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Trichopria anastrephae Lima, 1940 (Figs 1, 2, 16) Diagnosis. Body dark-brown to black, surface mostly smooth and polished except petiole. Fore wing fully developed, without closed cells; with complete, short, subcostal vein (=submarginal vein), ending in a short marginal vein (genus Trichopria). Female antennae 12-segmented, with 3-segmented clava; male antennae 14-segmented; flagellomeres long and pedunculate, swollen apically with long setae. Scutellum smooth, median carina absent; scutellar sulcus smooth and shallow. Body length 1.8���2.0 mm. Considering the Neotropical fauna, Tr. anastrephae is similar to Trichopria peraffinis (Ashmead), which is a much smaller species (~1.0 mm long), and presents a small pit on scutellum anteriorly (absent in Tr. anastrephae). Taxonomy. Trichopria is one of the largest genera in Diapriidae, and no revision including the Neotropical fauna has been published, making identification to species level difficult. For instance, there are at least two unidentified species of Trichopria that parasitize species of Anastrepha in the New World (USA and Costa Rica) (Ovruski et al. 2000). Of the 12 species of Trichopria in Brazil (Margar��a 2020), nine can be keyed out using Kieffer���s (1910; 1916) identification keys in combination with original descriptions (Fouts 1926) or recent diagnosis (Notton 2014). Trichopria catarinensis Ferri��re is discarded because of its specialized biology, as parasitoid on Ecitonini (Hymenoptera, Formicidae). The remaining species, Trichopria lamellifera Ogloblin could also be discarded based on host association with Micropezidae (Diptera, Nerioidea), and morphological differences such as the length of antennomeres and its setae in males being much longer in T. anastrephae (antennomere 4 ~ 200 ��m and longest setae ~ 300 ��m) than in T. lamellifera (91 ��m and 98 ��m respectively), and females with much larger compound eyes (eye diameter ~5x longer than malar space in T. anastrephae compared to ~2.5x in T. lamellifera) (Ogloblin 1934). Trichopria is a diverse genus, with possibly a high number of undescribed species in the neotropics (Masner & Garc��a 2002). Therefore, caution is advised when using the identification key presented below. A revision of the genus is badly needed, for the Neotropical region. Biology. Trichopria anastrephae is an endoparasitoid koinobiont on pupae of Tephritidae and less frequently on Drosophilidae, for example Drosophila suzukii (Matsumura) (Yoder 2007). Known Tephritidae hosts are A. serpentina and A. fraterculus in Brazil (Lima 1940 and Aguiar-Menezes et al. 2001, respectively) and Ce. capitata in Argentina (Turica & Mallo 1961). Biological control. The potential of Tr. anastrephae as a biological agent has not been investigated in detail, although it is likely to be an important natural enemy of tephritids in Brazil, being the most common parasitoid species in star fruits (Silva et al. 2003). Distribution. Brazil and Argentina. Distribution in Brazil (associated with tephritid species). BA (Souza-Filho et al. 2007), CE (Silva et al. 2020), GO (Marchiori & Penteado-Dias 2001), MG (Silva et al. 2003), RJ (Lima 1940), SC (Garcia & Corseuil 2004), RS (Cruz et al. 2011)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on pages 58-60, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Lima, A. M. C. (1940) Alguns parasitos de moscas das frutas. Anais da Academia Brasileira de Ciencias, 12, 17 - 20.","Ovruski, S., Aluja, M., Sivinski, J. & Wharton, R. A. (2000) Hymenopteran parasitoids on fruit-infesting Tephritidae (Diptera) in Latin America and the southern United States: Diversity, distribution, taxonomic status and their use in fruit fly biological control. Integrated Pest Management Reviews, 5, 81 - 107. https: // doi. org / 10.1023 / A: 1009652431251","Margaria, C. (2020) Diapriidae in Catalogo Taxonomico da Fauna do Brasil. PNUC. Available in http: // fauna. jbrj. gov. br / fauna / faunadobrasil / 14922 (access 11 August 2020)","Kieffer, J. J. (1910) Description de nouveaux microhymenopteres du Bresil. Annales de la Societe Entomologique de France, 78, 287 - 348.","Kieffer, J. J. (1916) Diapriidae. Das Tierreich, 44, 1 - 627. http: // biostor. org / reference / 80288","Fouts, R. M. (1926) Notes on Serphoidea with descriptions of new species (Hymenoptera). Proceedings of the Entomological Society of Washington, 28 (8), 167 - 179.","Notton, D. G. (2014) A catalogue of the types of Diapriinae (Hymenoptera, Diapriidae) at the Natural History Museum, London. European Journal of Taxonomy, 75, 1 - 123. https: // doi. org / 10.5852 / ejt. 2014.75","Ogloblin, A. (1934) Una especie nueva del genero Trichopria del Brasil (Hym. Diapriidae). Revista de Entomologia, 4 (1), 60 - 65.","Aguiar-Menezes, E. L., Menezes, E. B., Silva, P. S., Bittar, A. C. R. & Cassino, P. C. R. (2001) Native hymenopterous parasitoids associated with Anastrepha spp. (Diptera: Tephritidae) in Seropedica city, Rio de Janeiro, Brazil. Florida Entomologist, 84, 706 - 711. https: // doi. org / 10.2307 / 3496405","Turica, A. & Mallo, R. G. (1961) Observaciones sobre la poblacioin de las ' Tephritidae' y sus endoparasitos en algunas regiones citricolas argentinas. IDIA, 6, 145 - 161.","Silva, C. G., Marchiori, C. H., Fonseca, A. R. & Torres, L. C. (2003) Himenopteros parasitoides de larvas de Anastrepha spp. em frutos de carambola (Averrhoa carambola L.) na regiao de Divinopolis, Minas Gerais, Brasil. Ciencia e Agrotecnologia, 27 (6), 1264 - 1267. https: // doi. org / 10.1590 / S 1413 - 