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Pteromalidae; Temporal range: Aptian–Present PreꞒ Ꞓ O S D C P T J K Pg N
	Catolaccus grandis
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Superfamily:	Chalcidoidea
Family:	Pteromalidae; Dalman, 1820
Subfamilies
	Many (see text)

The Pteromalidae are a large family of wasps, the majority being parasitoids of other insects. They are found throughout the world in virtually all habitats, and many are important as biological control agents. The oldest known fossil is known from the Early Cretaceous.^[1]

Prior to 2022, the subfamily-level divisions of the family were highly contentious and unstable, and the family was thought to be "artificial", composed of numerous, distantly related groups (polyphyletic). In essence, a "pteromalid" was any member of the Chalcidoidea that had five-segmented tarsi and did not have the defining features of any of the remaining families with five-segmented tarsi.

In 2022, the Pteromalidae was split into 24 families.^[2]

Description

Pteromalidae are usually metallic chalcidoids of varying body size (from 1–48 mm long) and build (slender to quite robust), with the tarsi of the fore and hind legs consisting of five segments. They carry antennae consisting of eight to thirteen segments (including up to 3 anelli); in fully winged forms have in the fore wing a marginal vein that is at least several times longer than broad; very often have well-developed postmarginal and stigmal veins, although these are occasionally quite short; and nearly always have a distinct speculum.^[3]

Ecology

The life strategies of the species in this family vary greatly. There are both solitary and gregarious species, living outside (ectoparasitoid) or inside their prey (endoparasitoid), koinobionts and idiobionts, primary parasitoids and hyperparasitoids and even predators that kill and consume the prey immediately;^[3] they also include fig wasp genera. Because of their parasitoid nature, wasps of this family are often used as biological control agents for destructive pests such as American serpentine leafminers.^[4] Pteromalus cassotis is a parasitoid of the monarch butterfly.^[5]

Subfamilies, tribes, and genera

Historically, as many as 33 subfamilies of Pteromalidae were recognised.^[6] In this arrangement, and other similar ones, Pteromalidae had been recognized as paraphyletic, with many of these subfamilies and their genera being only distantly related.^[7]^[3] A large body of molecular and morphological research was considered in a new classification of these taxa, published in 2022; many taxa were removed from Pteromalidae and placed in other chalcidoid families, mostly new ones created from former subfamilies, leaving Pteromalidae with only 8 subfamilies.^[2]

The results of Burks et al.^[2] suggested the following arrangement of the remaining genera (in subfamilies and tribes):

Colotrechninae

Amerostenini: Amerostenus, Errolia, Glorimontana, Yrka;
Colotrechnini: Baridobius, Bofuria, Bomburia, Cameronella, Colotrechnus, Dipachystigma, Dvalinia, Elachertodomyia, Pachyneuronella, Podivna, Uriellopteromalus, Uzka;
Divnini: Divna;
Trigonoderopsini: Bugacia, Trigonoderopsis.

Erixestinae

Erixestus.

Miscogastrinae

Diconocarini: Diconocara;
Miscogastrini: Collentis, Drailea, Glyphognathus, Lamprotatus, Miscogaster, Neoskeloceras, Paralamprotatus, Seladerma, Sphaeripalpus, Stictomischus, Telepsogina, Thektogaster, Tumor, Xestomnaster;
Sphegigastrini: Acroclisis, Ammeia, Andersena, Ardilea, Bairamlia, Bubekia, Bubekiana, Callicarolynia, Callimerismus, Ceratetra, Cryptoprymna, Cyrtogaster, Haliplogeton, Halticoptera, Harrizia, Kazina, Maorita, Mauleus, Merismus, Notoglyptus, Notoprymna, Novitzkyanus, Paracroclisis, Ploskana, Polstonia, Rhicnocoelia, Schimitschekia, Sorosina, Sphegigaster, Syntomopus, Thinodytes, Toxeuma, Tricyclomischus, Trigonogastrella, Vespita.

Ormocerinae

Blascoa, Cecidoxenus, Monazosa, Nodisoplata, Ormocerus.

