From hell's heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera) Author Burks, Roger https://orcid.org/0000-0003-3032-7939 Department of Entomology, University of California Riverside, Riverside, CA, USA burks.roger@gmail.com Author Mitroiu, Mircea-Dan https://orcid.org/0000-0003-1368-7721 Faculty of Biology, Alexandru Ioan Cuza University, Iasi, Romania Author Fusu, Lucian https://orcid.org/0000-0003-0819-026X Faculty of Biology, Alexandru Ioan Cuza University, Iasi, Romania Author Heraty, John M. https://orcid.org/0000-0002-9246-5651 Department of Entomology, University of California Riverside, Riverside, CA, USA Author Jansta, Petr https://orcid.org/0000-0001-6409-3603 Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic & Department of Entomology, State Museum of Natural History, Stuttgart, Germany Author Heydon, Steve Bohart Museum of Entomology, University of California, Davis, CA, 95616, USA Author Papilloud, Natalie Dale-Skey https://orcid.org/0000-0001-7582-0386 Insects Division, Natural History Museum, London, UK Author Peters, Ralph S. Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for the Analysis of Biodiversity Change, Bonn, Germany Author Tselikh, Ekaterina V. https://orcid.org/0000-0002-9184-043X Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia Author Woolley, James B. Department of Entomology, Texas A & M University, College Station, TX, USA Author van Noort, Simon https://orcid.org/0000-0001-6930-9741 Research and Exhibitions Department, South African Museum, Iziko Museums of South Africa, PO Box 61, Cape Town 8000 South Africa & Department of Biological Sciences, University of Cape Town, Private Bag, Rondebosch, 7701, South Africa Author Baur, Hannes https://orcid.org/0000-0003-1360-3487 Department of Invertebrates, Natural History Museum Bern, Bern, Switzerland & Institute of Ecology and Evolution, University of Bern, Bern, Switzerland Author Cruaud, Astrid https://orcid.org/0000-0001-8932-4199 CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, University of Montpellier, Montpellier, France Author Darling, Christopher Department of Natural History, Royal Ontario Museum, Toronto, ON, M 5 S 2 C 6, Canada & Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON, M 5 S 1 A 1, Canada Author Haas, Michael https://orcid.org/0000-0001-6869-6698 Department of Entomology, State Museum of Natural History, Stuttgart, Germany Author Hanson, Paul Escuela de Biologia, Universidad de Costa Rica, San Pedro de Montes de Oca, San Jose 11501 - 2060, Costa Rica Author Krogmann, Lars https://orcid.org/0000-0002-3724-1735 Department of Entomology, State Museum of Natural History, Stuttgart, Germany & Institute of Biology, Biological Systematics (190 n) University of Hohenheim, Stuttgart, Germany Author Rasplus, Jean-Yves https://orcid.org/0000-0001-8614-6665 CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, University of Montpellier, Montpellier, France text Journal of Hymenoptera Research 2022 2022-12-20 94 13 88 http://dx.doi.org/10.3897/jhr.94.94263 journal article http://dx.doi.org/10.3897/jhr.94.94263 1314-2607-94-13 6CB807239A47403FABEC9AF8AE7F417F ADCFB8021287566FB2D7E8A8711D5CAE 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 Boucek 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 Boucek 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 Boucek ( 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 Kocak & Kemal and Ksenoplata Boucek 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