b)”. Consequently, here we do not consider Madagascar as a country where this species is found.

Apanteles brachmiae Bhatnagar, 1950

Apanteles brachmiae Bhatnagar, 1950.

Type information. Holotype female, INPC (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Notes. The year of publication of the Bhatnagar paper was until recently commonly cited as 1948 and/or 1950 (e.g., Chen and Song 2004, Yu et al. 2016), probably following Shenefelt (1972) who referred to this paper as “Bhatnagar (1948) 1950”. While the intended year for Volume X, Parts I & II of the Indian Journal of Entomology was 1948, the actual dates of publication were June 1950 (Part I) and October 1950 (Part II), as clearly shown on the cover page of the Volume, which we have checked. Because the dates of publication are the ones to be considered, and for the sake of clarity, we hereby revise the species year of description to 1950.

Apanteles braziliensis (Kieffer & Tavares, 1925), new combination

Cecidobracon braziliensis Kieffer & Tavares, 1925.

Type information. Type and depository unknown (not examined but original description checked). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (BA).

Notes. The type depository was not given in the original description, and the specimen has never been located (Shenefelt 1973, Mason 1981). See comments at the beginning of Apanteles for details on the decision to transfer this species to Apanteles (p 74, 75).

Apanteles bredoi de Saeger, 1941

Apanteles bredoi de Saeger, 1941.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Senegal.

Apanteles brethesi Porter, 1917

Apanteles brethesi Porter, 1917.

Type information. Type and depository unknown (not examined). Country of type locality: Chile.

Geographical distribution. NEO.

NEO: Chile.

Apanteles brevicarinis Song, 2002

Apanteles brevicarinis Song, 2002.

Type information. Holotype female, FAFU (not examined but subsequent treatment of the species checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (HB).

Notes. Our concept of this species is based on Chen and Song (2004).

Apanteles brevimetacarpus Hedqvist, 1965

Apanteles brevimetacarpus Hedqvist, 1965.

Illidops metacarpus Hedqvist, 1965 [subsequent misspelling (Papp 2003)].

Type information. Holotype female, MZH (examined). Country of type locality: Cape Verde.

Geographical distribution. AFR, PAL.

AFR: Cape Verde; PAL: Tunisia.

Notes. Papp (2003: 145) transferred this species to Illidops (although he misspelled the species name as metacarpus). A subsequent paper, also treating the species and reporting it for the first time from Tunisia, continued to place it within Illidops (Papp 2014). We examined the female holotype and a male paratype, and they clearly are not Illidops. The only feature that would suggest placement in that genus is the short vein R1 (metacarp), but that is known in several species of both Apanteles and Dolichogenidea. The posteromedian band of the scutellum is smooth. The propodeum, although without an areola, has a weak impression in its place, and its overall weak sculpture is not like that found in Illidops. Based on the hind wing, with a slightly concave vannal lobe lacking setae, the best generic placement for this species is Apanteles. This concurs with Forshage et al. (2016), although those authors were probably not aware of the two papers by Papp and were following the treatment of the original description. In any case, the statement by Forshage et al. (2016) that the holotype and paratype were missing is here updated, as in 2018 we found the specimens in the MZH.

Apanteles brevivena Liu & Chen, 2015

Apanteles brevivena Liu & Chen, 2015.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. PAL.

PAL: China (XJ, LN, JL, NM, SD).

Apanteles bruchi Blanchard, 1941

Apanteles bruchi Blanchard, 1941.

Type information. Type lost (not examined but subsequent treatment of the species checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina, Peru.

Notes. Our concept of this species is based on Aquino et al. (2010), including details on the fate of the type material.

Apanteles brunnistigma Abdinbekova, 1969

Apanteles brunnistigma Abdinbekova, 1969.

Apanteles sotades Nixon, 1976.

Type information. Holotype female, ZIN (not examined but authoritatively identified specimens examined). Country of type locality: Azerbaijan.

Geographical distribution. NEA, PAL.

NEA: Canada (MB, NL, NT, ON, YT); PAL: Azerbaijan, Canary Islands, Czech Republic, Finland, France, Germany, Hungary, Iran, Italy, Korea, Lithuania, Russia (ZAB, PRI, TOM), Sweden, Switzerland, Turkey, United Kingdom, Ukraine.

Notes. Our concept of this species is based on Fernandez-Triana et al. (2014c). We have also examined the type of Apanteles sotades Nixon. New data from specimens with sequences in BOLD expand the species distribution within the Nearctic (northwestern Canada) as well as the Palearctic (Germany, Ukraine).