70542003000600009","Souza-Filho, Z. A., Araujo, E. L., Guimaraes, J. A. & Silva, J. G. (2007) Endemic parasitoids associated with Anastrepha spp. (Diptera: Tephritidae) infesting guava (Psidium guajava L.) in southern Bahia, Brazil. Florida Entomologist, 90 (4), 83 - 85. https: // doi. org / 10.1653 / 0015 - 4040 (2007) 90 [783: EPAWAS] 2.0. CO; 2","Silva, B. K. A., Silva, H. M., Fernandes, C. E., Costa, V. A. & Araujo, E. L. (2020) Pupal parasitoids associated with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in a semiarid environment in Brazil. Revista Brasileira de Entomologia, 64 (2), e 20190002. https: // doi. org / 10.1590 / 1806 - 9665 - rbent- 2019 - 0002","Marchiori, C. H. & Penteado-Dias, A. M. (2001) Trichopria anastrephae (Hymenoptera: Diapriidae), parasitoide de Diptera, coletadas em area de mata nativa e pastagem em Itumbiara, Goias, Brasil. Arquivos do Instituto Biolgico, 68 (1), 123 - 124.","Garcia, F. R. M. & Corseuil, E. (2004) Native hymenopteran parasitoids associated with fruit flies (Diptera: Tephritidae) in Santa Catarina State, Brazil. Florida Entomologist, 87, 517 - 521. https: // doi. org / 10.1653 / 0015 - 4040 (2004) 087 [0517: NHPAWF] 2.0. CO; 2","Cruz, P. P., Neutzling, A. S. & Garcia, F. R. M. (2011) Primeiro registro de Trichopria anastrephae, parasitoide de moscas-dasfrutas, no Rio Grande do Sul. Ciencia Rural, 41 (8), 1297 - 1299. https: // doi. org / 10.1590 / S 0103 - 84782011000800001"]}

Pachycrepoideus vindemmiae (Rondani, 1875) (Figs 6���8) Diagnosis. Small wasps, black with slight metallic-green iridescence; head and mesosoma densely punctate to mostly reticulate, including mesopleuron; antennae with basal 2 or 3 segments of flagellum annular, subsequent funicular segments nearly as wide as long; metasoma appearing sessile, smooth, with petiole bordered on either sides by flange extending anteriorly from first sternite; marginal vein uniformly thickened, proportion of marginal: stigmal:posmarginal veins = 1:1:2; legs yellowish beyond coxae. Taxonomy. This species is treated in modern taxonomic revisions that include well-illustrated keys, such as those of Rueda & Axtell (1985) and Bouček & Heydon (1997). Pachycrepoideus vindemmiae is the only species of the genus known to occur in Brazil (Costa 2019). Biology. Pupal idiobiont ectoparasitoid, recorded in a wide range of Diptera species (at least 60 species in several families). It may also act as a facultative hyperparasitoid (Wang & Messing 2004). In Brazil, it is frequently associated with fruit flies (Tephritidae), namely A. fraterculus, A. obliqua and A. sororcula (Aguiar-Menezes & Menezes 2002; Aguiar-Menezes et al. 2003; Marchiori et al. 2000), although parasitism on species of other families is also common. Biological control. This parasitoid has been used as a biological control agent for filth flies (Diptera: Muscidae) (e.g. Meyer et al. 1990; Wharton 1989), and introduced in several countries in the Americas for classical and augmentative biological control of fruit-infesting tephritids (e.g. Gilstrap & Hart 1987; Purcell 1998), but usually established populations prior to intentional introductions. The efficacy of this species in augmentative programs is criticized, since it has been used for decades without evidence of success (Ovruski et al. 2000). Even though natural parasitism may aid in regulating population levels of some pests observed rates are usually low, and efforts on introducing or mass-rearing P. vindemmiae are discouraged, due to its polyphagy, resulting in negative effects on non-target species, and lack of efficacy (Ovruski et al. 2000; Wang & Messing 2004; Lee et al. 2019). Distribution. Cosmopolitan. The introduction of P. vindemmiae in Brazil, as in many other countries, is not well documented. For instance, Ovruski & Schliserman (2012) comment that occurrence in 11 American countries could be a result of intentional or unintentional introduction, or even of the natural distribution of P. vindemmiae. The first well documented introduction is reported for Hawaii, in 1914, for specimens brought from the Philippines by Mr. D. T. Fullaway (Pemberton & Willard 1918). The same researcher, working as an employee of the Bureau of Entomology and Plant Quarantine of Puerto Rico, was responsible for sending parasitoid specimens from Brazil to Puerto Rico, including P. vindemmiae, between years 1935 and 1937 (Bartlett 1941). Additionally, there are some reports of specimens of P. vindemmiae being redistributed from Hawaii to Latin American countries (e.g. Purcell 1998, for Costa Rica). Therefore, this species could have been introduced in Brazil between years 1914 and 1937, from Hawaii, by Mr. D. T. Fullaway, a possibility that we could not confirm by the lack of historical records. The first citation of P. vindemmiae in Brazil is given by Souza (1943), as a parasitoid in pupae of Billaea claripalpis (Wulp) (formerly Paratheresia brasiliensis Townsend) (Diptera, Tachinidae), which in turn was parasitizing Diatraea saccharalis (Fabricius) (Lepidoptera, Crambidae) in sugarcane. The specimens were obtained from samples taken in 1940-1941 (Souza 1943), but P. vindemmiae has possibly established populations in Brazil before 1937 (see Bartlett 1941). Distribution in Brazil (associated with tephritid species). CE and RN (Silva et al. 2020), GO (Marchiori et al. 2000), RJ (Aguiar-Menezes & Menezes 2002; Aguiar-Menezes et al. 2003), RS (Salles 1996), SP (Montes et al. 2011)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 62, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Rueda, L. M. & Axtell, R. C. (1985) Guide to common species of pupal parasites (Hymenoptera: Pteromalidae) of the house fly and other muscoid flies associated with poultry and livestock manure. North Carolina Agricultural Research Service, Technical Bulletin, 278, 1 - 88.","Boucek, Z. & Heydon, S. (1997) Pteromalidae. In: Gibson, G. A. P., Huber, J. T. & Wooley, J. B. (Eds.), Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, pp. 541 - 692.","Costa, V. A. (2019) Pteromalidae in Catalogo Taxonomico da Fauna Brasileira. PNUD. Available from: http: // fauna. jbrj. gov. br / fauna / faunadobrasil / 1054 (accessed 21 July 2019)","Wang, X. G. & Messing, R. H. (2004) The ectoparasitic pupal parasitoid, Pachycrepoideus vindemmiae (Hymenoptera: Pteromalidae), attacks other primary tephritid fruit fly parasitoids: host expansion and potential non-target impact. Biological Control, 31, 227 - 236. https: // doi. org / 10.1016 / j. biocontrol. 2004.04.019","Aguiar-Menezes, E. L. & Menezes, E. B. (2002) Effect of time of permanence of host fruits in the field on natural parasitism of Anastrepha spp. (Diptera: Tephritidae). Neotropical Entomology, 31, 589 - 595. https: // doi. org / 10.1590 / S 1519 - 566 X 2002000400011","Aguiar-Menezes, E. L., Menezes, E. B. & Loiacono, M. S. (2003) First record of Coptera haywardi Loiacono (Hymenoptera: Diapriidae) as a parasitoid of fruit-infesting Tephritidae (Diptera) in Brazil. Neotropical Entomology, 32, 355 - 358. https: // doi. org / 10.1590 / S 1519 - 566 X 2003000200025","Marchiori, C. H., Oliveira, A. M. S., Martins, F. F., Bossi, F. S. & Oliveira, A. T. (2000) Ocorrencia de moscas frugivoras (Diptera: Tephritidae e Lonchaeidae) e seus parasitoides em Itumbiara-GO. Arquivos do Instituto Biologico, 67 (1), 139 - 141.","Meyer, J. A., Mullens, B. A. & Cyr, T. L. (1990) Commercial and naturally occurring fly parasitoids (Hymenoptera: Pteromalidae) as biological control agents of stable flies and house flies (Diptera: Muscidae). Journal of Economic Entomology, 83 (3), 799 - 806. https: // doi. org / 10.1093 / jee / 83.3.799","Wharton, R. A. (1989) Classical biological control of fruit Tephritidae. In: Robinson, A. & Harper, G. (Eds.), World Crop Pests, Fruit flies: Their Biology, Natural Enemies, and Control. Vol. 3 b. Elsevier Science, Amsterdam, pp. 303 - 313.","Gilstrap, F. E. & Hart, W. G. (1987) Biological control of the Mediterranean fruit fly in the United States and Central America. USDA, Agricultural Research Service, ARS, 56, 1 - 64. https: // doi. org / 10.5962 / bhl. title. 133985","Purcell, M. F. (1998) Contribution of biological control to integrated pest management of tephritid fruit flies in the tropics and subtropics. Integrated Pest Management Reviews, 3, 63 - 83. https: // doi. org / 10.1023 / A: 1009647429498","Ovruski, S., Aluja, M., Sivinski, J. & Wharton, R. A. (2000) Hymenopteran parasitoids on fruit-infesting Tephritidae (Diptera) in Latin America and the southern United States: Diversity, distribution, taxonomic status and their use in fruit fly biological control. Integrated Pest Management Reviews, 5, 81 - 107. https: // doi. org / 10.1023 / A: 1009652431251","Lee, J. C., Wang, X., Daane, K. M., Hoelmer, K. A., Isaacs, R., Sial, A. A. & Walton, V. M. (2019) Biological control of spottedwing drosophila (Diptera: Drosophilidae) - Current and pending tactics. Journal of Integrated Pest Management, 10 (1), 13. https: // doi. org / 10.1093 / jipm / pmz 012","Ovruski, S. M. & Schliserman, P. (2012) Biological Control of Tephritid Fruit Flies in Argentina: Historical Review, Current Status, and Future Trends for Developing a Parasitoid Mass-Release Program. Insects, 3, 870 - 888. https: // doi. org / 10.3390 / insects 3030870","Pemberton, C. E. & Willard, H. F. (1918) A contribution to the biology of fruit-fly parasites in Hawaii. Journal of Agricultural Research, 15 (8), 419 - 467.","Bartlett, K. A. (1941) The introduction and colonization in Puerto Rico of Beneficial insects parasitic on West Indian fruitflies. The Journal of Agriculture of the University of Puerto Rico, 25 (1), 25 - 31. https: // revistas. upr. edu / index. php / jaupr / article / view / 3517","Souza, H. D. (1943) A broca da cana de acucar e seus parasitos em Campo, Estado do Rio de Janeiro. Boletim do Instituto de Ecologia e Experimentacao Agricolas, 4 (1942), 1 - 22.","Silva, B. K. A., Silva, H. M., Fernandes, C. E., Costa, V. A. & Araujo, E. L. (2020) Pupal parasitoids associated with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in a semiarid environment in Brazil. Revista Brasileira de Entomologia, 64 (2), e 20190002. https: // doi. org / 10.1590 / 1806 - 9665 - rbent- 2019 - 0002","Salles, L. A. B. (1996) Parasitismo de Anastrepha fraterculus (Wied.) (Diptera: Tephritidae) por Hymenoptera, na regiao de Pelotas, RS. Pesquisa Agropecuaria Brasileira, 31 (11), 769 - 774","Montes, S. M. N. M., Raga, A., Boliani, A. C. & Santos, P. C. (2011) Dinamica populacional e incidencia de moscas-das-frutas e parasitoides em cultivares de pessegueiros (Prunus persica L. Batsch) no municipio de Presidente Prudente-SP. Revista Brasileira de Fruticultura, 33 (2), 402 - 411. https: // doi. org / 10.1590 / S 0100 - 29452011005000052"]}

