Pteromalidae, Dalman, 1820

Burks, Roger, Mitroiu, Mircea-Dan, Fusu, Lucian, Heraty, John M., Jansta, Petr, Heydon, Steve, Papilloud, Natalie Dale-Skey, Peters, Ralph S., Tselikh, Ekaterina V., Woolley, James B., van Noort, Simon, Baur, Hannes, Cruaud, Astrid, Darling, Christopher, Haas, Michael, Hanson, Paul, Krogmann, Lars & Rasplus, Jean-Yves, 2022, From hell's heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera), Journal of Hymenoptera Research 94, pp. 13-88 : 13

Pteromalidae

Pteromalini Dalman, 1820. Type genus: Pteromalus Swederus, 1795. Treated as Pteromalidae by Walker (1834).

Diagnosis.

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 4th 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).

Discussion.

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.

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.

Subfamilies and tribes of Pteromalidae

Figures

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.

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 Boucek (Pteromalidae, Colotrechninae, Divnini): mesosoma dorsal view 89 Bugacia sp. (Pteromalidae, Colotrechninae, Trigonoderopsini) 89 mesosoma dorsal view 90 Erixestus sp. (Pteromalidae: Erixestinae): antenna.