Apanteles brunnus Rao & Chalikwar, 1976

Apanteles brunnus Rao & Chalikwar, 1976.

Type information. Holotype female, BAMU (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles burunganus de Saeger, 1944

Apanteles burunganus de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo.

Notes. The original description does not provide enough detail to place this species in a genus unambiguously (it could be Apanteles but also Dolichogenidea). Until the type series is studied, we retain it in the genus in which it was originally described.

Apanteles caesar Wilkinson, 1938

Apanteles caesar Wilkinson, 1938.

Type information. Holotype female, NHMUK (examined). Country of type locality: Namibia.

Geographical distribution. AFR.

AFR: Namibia, South Africa.

Notes. This species bears some resemblance to the two described species currently placed within Napamus. It shares with them the dark colour, infumate wings, elongate mouth parts (especially very long glossa and galea), and relatively short fore wing vein R1 (although not as short as in the two described Napamus). However, we retain caesar within Apanteles because it has some differences in propodeum sculpture (which is mostly smooth, having only small carinae near the nucha), metatibial spines (which are not as long as in Napamus) and the disparate geographic distribution of the known species.

Apanteles calixtomoragai Fernandez-Triana, 2014

Apanteles calixtomoragai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles calycinae Wilkinson, 1928

Apanteles calycinae Wilkinson, 1928.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: India, Vietnam.

Apanteles camilla Nixon, 1965

Apanteles camilla Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles camirus Nixon, 1965

Apanteles camirus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles canarsiae Ashmead, 1898

Apanteles canarsiae Ashmead, 1898.

Apanteles housatannuckorum Viereck, 1917.

Apanteles maquinnai Viereck, 1917.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (ON, QC), USA (AR, CT, DC, IL, IN, IA, KS, NY, VA).

Notes. We examined the holotype female of housatannuckorum and the holotype male of maquinnai, both currently considered as synonyms of A. canarsiae. All three holotypes are in the USNM and not in INHS as stated in Yu et al. (2016).

Apanteles carloscastilloi Fernandez-Triana, 2014

Apanteles carloscastilloi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carlosguadamuzi Fernandez-Triana, 2014

Apanteles carlosguadamuzi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carlosrodriguezi Fernandez-Triana, 2014

Apanteles carlosrodriguezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carlosviquezi Fernandez-Triana, 2014

Apanteles carlosviquezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carloszunigai Fernandez-Triana, 2014

Apanteles carloszunigai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carolinacanoae Fernandez-Triana, 2014

Apanteles carolinacanoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles carpatus (Say, 1836)

Microgaster carpata Say, 1836.

Urogaster solitarius Ashmead, 1900.

Protapanteles hawaiiensis Ashmead, 1901.

Urogaster fuscicornis Cameron, 1910.

Apanteles piceoventris Muesebeck, 1921.

Apanteles igae Watanabe, 1932.

Apanteles sarcitorius Telenga, 1955.

Apanteles ultericus Telenga, 1955.

Type information. Holotype female, lost (not examined but subsequent treatment of the species checked). Country of type locality: USA.

Geographical distribution. AFR, AUS, NEA, NEO, OTL, PAL.

AFR: Democratic Republic of Congo, Ghana, Mozambique, South Africa, Tanzania; AUS: Australia (QLD), Fiji, Hawaiian Islands, New Zealand; NEA: Canada (AB, BC, NB, NL, ON, PE, QC, SK), USA (CO, CT, DE, IL, IN, MD, MA, MI, MO, NH, NJ, NY, SC, TX, VA); NEO: Argentina, Bermuda, Brazil (SP), Cuba, Grenada, Peru, Puerto Rico; OTL: China (SN, TW, ZJ), Malaysia, Vietnam; PAL: Armenia, Croatia, Finland, France, Germany, Greece, Hungary, Iran, Israel, Japan, Kazakhstan, Latvia, Lithuania, Malta, Moldova, Mongolia, Poland, Romania, Russia (AMU, AST, KHA, PRI, SAK), Serbia, Spain, Switzerland, Turkey, Turkmenistan, United Kingdom, Uzbekistan.