Family Pteromalidae (Hymenoptera, Chalcidoidea) There are 3,506 species in 588 genera of Pteromalidae worldwide (Noyes 2019). The number of subfamilies varies depending on the authority, but the most widely accepted classification comprises 31 (Noyes 2002; 2019). This large number reflects its polyphyletic nature on the one hand, and the high proportion of monotypic subfamilies on the other (Peters et al. 2018). In the Neotropics, there are 476 species in 155 genera (Noyes 2019), of which 156 species in 65 genera are found in Brazil (Costa 2019). Knowledge of the Neotropical fauna is only incipient, with most species yet to be described (Hanson & Heydon 2006). No combination of characters exists to distinguish this family. The most recent phylogenies are consistent in recovering the family as polyphyletic, which to a great extent is responsible for the lack of morphological cohesion (Munro et al. 2011; Heraty et al. 2013) Since Pteromalidae is not a natural group, generalizations on its biology are difficult, and meaningless in an evolutionary perspective. Nevertheless, it is possible to say that pteromalids are in most cases ectoparasitoid idiobionts on pupae of Coleoptera and Diptera, developing inside the puparium case (Hanson & Heydon 2006). Most of the knowledge on biology was acquired through examination of two unrelated subfamilies, Pteromalinae and Spalangiinae (Peters et al. 2018). Even though the two groups have relatively distant evolutionary histories and very distinct morphologies, they possess several biological traits in common, and all the representatives of parasitoids on fruit-infesting tephritids in Brazil belong to these subfamilies. Pteromalids are usually synovigenic, requiring the female to feed, usually on the host, to acquire the necessary nutrients for oogenesis (Simmonds 1953; Doutt 1964; Legner & Gerling 1967; Keller & Sullivan 1976), but also as water-intake strategy (Bezerra da Silva et al. 2019a). Host-feeding by Pachycrepoideus vindemmiae (Pteromalinae), for instance, usually occurs in young pupae (Phillips 1993), and may increase host mortality rate (Bezerra da Silva et al. 2019b). This is the only Pteromalinae known to attack fruit flies in Brazil. For the Spalangiinae, four species have been recorded in Brazil attacking tephritids, all in the genus Spalangia. It is likely that many species in that genus occasionally parasitize fruit flies due to their polyphagous habit, as demonstrated by Silva et al. (2020), who obtained five species of Spalangia using sentinel puparia of Ce. capitata. For two of the five species (i.e. S. leiopleura and S. impunctata), there are no previous association records with fruit flies. The two species were not included in our results by the absence of association with host plants. For any species of Spalangia reared from tephritids, checking the identification key provided by Gibson (2009) is strongly recommended., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 61, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Noyes, J. S. (2019) Universal Chalcidoidea Database. World Wide Web electronic publication. Available from: http: // www. nhm. ac. uk / chalcidoids (accessed 16 July 2019)","Noyes, J. S. (2002) Interactive catalogue of world Chalcidoidea 2001. 2 nd Edition. Taxapad and the Natural History Museum, London, CD-ROM.","Peters, R. S., Niehuis, O., Gunkel, S., Blasere, M., Mayer, C., Podsiadlowski, L., Kozlov, A., Donath, A., Noort, S., Liuj, X., Zhou, X., Misof, B., Heraty, J. & Krogmann, L. (2018) Transcriptome sequence-based phylogeny of chalcidoid wasps (Hymenoptera: Chalcidoidea) reveals a history of rapid radiations, convergence, and evolutionary success. Molecular Phylogenetics and Evolution, 120 (2018), 286 - 296. https: // doi. org / 10.1016 / j. ympev. 2017.12.005","Costa, V. A. (2019) Pteromalidae in Catalogo Taxonomico da Fauna Brasileira. PNUD. Available from: http: // fauna. jbrj. gov. br / fauna / faunadobrasil / 1054 (accessed 21 July 2019)","Hanson, P. E. & Heydon, S. (2006) Pteromalidae. In: Hanson, P. E. & Gauld, I. D. (Eds.), Hymenoptera de la Region Neotropical. Memoirs of the American Entomological Institute, 77, pp. 403 - 420.","Munro, J. B., Heraty, J. M., Burks, R. A., Hawks, D. & Mottern, J. (2011) A Molecular Phylogeny of the Chalcidoidea (Hymenoptera). PLoS ONE, 6 (11), e 27023. https: // doi. org / 10.1371 / journal. pone. 0027023","Heraty, J. M., Burks, R. A., Cruaud, A., Gibson, G. A., Liljeblad, J., Munro, J., Rasplus, J. Y., Delvare, G., Jansta, P., Gumovsky, A. & Huber, J. (2013) A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics, 29, 466 - 542. https: // doi. org / 10.1111 / cla. 12006","Simmonds, F. J. (1953) Observations on the biology and mass-breeding of Spalangia drosophilae Ashm. (Hymenoptera: Spalangiidae), a parasite of the frit-fly, Oscinella frit (L.). Bulletin of Entomological Research, 44, 773 - 778. https: // doi. org / 10.1017 / S 0007485300024718","Doutt, R. L. (1964) Biological characteristics of entomophagous adults. In: DeBach, P. & Schlinger, E. I. (Eds.), Biological control of insect pests and weeds. Chapman & Hall, London, pp. 145 - 167.","Legner, E. F. & Gerling, D. (1967) Host-Feeding and oviposition on Musca domestica by Spalangia cameroni, Nasonia vitripennis, and Muscidifurax raptor (Hymenoptera: Pteromalidae) influences their longevity and fecundity. Annals of the Entomolological Society of America, 60 (3), 678 - 691. https: // doi. org / 10.1093 / aesa / 60.3.678","Keller, L. J. & Sullivan, D. J. (1976) Oviposition Behavior and Host Feeding of Asaphes lucens an Aphid Hyperparasitoid. Journal of the New York Entomological Society, 84, 206 - 211.","Bezerra da Silva, C. S., Price, B. E. & Walton, V. M. (2019 a) Water-Deprived Parasitic Wasps (Pachycrepoideus vindemmiae) Kill More Pupae of a Pest (Drosophila suzukii) as a Water-Intake Strategy. Scientific Reports, 9, 3592. https: // doi. org / 10.1038 / s 41598 - 019 - 40256 - 8","Phillips, D. S. (1993) Host-feeding and egg maturation by Pachycrepoideus vindemiae [sic]. Entomologia Experimentalis et Applicata, 