Antenna with 8 flagellomeres, including a single anellus and an undivided clava. Eyes ventrally divergent. Clypeus without transverse subapical groove. Labrum hidden behind clypeus. Mandibles with ventral tooth and large dorsal truncation. Mesoscutellum with frenum set off by complete frenal groove, and with axillular sulcus (Fig. 1). Mesopleural area without expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; metafemur with ventral lobe or subapical teeth (Fig. 2). Metasoma with epipygium (Fig. 3), or with syntergum (Chalcidiscelis Ashmead).

Figures 1–6. 

1–3 Boucekius sp. (Boucekiidae) 1 metascutellum, axillula and propodeum 2 hind femur 3 epipygium (epg) and metasomal terga VIII (Mt8) 4, 5 Spalangiopelta sp. (Ceidae) 4 clypeus, labrum and mandible 5 metascutellum, axillula and propodeum, arrow shows the propodeal spiracle far separated from the anterior propodeal margin 6 Muesebeckisia mandibularis Hedqvist (Cerocephalidae): head and antenna in lateral view, arrow indicating intertorular prominence.

Gibson (2003) described Boucekiidae as a new tribe, although it differed from many species in Cleonyminae (as then defined) in the frenum and potentially the labrum. In habitus boucekiids do resemble former Cleonyminae and other large-bodied Chalcidoidea that have metallic coloration. Out of former Cleonyminae, those with an unambiguous frenal arm (= mesoscutellar arm) laterally are now classified in Solenurinae (Lyciscidae), which differ most notably in having an incomplete frenal line and a flagellum with 2 or 3 clavomeres. Chalcedectidae and Heydeniidae can have either an indistinct frenal groove, a small frenum, or a strongly expanded marginal rim of the mesoscutellum that may resemble a frenum; however, both have a different clava from Boucekiidae, with multiple clavomeres instead of an undivided clava and, in Chalcedectidae, an apical spine in females. Chalcedectidae have a syntergum that is not crossed by a transverse sulcus and otherwise does not have an epipygium. Heydeniidae have a long prepectus that is enlarged both laterally and ventrally. The elongate ovipositor and more or less elongate cerci in females may cause confusion with Torymidae or Megastigmidae, both of which have multiple clavomeres and more than 8 flagellomeres. The narrow, essentially parallel-sided flagellomeres may invite confusion with the antenna in Ceidae or Macromesidae; however, members of both these taxa have multiple clavomeres and much narrower mandibles with no dorsal truncation, and Macromesidae lack a frenum. Pteromalidae and Pelecinellidae have more than 1 clavomere in nearly all cases, but Pteromalidae with apparently 1 clavomere (some males) have more than one anelliform basal flagellomere.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Eyes not ventrally divergent. Clypeus with transverse subapical groove. Labrum subrectangular and exposed, with marginal setae in a row (Fig. 4). Mandibles with 2 teeth (Fig. 4). Subforaminal bridge with postgena separated by lower tentorial bridge except for a small postgenal bridge dorsal to the hypostoma. Mesoscutellum with frenum indicated at least laterally, and with axillular sulcus. Mesopleural area without an expanded acropleuron. Propodeum with small and circular spiracle separated by more than its own length from the anterior propodeal margin (Fig. 5). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Ceidae differs from most other Chalcidoidea in having the propodeal spiracle separated from the anterior propodeal margin by more than its own length. Exceptions to this statement occur in numerous species across many families, including some Pteromalidae. Pteromalidae differ in having more than 2 mandibular teeth except in some fig associates which differ from Ceidae in many other ways. Pteromalidae also lack a transverse subapical clypeal groove, and have a hidden labrum with an elongate median lobe, instead of a subrectangular and exposed labrum. While the subforaminal bridge in Pteromalidae and Ceidae is different, the difference is so slight in many Pteromalidae (such as Colotrechninae and Miscogastrinae) that it should not be relied upon too heavily. Hetreulophidae and the single genus of Macromesidae also have propodeal spiracle separated far from the anterior propodeal margin. Hetreulophidae differ in having distinctly fewer antennal flagellomeres (9), with a single anellus and united clava, and by having 3 mandibular teeth. Macromesidae differ in having only 4 mesotarsomeres in females, at most 11 antennal flagellomeres, and 3 mandibular teeth.

Antenna with at most 10 flagellomeres and at most 3 clavomeres. Intertorular prominence present (Fig. 6). Eyes not ventrally divergent. Clypeus without transverse subapical groove. Labrum hidden behind clypeus, flexible. Mandibles with 2 or more teeth. Subforaminal bridge with a postgenal bridge occurring dorsal to the hypostoma. Mesoscutellum with frenum indicated at least laterally, although this may be very subtle. Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron (Fig. 7). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Figures 7–12. 

7 Neocalosoter sp. (Cerocephalidae): mesosoma in lateral view 8–10 Chalcedectus sp. (Chalcedectidae) 8 antenna 9 mesosoma in lateral view 10 hind leg 11 Cleonymus sp. (Cleonymidae): head in frontal view 12 Agrilocida ferrieri Steffan (Cleonymidae): hind leg.

Cerocephalidae differ from most other Chalcidoidea in having an intertorular prominence, although a few exceptional taxa exist across the superfamily that have a similar prominence, such as some Haltichellinae (Chalcididae). However, these exceptions can be distinguished from Cerocephalidae using other features mentioned in the diagnosis. Otherwise, Cerocephalidae bear little resemblance to other families, being somewhat similar to Spalangiidae, Storeyinae, and some Eulophidae, but without most diagnostic features of those families.

Antenna with 11 flagellomeres, including 3 clavomeres, clava with apical spine in females (Fig. 8). Eyes ventrally divergent. Labrum exposed, sclerotized. Mandibles with 3 teeth. Subforaminal bridge with a postgenal bridge occurring dorsal to the hypostoma, with convergent hypostomal carina, without a postgenal groove or postgenal lamina. Prepectus with dorsal margin at least as long as tegula. Notauli complete; tegula not covering most of humeral plate. Mesoscutellum with variable frenal area: either without a frenum, or having an expanded marginal rim of the mesoscutellum, or with either a frenum indicated mainly by a frenal arm and an indistinct frenal groove, or an ambiguous frenum that can be difficult to interpret; and with axillular sulcus or carina (Fig. 9). Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron (Fig. 9). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; metafemur with ventral teeth (Fig. 10), with metatibial spurs arising from a ventroapical projection (not shown) or absent (Fig. 10). Metasoma with syntergum, therefore without epipygium.

Chalcedectidae are most likely to be confused with other Chalcidoidea that have a metafemur with ventral teeth, which occurs in various families and isolated genera across the superfamily. Chalcididae differ in having a small prepectus, the dorsal margin of which is shorter than the tegula, and in that the tegula covers most or all of the humeral plate. In Lyciscidae, the metatibial spurs arise from a truncate apical margin of the metatibia. Leucospidae have, in females, unusual ovipositor sheaths that recurve over the gaster and fit in a notch and, in males, a carapace-like gaster with at most 3 separate terga. Pelecinellidae have an elongate petiole with many lateral setae at a right angle to the longitudinal axis of the petiole, whereas in Chalcedectidae the petiole is small and hardly visible from dorsal view. Boucekiidae have a single clavomere, and either an epipygium or a transverse sulcus across the syntergum immediately anterior to the cerci. A few Melanosomellidae have a toothed metafemur, but they either lack an axillular sulcus or carina or have a reduced and incomplete one, and do not have ventrally divergent eyes. Cleonymidae have incomplete notauli. Some Torymidae have ventral metafemoral teeth, but these have a separate epipygium in females and do not have ventrally divergent eyes. In Liepara Bouček (Coelocybidae), the frenum is unambiguously visible dorsally, with a pair of strong setae adjacent to the frenal groove. A few Eulophidae can have ventral teeth on the metafemur, but they have 4 tarsomeres on all legs.

Antenna with 9 flagellomeres, including usually a single clavomere, which is sometimes vaguely divided into 3 clavomeres in males, and with a subapical finger-like process or spine extending alongside the clava and/or with an additional apical spine in females. Eyes ventrally divergent. Clypeus with transverse subapical groove. Labrum exposed, sclerotized. Mandibles with 2 or 3 teeth (Fig. 11), sometimes with a truncation in place of the dorsal teeth. Subforaminal bridge with a postgenal bridge dorsal to the hypostoma and separating the lower tentorial bridge from the convergent hypostomal carina, and without a postgenal groove or postgenal lamina. Pronotum without a smooth median longitudinal line or carina. Prepectus with dorsal margin at least as long as tegula. Notauli incomplete. Tegula not covering most of humeral plate. Mesoscutellum without a frenum, although frenal arm visible only laterally immediately anterior to marginal rim of mesoscutellum; without axillular sulcus. Mesopleural area without an expanded acropleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; metafemur with or without ventral teeth, with apical spurs arising from a truncate metatibial apex when the metafemur has ventral teeth (Fig. 12). Metasoma with syntergum, therefore without epipygium.

Cleonymidae in its current, narrow sense represents the former Cleonymini, as characterized by Gibson (2003). It includes Agrilocida Steffan new placement, previously placed in Chalcedectini, but which is distinct morphologically and consistently is placed in Cleonymidae in next-generation molecular analyses. This is now a relatively small and narrowly-defined group in comparison with the previous sense of Cleonyminae, and may be confused with other relatively large-bodied taxa that have ventrally divergent eyes.

Females of Eupelmidae, Metapelmatidae, Neanastatidae, and Eopelma Gibson have an enlarged, convex and pad-like acropleuron that covers most or all of the mesopleural area. Lyciscidae, Chalcedectidae, Ooderidae, Pelecinellidae, and Boucekiidae differ from Cleonymidae in having complete notauli. Heydeniidae have a long prepectus with large lateral and ventral surfaces. While some Cleonymidae have ventral teeth on the metafemur, they do not strongly resemble Chalcididae, especially because of the metallic coloration of most Cleonymidae versus the usually non-metallic coloration of Chalcididae, but also because Cleonymidae have incomplete notauli, a larger prepectus, and a smaller tegula that does not cover most of the humeral plate. Coelocybidae usually have non-metallic coloration but also have a distinctive frenum with at least one pair of strong mesoscutellar setae on or nearly adjacent to the frenal groove, whereas the mesoscutellum in Cleonymidae is evenly covered with short and decumbent setae. Additionally, Coelocybidae do not have any spine or finger-like projection on or extending alongside the clava from a previous segment in females.

Antenna with 11 flagellomeres. Eyes ventrally divergent. Clypeus with transverse subapical groove (extending from one anterior tentorial pit to the other). Labrum hidden behind clypeus, flexible, subrectangular, with marginal setae in a row. Mandibles with 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge. Mesoscutellum with frenum indicated and with a pair of strong setae on or adjacent to frenal groove, and with axillular sulcus (Fig. 13). Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron (Fig. 14). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Figures 13–18. 

13 Coelocyboides sp. (Coelocybidae): mesosoma in dorsal view, arrow indicating setae on or adjacent to frenal groove 14 Ormyromorpha trifasciata Girault (Coelocybidae): mesosoma in lateral view 15, 16 Lelaps sp. (Diparidae) 15 head posterior view 16 cercal brush 17 Eufroggattisca polita (Ashmead) (Epichrysomallidae): mesosoma lateral view 18 Odontofroggattia sp. (Epichrysomallidae): propodeum female dorsal view.

As mentioned by Bouček (1988), Coelocybidae closely resemble what was then known as Cleonyminae, which is now multiple families. Among these, Coelocybidae bear greater resemblance to Cleonymidae and Lyciscidae. Cleonymidae differ from Coelocybidae in lacking a frenum and the accompanying strong setae, the setae also being absent from Lyciscidae. Otherwise, taxa with ventrally divergent eyes have an expanded acropleuron or fewer flagellomeres. Many other taxa have a pair of strong setae on the frenal groove, but not together with ventrally divergent eyes, except Cecidellis Hanson (Pirenidae) which has 9 antennal flagellomeres. Nearly all Coelocybidae are from the Southern Hemisphere, with the exception of a single species from India (Narendran 2001).

Lieparini new synonym is hereby abolished, and Liepara Bouček new placement is transferred here from its incertae sedis status (Heraty et al. 2013) because it has consistently been part of the new concept of a monophyletic Coelocybidae in next-generation molecular analyses (Cruaud et al., submitted). While the ventrally toothed metafemur of Liepara is distinctive, it is a well-known homoplastic feature in Chalcidoidea. Otherwise, the slightly ventrally divergent eyes, strong setae adjacent to the frenal groove, and non-metallic coloration of Liepara indicate that morphology agrees with molecules in this instance. Lieparini does not seem to be a useful tribe to keep as distinct from other coelocybines, although further study may lead to its resurrection once more coelocybids have been analyzed phylogenetically.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Eyes not ventrally divergent. Labrum hidden behind clypeus, flexible, subrectangular, with marginal setae in a row. Mandibles with 3 or 4 teeth. Subforaminal bridge with postgenal bridge occurring dorsal to the hypostoma (Fig. 15). Mesoscutellum with frenum indicated at least laterally, and with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron (except in Diparisca Hedqvist). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium; cercal brush present anterior to cercus (Fig. 16).

As discussed by Desjardins (2007), Diparinae have a cercal brush anterior to the cercus (Fig. 16). This feature is not perfectly diagnostic however, because it can also be present in Spalangiidae and Neapterolelapinae, which are recovered as the sister group to Lyciscidae, and Herbertiidae (Cruaud et al., submitted). Spalangiidae differ in having the mesepimeron extending over the anterior margin of the metapleuron. Lyciscidae differ in having an exposed, rigidly sclerotized labrum, and in lacking a frenum (except in Solenurinae). Herbertiidae differ in having at most 10 antennal flagellomeres, and in having an exposed, rigidly sclerotized labrum. In the features listed in the diagnosis, Cerocephalidae may appear similar to Diparidae, although they differ in that Cerocephalidae have an intertorular prominence and at most 10 antennal flagellomeres. Diparisca remains in Diparidae as a genus of uncertain placement (Mitroiu 2016), with distinction from Ceinae discussed therein.