Notes. We examined the types of two of the seven currently accepted synonyms of carpatus: hawaiiensis (in USNM) and solitarius (in NHMUK). If Apanteles carpatus is ever going to be split into several species, the type of hawaiiensis would be a candidate to be considered as a different species, supported by morphological differences when compared to other Apanteles carpatus specimens and also through different host associations. We also examined one female (in EIHU, identified by Muesebeck) which also looks different to the traditional carpatus and could represent yet another species. Determining the limits of A. carpatus is beyond the scope of this paper and at present we leave all of the examined specimens as a single species.

Apanteles cassiae Chalikwar & Rao, 1982

Apanteles cassiae Chalikwar & Rao, 1982.

Type information. Type and depository unknown (not examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles cato de Saeger, 1944

Apanteles cato de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Rwanda.

Apanteles cavatiptera Chen & Song, 2004

Apanteles cavatiptera Chen & Song, 2004.

Type information. Holotype female, FAFU (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ, YN).

Apanteles cavatithoracicus Chen, 2001

Apanteles cavatithoracica Chen, 2001.

Type information. Holotype female, FAFU (not examined but subsequent treatment of the species checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ, HB).

Notes. For the generic placement of this species we follow Chen and Song (2004).

Apanteles cavifrons Nixon, 1965

Apanteles cavifrons Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles cebes Nixon, 1965

Apanteles cebes Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles cecidiptae (Brèthes, 1916)

Allapanteles cecidiptae Brèthes, 1916.

Type information. Syntypes female and male, MACN (not examined). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles cerberus Nixon, 1965

Apanteles cerberus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles cestius Nixon, 1965

Apanteles cestius Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles chalcomelas Nixon, 1965

Apanteles chalcomelas Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles changhingensis Chu, 1937

Apanteles changhingensis Chu, 1937.

Type information. Holotype female, depository unknown (not examined but subsequent treatment of the species checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ, ZJ).

Notes. For the generic placement of this species we follow Chen and Song (2004).

Apanteles characomae Risbec, 1951

Apanteles characomae Risbec, 1951.

Type information. Holotype male, depository unknown (not examined but original description checked). Country of type locality: Ivory Coast.

Geographical distribution. AFR.

AFR: Ivory Coast.

Apanteles chatterjeei Sharma & Chatterjee, 1970

Apanteles chatterjeei Sharma & Chatterjee, 1970.

Type information. Holotype female, IFRI (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles chloris Nixon, 1965

Apanteles chloris Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines, Vietnam.

Apanteles christianzunigai Fernandez-Triana, 2014

Apanteles christianzunigai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles cingulicornis Granger, 1949

Apanteles cingulicornis Granger, 1949.

Type information. Syntypes female, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Apanteles cinthiabarrantesae Fernandez-Triana, 2014

Apanteles cinthiabarrantesae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles ciriloumanai Fernandez-Triana, 2014

Apanteles ciriloumanai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles clita Nixon, 1965

Apanteles clita Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: China (FJ), India, Vietnam.

Apanteles cockerelli Muesebeck, 1921

Apanteles cockerelli Muesebeck, 1921.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: USA.

Geographical distribution. NEA.

NEA: USA (CA, ID, IA, MI, MO, NE, NM, OR, SD, TX).

Apanteles cocotis Wilkinson, 1934

Apanteles cocotis Wilkinson, 1934.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: Indonesia, Vietnam.

Apanteles coedicius Nixon, 1965

Apanteles coedicius Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines, Vietnam.

Apanteles coffeellae Muesebeck, 1958

Apanteles coffeellae Muesebeck, 1958.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: Guadeloupe.

Geographical distribution. NEO.

NEO: Guadeloupe, Puerto Rico.

Apanteles coilus Nixon, 1965

Apanteles coilus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles conanchetorum Viereck, 1917

Apanteles conanchetorum Viereck, 1917.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (NS, ON), USA (AR, CT, DC, IL, IA, KS, MI, MO, NY, OH, PA, SC, SD, WV, WI).

Notes. Specimens of Apanteles conanchetorum that have rendered DNA barcodes comprise two BINS: BOLD:AAC5506 (eastern North America) and BOLD:AAC5507 (principally western Canada, but some records from ON, PE). Whether they represent two different species or not has been mentioned in the past (Fernandez-Triana et al. 2014b), but for the time being all known specimens are kept as one species.

Apanteles concordalis Cameron, 1911

Apanteles concordalis Cameron, 1911.

Type information. Holotype female, NHMUK (examined). Country of type locality: Guyana.

Geographical distribution. NEO.

NEO: Guyana, Peru.