Pachyneurinae

Acroclisoides, Amblyharma, Austroterobia, Canada, Coruna, Euneura, Fusta, Goidanichium, Golovissima, Inkaka, Metastenus, Nazgulia, Neotoxeumorpha, Oomara, Oricoruna, Ottaria, Pachycrepoideus, Pachyneuron, Parabruchobius, Platecrizotes, Teasienna, Toxeumorpha.

Pteromalinae

Otitesellini: Adiyodiella, Apocrypta, Arachonia, Bouceka, Comptoniella, Crossogaster, Diaziella, Dobunabaa, Eujacobsonia, Grandiana, Grasseiana, Guadalia, Lipothymus, Marginalia, Micranisa, Micrognathophora, Otitesella, Philosycella, Philosycus, Philoverdance, Philotrypesis, Robertsia, Seres, Sycoecus, Sycoryctes, Sycoscapter, Walkerella, Watshamiella;
Pteromalini: Ablaxia, Abomalus, Acaenacis, Acroclisella, Acroclisissa, Acroclypa, Acrocormus, Aepocerus, Afropsilocera, Aggelma, Agiommatus, Aiemea, Allocricellius, Alticornis, Amandia, Amblypachus, Amphidocius, Angulifrons, Anisopteromalus, Ankaratrella, Anogmoides, Anogmus, Anorbanus, Apelioma, Apsilocera, Apycnetron, Arachnopteromalus, Arriva, Arthrolytus, Asoka, Atrichomalus, Boharticus, Bonitoa, Boucekina, Brachycaudonia, Bupronotum, Caenacis, Caenocrepis, Calliprymna, Callitula, Canberrana, Capellia, Catolaccus, Cecidolampa, Cecidostiba, Cheiropachus, Chlorocytus, Chrysoglyphe, Coelopisthia, Conigastrus, Conomorium, Cratomus, Critogaster, Cyclogastrella, Cyrtophagoides, Cyrtoptyx, Dasyneurophaga, Delisleia, Delucchia, Dibrachoides, Dibrachys, Diglochis, Dimachus, Dinarmoides, Dinarmolaelaps, Dinarmus, Dineuticida, Dinotiscus, Dinotoides, Diourbelia, Dirhicnus, Doganlaria, Dorcatomophaga, Elderia, Endomychobius, Epanogmus, Epicatolaccus, Epipteromalus, Erdoesina, Erythromalus, Eulonchetron, Eumacepolus, Eurydinota, Eurydinoteloides, Eurydinotomorpha, Euteloida, Ezgia, Fedelia, Ficicola, Fijita, Frena, Gbelcia, Genangula, Globimesosoma, Grissellium, Guancheria, Gugolzia, Guinea, Guolina, Gyrinophagus, Habritella, Habritys, Habromalina, Halomalus, Halticopterella, Halticopteroides, Helocasis, Heterandrium, Heteroprymna, Heteroschema, Hillerita, Hlavka, Hobbya, Holcaeus, Homoporus, Huberina, Hypopteromalus, Ischyroptyx, Isocyrtella, Isocyrtus, Isoplatoides, Jaliscoa, Kaleva, Klabonosa, Kratka, Kukua, Kumarella, Lampoterma, Lariophagus, Laticlypa, Lenka, Leodamus, Leptomeraporus, Licteria, Lomonosoffiella, Lonchetron, Longinucha, Lyrcus, Lysirina, Lyubana, Makaronesa, Mazinawa, Megadicylus, Merallus, Meraporus, Merismoclea, Merismomorpha, Merisus, Mesopolobus, Metacolus, Meximalus, Mimencyrtus, Mirekia, Miristhma, Mokrzeckia, Monoksa, Morodora, Muscidifurax, Nadelaia, Narendrella, Nasonia, Neanica, Nedinotus, Neocatolaccus, Neolyubana, Neopolycystus, Nephelomalus, Nikolskayana, Norbanus, Nuchata, Oaxa, Obalana, Olchon, Oniticellobia, Ottawita, Oxyharma, Oxysychus, Pandelus, Panstenon, Paracarotomus, Paradinarmus, Paraiemea, Paroxyharma, Pegopus, Peridesmia, Perilampidea, Perniphora, Pestra, Pezilepsis, Phaenocytus, Platneptis, Platypteromalus, Procallitula, Propicroscytus, Propodeia, Pseudanogmus, Pseudetroxys, Pseudocatolaccus, Psilocera, Psilonotus, Psychophagoides, Psychophagus, Pterapicus, Pterisemoppa, Pteromalus, Pterosemigastra, Pterosemopsis, Ptinocida, Pycnetron, Quercanus, Rakosina, Raspela, Rhaphitelus, Rhopalicus, Rohatina, Roptrocerus, Sceptrothelys, Schizonotus, Sedma, Sigynia, Sisyridivora, Spaniopus, Sphegigastrella, Sphegipterosema, Sphegipterosemella, Spilomalus, Spintherus, Spodophagus, Staurothyreus, Stenetra, Stenomalina, Stenoselma, Stichocrepis, Stinoplus, Strejcekia, Synedrus, Systellogaster, Szelenyinus, Tachingousa, Tanina, Tanzanicesa, Termolampa, Thureonella, Tomicobia, Toxeumella, Toxeumelloides, Trichargyrus, Trichokaleva, Trichomalopsis, Trichomalus, Tricolas, Trimeromicrus, Trinotiscus, Tritneptis, Trjapitzinia, Trychnosoma, Tsela, Uniclypea, Urolepis, Usubaia, Veltrusia, Vrestovia, Xiphydriophagus, Yanchepia, Yosemitea, Zdenekiana.