Antenna with 10–12 flagellomeres, including a small 4^th clavomere. Eyes not ventrally divergent. Labrum hidden behind clypeus, flexible. Mandibles with 3 teeth. Subforaminal bridge with postgenal bridge separating secondary posterior tentorial pits from hypostoma. Notauli complete. Mesoscutellum with frenum indicated laterally, without axillular sulcus (Fig. 17). Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron (Fig. 17). All legs with 5 tarsomeres in most, except tarsi 4-segmented in Odontofroggatia Ishii and Josephiella Narendran; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Epichrysomallidae mostly resemble Melanosomellidae in habitus, but do not have a linear mesopleural sulcus. They also have different fore wing venation with a stigmal vein arising at a right angle (excepted in Acophila Ishii) and a postmarginal vein that is shorter than the stigmal vein. Epichrysomallidae have a characteristic flap-like expansion of cuticle from the lateral edge of the propodeal spiracle, partially covering the spiracle in dorsal view (Fig. 18) that neither Melanosomellidae nor Ormyridae have. Ormyridae differ further from Epichrysomallidae by having a more conventional fore wing venation, with longer marginal and postmarginal veins, and iridescent coloration in most species.

Antenna with at most 11 flagellomeres. Eyes ventrally divergent. Clypeus with transverse subapical groove. Labrum either exposed and well-sclerotized (most species), or hidden behind clypeus (Epicopterus Westwood), subrectangular, with marginal setae in a row. Mandibles with 2 or rarely 3 teeth (Fig. 19). Subforaminal bridge with postgena separated by lower tentorial bridge. Pronotum transverse. Mesoscutellum with frenum indicated laterally, with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron; only one mesofurcal pit present (Fig. 20). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal (Fig. 21). Metasoma with syntergum, therefore without epipygium.

Figures 19–24. 

19–21 Eunotus sp. (Eunotidae) 19 mandible and labrum in frontal view 20 mesosoma ventral view 21 protibial spur and basitarsal comb 22 Herbertia brasiliensis Ashmead (Herbertiidae) head posterior view 23 Hetreulophus sp. (Hetreulophidae), mesosoma lateral view 24 Zeala walkerae Bouček (Hetreulophidae): propodeum.

Eunotidae, as defined herein, was previously known as Eunotini (Bouček 1988). Moranilidae, composed of species previously included in Eunotinae, differ in having an oblique basitarsal comb that crosses the area where the basitarsal notch would be, in having pits on the mesopleural area of the mesopectus, and in having two mesofurcal pits instead of the single pit usually found in Chalcidoidea. Idioporus affinis (Idioporidae) differs in having 4 tarsomeres. Aphelinidae differ in lacking any indication of a frenum, in having a flexible, hidden labrum, usually in having more advanced axilla, and the second phragma extending into the mesosoma. Despite apparent similarities between Aphelinidae and Eunotidae, the two taxa, are seldom confused because Eunotidae have a more strongly sclerotized body, which means that the two families have a very different habitus. Lyciscidae differ in having a subconical pronotum with a median longitudinal carina, whereas the pronotum of Eunotidae is transverse without a visible median carina in dorsal view.

Antenna with 10 flagellomeres, including 3 clavomeres. Clypeus with transverse subapical groove. Labrum exposed, well-sclerotized, subrectangular with marginal setae in a row. Mandibles with 2 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge; head posteriorly with postgenal lamina and postgenal groove (Burks et al. 2018) (Fig. 22). Mesoscutellum with frenum indicated only laterally by the frenal arm, without axillular sulcus. Mesopleural area without an expanded acropleuron, with or without pits. Fore wing marginal vein more than 1.5× stigmal vein length, without elongate uncus. Mesepimeron extending over anterior margin of metapleuron; two mesofurcal pits present. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb oblique. Metasoma with syntergum, therefore without epipygium.

The family Herbertiidae has uncertain placement based on both molecular (Cruaud et al., submitted) and morphological data, and is treated as incertae sedis in Chalcidoidea pending more consistent phylogenetic resolution. Micradelinae are similar to Herbertiidae in body shape, but differ in having an elongate uncus and much shorter marginal vein relative to the stigmal vein on the fore wing, in having an indicated axillular sulcus, in having only one mesofurcal pit, and in lacking a postgenal lamina and postgenal groove. Erotolepsiinae (Spalangiidae) are similar to Herbertiidae in habitus but differ in having a transverse anterior carina across Gt₁, and in most species having a long carina encircling most of the face. Eunotidae differ in having a longitudinal basitarsal comb, and in lacking a postgenal lamina and postgenal groove. The presence of two mesofurcal pits is unusual, a feature shared with Moranilidae, Enoggerinae, Asaphesinae, some Eurytominae (Krogmann and Vilhelmsen 2006) and Chalcididae (Haltichellinae and Smicromorphinae, G. Delvare, pers. comm). Moranilidae differ in having an axillular sulcus. Other small-bodied families such as Pirenidae and Systasidae differ in having more than 2 mandibular teeth, and Pirenidae differ in having a concealed, flexible labrum.

Antenna with 9 flagellomeres, including a 1-segmented clava. Clypeus without transverse subapical groove. Labrum flexible, hidden behind clypeus. Mandibles with 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge except for the small postgenal bridge dorsal to the hypostoma. Mesoscutellum with short frenum, with axillular sulcus, and expanded, convex axillula (Fig. 23). Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron (Fig. 23). Propodeum with spiracle small, oval, separated by more than its own length from anterior propodeal margin (Fig. 24). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal.

Ceidae, another family with propodeal spiracle separated far from the anterior margin of the propodeum, differ in having 12 antennal flagellomeres and only 2 mandibular teeth. Macromesidae share this feature as well, but have at least 10 flagellomeres including multiple clavomeres, 4 mesotarsomeres in females, and the mesepimeron does not extend over the anterior margin of the metapleuron. Otherwise, families that resemble Hetreulophidae in habitus have more flagellomeres and multiple clavomeres.

Omphalodipara Girault new placement is transferred from Colotrechninae, Amerostenini (Pteromalidae) to Hetreulophidae based on next generation molecular data (Cruaud et al., submitted). Given the 9 antennal flagellomeres with a 1-segmented clava, strongly convex axillula, short frenum, and posteriorly displaced propodeal spiracle shared between Omphalodipara and other Hetreulophidae, it is reasonable to say that morphology agrees with this placement.

Antenna with 10 or 11 flagellomeres, including 3 clavomeres. Eyes ventrally divergent. Clypeus without transverse subapical groove. Labrum exposed or hidden behind clypeus, sclerotized. Mandibles with 3 teeth. Pronotum expanded laterally and forming a subrectangular or laterally expanded structure from dorsal view (Fig. 67). Prepectus long, with large dorsal and ventral surfaces. Mesoscutellum either without a frenum, or with frenum laterally indicated by a frenal arm, with or without axillular sulcus (Fig. 68). Mesopleural area with acropleuron slightly expanded but occupying less than half its surface; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; profemur strongly (Fig. 69) or only mildly expanded; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium, although terga not well-sclerotized and often difficult to assess.

Some species of Heydeniidae are distinctive and resemble Ooderidae, while more generalized species are considerably more difficult to recognize. Ooderidae differ in having multiple rows of spine-like structures on the ventral surface of the always strongly expanded profemur. Heydeniidae have instead at most a single row of crest-like structures ventrally on the profemur. Otherwise, the ventrally elongate prepectus of Heydeniidae is distinctive. When the profemur is not strongly expanded and the pronotum is relatively short and not tent-like, species of Heydenia Förster can be more difficult to recognize. Given the presence of a convex mesoscutellum with a weakly distinct or indistinct frenum and ventrally divergent eyes, generalized Heydeniidae may be confused with Cleonymidae or Lyciscidae, both of which have a much shorter prepectus ventrally.

Antenna with 9 flagellomeres, including 4 distinct clavomeres (Fig. 25). Clypeus with transverse subapical groove. Labrum hidden behind clypeus, flexible, subrectangular, with marginal setae in a row. Mandibles with 2 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron (Fig. 26). All legs with 4 tarsomeres; protibial spur stout and slightly curved; basitarsal comb longitudinal (Fig. 27).

Figures 25–30. 

25–27 Idioporus affinis LaSalle & Polaszek (Idioporidae) 25 antenna 26 mesosoma lateral 27 protibial spur and basitarsal comb. 28 Lycisca nebulipennis Strand (Lyciscidae) head frontal view 29 Lycisca ignicaudata Westwood (Lyciscidae): pronotum and mesonotum dorsal view 30 Agamerion cleptideum (Westwood) (Lyciscidae): mesosoma lateral view.

Idioporus affinis LaSalle & Polaszek is a highly distinctive species in Chalcidoidea (LaSalle et al. 1997) that has been problematic in placement regardless of whether using morphology or molecules. Relative to most other families discussed here, it is distinct in tarsomere count; only Zebe La Salle (Pirenidae) has four tarsomeres, but Zebe differs in having most funiculars greatly reduced. Idioporus differs from other taxa with 4-segmented tarsi on all legs, such as Eulophidae and Calesidae, and in having a stout and slightly curved protibial spur.

Antenna with 8 or 7 flagellomeres, including usually a single clavomere but sometimes with 2 or (in males) 3 clavomeres. Eyes ventrally divergent (Fig. 28). Clypeus with or without transverse subapical groove. Labrum exposed or hidden behind clypeus, sclerotized and with a row of submarginal setae extending anteriorly (Fig. 28). Mandibles with 2 or 3 teeth, sometimes with a truncation in place of the dorsal teeth. Subforaminal bridge with elongate lower tentorial bridge and secondary tentorial pits that extend to the convergent hypostomal carina, with or without a postgenal groove and postgenal lamina, or (Solenurinae) with a postgenal bridge that externally separates the lower tentorial bridge from the convergent hypostomal carina. Pronotum with a smooth median longitudinal line or carina (Fig. 29). Notauli complete. Mesoscutellum usually without a frenum (Fig. 30) or (Solenurinae) with a frenum indicated by lateral frenal arms (Fig. 32); without or (rarely) with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; metafemur with or without ventral teeth or expansion, with apical spurs arising from a truncate metatibial apex when the metafemur has ventral teeth. Metasoma with syntergum, therefore without epipygium, although a complex set of carinae sometimes present on syntergum.

Figures 31–36. 

31, 32 Solenura sp. (Lyscicidae, Solenurinae) 31 antenna 32 mesosoma lateral 33 Macromesus sp. (Macromesidae): head frontal view 34 Macromesus amphiretus Walker (Macromesidae), propodeum 35, 36 Trichilogaster acaciaelongifoliae (Froggatt) (Melanosomellidae) 35 head frontal 36 mesosoma lateral view.

The family Lyciscidae was potentially a major part of what Bouček (1988) had in mind when describing his concept of Cleonyminae as a monophyletic lineage, being “certainly one of the oldest in Pteromalidae, as seems to be supported by their association with wood-boring beetles.” However, Lyciscidae itself appears to be relatively young and separate from Cleonymidae and all other members of the former sense of Cleonyminae, based upon next-generation molecular data (Cruaud et al., submitted).

Lyciscidae are relatively generalized and are therefore easily confused with many other large-bodied Chalcidoidea. While the longitudinal median smooth strip or carina of the pronotum is distinctive, it can be difficult to assess in some taxa depending on the position of the head. However, Neapterolelapinae differ from Lyciscidae chiefly in the lack of this feature. Many Eupelmidae are similar to Lyciscidae but females and some males have an expanded, convex and pad-like acropleuron that covers most or all of the mesopleural area. In all Chalcedectidae the metafemur has ventral teeth, a feature also present in some Lyciscidae, but in Lyciscidae the metatibia is truncate where the metatibial spurs insert, whereas in Chalcedectidae the spurs are either absent or placed on a ventroapical projection.

Lyciscidae differ from many other large-bodied Chalcidoidea in lacking a frenum. In Cleonymidae the notauli are incomplete. Pelecinellidae differ in having an elongate petiole with long setae perpendicular to its longitudinal axis. Macromesidae do not have ventrally divergent eyes, and often instead have ventrally convergent eyes. Eunotidae have a much shorter pronotum without a distinctive anterior neck.

For Solenurinae, identification can be more difficult due to the presence of a frenal arm, which is shared with a greater number of other chalcidoids. While Solenura Westwood itself is a highly distinctive genus with an elongate gaster, Grooca Sureshan & Narendran has a shorter gaster similar to that of many other Chalcidoidea. Confusion is most likely with other groups that have ventrally divergent eyes, such as Coelocybidae which differ in having strong setae on or nearly adjacent to the frenal groove. Herbertiidae and Micradelinae have a different antenna with 10 or 11 flagellomeres. Ditropinotellinae differ in having a distinctive T-shaped and elongate syntergum that resembles an elongate epipygium. Moranilidae may appear similar to Lyciscidae when comparing the lists of features, but in practice are easily distinguished. Some Moranilidae do not have ventrally divergent eyes (Asaphesinae), while others have a much smaller body with a different antennal flagellum that is strongly clavate and or with transverse funiculars.

Solenura Westwood, 1868.

Antenna with 2 or 3 clavomeres (Fig. 31). Clypeus without transverse subapical groove. Labrum exposed small and subrectangular, with marginal setae. Mandibles with 2 similarly-sized teeth. Subforaminal bridge with a postgenal bridge that externally separates the lower tentorial bridge from the convergent hypostomal carina; postgenal groove and postgenal lamina absent. Frenum indicated laterally by frenal arm that is well-separated from the marginal rim of the mesoscutellum (Fig. 32). Metafemur withoutventral teeth or expansion. Other features as in Lyciscinae.