Notes. Based on the carination and sculpture pattern of propodeum and fore wing venation, this species belongs to the leucostigmus group (sensu Fernandez-Triana et al. 2014e).

Apanteles conon Nixon, 1965

Apanteles conon Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL, PAL.

OTL: China (HB, TW), Indonesia, Philippines; PAL: Korea.

Notes. It is possible this is actually a Dolichogenidea species. The hind wing vannal lobes are not clearly visible (they are folded in both wings in the holotype) but what can be seen suggests they may be setose. Additionally, the anteromesoscutum punctures near the scutoscutellar sulcus do not fuse. However, because we cannot see the vannal lobe clearly, we refrain from transferring the species in this paper.

Apanteles conspicabilis de Saeger, 1944

Apanteles conspicabilis de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Rwanda.

Apanteles contactus Papp, 1977

Apanteles contactus Papp, 1977.

Type information. Holotype female, HNHM (not examined but original description checked). Country of type locality: Mongolia.

Geographical distribution. PAL.

PAL: Mongolia, Russia (ZAB).

Apanteles contaminatus (Haliday, 1834)

Microgaster contaminatus Haliday, 1834.

Type information. Neotype female, NHMUK (examined). Country of type locality: United Kingdom.

Geographical distribution. PAL.

PAL: Ireland, Italy, Netherlands, United Kingdom.

Apanteles contemptus Nixon, 1965

Apanteles contemptus Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Singapore.

Geographical distribution. OTL.

OTL: Singapore.

Apanteles cordoi de Santis, 1980

Apanteles cordoi de Santis, 1980.

Type information. Holotype female, MLP (not examined but subsequent treatment of the species checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Notes. Our concept of this species is based on Aquino et al. (2010).

Apanteles cornicula Chen & Song, 2004

Apanteles cornicula Chen & Song, 2004.

Type information. Holotype female, FAFU (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ).

Apanteles cosmopterygivorus Liu & Chen, 2014

Apanteles cosmopterygivorus Liu & Chen, 2014.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (ZJ).

Apanteles coxalis Szépligeti, 1911

Apanteles coxalis Szépligeti, 1911.

Type information. Holotype female, ZMHB (not examined but subsequent treatment of the species checked). Country of type locality: Tanzania.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Malawi, Senegal, Tanzania.

Notes. Our species concept is based on the redescription provided by Wilkinson (1932a), who was able to study the holotype.

Apanteles crassicornis (Provancher, 1886)

Microgaster crassicornis Provancher, 1886.

Microgaster crassicornis Provancher [primary junior homonym of Microgaster crassicornis Ruthe, 1860].

Type information. Lectotype female, ULQC (examined). Country of type locality: Canada.

Geographical distribution. NEA.

NEA: Canada (AB, ON, QC, SK), USA (AZ, IL, IN, IA, KS, MD, MA, MI, MN, MO, NJ, NY, OH, PA).

Apanteles crates Nixon, 1965

Apanteles crates Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines, Vietnam.

Apanteles crispulae Blanchard, 1943

Apanteles crispulae Blanchard, 1943.

Type information. Holotype female, MACN (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles cristianalemani Fernandez-Triana, 2014

Apanteles cristianalemani Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles crius Nixon, 1965

Apanteles crius Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Notes. The ovipositor sheaths are short, and the hypopygium has only one median fold (not pleated), similar to that of Pholetesor. However, the species otherwise resembles Apanteles and thus we have decided to maintain this species in the latter genus.

Apanteles croceicornis Muesebeck, 1958

Apanteles croceicornis Muesebeck, 1958.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: Peru.

Geographical distribution. NEO.

NEO: Peru.

Apanteles crocidolomiae Ahmad, 1945

Apanteles crocidolomiae Ahmad, 1945.

Type information. Holotype female, INPC (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles crouzeli Blanchard, 1947

Apanteles crouzeli Blanchard, 1947.

Apanteles crouzelae de Santis, 1967 [unjustified emendation].

Type information. Holotype female, MACN (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles cuneiformis Song & Chen, 2004

Apanteles cuneiformis Song & Chen, 2004.

Type information. Holotype female, FAFU (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (FJ, YN).

Apanteles curvicaudatus Granger, 1949

Apanteles curvicaudatus Granger, 1949.

Type information. Syntypes female and male, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Apanteles cynthiacorderoae Fernandez-Triana, 2014

Apanteles cynthiacorderoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles cyprioides Nixon, 1965

Apanteles cyprioides Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. AFR, OTL.