69 (1), 75 - 82. https: // doi. org / 10.1111 / j. 1570 - 7458.1993. tb 01730. x","Bezerra da Silva, C. S., Price, B. E. & Walton, V. M. (2019 b) Factors affecting the biology of Pachycrepoideus vindemmiae (Hymenoptera: Pteromalidae), a parasitoid of spotted-wing drosophila (Drosophila suzukii). PLoS ONE, 14 (7), e 0218301. https: // doi. org / 10.1371 / journal. pone. 0218301","Silva, B. K. A., Silva, H. M., Fernandes, C. E., Costa, V. A. & Araujo, E. L. (2020) Pupal parasitoids associated with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in a semiarid environment in Brazil. Revista Brasileira de Entomologia, 64 (2), e 20190002. https: // doi. org / 10.1590 / 1806 - 9665 - rbent- 2019 - 0002","Gibson, G. A. P. (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae). Zootaxa, 2259 (1), 1 - 159. https: // doi. org / 10.11646 / zootaxa. 2259.1.1"]}

Key to species of Diapriidae, Eulophidae and Pteromalidae (Hymenoptera) parasitoids of fruit flies (Diptera, Tephritidae) in Brazil 1 Fore wing with submarginal, marginal and stigmal veins present (Figs 5, 8, 15); antennae inserted close to or below lower margin of eyes, not on a projecting ledge (Figs 3, 6) (Chalcidoidea)............................................. 2 - Fore wing at most with a short submarginal vein (Figs 16, 17); antennae inserted on a projecting ledge near upper margin of eyes (Figs 1, 2) (Diaprioidea)............................................................................ 6 2 Tarsi 4-segmented; scutellum with a pair of submedian grooves and a pair of sublateral grooves (Fig. 4) (Eulophidae)...................................................................................... Tetrastichus giffardianus - Tarsi 5-segmented; scutellum without longitudinal grooves (Fig. 7) (Pteromalidae)................................. 3 3 Antennae separated from clypeus, inserted nearly in the middle of face, slightly above lower margin of eyes (Fig. 6); marginal vein distinctly and uniformly thickened (Fig. 8)...................................... Pachycrepoideus vindemmiae - Antennae inserted just above oral margin, far below lower margin of eyes (Fig. 3); marginal vein not thickened (Fig. 15) (Spalangia).......................................................................................... 4 4 Pronotum smooth, with setae on tiny bumps (Fig. 14); median lobe of mesoscutum with a single median pit (Fig. 14)............................................................................................ Spalangia simplex - Pronotum punctate or reticulate-rugose (Figs 12, 13); medial lobe of mesoscutum with different sculpturing............. 5 5 Pronotum with circular punctures distinctly separated by smooth interstices (Fig. 12); malar sulcus present; propodeum with paramedian crenulate furrows sub parallel, all cells with similar size................................ Spalangia endius - Pronotum mostly reticulate-rugose (Fig. 13); malar sulcus absent; propodeum with paramedian crenulate furrows widened anteriorly, forming a Y, with anterior cells larger than posterior cells................................ Spalangia gemina 6 Fore wing lacking tubular veins, with a mid-longitudinal fold line, apex with apical notch (Fig. 17)....... Coptera haywardi - Fore wing with a tubular vein (submarginal), without fold-line, apex rounded (Fig. 15)...... Trichopria sp. aff. anastrephae

Tetrastichus giffardianus Silvestri, 1915 (Figs 4, 5) Diagnosis. Color varying from dark-brown to black, with metallic bluish-green iridescence, more intense on the mesosoma; legs brown with pale-yellow tarsi. The species belongs to the murcia group (Graham 1991), which includes five Old World species, distinguished mainly by the presence of long erect setae on the eyes, vertex and thorax. Within the murcia group, Te. giffardianus is distinct from other species in having a long speculum (area without setae) along the anterior margin of the fore wing, from its base to the stigmal vein or beyond (Graham 1991; LaSalle & Wharton 2002). Taxonomy. The species concept and its identification are relatively straightforward, thanks to the taxonomic revision of African species of Tetrastichus (LaSalle & Wharton 2002), which includes a key to species. Nevertheless, this revision is limited to three species from Africa, and the more comprehensive revision of Graham (1991) is helpful in the identification and confirmation of the taxonomic concept at the species-group level. Biology. This is a gregarious, larval-pupal, koinobiont endoparasitoid (LaSalle & Wharton 2002). Females search for hosts by penetrating fruits to oviposit in late-instar larvae of tephritid species (Pemberton & Willard 1918; Fernandes et al. 2019). Adults will emerge from the puparium of their host, although oviposition occurs in the larva, a unique parasitism strategy among the species treated here, and closer to the biology of most the common species of Braconidae and Figitidae that parasitize fruit flies. All of the several species recorded as hosts of Te. giffardianus are tephritids, and most of them in the genus Ceratitis in its native distribution area. In Brazil, Te. giffardianus was reared from A. obliqua (Ara��jo et al. 2016) and A. distincta (Abreu et al. 2019), however, a clear preference for Ce. capitata has been observed (Costa et al. 2005; Araujo et al. 2015) (Table 1). Biological control. This species possesses the same biological traits, except for being gregarious rather than solitary, found in the main natural enemies of fruit flies (Braconidae, Opiinae), making them excellent candidates for biological control programs, especially for