Antenna with 10 flagellomeres in females, 11 in males. Face between malar sulcus and torulus with a second longitudinal sulcus (Fig. 33). Clypeus without transverse subapical groove. Labrum subrectangular and hidden, with marginal setae in a row. Mandibles with 3 teeth (Fig. 33). Subforaminal bridge with postgena separated by lower tentorial bridge except for a small postgenal bridge dorsal to the hypostoma. Mesoscutellum with frenal arm indicated laterally, and with axillular carina or sulcus. Mesopleural area without an expanded acropleuron. Propodeum with spiracle separated by more than its own length from the anterior propodeal margin (Fig. 34). Fore and hind legs with 5 tarsomeres, middle legs in females with 4 tarsomeres. Protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Macromesus, the only genus of Macromesidae, differs from other Chalcidoidea inthe tarsomere count of females and the usually conspicuous second longitudinal sulcus on the lower face, although it otherwise bears some resemblance to other large-bodied chalcidoids with metallic coloration. The distance from the propodeal spiracle to the anterior margin of the propodeum may cause it to be confused with Ceidae or Hetreulophidae, but this feature is likely convergent, apparently occurring in Macromesidae because of its unusual propodeum.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Eyes not divergent ventrally (Fig. 35). Clypeus without transverse subapical groove. Labrum hidden, flexible. Mandibles with 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge, or with a short apparent postgenal bridge immediately dorsal to the hypostoma. Notauli complete. Mesoscutellum with frenum indicated laterally, either without axillular sulcus or carina, or with it greatly reduced and incomplete (Fig. 36). Mesopleural area without an expanded acropleuron (Fig. 36). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium, and rigidly convex.

Additionally, Melanosomellidae typically have a linear mesopleural sulcus that is more distinct than in most other Chalcidoidea (Fig. 36), although this feature also occurs in various species from other families. There is a strong chance of confusion of Melanosomellidae with Epichrysomallidae and Ormyridae, members of the Gall Clade (Cruaud et al., submitted, van Noort et al., in prep.). Epichrysomallidae are very similar to Melanosomellidae in habitus, but have different fore wing venation, with a nearly straight stigmal vein arising at a right angle from the wing margin. Furthermore, Epichrysomallidae are strictly associated with Ficus (Moraceae) as gall-makers either within figs or on leaves and twigs. Ormyridae are also very similar to Melanosomellidae but have an occipital carina. Pteromalidae almost always have a distinct and complete axillular sulcus or carina. In species where this may not be the case, such as Nikolskayana mirabilis Bouček, the notauli are incomplete.

Encyrtocephalus Ashmead is very similar to other genera classified in Melanosomellidae, but molecular data (Cruaud et al., submitted) indicate that it may not belong inside this group. However, the only morphological features that imperfectly separate it from most Melanosomellidae are a large supracoxal flange on the posterior margin of the propodeum (Fig. 37) and a distinctly curved stigmal vein. These features are shared with a few other melanosomellid genera such as Alyxiaphagus Riek, with intermediates that would make diagnosis either very difficult or impossible. Therefore, Encyrtocephalus is kept in Melanosomellidae.

Figures 37–42. 

37 Encyrtocephalus sp. (Melanosomellidae): propodeum and supracoxal flange 38 Hansonita pertusae Bouček (Melanosomellidae): venation 39 Moranila californica (Howard) (Moranilidae): antenna 40 Moranila viridivertex (Girault) (Moranilidae): mesosoma lateral view 41 Moranila californica (Howard) (Moranilidae): mesosoma ventral view 42 Neodipara masneri Bouček (Neodiparidae): head lateral view and antenna.

The fig associate species Hansonita pertusae Bouček new placement is transferred here because its fore wing venation resembles that of Melanosomellidae (Fig. 38) more strongly than that of Epichrysomallidae or other fig associates.

Antenna with 8 flagellomeres, clava undivided or incompletely divided (Fig. 39). Clypeus with transverse subapical groove (Fig. 56b). Labrum exposed, well-sclerotized, subrectangular with marginal setae in a row. Mandibles with 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge. Mesoscutellum with frenum indicated at least laterally, with axillular sulcus. Mesopleural area without an expanded acropleuron, with pits (Fig. 40); mesepimeron not extending over anterior margin of metapleuron; two mesofurcal pits present (Fig. 41). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb oblique. Metasoma with syntergum, therefore without epipygium.

Moranilidae contains two subfamilies: the former tribe Moranilinae new status and Tomocerodinae new placement, new status, based on morphological similarity since molecular data are absent for Tomocerodinae. Moranilidae differ from almost all other Chalcidoidea in having 2 mesofurcal pits instead of the usual single pit, but this feature appears to be homoplastic within Chalcidoidea. Indeed, some Eurytominae (Eurytomidae) (Krogmann and Vilhelmsen 2006) as well as Smicromorphinae and some Haltichellinae (G. Delvare comm. pers) also have two mesofurcal pits but differ from Moranilidae in many ways, including a different basitarsal comb and subforaminal bridge. These families are not easily confused with one another due to the very different habitus of the much more strongly sclerotized Eurytominae and Chalcididae.

Herbertiidae, Asaphesinae, and Enoggerinae new subfamily also share 2 mesofurcal pits with Moranilidae. Herbertiidae differ in lacking an axillular sulcus. Asaphesinae have 12 antennal flagellomeres instead of the maximum of 8 in Moranilidae. Enoggerinae lack a temple on the head, thus having the posterior margin of the eye coincident with that of the head dorsally. Micradelinae also resemble Moranilidae, but have only 1 mesofurcal pit instead of 2, and lack pits on the mesopleural area.

Tomocerodinae differ from Moranilinae in features discussed by Bouček (1988), most noticeably in the much shorter Gt₁, which is the longest tergum in Moranilinae but is much shorter than Gt₂ in Tomocerodes.

Antenna with 10 (Neodipara) or 11 (Elatoides) flagellomeres, including 4 clavomeres (Fig. 42) and an inconspicuous anellus. Clypeus without transverse subapical groove. Labrum hidden, flexible, subcordiform with a median lobe, with marginal setae projecting forward from the lateral lobes (Fig. 43). Mandibles with 2 teeth (Neodiparinae) or with 2 teeth in the left mandible and 3 in the right (Elatoidinae). Subforaminal bridge with postgena separated by lower tentorial bridge. Mesoscutellum with frenum indicated at least laterally, without axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb oblique. Metasoma with a separate epipygium.

Figures 43–48. 

43 Neodipara masneri Bouček (Neodiparidae): head frontal view 44, 45 Oodera formosa Giraud (Ooderidae) 44 mesosoma dorsal view 45 fore leg 46 Leptofoenus stephanoides (Roman) (Pelecinellidae): petiole 47 Doddifoenus rex Bouček (Pelecinellidae): head antero-lateral view 48 Nefoenus pilosus Bouček (Pelecinellidae, Nefoeninae): mesosoma lateral view.

Although similar in habitus to some species with a long petiole from other families, such as Spalangiidae or some Pteromalidae, Neodiparidae differ from these in having a small separate epipygium instead of a syntergum, an oblique basitarsal comb, and a relatively large 4^th clavomere. Elatoidinae new placement is transferred here, with its single genus Elatoides Nikol’skaya, differing from Neodiparinae in having a complete set of 11 flagellomeres instead of 10 and in the right mandible having 3 teeth.

Antenna with 11 flagellomeres, including 3 clavomeres. Eyes ventrally divergent. Clypeus with transverse subapical groove. Labrum exposed, sclerotized. Mandibles with 2 or 3 weakly separated teeth or essentially truncate. Pronotum elongate, with lateral surfaces divergent such that the pronotum islaterally expanded (Fig. 44). Notauli complete, linear except for a short distance anteriorly, and almost meeting posteriorly, forming a distinctive 4-pronged pattern with the also sublinear part of the transscutal articulation that occurs along the anterior edge of the axilla (although this is often broken by a transverse fracture across the sulci) (Fig. 44). Mesoscutellum without a frenum but with an expanded posterior rim of the mesoscutellum that can resemble a frenum; without a distinct axillular sulcus but with some longitudinal sculpture in the axillular area. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; profemur expanded and with multiple rows of ventral spine-like structures (described in detail by Gibson 2003) (Fig. 45); protibial spur stout and curved; basitarsal comb longitudinal; metafemur not expanded and lacking ventral teeth. Metasoma with syntergum, therefore without epipygium.

Oodera Westwood, the only genus in Ooderidae, is highly distinctive and does not resemble any other Chalcidoidea, especially in mesosomal features. The expanded profemur of Oodera can be compared with that of some Heydeniidae, which is also expanded but lacks the additional rows of spine-like structures of Oodera, instead having broad ventral crest-like projections. Additionally, the laterally expanded pronotum also occurs in some Heydeniidae, which can have a somewhat comparable pattern of sulci on the mesoscutal dorsum, even though they are often less distinct in Heydeniidae. Otherwise, Pelecinellinae (Pelecinellidae) have parascrobal crests as in Oodera, although the two groups are differ in many other features.

Antenna with 11 flagellomeres. Clypeus without transverse subapical groove. Mandibles with 3 teeth or with a broad apical truncation. Subforaminal bridge with postgenal bridge. Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Petiole elongate with a row of lateral setae (Fig. 46).

Leptofoeninae is here recognized as a junior synonym of Pelecinellidae, since Pelecinellinae (Ashmead 1895) was described earlier than Leptofoeninae (Handlirsch 1925) and because the situation does not qualify for preserving prevailing usage. Pelecinellidae are similar to other large-bodied chalcidoid parasitoids of wood-boring beetles, although they differ radically from all Chalcidoidea morphologically. Nefoeninae new placement is included here as a subfamily distinct from Pelecinellinae, on the strength of sharing the elongate petiole with lateral setae (Fig. 46) although it lacks the parascrobal crests present in Pelecinellinae (Fig. 47). An elongate petiole with lateral setae is unusual but not unique in Chalcidoidea, being present also in Polstonia Heydon (Pteromalidae: Miscogastrinae: Sphegigastrini), some Spalangia Latreille (Spalangiidae: Spalangiinae), and in some Orasema Cameron (Eucharitidae: Oraseminae) each of these differing greatly from Pelecinellidae in many other features. However, the form of the petiole in Nefoeninae (Fig. 48) is somewhat similar to that of Pelecinellinae, and the two groups share several other features, including elongation of the pronotum and certain other areas of the mesothoracic dorsum. While Ooderidae also have parascrobal crests, the pattern of sulci present on the mesothoracic dorsum in Ooderidae is unmistakable.

Antenna with at most 11 flagellomeres, including 1 or more visible anellus, not counting any indistinct anelli that are usually present (Fig. 49). Eyes either not ventrally divergent, or diverging linearly (Cecidellinae, Eriaporinae, Euryischiinae), instead of with a concave medial margin in their lower half as in Cleonyminae and others (the exceptions are some male Macroglenes Westwood with huge eyes). Clypeus without transverse subapical groove. Labrum hidden, flexible, subrectangular with marginal setae in a row. Mandibles with 3 or 4 teeth, splayed in a characteristic way (Bouček 1988) (Fig. 50). Subforaminal bridge with postgena separated by lower tentorial bridge. Notauli complete. Mesoscutellum with frenum indicated at least laterally, and with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres, except in Zebe La Salle with 4; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Figures 49–54. 

49 Macroglenes varicornis (Haliday) (Pirenidae): antenna 50 Gastrancistrus sp. (Pirenidae, Tridyminae): head frontal view 51 Cecidellis sp. (Pirenidae, Cecidellinae): petiole with lamina 52 Spathopus sp. (Pirenidae, Tridyminae): antenna 53 Spalangia alycia Gibson (Spalangiidae): head anterolateral view 54 Erotolepsia sp. (Spalangiidae, Erotolepsiinae): head frontal view.

The family Eriaporidae is synonymized with Pirenidae, with Eriaporinae and Euryischiinae retained as separate subfamilies. Cecidellinae is described for the unusual genus Cecidellis.

Pirenidae most strongly resemble those few Pteromalidae that have 11 antennal flagellomeres, otherwise differing from most in having 5 funiculars or fewer, without enough visible anelli to bring the total flagellomeres before the clava to the count of 8 that is present in nearly all Pteromalidae. Out of those Pteromalidae with 11 flagellomeres, Termolampa pinicola Bouček differs in having incomplete notauli, Andersena anomala Andersen differs in having no anelli. Bugacia Erdős differs in having the vertex with blunt carina or crest, and Trigonoderopsis Girault differs in having 8 flagellomeres between pedicel and clava. Eunotidae differ in having an exposed, rigid labrum and divergent eyes with a concave medial margin in their lower half. While this may make Eunotidae and Pirenidae sound very similar to one another, the habitus of Eunotidae is very different from most Pirenidae, being stout and flattened instead of being more moderate in body proportions and with a deeper mesosoma. Eriaporinae are the pirenids most likely to be confused with Eunotidae, but differ most conspicuously in having stout setae on the parastigma. Moranilidae differ in having 2 mesofurcal pits, in having pits on the mesopleural area of the mesopectus, and in having an oblique basitarsal comb. Herbertiidae and Systasidae differ from Pirenidae in having 2 mandibular teeth, and along with Micradelinae, an exposed, rigidly sclerotized labrum.

Cecidellis Hanson, 2005.

Body pale, white to yellowish or pale brown, without metallic luster. Antenna with 9 flagellomeres, including 4 funiculars and 2 anelli. Eyes linearly diverging in ventral half. Petiole with lamina that overlaps part of the propodeal margin (Heydon and Hanson 2005) (Fig. 51). Otherwise as in Pirenidae.