AFR: South Africa; OTL: China (FJ, HN), Philippines, Singapore.

Apanteles cypris Nixon, 1965

Apanteles cypris Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL, PAL.

OTL: Bangladesh, China (FJ, GD, GX, GZ, HI, HK, HB, HN, JS, JX, SH, SN, TW, YN, ZJ), India, Indonesia, Malaysia, Nepal, Pakistan, Philippines, Singapore, Sri Lanka, Vietnam; PAL: China (AH, HA, SN, SD), Japan.

Apanteles daimenes Nixon, 1965

Apanteles daimenes Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Fiji.

Geographical distribution. AUS.

AUS: Fiji.

Apanteles dakotae Muesebeck, 1921

Apanteles dakotae Muesebeck, 1921.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: USA.

Geographical distribution. NEA.

NEA: USA (ID, SD).

Apanteles decoloratus Granger, 1949

Apanteles decoloratus Granger, 1949.

Type information. Syntypes female, MNHN (not examined but original description checked). Country of type locality: Madagascar.

Geographical distribution. AFR.

AFR: Madagascar.

Apanteles deifiliadavilae Fernandez-Triana, 2014

Apanteles deifiliadavilae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles delhiensis Muesebeck & Subba Rao, 1958

Apanteles delhiensis Muesebeck & Subba Rao, 1958.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles dentatus Muesebeck, 1958

Apanteles dentatus Muesebeck, 1958.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: Brazil.

Geographical distribution. NEO.

NEO: Brazil (SP).

Apanteles deplanatus Muesebeck, 1957

Apanteles deplanatus Muesebeck, 1957.

Type information. Holotype female, USNM (not examined but subsequent treatment of the species checked). Country of type locality: Mexico.

Geographical distribution. NEO.

NEO: Mexico.

Notes. Our species concept is based on Austin and Dangerfield (1989) and Fernandez-Triana et al. (2014e).

Apanteles depressariae Muesebeck, 1931

Apanteles depressariae Muesebeck, 1931.

Type information. Holotype female, USNM (not examined but original description checked). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (NS, ON, QC), USA (IA, ME, MA, VT).

Apanteles derivatus Long, 2010

Apanteles derivatus Long, 2010.

Type information. Holotype female, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Apanteles desantisi Blanchard, 1947

Apanteles desantisi Blanchard, 1947.

Type information. Holotype female, MACN (not examined but original description checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles despectus Nixon, 1965

Apanteles despectus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Thailand.

Geographical distribution. OTL.

OTL: Thailand.

Apanteles diatraeae Muesebeck, 1921

Apanteles diatraeae Muesebeck, 1921.

Type information. Holotype female, USNM (not examined but subsequent treatment of the species checked). Country of type locality: Cuba.

Geographical distribution. NEA, NEO.

NEA: USA (AZ, FL); NEO: Colombia, Cuba, Dominican Republic, Grenada, Guatemala, Haiti, Honduras, Jamaica, Mexico, Nicaragua, Panama, Puerto Rico, Trinidad & Tobago, Venezuela.

Notes. Our species concept is based on Austin and Dangerfield (1989) and Fernandez-Triana et al. (2014e). Yu et al. (2016) recorded France as a country for the species, based on one reference (Paddock 1933). However, we have read that paper, and there is no mention of A. diatraea there. Because we have not found any other source citing or supporting the presence of this species for France, we consider that record to be incorrect. Other country records (for introductions of diatraea) can be found in Bartlett et al. (1978).

Apanteles dickyui Fernandez-Triana, 2014

Apanteles dickyui Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles dictys Nixon, 1965

Apanteles dictys Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: Indonesia.

Apanteles didiguadamuzi Fernandez-Triana, 2014

Apanteles didiguadamuzi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles dido Nixon, 1965

Apanteles dido Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles diegoalpizari Fernandez-Triana, 2014

Apanteles diegoalpizari Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles diegotorresi Fernandez-Triana, 2014

Apanteles diegotorresi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles diniamartinezae Fernandez-Triana, 2014

Apanteles diniamartinezae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles diocles Nixon, 1965

Apanteles diocles Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: India.

Geographical distribution. OTL.

OTL: China (HN), India, Indonesia, Philippines, Vietnam.

Apanteles diourbeli Risbec, 1951

Apanteles diourbeli Risbec, 1951.

Type information. Holotype male, depository unknown (not examined but original description checked). Country of type locality: Senegal.

Geographical distribution. AFR.

AFR: Senegal.