Ce. capitata. After confirmation that Te. giffardianus had established in Brazil, most successfully in the Northeast region, research on this parasitoid flourished. Currently, the efforts to mass-rearing this parasitoid, which started more than 70 years ago (Fonseca 1947), are being renewed (Fernandes et al. 2019). Distribution. Widely distributed in Africa (LaSalle & Wharton 2002). Introduced into several countries in the New World (Argentina, Brazil, Colombia and Uruguay), and established in Brazil (Costa et al. 2005) and Hawaii. Distribution in Brazil. BA, CE, RN (Costa et al. 2005), CE (Araujo et al. 2015; Fernandes et al. 2020, MG (Abreu et al. 2019), PI (Ara��jo et al. 2016; Coelho et al. 2020), RN (Fernandes et al. 2019), SP (Montes et al. 2011)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on pages 60-61, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Graham, M. W. R. V. (1991) A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae): revision of the remaining genera. Memoirs of the American Entomological Institute, 49, 1 - 322.","LaSalle, J. & Wharton, R. A. (2002) The identity and recognition of African Tetrastichus species (Hymenoptera: Eulophidae) associated with fruit flies (Diptera: Tephritidae). African Entomology, 10 (2), 297 - 304.","Pemberton, C. E. & Willard, H. F. (1918) A contribution to the biology of fruit-fly parasites in Hawaii. Journal of Agricultural Research, 15 (8), 419 - 467.","Fernandes, E., Araujo, E., Souza, I., Souza, M. & Nunes, G. (2019) Development and morphological characterization of the immature stages of Tetrastichus giffardianus Silvestri (Hymenoptera: Eulophidae). Revista Brasileira de Entomologia, 63 (3), 261 - 267. https: // doi. org / 10.1016 / j. rbe. 2019.05.005","Araujo, A. A. R., Silva, P. R. R., Querino, R. B. & Sousa, E. P. S. (2016) Tetrastichus giffardianus on pupae of Anastrepha in Brazil. Ciencia Rural, 46 (7), 1134 - 1135. https: // doi. org / 10.1590 / 0103 - 8478 cr 20150890","Abreu, B. R., Soares, D. P., Santos, T. P., Duraes, T. M., Conceicao, E. R. S., Brito, C. H., Matrangolo, C. A. R. & Alvarenga, C. D. (2019) Novos registros de parasitoides de moscas-das-frutas (Diptera: Tephritidae) em Minas Gerais. In: Bueno, A. F. (Pres.), Anais do 16 Simposio de Controle Biologico, Siconbiol, Londrina, 2019, pp. 209.","Costa, V. A., Araujo, E. L., Guimaraes, J. A., Nascimento, A. S. & LaSalle, J. (2005) Redescoberta de Tetrastichus giffardianus (Hymenoptera: Eulophidae) apos 60 anos da sua introducao no Brasil. Arquivos do Instituto Biologico, 72, 539 - 541. http: // www. biologico. sp. gov. br / uploads / docs / arq / v 72 _ 4 / costa. PDF","Araujo, E. L., Fernandes, E. C., Silva, R. I. R., Ferreira, D. C. L. & Costa, V. A. (2015) Parasitoides (Hymenoptera) de moscas-dasfrutas (Diptera: Tephritidae) no semiarido do estado do Ceara, Brasil. Revista Brasileira de Fruticultura, 37, 610 - 616. https: // doi. org / 10.1590 / 0100 - 2945 - 188 / 14","Fonseca, J. P. (1947) Criacao da vespinha africana no Instituto Biologico. O Biologico, 13 (9), 147 - 156.","Fernandes, E. C., Souza, M. M., Nava, D. E., Silva, J. G. & Araujo, E. L. (2020) Fertility life table and biology of Tetrastichus giffardianus (Hymenoptera: Eulophidae) in the larvae of Ceratitis capitata (Diptera: Tephritidae). Bulletin of Entomological Research 1 - 8.","Coelho, J. B., Araujo, E. S., Silva, L. B., Strikis, P. C., Zucchi, R. A. & Lopes, G. N. (2020) Frugivorous flies and their parasitoids associated with native fruits in an urban area. Semina Ciencias Agrarias, 41 (3), 1053 - 1060. https: // doi. org / 10.5433 / 1679 - 0359.2020 v 41 n 3 p 1053","Montes, S. M. N. M., Raga, A., Boliani, A. C. & Santos, P. C. (2011) Dinamica populacional e incidencia de moscas-das-frutas e parasitoides em cultivares de pessegueiros (Prunus persica L. Batsch) no municipio de Presidente Prudente-SP. Revista Brasileira de Fruticultura, 33 (2), 402 - 411. https: // doi. org / 10.1590 / S 0100 - 29452011005000052"]}

Spalangia gemina Bouček, 1963 (Figs 10, 13) Diagnosis. Head with dense circular setiferous punctures; gena with malar space distinctly shorter than eye height and malar sulcus absent; pronotal collar coarsely reticulate-rugose, except for a nearly triangular area close to the crenulate cross-line posteriorly; mesoscutal median lobe with median longitudinal carina separating two groups of irregular foveae posteriorly (Bouček 1963; Gibson 2009). Taxonomy. See S. endius. Biology. Most of the host records of this parasitoid are from tephritids. There are also reports of this species as hyperparasitoid, using tachinids (Diptera) as primary hosts, all in Lepidoptera. Other hosts include Musca domestica L. and species of Drosophila. Spalangia gemina has apparently a narrower host range than S. endius, and experiments showed that some hosts, including Drosophila virillis Sturtevant and A. ludens, are not parasitized by S. gemina (Sivinski et al. 1998). Tephritid hosts in Brazil include A. obliqua (Uch��a et al. 2003) and Ce. capitata (Silva et al. 2020). Biological control. See S. endius. Distribution. Mainly tropical, but reaching temperate zones of the New and Old World, and Australasia. Distribution in Brazil (associated with Tephritidae). MS (Uch��a et al. 2003) and RN (Silva et al. 2020)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 64, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Boucek, Z. (1963) A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Sbornik Entomologickeho Oddeleni Narodniho Musea v Praze, 35, 429 - 512.","Gibson, G. A. P. (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae). Zootaxa, 2259 (1), 1 - 159. https: // doi. org / 10.11646 / zootaxa. 