Body usually dark brownish or metallic. Antenna with at most 3 large flagellomeres and at least 2 anelli before clava (Fig. 49). Eyes usually not or only slightly diverging ventrally (except some males). Marginal vein at least 3.5× as long as the short and mostly straight stigmal vein. Petiole without dorsal lamina.

In this new, more restricted sense, Pireninae contains genera that are morphologically similar to Macroglenes. They are here distinguished from Tridyminae, which are generally more stout in body shape and differ in features mentioned in diagnoses of both subfamilies, but most prominently in the antennal flagellum and relative lengths of the marginal and stigmal veins of the fore wing.

Body usually metallic, except Calyconotiscus Narendran & Saleem. Antenna with 4 or 5 large flagellomeres and at least one anelliform flagellomere before clava (Fig. 52). Eyes not divergent ventrally. Marginal vein at most 3× as long as the relatively long stigmal vein. Petiole without dorsal lamina.

The subfamily Tridyminae is removed from synonymy with Pireninae to include Gastrancistrus Westwood new placement and related genera. Calyconotiscus Narendran & Saleem new placement, Ecrizotes Förster new placement, Epiterobia Girault new placement, Melancistrus Graham new placement, Oxyglypta Förster new placement, Premiscogaster Girault new placement, Sirovena Bouček new placement, Spathopus Ashmead new placement, and Spinancistrus Kamijo new placement are here confirmed to belong to this subfamily.

Antenna usually with 8 flagellomeres, including a 1-segmented clava, with 11 flagellomeres including 3 clavomeres in Eunotopsia Bouček. Clypeus without transverse subapical groove. Labrum exposed, well-sclerotized, subrectangular or semicircular with marginal setae in a row (Fig. 53). Mandibles with 2 or 3 teeth, or undivided (in Eunotopsia). Subforaminal bridge with postgenal bridge or with postgena separated by lower tentorial bridge. Mesoscutellum with frenum indicated at least laterally, and without axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

There are two distinctive subfamilies in Spalangiidae, Spalangiinae and Erotolepsiinae new placement, both comprised of parasitoids of Diptera. The antennal toruli are placed very low on the head in most species of both subfamilies (Fig. 54). In most Erotolepsiinae (except Eunotopsia where they are placed higher), the toruli are just above the very short clypeus, while in Spalangiinae the toruli are placed on lobes that overhang the clypeus and labrum, rendering them difficult to see. This distinctive antennal placement and the prognathous head make members of Spalangiinae easily identifiable. Most Erotolepsiinae (again, not Eunotopsia) are readily identified by the presence of a long carina that encircles most of the face, extending from near the median ocellus to the top of the clypeus (Fig. 54), and have a transverse carina across the anterior part of Gt₁ (Fig. 55). The enigmatic genus Eunotopsia shares the transverse carina on Gt₁ with other Erotolepsiinae and can be identified to subfamily using this feature. Erotolepsiinae strongly resemble Herbertiidae and Micradelinae, which differ in lacking the carinae of the face and Gt₁. Herbertiidae further differ in having an oblique basitarsal comb, although the basitarsal comb of Micradelinae is reduced and difficult to evaluate. Eunotidae are also somewhat similar to Erotolepsiinae, but differ in that the mesepimeron does not overlap the anterior margin of the metapleuron, and in lacking the Gt₁ sculpture of Erotolepsiinae.

Figures 55–60. 

55 Erotolepsia sp. (Spalangiidae, Erotolepsiinae): mesosoma and base of metasoma dorsal view 56a, 57 Systasis sp. (Systasidae) 56a apex of clypeus without subapical groove 57 mesosoma ventral, mesotrochantinal plate and mesofurcal pit 58 Trisecodes africanum Gumovsky (Pirenidae, Trisecodinae): antenna 56b, 59 Asaphes sp. (Asaphesinae, incertae sedis) 56b clypeal subapical groove 59 mesosoma ventral 60 Austrosystasis atricorpus Girault (Austrosystasinae, incertae sedis) 60 head frontal view, mesosoma lateral view.

Antenna with 7 or 11 flagellomeres, including 1 or more anellus and a small 4^th clavomere. Eyes not ventrally divergent. Clypeus without transverse subapical groove. Labrum exposed, well-sclerotized (Fig. 56a). Subforaminal bridge with postgena separated by lower tentorial bridge. Notauli complete. Mesoscutellum with short frenum indicated at least laterally, with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron; mesofurcal pit on mesotrochantinal plate directly between the mesocoxal insertions (Fig. 57). Protibial spur curved; basitarsal comb oblique; all legs with either 5 (Systasinae) or 3 (Trisecodinae) tarsomeres. Metasoma with syntergum, therefore without epipygium.

Systasinae are most likely to be confused with Pirenidae and Pteromalidae, which differ in having a flexible labrum that is concealed behind the protruding clypeus, whereas the clypeus in Systasinae recedes medially to expose the sclerotized labrum; they also have a longitudinal basitarsal comb, whereas it is oblique in Systasinae. The position of the mesofurcal pit in Systasidae is very unusual, although a leg may need to be removed to see it. Trisecodinae can be confused with Trichogrammatidae based on the 3-segmented tarsi, the head sulci, and the setal lines on the fore wing, and with some Eulophidae, based on the reduced number of flagellomeres, the head sulci, the setal lines on the fore wing, and the very short postmarginal and stigmal veins. From the former, Trisecodinae differ in the longer flagellum, the narrowly attached gaster with phragma restricted to mesosoma, the different pattern of head sulci, and the shape of the fore tibial spur. From the latter, although Trisecodes was preliminary placed in Entedoninae (Delvare & LaSalle, 2000), Trisecodinae differ in various features that do not fit with any current eulophid subfamily. While Trisecodes is easy to distinguish from other Systasidae due to the difference in tarsomere count, it is retained in this family to indicate the phylogenetic context provided by both the molecular and morphological data.

Trisecodes Delvare & LaSalle, 2000.

Antenna with 7 flagellomeres (Fig. 58); multiporous plate sensilla unusually long and curved; head except malar sulcus with frontal, scrobal and subtorular sulci; anterior tentorial pits absent; toruli at or below the lower ocular line; all legs with 3 tarsomeres.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Clypeus with transverse subapical groove. Head dorsally with temple separating posterior margin of eye from that of the head. Labrum exposed, well-sclerotized, subrectangular with marginal setae in a row. Mandibles with 2 or 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge; occipital carina present. Mesoscutellum with frenum indicated at least laterally, and with axillular sulcus. Mesopleural area without an expanded acropleuron, with pits; mesepimeron not extending over anterior margin of metapleuron; two mesofurcal pits usually present (Fig. 59) (absent in the fossil genus Coriotela). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb oblique. Metasoma with syntergum, therefore without epipygium.

The scope of Asaphesinae is much reduced with the removal of Enoggerinae new subfamily to a separate incertae sedis subfamily in Chalcidoidea, and Bairamlia Waterston to Sphegigastrini (Pteromalidae). Asaphesinae is part of a set of taxa with two mesofurcal pits and some other shared features, but which do not form a clade in molecular analyses (Cruaud et al, submitted), including Enoggerinae, Herbertiidae, and Moranilidae. Asaphesinae differ from Herbertiidae and Moranilidae in having 12 flagellomeres instead of a maximum of 10. Enoggerinae differ in lacking a temple, thus with the posterior margin of the eye dorsally meeting that of the head. Some Asaphesinae resemble Pteromalidae in habitus, differing in the clypeus, reduced mandibles, labrum, oblique basitarsal comb, and presence of two mesofurcal pits, but also having features that are rarely found in Pteromalidae, such as an occipital carina and the mesopleural area with pits.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Eyes slightly linearly divergent ventrally. Clypeus with transverse subapical groove. Labrum hidden, flexible, with marginal setae in a row (Fig. 60). Subforaminal bridge with postgena separated by lower tentorial bridge, with a short apparent postgenal bridge immediately dorsal to the hypostoma. Mesoscutellum with frenum indicated laterally, with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron (Fig. 61). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium, rigidly convex.

Figures 61–66. 

61 Austrosystasis atricorpus Girault (Austrosystasinae, incertae sedis): mesosoma lateral view 62–64 Ditropinotella sp. (Ditropinotellinae, incertae sedis) 62 antenna 63 mesosoma lateral view 64 metasoma 65 Enoggera reticulata Naumann (Enoggerinae, incertae sedis): mesosoma lateral view 66 Eopelma sp. (incertae sedis): apex of mesotibia and mesotarsus.

Austrosystasis atricorpus Girault, the sole species in this subfamily, has not been sequenced, and its place is uncertain given our incomplete knowledge of its morphology. It appears to be a member of the Gall Clade, and it is an associate of galls on Elaeocarpus (Elaeocarpaceae) in Australia. It has rough surface sculpture (Fig. 61) that causes it to resemble Eurytomidae (especially Rileyinae), although it has a shorter pronotum. Otherwise, it resembles Melanosomellidae, differing in having a distinct and complete axillular sulcus and distinctive sculpture on the axillula. It also resembles Ormyridae in overall body shape, but this could be attributed to gall association in both taxa. The differing sculpture of the axillula and the posterior surface of the head separate Austrosystasinae from Ormyridae. Given that A. atricorpus would complicate the diagnosis of whatever family it could be transferred to, the genus is dealt with here as incertae sedis in Chalcidoidea.

Antenna with 11 flagellomeres, without a 4^th clavomere (Fig. 62). Eyes slightly divergent ventrally. Clypeus without a transverse subapical groove, with a small median incision. Labrum hidden behind clypeus, flexible. Mandibles with 3 teeth. Occipital carina absent. Subforaminal bridge with a postgenal bridge. Notauli complete. Mesoscutellum with a densely setose frenum that is indicated laterally, without axillular sulcus (Fig. 63). Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with an elongate, T-shaped syntergum in females that may resemble an epipygium because of its shape (Fig. 64).

Ditropinotella Girault is a morphologically enigmatic Australasian genus of gall associates, transferred out of Torymidae and placed in its own subfamily in Pteromalidae by Bouček (1988). It renders Eupelmidae paraphyletic in next-generation molecular data (Cruaud et al., submitted), although it lacks the expanded acropleuron of that family and lacks the diagnostic features of all genera in Calosotinae. Ditropinotella has a broad membranous area posterior to its mesocoxae, although this also occurs in various other chalcidoids that are not related to Eupelmidae. The possibility remains that Ditropinotella may be a reduced eupelmid, but morphological evidence in support of this possibility is lacking. Because of the possible instability of this situation, Ditropinotellinae is removed from Pteromalidae to be treated as incertae sedis in Chalcidoidea.

The general habitus, setose frenum, and approximated, slightly advanced axilla of Ditropinotella invite comparison with Torymidae, which differ in having a true epipygium in females that is shorter and not so elongate. Males are more difficult to distinguish, differing in the slightly divergent eyes and incised clypeus of Ditropinotella, features that do not occur together in Torymidae. Most Megastigmidae also resemble Ditropinotella, although most Megastigmidae and Torymidae have an occipital carina. Megastigminae additionally differ from Ditropinotellinae in having an enlarged fore wing stigma and along with Chromeurytominae have a true epipygium in females, while Keiraninae have an occipital carina and do not have an elongate syntergum. Although some pteromalid fig associates have an elongate epipygium that resembles the syntergum of Ditropinotella, these differ from Ditropinotella in having a larger axillula with a distinct axillular sulcus. Male Eupelminae can strongly resemble those of Ditropinotella, but differ in having a distinct frenal arm laterally.

Enoggera Girault, 1926.

Antenna with 9–12 flagellomeres, including either an incompletely divided clava or up to 4 clavomeres, sometimes including a small 4^th clavomere. Temple absent, thus posterior margin of eye coincident with the posterior margin of the head dorsally (Fig. 65). Clypeus with transverse subapical groove. Labrum exposed, well-sclerotized, subrectangular, with marginal setae in a row. Mandibles with 3 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge. Mesoscutellum with frenum indicated at least laterally, with axillular sulcus. Mesopleural area without an expanded acropleuron, with pits; mesepimeron not extending over anterior margin of metapleuron; two mesofurcal pits present. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb oblique. Metasoma with syntergum, therefore without epipygium. Egg parasitoids of Chrysomelidae (Coleoptera).

Enoggerinae share a pair of mesofurcal pits with a number of other chalcidoid groups, including Herbertiidae, Moranilidae, and Asaphesinae. However, these groups are unstable in molecular analyses and do not form a clade (Cruaud et al., submitted), with Enoggerinae more often as the sister group of Coelocybidae. Therefore, Enoggerinae can be separated from all similar groups by the absence of the temple. With a different biology, Enoggerinae would also represent a discordant element if placed in any of the other groups.

Antenna with 8 flagellomeres, with an undivided clava. Eyes ventrally divergent. Clypeus without transverse subapical groove. Labrum hidden behind clypeus, flexible. Mandibles with 3 teeth or with a ventral tooth and dorsal slightly emarginate truncation. Axilla long, separated anteromedially. Axillular sulcus more or less distinct. Frenum absent. Acropleuron enlarged, convex and pad-like; covering most of mesopleural area, separated from mesocoxa by the lower mesepimeron, metapleuron, and a dorsal extension of the metasternal area. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; ventral membranous area anterior to mesocoxal attachment lacking; mesotibial spur stout; mesotarsus with 1 row of pegs anteroventrally (Fig. 66). Metasoma with syntergum, therefore without epipygium.

Eopelma is consistently the sister group of another incertae sedis taxon, Storeya Bouček (Storeyinae), in next-generation molecular analysis (Cruaud et al., submitted). These two groups do not strongly resemble each other in body shape. Storeya does not have an expanded acropleuron, and has not previously been indicated as a relative of any eupelmid. They do share an antennal flagellomere count, a long radicle, general coloration, and an undivided clava, but the list of shared features possessed by these two genera is much shorter than the list of differences between them. A new subfamily is not described for Eopelma because it is a single genus, the position of which in Chalcidoidea is still in question.