Apanteles dissimilis Nixon, 1965

Apanteles dissimile Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL, PAL.

OTL: China (FJ, HB), Philippines, Vietnam; PAL: China (JL).

Notes. Both versions of Taxapad (Yu et al. 2012, 2016) correctly spell the name of this species as Apanteles dissimilis. However, the original description of the species (Nixon 1965) and most of the subsequent, published references (e.g., Shenefelt 1972, Long et al. 2004, Chen and Song 2004) incorrectly refer to the species as Apanteles dissimile, which is the neuter rather than the masculine form of the adjective, and therefore violates ICZN Article 31.2. The Code-compliant spelling must be dissimilis, regardless of the original spelling (Doug Yanega, pers. comm.), and it is the one we follow here. The specimen is in poor condition, with detached metasoma and one pair of wings glued to a second card (underneath the point with the specimen). Because the wings were detached, the vannal lobe was torn and its shape and setation patterns can no longer be determined. But the punctures on the posterior margin of the anteromesoscutum are fused, thus we consider this species to belong to Apanteles.

Apanteles dores Nixon, 1965

Apanteles dores Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Malaysia.

Geographical distribution. OTL.

OTL: Malaysia, Vietnam.

Apanteles dotus Nixon, 1965

Apanteles dotus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Sri Lanka.

Geographical distribution. OTL.

OTL: Philippines, Sri Lanka, Vietnam.

Apanteles drupes Nixon, 1965

Apanteles drupes Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles dumosus Liu & Chen, 2014

Apanteles dumosus Liu & Chen, 2014.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. PAL.

PAL: China (JL, LN).

Apanteles duniagarciae Fernandez-Triana, 2014

Apanteles duniagarciae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles duplicatus Brèthes, 1922

Apanteles duplicatus Brèthes, 1922.

Type information. Holotype female, MACN (not examined). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles duvalierbricenoi Fernandez-Triana, 2014

Apanteles duvalierbricenoi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles edgarjimenezi Fernandez-Triana, 2014

Apanteles edgarjimenezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles edithlopezae Fernandez-Triana, 2014

Apanteles edithlopezae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eduardoramirezi Fernandez-Triana, 2014

Apanteles eduardoramirezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles edwardsii Riley, 1889

Apanteles edwardsii Riley, 1889.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (ON, QC), USA (CT, IL, IA, ME, MA, MI, MN, NY, OH).

Apanteles edwinapuii Fernandez-Triana, 2014

Apanteles edwinapuii Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles elagabalus Nixon, 1965

Apanteles elagabalus Nixon, 1965.

Type information. Holotype female, AEIC (not examined but original description checked). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles eldarayae Fernandez-Triana, 2014

Apanteles eldarayae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eliethcantillanoae Fernandez-Triana, 2014

Apanteles eliethcantillanoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles epiblemae Muesebeck, 1935

Apanteles epiblemae Muesebeck, 1935.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: USA (CA, DE, FL, GA, KS).

Apanteles epijarbi Rao, 1953

Apanteles epijarbi Rao, 1953.

Type information. Holotype female, SJCA (not examined but original description checked). Country of type locality: India.

Geographical distribution. OTL.

OTL: India.

Apanteles epinotiae Viereck, 1912

Apanteles epinotiae Viereck, 1912.

Type information. Holotype male, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (ON), USA (CT, FL, IL, KS, KY, ME, MD, MA, MO, NE, NJ, NY, OH, OK, PA, SC, TX, VA, WV).

Apanteles erickduartei Fernandez-Triana, 2014

Apanteles erickduartei Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eriphyle Nixon, 1965

Apanteles eriphyle Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles erse Nixon, 1965

Apanteles erse Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Indonesia.

Geographical distribution. OTL.

OTL: Indonesia, Malaysia.

Apanteles esthercentenoae Fernandez-Triana, 2014

Apanteles esthercentenoae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eublemmae Nixon, 1965

Apanteles eublemmae Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Tanzania.

Geographical distribution. AFR.

AFR: Kenya, South Africa, Tanzania.

Apanteles eugeniaphilipsae Fernandez-Triana, 2014

Apanteles eugeniaphilipsae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eulogiosequeirai Fernandez-Triana, 2014

Apanteles eulogiosequeirai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles eupolis Nixon, 1965

Apanteles eupolis Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: South Africa.

Geographical distribution. AFR.

AFR: South Africa.

Apanteles eurynome Nixon, 1965

Apanteles eurynome Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Fiji.

Geographical distribution. AUS.