2259.1.1","Sivinski, J., Vulinec, K., Meneses, E. & Aluja, M. (1998) The bionomics of Coptera haywardi (Ogloblin) (Diapriidae: Hymenoptera) and other pupal parasitoids of tephritid fruit flies (Diptera). Biological Control, 11, 193 - 202. https: // doi. org / 10.1006 / bcon. 1997.0597","Uchoa, M. A., Molina, R. M. S., Oliveira, I., Zucchi, R. A., Canal, N. A. D. & Diaz, N. B. (2003) Larval endoparasitoids (Hymenoptera) of frugivorous flies (Diptera, Tephritoidea) reared from fruits of the cerrado of the State of Mato Grosso do Sul, Brazil. Revista Brasileira de Entomologia, 47 (2), 181 - 186. https: // doi. org / 10.1590 / S 0085 - 56262003000200005","Silva, B. K. A., Silva, H. M., Fernandes, C. E., Costa, V. A. & Araujo, E. L. (2020) Pupal parasitoids associated with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in a semiarid environment in Brazil. Revista Brasileira de Entomologia, 64 (2), e 20190002. https: // doi. org / 10.1590 / 1806 - 9665 - rbent- 2019 - 0002"]}

Spalangia endius Walker, 1839 (Figs 9, 12) Diagnosis. Head and pronotal collar with sparse and circular setiferous punctures, interstices smooth and shiny; gena with malar sulcus present; posterior half of mesoscutal median lobe punctate-rugose medially, with punctures varying in size and shape (Bouček 1963; Gibson 2009). Biology. Recorded from about 50 species in nine families of Diptera (Noyes 2019), developing as solitary ectoparasitoid idiobionts on pupae. Spalangia endius is also recorded as a hyperparasitoid on Lepidoptera, probably via Tachinidae (Diptera) (Gibson 2009). Tephritid hosts in Brazil include A. alveatoides (Nic��cio et al. 2011), A. fraterculus, A. obliqua (Aguiar-Menezes & Menezes 2002; Uch��a et al. 2003) and A. sororcula (Aguiar-Menezes et al. 2003). Taxonomy. The genus Spalangia was revised by Gibson (2009) and recognition of species using the illustrated identification key is simple and straightforward. The three species listed here, are discussed in Gibson���s revision (2009), which should be consulted for further information and identification of species of Spalangia. Biological control. Species of Spalangia are the most commonly used biological control agents for filth flies (Diptera: Muscidae) on animal manure in several countries (e.g. Tormos et al. 2018 and references within), where companies mass-rear and sell parasitoids for augmentative biological control, with variable success (Machtinger et al. 2015a; 2015b). These are polyphagous species, not restricted to one dipteran family, and the same species are known to use Tephritidae and Muscidae species as hosts (Gibson 2009), with no observed preference for one family over the other (Tormos et al. 2018). Use of Spalangia spp. in biological control of tephritids is still restricted, and considered inefficient (Pe��a et al. 2002) Distribution. This is a cosmopolitan species, found in all biogeographical regions. Distribution in Brazil (associated with Tephritidae). MG (Silva et al. 2003), MS (Uch��a et al. 2003, Nic��cio et al. 2011) and RJ (Aguiar-Menezes & Menezes 2002; Aguiar-Menezes et al. 2003), Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on pages 62-64, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Boucek, Z. (1963) A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Sbornik Entomologickeho Oddeleni Narodniho Musea v Praze, 35, 429 - 512.","Gibson, G. A. P. (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae). Zootaxa, 2259 (1), 1 - 159. https: // doi. org / 10.11646 / zootaxa. 2259.1.1","Noyes, J. S. (2019) Universal Chalcidoidea Database. World Wide Web electronic publication. Available from: http: // www. nhm. ac. uk / chalcidoids (accessed 16 July 2019)","Nicacio, J. N., Uchoa, M. A., Faccenda, O., Guimaraes, J. A. & Marinho, C. F. (2011) Native larval parasitoids (Hymenoptera) of frugivorous Tephritoidea (Diptera) in South Pantanal Region, Brazil. Florida Entomologist, 94 (3), 407 - 419. https: // doi. org / 10.1653 / 024.094.0305","Aguiar-Menezes, E. L. & Menezes, E. B. (2002) Effect of time of permanence of host fruits in the field on natural parasitism of Anastrepha spp. (Diptera: Tephritidae). Neotropical Entomology, 31, 589 - 595. https: // doi. org / 10.1590 / S 1519 - 566 X 2002000400011","Uchoa, M. A., Molina, R. M. S., Oliveira, I., Zucchi, R. A., Canal, N. A. D. & Diaz, N. B. (2003) Larval endoparasitoids (Hymenoptera) of frugivorous flies (Diptera, Tephritoidea) reared from fruits of the cerrado of the State of Mato Grosso do Sul, Brazil. Revista Brasileira de Entomologia, 47 (2), 181 - 186. https: // doi. org / 10.1590 / S 0085 - 56262003000200005","Aguiar-Menezes, E. L., Menezes, E. B. & Loiacono, M. S. (2003) First record of Coptera haywardi Loiacono (Hymenoptera: Diapriidae) as a parasitoid of fruit-infesting Tephritidae (Diptera) in Brazil. Neotropical Entomology, 32, 355 - 358. https: // doi. org / 10.1590 / S 1519 - 566 X 2003000200025","Loiacono, M. S. (1981) Notas sobre Diapriinae neotropicales (Hymenoptera, Diapriidae). Revista de la Sociedad Entomologica Argentina, 40, 237 - 241.","Tormos, J., Beitia, F., Asisa, J. D. & de Pedro, L. (2018) Natal host and learning as factors in host preference by Spalangia cameroni Perkins (Hymenoptera: Pteromalidae). Crop Protection, 110, 155 - 159. https: // doi. org / 10.1016 / j. cropro. 2017.11.014","Machtinger, E. T., Geden, C. J., Kaufman, P. E. & House, A. M. (2015 a) Use of Pupal Parasitoids as Biological Control Agents of Filth Flies on Equine Facilities. Journal of Integrated Pest Management, 6 (1), 16. https: // doi. org / 10.1093 / jipm / pmv 015","Machtinger, E. T., Geden, C. J. & Leppla, N. C. (2015 b) Linear Dispersal of the Filth Fly Parasitoid Spalangia cameroni (Hymenoptera: Pteromalidae) and Parasitism of Hosts at Increasing Distances. PLoS ONE, 10 (6), e 0129105. https: // doi. org / 10.1371 / journal. pone. 0129105","Pena, J. E., Sharp, J. L. & Wysoki, M. (2002) Tropical fruit pests and pollinators. CAB International Publishing, New York, 448 pp.","Silva, C. G., Marchiori, C. H., Fonseca, A. R. & Torres, L. C. (2003) Himenopteros parasitoides de larvas de Anastrepha spp. em frutos de carambola (Averrhoa carambola L.) na regiao de Divinopolis, Minas Gerais, Brasil. Ciencia e Agrotecnologia, 27 (6), 1264 - 1267. https: // doi. org / 10.1590 / S 1413 - 70542003000600009"]}