Eopelma vaguely resembles Neanastatus Girault in body shape and coloration. Neanastatus differs in having a much smaller axilla, which may not be clearly visible.

Antenna with 10 flagellomeres, including a single anellus (Fig. 70). Clypeus with lateral sulci but without a dorsal sulcus; ventral margin protruding and slightly convex but not bilobed (Fig. 70). Toruli slightly above center of face, immediately below a forked sulcus that exposes conjunctiva. Trabeculae absent (although sulcus present in middle of upper face). Notauli complete, incised. Frenum present, indicated by a complete frenal sulcus (Fig. 71). Axillular sulcus present. Acropleuron not enlarged (Fig. 71). Fore wing densely setose, without speculum; basal fold pigmented. Legs with 4 tarsomeres on all legs; protibial spur short and straight. Cerci surrounded by conjunctiva.

Figures 67–72. 

67–69 Heydenia longicollis (Cameron) (Heydeniidae) 67 mesosoma dorsal view 68 mesosoma lateral view 69 fore leg 70, 71 Kerya antennalis Bouček (Keryinae, incertae sedis) 70 head frontal view and antenna 71 mesosoma lateral view 72 Callimomoides sp. (Louriciinae, incertae sedis): head frontal view.

The combination of having just 4 tarsomeres on all legs, 10 flagellomeres, and a frenum, together with a lack of trabeculae, makes Kerya antennalis Bouček distinct from all other chalcidoids. It was once placed in Eulophinae (Bouček 1988), although analysis of 28S D2 ribosomal DNA indicated that it is not a eulophid and it was moved to Ormocerinae based on the placement of the toruli and the arched body (Gauthier, et al. 2000). Indeed some Melanosomellidae resemble K. antennalis in body shape, color, and smooth surface sculpture, but they have 11 or more flagellomeres, a bilobed clypeal margin, 5 tarsomeres, a stout and curved protibial spur, cerci surrounded by cuticle, no axillular sulcus, and no forked sulcus on the face. Eulophidae and some Aphelinidae also have 4 tarsomeres and are relatively generalized (unlike Calesidae and Idioporidae), but have fewer flagellomeres and no frenum. Given the strong differences between K. antennalis and all the taxa with which it has been previously placed, and because it has not been analyzed using next-generation molecular data, Keryini is here elevated to subfamily status and placed as incertae sedis in Chalcidoidea.

Antenna with 8 flagellomeres, including an undivided clava and 2 anelli; radicle elongate. Eyes ventrally divergent. Face with a network of grooves that is usually concealed by the strongly collapsing head (Fig. 72). Clypeus without transverse subapical groove. Labrum subrectangular and exposed, with marginal setae in a row. Mandibles with 3 teeth. Subforaminal bridge with a postgenal bridge separating the secondary tentorial pits from the hypostoma. Pronotum long, with a slightly expanded lateral surface and therefore somewhat expanded laterally. Notauli complete, linear and incised. Axilla strongly advanced (Fig. 73). Mesoscutellum with frenum indicated at least laterally or by an abrupt transition to smooth surface sculpture, without axillular sulcus (Fig. 73). Mesopleural area without an expanded acropleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma in females with elongate syntergum extending over the exserted ovipositor (Fig. 74).

Figures 73–78. 

73, 74 Callimomoides sp. (Louriciinae, incertae sedis) 73 mesosoma dorsal view 74 extremity of metasoma, elongated syntergum 75, 76 Micradelus acutus Graham (Micradelinae, incertae sedis) 75 head frontal view 76 head posterior view 77 Neapterolaelaps sp. (Neapterolelapsinae, incertae sedis): antenna 78 Pseudoceraphron albifrons (Bouček) (Neapterolelapsinae, incertae sedis): head posterior view.

Next-generation molecular analysis (Cruaud et al., submitted) consistently recovers the distinctive Callimomoides Girault as a member of a strongly supported clade that also includes Neanastatus and Lambdobregma, with Callimomoides as the sister group of Neanastatus, which therefore renders Neanastatidae paraphyletic. Morphologically, this relationship is highly debatable and no unique character supports it. However, this strong morphological disparity may be due to difference in life history as Callimomoides is an egg parasitoid of Cerambycidae while Neanastatus is parasitic in galls of Cecidomyiidae (Diptera) and Lambdobregma could be parasitoids of cricket eggs (Gibson 1989). While Callimomoides has an enlarged mesotibial spur and a large membranous area posterior to the mesocoxa, these features are not conclusive proof of relationship with Neanastatidae because they are found in various other taxa as well. There are no diagnostic features shared with either Neanastatus or Lambdobregma.

The highly unusual gestalt of Callimomoides, together with the combination of linear notauli, long pronotum, and stout mesotibial spur, prevent it from being easily confused with other Chalcidoidea. Eulophidae can have similar notauli and a weakly sclerotized, collapsing head and body, but differ in having 4 tarsomeres on all legs.

Antenna with 10 or 11 flagellomeres, including 3 or 4 clavomeres. Eyes slightly ventrally divergent. Clypeus transverse, with transverse subapical groove. Labrum exposed, well-sclerotized, subrectangular with marginal setae in a row (Fig. 75). Mandibles with 2 teeth or with a small 3^rd dorsal tooth (Fig. 75). Subforaminal bridge with postgena separated by lower tentorial bridge except for a small postgenal bridge dorsal to the hypostoma; posterior surface of head without (Fig. 76) postgenal lamina or postgenal groove. Pronotum transverse in dorsal view. Mesoscutellum with frenum reduced, with frenal arm laterally but sometimes hardly separated from marginal rim of mesoscutellum; axillular sulcus present. Mesopleural area without expanded acropleuron, without pits; mesepimeron extending over anterior margin of metapleuron; one mesofurcal pit present. Fore wing marginal vein subequal to stigmal vein in length; uncus elongate. All legs with 5 tarsomeres; protibial spur stout and curved. Metasoma with syntergum, therefore without epipygium.

The former tribe Micradelini was treated as a subfamily by Vago (2006) and we preserve that rank here, although we remove it from Pteromalidae based on both molecular and morphological data (Cruaud et al., submitted). While Micradelinae are very similar to Herbertiidae in body shape, they only occasionally form a monophyletic group and Micradelinae are particularly unstable in phylogenetic placement.

Morphologically, Micradelinae strongly resemble Herbertiidae and the incertae sedis genus Rivasia Askew & Nieves-Aldrey in habitus and other features. The basitarsal comb of Micradelus Walker is reduced and difficult to evaluate, although it may be oblique like that of Herbertiidae. Herbertiidae differ from Micradelinae in having a much shorter stigmal vein relative to the marginal vein, in lacking an axillular sulcus, in lacking the postgenal groove and postgenal sulcus on the back of the head, and in having 2 mesofurcal pits instead of the single pit of Micradelinae. Rivasia is very similar to Micradelinae in most features, differing in having a more elongate body including the pronotum, and in having metallic green coloration instead of the short, stout body and brownish coloration of Micradelinae. Eunotidae may appear similar to Micradelidae when comparing lists of features, but in practice are easily distinguished in their different habitus and in having a short fore wing uncus.

Neapterolelaps Girault, 1913.

Antenna with 11 flagellomeres, with 3 clavomeres (Fig. 77). Eyes ventrally divergent. Clypeus with a transverse subapical groove. Labrum exposed, sclerotized, subtriangular with setae. Mandibles with 2 or 3 narrow teeth. Subforaminal bridge with elongate lower tentorial bridge and secondary tentorial pits that extend to the convergent hypostomal carina, with or without a postgenal groove and postgenal lamina (Fig. 78). Pronotum without a smooth median longitudinal line or carina. Mesoscutellum without frenum, at least sometimes with a small axillula indicated by an axillular sulcus or carina. Mesopleural area without an expanded acropleuron; mesepimeron not extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; metafemur without ventral teeth or expansion. Metasoma with syntergum, therefore without epipygium; cercal brush present.

Neapterolelaps and Pseudoceraphron Dodd form the sister group of Lyciscidae in next-generation molecular data, although they were previously classified in Diparinae (Bouček 1988; Desjardins 2007; Desjardins et al. 2007). While Desjardins (2004) mentioned the name Neapterolelapini in his doctoral dissertation, it was not mentioned in the two resulting publications. Additionally, it was mentioned by Heraty et al. (2013), although it was not diagnosed in that publication and therefore was not described there. Janšta (2014) also mentioned Neapterolelapini in a doctoral dissertation, but did not diagnose it. None of these previous usages satisfy article 13.1.1 of the ICZN Code, and therefore Neapterolelapinae is described as new here and it is removed from Diparidae to be treated as incertae sedis in Chalcidoidea. Nosodipara Bouček is also placed here based on morphology.

Given the characters described here, confusion with Lyciscidae is most likely, which differ in having a longitudinal median pronotal carina. However, a lack of metallic coloration on the mesosoma (but sometimes not of the head) of females makes Neapterolelapinae much more likely to be confused with Diparidae, which contains numerous species that resemble this group. Diparidae differ in having a conspicuous frenum in most species, although this may be indistinct or absent in highly derived brachypterous species. These must be distinguished using features of the head, such as the hidden labrum, convex clypeal margin, absence of clypeal subapical groove (excepted in Dipara) and striation of metacoxa of Diparidae versus the exposed labrum, concave clypeal margin, presence of clypeal subapical groove and absence of striation on metacoxa in Neapterolelapinae.

The placement of Neapterolelapinae does not conflict with the findings of Desjardins et al. (2007), who also placed Neapterolelaps as the sister group of what is now Lyciscidae in molecular analyses. This placement was discussed as “difficulty in uniting” [Diparinae] by Desjardins (2007).

Antenna with 11 flagellomeres, including 1 anellus and 7 funiculars (Fig. 79a). Eyes ventrally not or slightly divergent. Clypeus without transverse subapical groove. Labrum hidden (Fig. 80). Mandibles with 2 teeth. Mesoscutellum with frenum not distinctly indicated, without axillular sulcus. Mesopleural area without an expanded acropleuron, without pits; mesepimeron with posterior margin deeply concave and extending over anterior margin of metapleuron. All legs with 5 tarsomeres; protibial spur stout and curved. Marginal and postmarginal veins widened adjacent to stigmal vein (Fig. 79b).

Figures 79–84. 

79 Parasaphodes afer Mitroiu (Parasaphodinae, incertae sedis) 79 a antenna b venation 80 P. iceryae (Ashmead) (Parasaphodinae): head frontal view 81, 82 Storeyia sp. (Storeyinae, incertae sedis) 81 head lateral view and antenna 82 venation 83 Caenocrepis sp. (Pteromalidae, Pteromalinae): lower face 84 Nasonia vitripennis (Walker) (Pteromalinae): head dorsal view.

No molecular data are available for Parasaphodinae; moreover, because of the poor preservation of most available specimens, several characters such as the basitarsal comb, the structure of the back of the head, or the mesofurcal pits could not be observed. It is thus very difficult to assess the taxonomic position of Parasaphodinae in relation to other chalcids, therefore we treat it as incertae sedis in Chalcidoidea. Similarities with Asaphesinae, Elatoidinae and Herbertiidae have been discussed (Mitroiu 2017), but all these groups differ from Parasaphodinae in many respects.

Antenna with 8 flagellomeres, including an undivided or incompletely divided clava. Eyes slightly ventrally divergent. Clypeus transverse. Labrum exposed, well-sclerotized. Mandibles with 2 teeth. Subforaminal bridge with postgena separated by lower tentorial bridge except for a small postgenal bridge dorsal to the hypostoma; posterior surface of head without postgenal lamina or postgenal groove. Pronotum subconical and elongate in dorsal view. Mesoscutellum with frenum; mesopleural area without an expanded acropleuron, without pits; mesepimeron extending over anterior margin of metapleuron; one mesofurcal pit present. Fore wing marginal vein subequal to stigmal vein in length; uncus elongate. All legs with 5 tarsomeres; protibial spur stout and curved. Metasoma with syntergum, therefore without epipygium.

Rivasia is usually the sister group of fellow incertae sedis taxon Asaphesinae in molecular analyses (Cruaud et al., submitted), although the two groups share few distinctive features morphologically. Instead, Rivasia more strongly resembles the subfamily Micradelinae, though never forming a clade with it. Rivasia shares no particularly distinctive features with Ormocerinae (Pteromalidae) nor any former part of it, differing from them in numerous features listed in the diagnosis.

Micradelinae can be distinguished from Rivasia by having a shorter, transverse pronotum and brownish coloration, in contrast with the longer pronotum and metallic coloration of Rivasia. Asaphesinae differ from Rivasia in a number of features, including the presence of 8 flagellomeres instead of 12, the lack of an occipital carina, and from extant species in the lack of pits on the mesopleural area.

Antenna with 8 flagellomeres, including an undivided clava; radicle elongate (Fig. 81). Intertorular prominence absent (Fig. 81). Eyes slightly divergent ventrally. Clypeus without transverse subapical groove. Labrum hidden behind clypeus, flexible, subrectangular and with marginal setae in a row. Mandibles with 3 teeth (may be 2 in some specimens in which it is difficult to see). Subforaminal bridge with a postgena separated by lower tentorial bridge. Notauli incomplete. Mesoscutellum with frenum indicated; with axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron. Fore wing with a tuft of thickened leaf-shaped setae on the parastigma (Fig. 82). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Metasoma with syntergum, therefore without epipygium.