AUS: Fiji.

Apanteles eurytergis de Saeger, 1941

Apanteles eurytergis de Saeger, 1941.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Democratic Republic of Congo.

Geographical distribution. AFR.

AFR: Cape Verde, Democratic Republic of Congo.

Apanteles evadnix Shenefelt, 1972

Apanteles evadnix Shenefelt, 1972.

Apanteles evadne Nixon, 1965 [primary junior homonym of Apanteles evadne Nixon, 1955].

Type information. Holotype female, NHMUK (examined). Country of type locality: Uganda.

Geographical distribution. AFR.

AFR: Uganda.

Apanteles evanidus Papp, 1975

Apanteles evanidus Papp, 1975.

Apanteles calpurnia Nixon, 1976.

Type information. Holotype female, HNHM (not examined but original description checked). Country of type locality: Hungary.

Geographical distribution. PAL.

PAL: Finland, Hungary, Moldova, Russia (S), Sweden, Turkey, Ukraine.

Apanteles evansi Nixon, 1971

Apanteles evansi Nixon, 1971.

Type information. Holotype female, NHMUK (examined). Country of type locality: Kenya.

Geographical distribution. AFR.

AFR: Cape Verde, Kenya.

Apanteles faustina Nixon, 1965

Apanteles faustina Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Mauritius.

Geographical distribution. AFR.

AFR: Mauritius.

Apanteles federicomatarritai Fernandez-Triana, 2014

Apanteles federicomatarritai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles felipechavarriai Fernandez-Triana, 2014

Apanteles felipechavarriai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles felixcarmonai Fernandez-Triana, 2014

Apanteles felixcarmonai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles feltiae Viereck, 1912

Apanteles feltiae Viereck, 1912.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (SK), USA (AZ, CA, ID, IN, IA, MI, OH, SD, UT).

Apanteles fernandochavarriai Fernandez-Triana, 2014

Apanteles fernandochavarriai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles firmus Telenga, 1949

Apanteles firmus Telenga, 1949.

Apanteles firmus rufipes Telenga, 1955.

Type information. Holotype and depository unknown not examined but authoritatively identified specimens examined). Country of type locality: Tajikistan.

Geographical distribution. PAL.

PAL: Armenia, Azerbaijan, France, Hungary, Kazakhstan, Korea, Mongolia, Romania, Russia (YAR), Tajikistan, Ukraine, Yugoslavia.

Notes. We have examined a female paratype, donated to the CNC. That specimen looks like Dolichogenidea (the vannal lobes in both hind wings are broken but they appear to be setose, although it is not entirely clear). The descriptions and comments by Nixon (1973) and Papp (1984a) also suggest this species could be placed in Dolichogenidea. However, without examining more specimens we refrain to transfer the species in this paper and prefer to maintain it in Apanteles for the time being.

Apanteles flavicapus Liu & Chen, 2014

Apanteles flavicapus Liu & Chen, 2014.

Type information. Holotype female, ZJUH (not examined but original description checked). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (GD).

Apanteles flavicentrus Long, 2010

Apanteles flavicentrus Long, 2010.

Type information. Holotype female, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Notes. This species might not be Apanteles, but the original description does not provide enough details to determine its placement, so we retain it within Apanteles.

Apanteles flavigaster Long, 2010

Apanteles flavigaster Long, 2010.

Type information. Holotype female, IEBR (not examined but original description checked). Country of type locality: Vietnam.

Geographical distribution. OTL.

OTL: Vietnam.

Notes. This species might not be Apanteles, but the original description does not provide enough details to determine its placement, so we retain it within Apanteles.

Apanteles floralis Tobias, 1966

Apanteles floralis Tobias, 1966.

Type information. Holotype female, ZIN (not examined but subsequent treatment of the species checked). Country of type locality: Turkmenistan.

Geographical distribution. PAL.

PAL: Kazakhstan, Mongolia, Turkmenistan.

Notes. Our species concept is based on Papp (1984a) and Tobias (1986).

Apanteles flormoralesae Fernandez-Triana, 2014

Apanteles flormoralesae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles florus Nixon, 1965

Apanteles florus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: China.

Geographical distribution. OTL.

OTL: China (GD, HN).

Apanteles fluitantis de Santis, 1980

Apanteles fluitantis de Santis, 1980.

Type information. Holotype female, MLP (not examined but subsequent treatment of the species checked). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Notes. Our concept of this species is based on Aquino et al. (2010).