Sycophaginae

Anidarnes, Conidarnes, Eukoebelea, Idarnes, Neoeukobelea, Pseudidarnes, Sycidiphaga, Sycophaga.

Trigonoderinae

Erdoesia, Eutelisca, Gastracanthus, Janssoniella, Miscogasteriella, Neolelaps, Ogloblinisca, Platygerrhus, Plutothrix, Trigonoderus.

Incertae sedis (unplaced to subfamily)

Calolelaps, Hemitrichus, Ksenoplata, Mesolelaps, Stictolelaps, Yusufia.

References

^ Barling, Nathan; Heads, Sam W.; Martill, David M. (October 2013). "A new parasitoid wasp (Hymenoptera: Chalcidoidea) from the Lower Cretaceous Crato Formation of Brazil: The first Mesozoic Pteromalidae". Cretaceous Research. 45: 258–264. Bibcode:2013CrRes..45..258B. doi:10.1016/j.cretres.2013.05.001. ISSN 0195-6671.
^ ^a ^b ^c Roger A. Burks; Mircea-Dan Mitroiu; Lucian Fusu; et al. (20 December 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: 13–88. doi:10.3897/JHR.94.94263. ISSN 1070-9428. Wikidata Q115923766.
^ ^a ^b ^c B.R. Pitkin (2003). "Pteromalidae". Universal Chalcidoidea Database. Natural History Museum, London.
^ "American serpentine leafminer - Liriomyza trifolii (Burgess)". entnemdept.ufl.edu. Retrieved 2019-11-20.
^ Stenoien, Carl; McCoshum, Shaun; Caldwell, Wendy; Anda, Alma De; Oberhauser, Karen (1 January 2015). "New reports that Monarch butterflies (Lepidoptera: Nymphalidae, Danaus plexippus Linnaeus) are hosts for a pupal parasitoid (Hymenoptera: Chalcidoidae, Pteromalus cassotis Walker)". Journal of the Kansas Entomological Society. 88 (1): 16–26. doi:10.2317/JKES1402.22.1. S2CID 52231552.
^ Noyes, J.S. (March 2019). "Universal Chalcidoidea Database". The Natural History Museum. Retrieved 5 April 2023.
^ Gibson, G.A.P. (1993) Superfamilies Mymarommatoidea and Chalcidoidea (pp. 570-655). In Goulet, H. & Huber, J. (eds). Hymenoptera of the World: an identification guide to families. Research Branch, Agriculture Canada, Ottawa, Canada, 668 pp. PDF