Molecular and morphological data do not provide a clear signal on where this subfamily should be placed, and therefore we treat it as incertae sedis in Chalcidoidea. The elongate antennal radicle is an unusual feature also shared by its sister group (Eopelma) in molecular analyses and, in comparable taxa, is only found in Louriciinae. The enlarged axillula, easily visible dorsally, invites comparison with Colotrechninae and Sycophaginae in Pteromalidae, and with Hetreulophidae, although each of these groups differs from Storeyinae in having complete notauli and in many other features as well. The low placement of the toruli may cause Storeyinae to key near Spalangiinae in family keys, although the toruli are not on lobes and the two groups differ in many features. The body coloration (excepted for one undescribed iridescent species) and the tuft of thickened, leaf-shaped setae on the parastigma recall Cerocephalidae, although Storeyinae lack an intertorular prominence and are not placed with cerocephalids using molecular data.

Antenna with 12 flagellomeres in nearly all cases (except in some fig associates, Amphidocius, Andersena anomala, Bugacia, Trigonoderopsis, and possibly Termolampa pinicola), including a small 4^th clavomere; with at least 5 funiculars, and if with 5 then with 2 or more anelli. Eyes not ventrally divergent (although some genera have modified eyes that are difficult to evaluate). Clypeus subquadrate and without ventral transverse groove (Fig. 83). Labrum flexible (Darling 1988) and hidden behind clypeus. Mandibles with 3 or 4 teeth each (except in some fig associates (1–3 teeth) and in Austroterobia Girault where the left mandible has 2 teeth but is falcate). Subforaminal bridge with postgena separated by lower tentorial bridge; posterior surface of the head without postgenal lamina or postgenal groove (Fig. 84). Mesoscutellum with frenum indicated at least laterally, with axillular sulcus (except in some fig associates and Nikolskayana mirabilis). Mesopleural area without an enlarged acropleuron. All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Gaster, while sometimes rigidly convex, not strongly sclerotized; metasomal apex in most species with a syntergum and therefore without an epipygium (except in some fig associates).

Agaonidae are similar to fig-associated Pteromalidae, differing from all in having a mandibular appendage bearing rows of spurs or lamellae in females (Seres Waterston, which may appear especially similar to Agaonidae, have an enlarged plate-like mandible, see figs 14–16 in van Noort and van Arten 2006, but do not have a mandibular appendage), and in more or less fused anelli, the last one mostly spine-like (Fig. 85). Male Agaonidae differ from male pteromalid fig wasps in that the metasomal apex is telescoped in a U-shaped arrangement under the body. Eucharitidae differ from most Pteromalidae in that the pronotum is not visible from dorsal view, but also in the flattened marginally digitate labrum. Likewise, most of the members of the Planidial Clade (Zhang et al. 2022) such as Chrysolampidae (digits sometimes absent in Chrysolampinae and labrum plate-like in Philomidinae), Eutrichosomatidae, and Perilampidae have a digitate labrum. Eulophidae differ in having 4-segmented tarsi and a short, straight protibial spur. Some Eurytomidae, such as Rileyinae and Buresium Bouček can be similar to a few Pteromalidae, even though nearly all Pteromalidae have a much smaller pronotum; in a few cases where the pronotum is long, e.g. Trigonoderinae, it is not subrectangular in dorsal view, but instead is gradually narrowing anteriorly. Rileyinae and Buresium differ from Pteromalidae in having a postgenal groove and a strongly sclerotized gaster. In practice, Rileyinae and Buresium are easily recognized once they are familiar, and no Pteromalidae have the combination of a long pronotum with a rigidly convex gaster. Ormyridae also differ from Pteromalidae in having a carapace-like gaster. Other members of the Gall Clade, such as Epichrysomallidae and Melanosomellidae, can be very similar to those pteromalids with complete notauli, but they differ in having either no indication of an axillular sulcus or carina, or in having a reduced and incomplete one. Pirenidae differ in having fewer flagellomeres: most especially if they have 5 funiculars, then they only have 1 anellus. Torymidae and Megastigmidae differ from most Pteromalidae in having an epipygium in females (except in Keirana Bouček which has a transverse sulcus across its syntergum immediately anterior to the cerci), but some pteromalid fig associates have a separate epipygium and a long, exserted ovipositor, therefore strongly resembling both of these families. Torymidae differ from all Pteromalidae in the form of their postgenal bridge, which occurs in the form of postgenal lobes meeting each other over the hypostoma (Burks and Heraty 2015). Most fig-associated Pteromalidae have the postgena separated by the lower tentorial bridge (Rasplus et al. 1998), additionally differing in ways described by Grissell (1995). Those with an elongate head, such as Sycoecus Waterston, have rederived a true postgenal bridge but differ from Torymidae in the shape of the highly modified head itself and in the vast number of other unusual features of their highly modified bodies.

Figures 85–90. 

85 Blastophaga psenes (L.) (Agaonidae): antenna part 86 Bofuria sp. (Pteromalidae, Colotrechninae): mesosoma dorsal view 87 Yrka sp. (Pteromalidae, Colotrechninae, Amerostenini): antenna 88 Divna hirsuta Bouček (Pteromalidae, Colotrechninae, Divnini): mesosoma dorsal view 89 Bugacia sp. (Pteromalidae, Colotrechninae, Trigonoderopsini) 89 mesosoma dorsal view 90 Erixestus sp. (Pteromalidae: Erixestinae): antenna.

From the new families diagnosed above, Pteromalidae can be distinguished using the given diagnoses. Species with 2 mandibular teeth differ in this count from nearly all Pteromalidae except for in the left mandible of Austroterobia, but in Austroterobia the mandibles are additionally very different in that they are falcate, whereas the bidentate mandibles of other families, such as Ceidae, are small and narrow. A partial exception to this is Neodiparidae, where the mandibles are relatively large but still not the same shape as in Austroterobia and are not falcate.

Yusufia Koçak & Kemal and Ksenoplata Bouček are kept as unplaced to subfamily in Pteromalidae new placement, because of uncertainty over their molecular placement and difficulty in assigning them to a subfamily morphologically. They would render any other subfamily more difficult to diagnose, although they bear some similarity to Miscogastrinae and Trigonoderinae in having a clypeal margin with a single median tooth.

Mandibles not falcate. Scapula not exposed anteriorly by pronotum. Axilla strongly advanced (Fig. 86). Axillula enlarged and convex (Fig. 86). Propodeum with or without plicae. Petiole simple, usually transverse and hardly visible.

Antennal flagellum with 4 anelli and 4 funiculars (Fig. 87). Pronotum not medially divided. Ovipositor sheaths not expanded.

Molecular data indicate that the previously mentioned (Bouček 1988) unusual morphology of Yrka dahmsi Bouček new placement suggests that it belongs in the tribe Amerostenini in Colotrechninae instead of in Coelocybidae.

Morphology is consistent with this change, given that Yrka and some Amerostenini share a count of 4 antennal anelli.

Antennal flagellum usually with 3 anelli and 5 funiculars, but sometimes with 2 anelli and 6 funiculars. Pronotum not medially divided. Ovipositor sheaths not expanded.

Uzkini new syn. is treated as a synonym of Colotrechnini due to the lack of distinguishing features between them. Uzka Bouček has not been analyzed molecularly, but morphologically it is distinguished by a differently shaped head, and therefore seems to be a derived element within Colotrechnini instead of being part of a distinct lineage.

Antennal flagellum with 2 anelli and 5 funiculars. Pronotum medially divided (Fig. 88). Ovipositor sheaths expanded and densely setose.

The sole species in Divnini, Divna hirtura Bouček, has not been analyzed molecularly, and has only a dubious membership in Colotrechninae or even in Pteromalidae. Divnini is retained in Colotrechninae because we are not aware of any definitive reason to remove it from Pteromalidae, but molecular data or new morphological data may well suggest a change in its placement.

Trigonoderopsis Girault, 1915.

Antenna variously with 11 flagellomeres, either with 1 anellus and 5 funiculars, 2 anelli and 5 funiculars, 1 anellus and 6 funiculars, or with 1 anellus and 7 funiculars in some males but then with only 2 clavomeres. Pronotum not medially divided (Fig. 89). Ovipositor sheaths not expanded.

Next generation molecular data (Cruaud et al., submitted) place Trigonoderopsis Girault and Bugacia Erdős new placement in Colotrechninae, as the sister group of Colotrechnini, instead of their previous placements in Pireninae or (in the case of Bugacia) dubiously in Ormocerinae. The reduced antennal flagellum helps diagnose this emergent group, and the relatively large mandible with 4 mandibular teeth may also help diagnose the tribe, although this feature is variable in Pteromalidae.

Erixestus Crawford, 1910.

Antenna with 11 flagellomeres, with 2 anelli and 5 funiculars (Fig. 90). Mandibles falcate. Scapula not anteriorly exposed by pronotum. Axilla not strongly advanced. Axillula not enlarged (Fig. 91). Propodeum with plicae. Petiole transverse.

Figures 91–96. 

91 Erixestus winnemana Crawford (Pteromalidae, Erixestinae): mesosoma lateral view 92, 93 Diconocara petiolata Dzhanokmen (Pteromalidae, Miscogastrinae, Diconocarini) 92 head frontal view 93 mesosoma dorsal view 94, 95 Thektogaster aberlenci Delvare (Pteromalidae, Miscogastrinae, Miscogastrini) 94 clypeus 95 mesosoma dorsal view 96 Sphegigaster obliqua Graham (Pteromalidae, Miscogastrinae, Sphegigastrini): lower face frontal view.

Erixestus, containing egg parasitoids of Chrysomelidae, had persisted in uncertain placement since its description, but had apparent affinities with Ormocerini due to the high dorsal placement of the toruli. Next-generation molecular data (Cruaud et al., submitted) suggest that it is the sister group of Pteromalinae plus Pachyneurinae. The falcate mandibles of Erixestinae and Pachyneurinae indicate that this may be a synapomorphy of this entire clade, rather than a synapomorphy of Pachyneurinae alone. Otherwise, features such as the subforaminal bridge do not reject the placement of Erixestinae, but the reduced flagellomere count would be very unusual, though not unknown in the clade. It is likely that divergent features of Erixestinae are due to a life history as egg parasitoids of Chrysomelidae.

Antenna nearly always with 12 flagellomeres (exception: Andersena Bouček). Mandibles usually not falcate, but sometimes enlarged (Diconocara Dzhanokmen). Scapula not anteriorly exposed by pronotum. Axilla not strongly advanced. Axillula rarely enlarged, but if so then they are not convex. Propodeum with or without plicae. Petiole variable, either with anterolateral flange extending from ventral surface to form lateral tooth-like protrusions (Fig. 97), or simple. If petiole simple, then clypeal margin asymmetrical (Fig. 94), or concave ventrally and then clypeus enlarged and nearly adjacent to toruli dorsally (Fig. 92).

Figures 97–102. 

97 Halticoptera sp. (Sphegigastrini): petiole 98 Ormocerus latus Walker (Pteromalidae, Ormocerinae): head frontal view 99 Acroclisoides sinicus (Huang & Liao) (Pteromalidae, Pachyneurinae): head frontal and mandibles 100, 101 Pachycrepoideus sp. (Pachyneurinae) 100 venation 101 propodeum and petiole dorsal view 102 Chlorocytus scandolensis Rasplus (Pteromalidae, Pteromalinae): mesosoma dorsal view.

Distinction of Miscogastrinae from Pteromalinae is best done by tribe, using either the distinctive petiole of Sphegigastrini or the distinctive clypeus of Diconocarini or Miscogastrini. We propose keeping these tribes together as Miscogastrinae, because they form a stable clade in molecular analyses (Cruaud et al., submitted), and because Sphegigastrini also can have an asymmetrical clypeus as in Miscogastrini.

Diconocara Dzhanokmen, 1986.

Clypeal margin concave, symmetrical (Fig. 92). Mandibles enlarged. Petiole small but subquadrate, without anterolateral carina (Fig. 93), ventrally narrowly connected and without a flange.

Diconocara was described as an enigmatic member of Pteromalinae from the Russian Far East (Dzhanokmen 1986). Its position was unclear based on morphology (Heraty et al., 2013), although next-generation molecular data indicate that it is the sister group of all other Miscogastrinae (Cruaud et al., submitted). It does not have any obvious near relatives within Pteromalidae but also bears no resemblance to any other taxa outside Pteromalidae.

Clypeal margin asymmetrical, with 2 or 3 teeth (Fig. 94). Mandibles not enlarged. Petiole variable in length but usually elongate, without the anterolateral carina of Sphegigastrini but often with a small lateral process that does not extend anteriorly (Fig. 95), ventrally either with a membranous gap medially or fused but without a flange.

Miscogastrini comprise some of the most easily recognizable miscogastrines, large-bodied and usually with an expanded fore wing stigma.

Clypeal margin symmetrical or asymmetrical, with 2 or 3 denticles or uniformly convex (Fig. 96). Mandibles not enlarged. Petiole usually elongate (exception: Tricyclomischus Graham), with anterolateral flange that extends anteriorly to flank petiolar insertion (Fig. 97), ventrally closed by the continuation of this flange.

The current concept of Sphegigastrini agrees with that presented by Heydon (1988). Specimens of Bairamlia fuscipes Waterston new placement were examined morphologically in the course of this study, and were found to belong unambiguously to Sphegigastrini based on the anterolateral petiolar carina. Other genera that we place in Sphegigastrini, based on molecules and/or morphology, are listed in Appendix I.

Antenna with 12 flagellomeres. Clypeal margin symmetrical, truncate or uniformly slightly convex (Fig. 98). Mandibles not enlarged. Scapula not anteriorly exposed by pronotum. Notauli complete. Axilla not strongly advanced. Axillula not enlarged. Propodeum without plicae. Petiole transverse, without anterolateral carina.