Apanteles fontinalis de Saeger, 1944

Apanteles fontinalis de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Rwanda.

Geographical distribution. AFR.

AFR: Democratic Republic of Congo, Réunion, Rwanda.

Apanteles forbesi Viereck, 1910

Apanteles forbesi Viereck, 1910.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA.

NEA: Canada (MB, NS, ON), USA (AZ, CT, FL, IL, IN, IA, KS, KY, MD, MA, MO, NY, OR, SD).

Apanteles franciscopizarroi Fernandez-Triana, 2014

Apanteles franciscopizarroi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles franciscoramirezi Fernandez-Triana, 2014

Apanteles franciscoramirezi Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles freddyquesadai Fernandez-Triana, 2014

Apanteles freddyquesadai Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles freddysalazari Fernandez-Triana, 2014

Apanteles freddysalazari Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles fredi Austin & Dangerfield, 1989

Apanteles fredi Austin & Dangerfield, 1989.

Type information. Holotype female, NHMUK (not examined but original description checked). Country of type locality: Guatemala.

Geographical distribution. NEO.

NEO: Guatemala.

Apanteles frersi (Brèthes, 1917)

Coelothorax frersi Brèthes, 1917.

Type information. Holotype female, MACN (not examined). Country of type locality: Argentina.

Geographical distribution. NEO.

NEO: Argentina.

Apanteles fumiferanae Viereck, 1912

Apanteles fumiferanae Viereck, 1912.

Type information. Holotype female, USNM (examined). Country of type locality: USA.

Geographical distribution. NEA, PAL.

NEA: Canada (BC, MB, NB, NL, NT, ON, QC), USA (AK, CO, ID, ME, MA, MI, MN, MT, NM, NY, OR, SC, SD, WA, WI); PAL: Poland.

Notes. We consider A. fumiferanae as a Nearctic species (Fernandez-Triana 2010, Fernandez-Triana and Huber 2010). The single record from the Palearctic is based on one publication compiling the Hymenoptera from Poland (Huflejt 1997), and it is very likely to be incorrect; however, we refrain to remove that record until more studies are done. The generic placement of this species is also somewhat controversial as the female holotype has the hind wings with a straight vannal lobe with small setae which are just slightly sparser than proximal and distal margins of lobe, and the (shallow and sparse) punctures on the anteromesoscutum are not fused near posterior margin. These two features are borderline with Dolichogenidea and more studies, combining morphology, biology, and molecular data, will be needed.

Apanteles fundulus Nixon, 1965

Apanteles fundulus Nixon, 1965.

Type information. Holotype female, NHMUK (examined). Country of type locality: Australia.

Geographical distribution. AUS, OTL.

AUS: Australia (QLD); OTL: Vietnam.

Apanteles gabrielagutierrezae Fernandez-Triana, 2014

Apanteles gabrielagutierrezae Fernandez-Triana, 2014.

Type information. Holotype female, CNC (examined). Country of type locality: Costa Rica.

Geographical distribution. NEO.

NEO: Costa Rica.

Apanteles galatea Nixon, 1965

Apanteles galatea Nixon, 1965.

Type information. Holotype female, USNM (examined). Country of type locality: Philippines.

Geographical distribution. OTL.

OTL: Philippines.

Apanteles galleriae Wilkinson, 1932

Apanteles galleriae Wilkinson, 1932.

Type information. Holotype female, NHMUK (examined). Country of type locality: France.

Geographical distribution. AFR, NEA, NEO, OTL, PAL.

AUS: Hawaiian Islands, New Zealand; AFR: Mauritius, Réunion; NEA: Canada (BC), USA (GA, NC, OH, SC); NEO: Argentina, Brazil (SP); OTL: China (GZ, HN, TW, ZJ), India, Pakistan; PAL: Armenia, Bulgaria, France, Greece, Hungary, Iran, Italy, Japan, Malta, Romania, Russia (PRI), Spain, Turkey, United Kingdom.

Notes. Distribution in Brazil based on de Santis (1964) and Shimbori (pers. comm.).

Apanteles gandoensis de Saeger, 1944

Apanteles gandoensis de Saeger, 1944.

Type information. Holotype female, RMCA (not examined but original description checked). Country of type locality: Rwanda.

Geographical distribution. AFR.

AFR: Rwanda.

Apanteles garygibsoni Fernandez-Triana, 2014

Apanteles garygibsoni Fernandez-Triana, 2014.

Type information. Holotype female,