The previous concept of Ormocerinae contained three additional tribes that are now recognized as separate groups in diverse lineages: Melanosomellidae, Systasidae, and the incertae sedis tribe Keryini (Bouček 1988; Gauthier et al. 2000). While there was some morphological resemblance between these groups, especially in the position of the toruli and in general gestalt, it became clear that they differed in many ways morphologically, as described in their given sections. Molecular data have suggested that the differences between these groups outweigh their similarities, and indeed Melanosomellidae is morphologically (see discussion of Melanosomellidae in its section above) and molecularly more similar to other members of the Gall Clade (including Cynipencyrtidae, Epichrysomallidae, Ormyridae, and Tanaostigmatidae) than to other former Ormocerinae. Additionally, Ormocerus Walker is much more similar to Pteromalinae than to the aforementioned tribes. This resemblance is strongest with relatively generalized taxa such as Nodisoplata Graham new placement that is transferred here from Miscogastrini. The features comprising this resemblance are largely those of gestalt, including enlarged but flat axillula and the nearly equal lengths of fore wing marginal and stigmal veins. The other former members of Ormocerinae, Systasidae and Keryini, are more enigmatic in placement, but differ from the new concept of Ormocerini in features covered in their sections.

Cecidoxenus Ashmead new placement is transferred here from its former position in what is now Melanosomellidae, because of its strong morphological resemblance to Ormocerus although with a slightly longer marginal vein. Additionally, Blascoa Askew new placement and Monazosa Dzhanokmen new placement are transferred here based on morphology.

Ormocerinae strongly resemble some genera of Pteromalini (Pteromalinae) that are retained with some doubt in their current classification because they have not been analyzed molecularly, such as Fijita Bouček and Huberina Bouček. They differ from Ormocerinae as characterized here by either having features that are conspicuously different from those of ormocerines, such as a median clypeal tooth or pair of teeth, a propodeum with plicae, or an elongate petiole. Other Pteromalinae with complete notauli are fig associates (Otitesellini) that have very different fore wing venation from Ormocerinae.

Antenna with 12 flagellomeres. Mandibles falcate (Fig. 99). Scapula not anteriorly exposed by pronotum. Axilla not strongly advanced. Axillula not enlarged. Fore wing marginal vein abruptly thickened at parastigmal break relative to submarginal vein thickness (Fig. 100). Propodeum with or without plicae. Petiole (when distinct) anteroventrally either braced by a flange extending from the 1^st gastral sternum that reaches anteriorly under the petiolar attachment, e.g. Pachycrepoideus Ashmead (Fig. 101), or with a more or less developed lateral tooth-like process that extends perpendicular with the longitudinal axis of the petiole, e.g. Pachyneuron Walker.

Austroterobiinae new syn. was described to contain morphologically divergent parasitoids of margarodid scales (Hemiptera). However, during this study it was observed that they have a recessed subforaminal bridge as in Pteromalinae and Pachyneurinae. Next generation molecular data (Cruaud et al., submitted) place austroterobiines as a derived group inside Pachyneurinae. Although Austroterobia Bouček and Teasienna Heydon are morphologically divergent from other Pachyneurinae in having complete notauli, this can be attributed to the switch to Margarodidae as hosts. Austroterobia and Teasienna agree with other Pachyneurinae in having falcate mandibles. Given the derived position of these genera within Pachyneurinae, we see no value in preserving the group as a tribe. Other genera transferred to Pachyneurinae, based on molecules and/or morphology, are listed in Appendix 1.

Antennal nearly always with 12 flagellomeres (exceptions: some Otitesellini, Amphidocius Dzhanokmen, possibly Termolampa). Mandibles usually not falcate (exceptions include some Apsilocera Bouček and Kaleva Graham). Scapula not anteriorly exposed by pronotum. Notauli usually incomplete, but if complete then clypeus with median tooth or teeth, propodeum with plicae, or petiole distinct and not strongly transverse, e.g., Fijita. Axilla variable but usually not strongly advanced (exceptions: Manineura Bouček). Axillula usually not enlarged (Fig. 102), but if enlarged then not convex. Fore wing with marginal vein usually slender, if distinctly thickened, e.g. Rhaphitelus Walker, then mandibles not falcate. Petiole simple or with small anterolateral processes.

Pachyneurinae differ from Pteromalinae in having a combination of falcate mandibles and an abruptly thickened marginal vein base immediately apical to the parastigmal break. Ormocerinae resemble some genera of Pteromalini that are retained with some doubt, but which differ from ormocerines in a number of respects discussed above. The few petiolate Pteromalinae differ from Miscogastrinae as follows: from Diconocarini in having small mandibles; from Miscogastrini in the symmetric clypeus; and from Sphegigastrini in the petiole structure, such as the lack of the characteristic anterolateral petiolar carina.

Antenna with 10 flagellomeres (Diaziella Grandi, Marginalia Priyadarsanan, Robertsia Bouček, Seres in part, Sycoecus), 11 flagellomeres (Apocrypta Coquerel, Arachonia Joseph, Seres in part, Otitesella, Walkerella Dalla Torre) or 12 flagellomeres (other genera). Notauli mostly complete (Fig. 103) (absent in Seres). Propodeal spiracle usually separated from the anterior propodeal margin by about their own length, or more (except Marginalia, some Walkerella and Robertsia) (Fig. 104). Males usually apterous (except in most members of previous Sycoecinae, some Watshamiella Wiebes, and Sycoryctes Mayr).

Figures 103–108. 

103 Apocrypta caudata (Girault) (Pteromalidae, Otitesellini): mesosoma dorsal view 104 Philotrypesis caricae L. (Otitesellini): propodeum 105 Seres wardi van Noort (Otitesellini): fore tibia 106 Grandiana sp. (Otitesellini): male dorsal view 107 Pseudidarnes minerva Girault (Pteromalidae, Sycophaginae): metasoma dorsal view 108 Sycophaga sp. (Pteromalidae, Sycophaginae): male lateral view.

This tribe of Pteromalinae contains the previously recognized subfamilies Otitesellinae, Sycoecinae new syn. and Sycoryctinae new syn., all fig associates occurringin the Old World (about 30 genera). It does not contain all the fig associates in Pteromalidae, since Sycophaginae is a separate subfamily of fig associates, Podivna Bouček is classified in Colotrechninae, and some New World genera (Critogaster Mayr, Aepocerus Mayr, Heterandrium Mayr, and similar genera) belong to another group of Pteromalinae. Otitesellini possess an amazing morphological disparity. Some genera that enter the fig through the ostiole have a flattened head and smooth cuticle, as well as adaptations to crawl through the bracts filling the fig aperture (mandible lengthened and covered with multiple small teeth, spurs on fore- and hind legs, enlarged protibial spur etc.) (Fig. 105). Some others oviposit within flowers from the outside once the fig is enlarged and therefore have elongated valves, or with last two tergites lengthened. Most males are greatly transformed, often apterous and sometimes with enlarged mandibles and scapes that are used for fighting for females (Fig. 106). Other genera that we place in Otitesellini based on molecules and/or morphology are listed in Appendix 1.

The previously recognized tribe Termolampini new syn. is abolished here, because it can only be distinguished from other Pteromalini using features that define the genus, and because it seems to be a derived genus within Pteromalini, instead of the sister group of another tribe or set of tribes in Pteromalinae. To resolve previous confusion, Boucekina Szelényi new placement and Morodora Gahan new placement belong here according to their morphological resemblance to other Pteromalini, respectively Neanica Erdős and Perniphora Ruschka. The complete list of genera of Pteromalini are listed in Appendix I, based on molecules and/or morphology. At present the tribe is not diagnosed, because it is the remainder of Pteromalinae excluding Otitesellini. The vast remaining number of genera in Pteromalini makes it unwieldy, and future analysis will be needed to break it up into useful natural tribes that can be more easily diagnosed.

Antennal variable in flagellomere count. Mandibles not falcate. Scapula anteriorly exposed by narrow pronotum. Axilla not strongly advanced. Axillula enlarged and convex (Fig. 107). Petiole simple, transverse. Males usually apterous (except in a few genera), residing inside figs (Fig. 108).

The taxonomic placement of the subfamily Sycophaginae has long been controversial. Sycophaginae were previously classified in Torymidae (Wiebes, 1967) and Agaonidae (Bouček 1988; Heraty et al. 2013) as the family shares at least few features with these families. Different strategies to reduce biases in our phylogenetic inference (Cruaud et al., submitted) stabilized the position of Sycophaginae as sister to all other Pteromalidae. This position is corroborated by several features shared with other members of the family and a few others shared with Colotrechninae (large and convex axillula), the next lineage in the Pteromalidae topology. Sycophaginae and Colotrechninae also share a gall-associated biology, indeed most Sycophaginae are either gall-makers or parasitoids of gall-makers within figs. We therefore propose to include Sycophaginae in Pteromalidae new placement.

Antenna with 12 flagellomeres. Mandibles often large but not falcate (Fig. 109). Scapula anteriorly exposed by narrow pronotum. Axilla hardly advanced. Axillula usually not enlarged, but if so (Plutothrix Förster), then it is not convex (Fig. 110). Petiole simple, usually transverse and hardly visible.

Figures 109–114. 

109, 110 Gastracanthus acutus (Kamijo) (Pteromalidae, Trigonoderinae) 109 lower face frontal view 110 mesosoma dorsal view 111 Cales noacki Howard (Calesidae): mesosoma lateral view 112 Eusandalum flavipenne Ruschka: mesosoma dorsal view 113 Pentacladia elegans Westwood: Mt8 and Mt9 fused, but delimited by an oblique suture below cercus 114 Chromeurytoma sp. (Megastigmidae, Chromeurytominae): head and mesosoma dorsal view.

Trigonoderinae are relatively distinctive pteromalids that differ from most others in the family in their long, large bodies in combination with complete notauli and a well-indicated frenal groove (displaced posteriorly in Platygerrhus Thomson). Many Pteromalinae are also large-bodied and can be confused superficially with Trigonoderinae, although they have incomplete notauli.

The family group name Baeomorphinae was established by Yoshimoto (1975). Rotoitidae was established by Bouček and Noyes (1987). When the two groups were synonymized by Gumovsky et al. (2018), they invoked article 35.5 of the ICZN to conserve the family name Rotoitidae. However, as Baeomorphinae cannot be recognized as a subfamily independent of a Rotoitinae, the family name must be treated with priority and Baeomorphidae recognized over Rotoitidae.

Antenna with radicle 2–4 times as long as broad; 3 or 4 flagellomeres, clava fused and longer than preceding flagellomeres; most species with multiporous plate sensilla raised and not attached along length to flagellomeres. Protibial spur short and straight. All legs with 4 tarsomeres. Axilla extending forward as distinct narrow scapula (Fig. 111). Mesosoma broadly joined to metasoma, with second phragma extending into metasoma. Hind wing with strongly curved marginal vein. Body pale yellowish or yellowish brown in color.

Calesinae has been treated as an unusual component of Aphelinidae with no direct affinities to other Aphelinidae (Mottern et al. 2011; Heraty et al. 2013). Next generation molecular data conclusively place the only genus Cales Howard as part of a sister group to most other lineages of Chalcidoidea (Cruaud et al., submitted) and not as a member of Aphelinidae. Given that Calesidae is divergent from Aphelinidae both morphologically (Fig. 111) and molecularly, we raise it to family status. Calesidae resemble many Trichogrammatidae in features of the fore wing venation and antennae, but differ from this family by having 4-segmented tarsi. Euderomphalini (Eulophidae) are similar whitefly parasitoids, but these can be distinguished by their narrow petiole with the second phragma restricted to the mesosoma.

The family Eulophidae is not diagnosed here since its limits are not being adjusted, but some taxonomic changes have been carried out as a result of our studies. The Australasian tribe Platytetracampini was described in Entedoninae by Bouček (1988), placed there because of general resemblance to some Euderomphalini. Its placement has been treated as debatable in part because of the relatively large flagellomere count (Bouček 1988; Gauthier, et al. 2000), and it was found to be the sister group of Anselmellini, in what is now Opheliminae, a phylogenetic analysis of 28S D2 ribosomal DNA by Gauthier et al. (2000) and Gumovsky (2002). We suggest that Platytetracampini should be treated as a tribe in Opheliminae new placement, because Platytetracampe Girault shares with other ophelimines the placement of fore wing admarginal setae on the ventral side of the marginal vein, and because this transfer renders Entedoninae more definable morphologically, agreeing better with Opheliminae. Although this placement of the admarginal setae is not unique in Eulophidae, we suggest that it may be locally informative for Opheliminae. More definitive placement awaits analysis of next-generation molecular data for Platytetracampe, which so far has not been available.

Boucekelimini is a morphologically distinctive group that was treated as an unplaced tribe in Eulophidae by Kim & La Salle (2005). Molecular data for this tribe have not been available, but based again on the placement of the admarginal setae on the ventral side of the marginal vein, and upon the similarity of the antenna of Boucekelimus Kim & La Salle (cf. fig. 2) and fore wing stigma shape (Kim and La Salle 2005: figs 11, 12, 19, 20) with that of Ophelimus Haliday, we treat Boucekelimini as a tribe within Opheliminae new placement, again awaiting next-generation molecular data for further evaluation of the phylogenetic value of morphological features mentioned here.

Elasmus Westwood was treated in a separate family until molecular data (Gauthier et al. 2000) suggested that it is part of Eulophinae. Elasmini was therefore reduced to tribe rank within Eulophinae. Rasplus et al. (2020) later found using next-generation UCE molecular data that Sympiesis Foerster is the sister group of Elasmus, rendering recognition of Elasmini problematic versus the now paraphyletic Eulophini, especially since Sympiesis is very similar to many other genera of Eulophini morphologically. Given that males of Elasmus have branched flagellomeres very much like those of Eulophini, we find the recognition of a separate Elasmini to be an unjustifiable misrepresentation of the evolutionary timing of the distinctive traits of Elasmus relative to the traits that are representative of Eulophini, and therefore Elasmini new syn. is a synonym of Eulophini.

Gyrolasomyiini was described as a separate tribe of Tetrastichinae by Bouček (1988). Molecular data have so far placed Gyrolasomyia Girault within Tetrastichini, with instead a Tetrastichus-group (Rasplus et al. 2020) being the sister group of remaining Tetrastichinae: the Aprostocetus-group including Gyrolasomyia. Therefore, recognition of Gyrolasomyiini would necessitate recognition of a separate tribe for the Aprostocetus-group or synonymy of Gyrolasomyiini with the oldest available tribe name of the Aprostocetus-group. However, this is untenable due to the highly problematic morphological diagnosability of tetrastichine genera and potential tribes. We suggest that both uncertain phylogenetic relationships and the practical difficulties of diagnosing subordinate taxa make recognition of tribes in Tetrastichinae currently inadvisable, and therefore Gyrolasomyiini new syn. is a junior synonym of Tetrastichini.

Eusandalum Ratzeburg, 1852.

Antennal flagellum with 9 flagellomeres, clava undivided. Eyes ventrally divergent. Clypeus with truncate apical margin. Labrum subquadrate, exposed. Subforaminal bridge with median area flanked by elongate posterior tentorial attachments; postgenal bridge externally separates the lower tentorial bridge from the hypostomal carina; postgenal groove and postgenal lamina absent. Anterolateral mesoscutal corners projecting shoulder-like on either side of the pronotum (a feature shared with Calosotinae sensu stricto). Notauli superficial and convergent, and except for Archaeopelma, ending about halfway across mesoscutum (Fig. 112). Axilla approximated (Archaeopelma Gibson and Paraeusandalum Gibson) or widely separated (Eusandalum Ratzeburg, Licrooides Gibson and Pentacladia Westwood) medially. Axillular groove or carina absent. Frenum absent. Acropleuron expanded and forming the largest surface of the mesopleuron, either comparatively small and not reaching metapleuron and metacoxa (Archaeopelma, Licrooides) or occupying most of the visible part of the mesopleuron and extending to the metapleuron (the other three genera). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; ventral membranous area anterior to mesocoxal attachment present; mesotibial spur stout except only slightly thickened in Archaeopelma. Fore leg with protibial dorsal spicules (except in Licrooides). Mesotarsus almost never with row of pegs along both sides, the exception being Licrooides: with spine-like setae on both antero- and posteroventral margins (Archaeopelma) or a row of pegs on posteroventral margin and a row of spine-like setae along anteroventral margin (remaining three genera). In Licrooides there are robust spines on both margins that almost appear as pegs. Metasoma in females with separate Mt8 and Mt9, hence without syntergum (Eusandalum) or with Mt8 and Mt9 fused but delimited by a transverse suture between cerci (Archaeopelma) or below each cercus (remaining genera; Fig. 113). Sexual dimorphism reduced, limited mainly to primary sexual features and antennal structure.

Eusandalinae new subfamily were treated until now as part of Calosotinae, however in next-generation molecular analyses (Cruaud et al., submitted) they were never recovered as monophyletic with the other Calosotinae. Instead, a reduced group of Calosotinae that includes Balcha Walker, Calosota Curtis and Tanythorax Gibson (Calosotinae sensu stricto) are sister to Heydenia (Heydeniidae) in all final analyses and closer to Eupelminae than to Eusandalinae. Eusandalinae are part of the same large clade containing also the Eupelminae and Calosotinae, but the three Eupelmidae subfamilies never form a monophyletic group since the clade also includes Ditropinotellinae, Heydeniidae and Solenurinae; Eusandalinae are the basal group. Beside the three genera included in these molecular analyses (Cruaud et al., submitted) we also include in Eusandalinae Archaeopelma and Licrooides based on a UCE analysis with a larger sampling (unpublished data). A possibly biphyletic Calosotinae sensu lato was also recognised by Gibson (1989), however with Licrooides hypothesized as closer to Calosota and allied genera and not to Eusandalum, and Archaeopelma as the most basal lineage of all Calosotinae. From the morphologically close Calosotinae sensu stricto, the Eusandalinae differ mainly in having an undivided clava, convergent notauli, scutellum without axillular groove or carina, mesotarsus almost never with two fully developed rows of pegs and a metasoma with incompletely fused or independent Mt8 and Mt9. In Calosotinae: clava with three clavomeres, notauli present as paramedially parallel lines, axillular groove present and continuing seamlessly with the scuto-scutellar suture and as a result scutellum with a carinated groove extending from base to apex, mesotarsus with a row of pegs on either side, and Mt8 and Mt9 completely fused to form the syntergum. However, all the characteristic features of Eusandalinae are either plesiomorphic or homoplastic. A thorough comparative analysis of all these characters can be found in Gibson (1989).

Khutelchalcididae was described as a new family of Chalcidoidea from a compression fossil at an estimated age of around the Jurassic/Cretaceous boundary (Rasnitsyn et al. 2004). Gibson et al. (2007) rejected the placement of Khutelchalcididae in Chalcidoidea on the grounds that the position of the mesothoracic spiracle is not in the location that is apomorphic for Chalcidoidea, but is instead in a position similar to that in Serphitidae. We follow the conclusion from Gibson et al. (2007) here, and officially exclude Khutelchalcididae new placement from Chalcidoidea, to be retained in Apocrita as an incertae sedis taxon that is not placed to superfamily.

Antenna with 10 (Chromeurytoma) or 12 (Megastigminae and Keiraninae) flagellomeres, including a 4^th clavomere. Eyes not or only slightly divergent ventrally. Clypeus bilobed or medially produced, without transverse subapical groove. Labrum flexible and mostly or entirely hidden behind clypeus, divided into several small lobes or subtriangular. Mandibles with 3 teeth. Occipital carina present, at least dorsally in Chromeurytoma (Fig. 114), usually high on the head. Pronotum usually elongate, but about as long as mesoscutum or shorter in Chromeurytominae. Notauli complete. Mesoscutellum with frenum defined by a distinct frenal groove, with or without axillular sulcus (Fig. 114). Postmarginal vein longer than stigmal vein (excepted in Patiyana Bouček) and frequently longer than marginal vein. Basal fold usually pigmented, in a few genera developed into a basal vein curved outwards (Megastigminae and Keiraninae) (Fig. 115) or rarely basal fold hyaline (Chromeurytominae). All legs with 5 tarsomeres, protibial spur stout and curved; basitarsal comb longitudinal. Gaster, while sometimes rigidly convex, not strongly sclerotized; metasomal apex usually with a separate epipygium in females (in Chromeurytominae and Megastigminae) or with a syntergum that is crossed by a sulcus immediately anterior to the cerci (Keiraninae). Cerci elongate. Ovipositor sheaths long and upcurved in females (Fig. 116).

Figures 115–120. 

115, 116 Keirana sp. (Megastigmidae, Keiraninae) 115 wing 116 metasoma lateral view 117, 118 Metapelma sp. (Metapelmatidae) 117 mesosoma lateral view 118 apex of mesotibia and mesotarsus 119, 120 Neanastatus sp. (Neanastatidae) 119 mesosoma lateral view 120 mesosoma ventral view.

Megastigminae had long been recognized as a monophyletic subfamily of Torymidae and has recently been upgraded to family rank (Janšta et al. 2018). Next-generation molecular analyses (Cruaud et al., submitted) have consistently suggested that two previous subfamilies of Pteromalidae (Chromeurytominae and Keiraninae) were closely related to Megastigmidae. Morphological examination of species of Chromeurytoma Cameron and Keirana Bouček has confirmed that these clades share several diagnostic features with Megastigmidae and belong to this family. Adding these two entities has increased the morphological disparity within Megastigmidae and has decreased the already low number of apomorphies that define the family. Consequently, Megastigmidae now comprises three subfamilies: Megastigminae including most megastigmid genera (Böhmová et al. 2022), Chromeurytominae new placement (including Chromeurytoma, Asaphoideus Girault and Patiyana) and Keiraninae new placement (with only Keirana).

Antenna with 11 flagellomeres, including 3 clavomeres. Eyes ventrally divergent. Clypeus with truncate apical margin. Labrum subquadrate, exposed. Mandibles with a ventral tooth and a dorsal weakly emarginate truncation or with 3 teeth. Subforaminal bridge with postgenal bridge separating secondary posterior tentorial pits from hypostoma; hypostomal carina convergent. Axilla transverse, approximated medially. Mesoscutellum with apex rounded; axillular groove or carina present. Frenum absent. Acropleuron enlarged, convex and pad-like, covering most of mesopleural area but separated from metacoxa by mesepimeron and metapleuron (Fig. 117). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; ventral membranous area anterior to mesocoxal attachment absent; mesotibial spur stout; mesotarsus with 1 row of pegs anteroventrally (Fig. 118). Metasoma with separate Mt8 and Mt9 in females, without syntergum.

Next-generation molecular data (Cruaud et al., submitted) consistently place Metapelma Westwood far from its former position in what is now Neanastatidae, instead most frequently as the sister group of Macromesidae + Cleonymidae. There is only minor, and not consistently diagnostic, resemblance between these three groups, and therefore Metapelmatidae revised status is restored from synonymy as a separate family. The difference between Metapelmatidae and Neanastatidae is subtle but present, in that Neanastatidae do not have the mesopleural area separated from the metacoxa by the mesepimeron and metapleuron. Eopelma also differs in several respects, including having fewer flagellomeres with an undivided clava, and does not bear any resemblance to Metapelma. Eupelmidae differ in ways explained by Gibson (1989, 1995), but notably by having a membranous area anterior to the mesocoxal attachment ventrally. Male Eupelminae additionally do not have an expanded acropleuron and all the associated modifications of the mesosoma and mid legs.

Antenna with 8, or sometimes apparently 7 flagellomeres in Neanastatus, or with 11 in Lambdobregma Gibson. Eyes ventrally divergent. Clypeus without transverse subapical groove. Labrum hidden behind clypeus, flexible. Mandibles with 3 teeth. Subforaminal bridge with postgenal bridge separating secondary posterior tentorial pits from hypostoma. Axilla transverse, approximated (Lambdobregma) or widely separated (Neanastatus) medially. Mesoscutellum with a downwards-projecting hook-like apex; axillular groove or carina present. Frenum apparently absent though the marginal rim of the mesoscutellum may be greatly expanded. Acropleuron enlarged, convex and pad-like; covering mesopleural area (Fig. 119). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; ventral membranous area anterior to mesocoxal attachment absent (Fig. 120); mesotibial spur stout; mesotarsus with 1 row of pegs anteroventrally. Metasoma with syntergum, therefore without epipygium.

Neanastatidae, with only Neanastatus and Lambdobregma remaining, is not related to the formerly included genera Metapelma (Metapelmatidae) or Eopelma in next-generation molecular analysis (Cruaud et al., submitted). Even then, Neanastatidae is rendered paraphyletic by the incertae sedis taxon Callimomoides (Louriciinae). Metapelmatidae differ from Neanastatidae in a number of features, including the separate Mt9 in females and the separation of the acropleuron from the mesocoxa by the metapleuron and small mesepimeron in Metapelmatidae. The acropleuron is also separated from the mesocoxa in Eopelma, which nevertheless bears the greatest gestalt resemblance to Neanastatus out of all these taxa due to size, coloration, and flagellomere count. Each of these genera differs from Eupelmidae in lacking a membranous area anterior to the mesocoxal attachment ventrally.

Antenna with 12 flagellomeres, including a small 4^th clavomere. Eyes not ventrally divergent. Clypeus bilobed, without transverse subapical groove. Labrum hidden behind clypeus, flexible, subrectangular with marginal setae in a row. Mandibles with 2 or 3 teeth. Subforaminal bridge with postgenal lobe separating the secondary posterior tentorial pit from the hypostoma and restricting it to the vicinity of the occipital foramen; postgenal bridge present or separated (and therefore lower tentorial bridge reaching or not reaching hypostoma); postgenal lamina usually absent; hypostomal carina usually (but not always) convergent; occipital carina present (Fig. 121). Axilla advanced (Fig. 123). Mesoscutellum with frenum indicated at least laterally, without axillular sulcus. Mesopleural area without an expanded acropleuron; mesepimeron extending over anterior margin of metapleuron (Fig. 122). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal. Fore wing stigmal vein not at a right angle with anterior fore wing margin. Metasoma with syntergum and therefore without an epipygium, convex or (more frequently) strongly sclerotized and carapace-like (Fig. 124).

Figures 121–124. 

121, 122 Ormyrus sp. (Ormyridae) 121 head dorsal view 122 mesosoma lateral view 123 Hemadas nubilipennis (Ashmead) (Ormyridae): mesosoma dorsal view 124 Asparagobius sp. (Ormyridae): metasoma lateral view.

Asparagobius Mayr has been consistently recovered as the sister group of Ormyrus Westwood /Ormyrulus Bouček with strong support in next-generation molecular analysis (Cruaud et al., submitted). In analysis of morphological characters (van Noort et al., in prep.), we acknowledged the close relationship between Asparagobius new placement and Ormyridae. We also propose the inclusion of Hemadas Crawford new placement (previously classified in Ormocerinae) in the newly defined Ormyridae.

Ormyridae are most frequently confused with Torymidae because both groups tend to have an arched body and enlarged metacoxa, although torymids have a separate epipygium and usually a long and exserted ovipositor in females whereas that of ormyrids is short. Males of the two groups are usually distinguished using habitus features of genera, and in practice can be easily recognized once the habitus of the two families is learned. Pteromalidae have an axillular sulcus in almost all species, but not in some fig associates that are otherwise highly divergent morphologically and not similar to Ormyridae. Epichrysomallidae have distinctly different fore wing venation from Ormyridae, with a longer stigmal vein that is at approximately a right angle with the anterior fore wing margin, and have a shorter marginal and postmarginal vein. Melanosomellidae lack an occipital carina, and otherwise nearly all species differ in having non-metallic coloration versus the usually metallic Ormyridae (except Hemadas).