Coptera haywardi Loi��cono, 1981 (Fig. 17) Diagnosis. Black with reddish-brown legs; head in dorsal view nearly as wide as long, vertex and occipital margin marked with large punctures; fore wing without distinct veins, but with longitudinal fold line, ending apically in a distinct notch in both sexes; metasomal petiole with complete median carina. This species is similar to Coptera pomonellae Muesebeck, differing mainly in the proportion between the eye and the malar space (larger in Co. pomonellae) and the complete median carina on the petiole (incomplete in Co. pomonellae). For the Neotropical species, Co. haywardi resembles Coptera sexpunctata (Ashmead), from which it can be distinguished by the smaller length/width ratio of head and petiole (Loi��cono 1981). Taxonomy. In the Neotropics, three species are known, Coptera brevipes (Kieffer), Co. haywardi and Co. sexpunctata (Johnson 1992; Notton 2014). The only key including Neotropical species was published more than a century ago (Kieffer 1916), species identification is therefore problematic, and the revision of Nearctic species by Muesebeck (1980) is helpful to some extent. In addition to those keys, a detailed, illustrated description is available for Co. haywardi, which allows comparison with reared material and more reliable identification (Loi��cono 1981). The Neotropical fauna is in urgent need of revision, and molecular characterization of the species seems to be important, since cryptic species have already been identified in this genus (Forbes et al. 2012). Biology. Endoparasitod koinobiont on pupae of tephritids (Sivinski et al. 1998), such as A. fraterculus, A. schultzi Blanchard (Loi��cono 1981) and A. ludens (Loew), infesting citrus (L��pez 1996); A. serpentina and A. striata Schiner in Venezuela (Garc��a & Montilla 2001); and A. fraterculus and A. sororcula in Brazil (Aguiar-Menezes et al. 2003) (Table 1). In the laboratory, Co. haywardi has been reared from A. obliqua (Garc��a & Montilla 2001), A suspensa (Loew, 1862), A. ludens, A. curvicauda (Gerstaecker) (as Toxotrypana curvicauda) and Ce. Capitata (Sivinski 1998). Biological control. This species has been considered for augmentative biological control programs because it possesses suitable biological traits, most importantly its specificity for fruit-infesting tephritids (Sivinski et al. 1998; Baeza-Larios et al. 2002). Distribution in the Neotropics. Recorded in Argentina, Brazil, Mexico and Venezuela. Distribution in Brazil: RJ (Aguiar-Menezes et al. 2003) (Table 1)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 58, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Loiacono, M. S. (1981) Notas sobre Diapriinae neotropicales (Hymenoptera, Diapriidae). Revista de la Sociedad Entomologica Argentina, 40, 237 - 241.","Johnson, N. (1992) Catalog of world Proctotrupoidea excluding Platygastridae. Memoirs of the American Entomological Institute, 51, 1 - 825.","Notton, D. G. (2014) A catalogue of the types of Diapriinae (Hymenoptera, Diapriidae) at the Natural History Museum, London. European Journal of Taxonomy, 75, 1 - 123. https: // doi. org / 10.5852 / ejt. 2014.75","Kieffer, J. J. (1916) Diapriidae. Das Tierreich, 44, 1 - 627. http: // biostor. org / reference / 80288","Muesebeck, C. F. W. (1980) The Nearctic parasitic wasps of the genera Psilus Panzer and Coptera Say (Hymenoptera, Proctotrupoidea, Diapriidae). U. S. Department of Agriculture, Technical Bulletin 1617. United States Department of Agriculture, Economic Research Service, Washington, D. C., 71 pp.","Forbes, A. A., Satar, S., Hamerlink, G., Nelson, A. E. & Smith, J. J. (2012) DNA Barcodes and targeted sampling methods identify a new species and cryptic patterns of host specialization among North American Coptera (Hymenoptera: Diapriidae). Conservation Biology and Biodiversity, 105 (4), 608 - 612. https: // doi. org / 10.1603 / AN 12012","Sivinski, J., Vulinec, K., Meneses, E. & Aluja, M. (1998) The bionomics of Coptera haywardi (Ogloblin) (Diapriidae: Hymenoptera) and other pupal parasitoids of tephritid fruit flies (Diptera). Biological Control, 11, 193 - 202. https: // doi. org / 10.1006 / bcon. 1997.0597","Lopez, M. (1996) Padrones de parasitismo en moscas de la fruta del genero Anastrepha (Diptera: Tephritidae), en frutos nativos (Spondias mombin L. y Psidium guajava L.) y exocotos (Mangifera indica L. y Citrus sinensis L.). MSc thesis, Universidad de Veracruz, Xalapa, 109 pp.","Garcia, J. L. & Montilla, R. (2001) Coptera haywardi Loiacono (Hymenoptera: Diapriidae) parasitoide de pupas de Anastrepha spp. (Diptera: Tephritidae) en Venezuela. Entomotropica, 16, 191 - 195","Aguiar-Menezes, E. L., Menezes, E. B. & Loiacono, M. S. (2003) First record of Coptera haywardi Loiacono (Hymenoptera: Diapriidae) as a parasitoid of fruit-infesting Tephritidae (Diptera) in Brazil. Neotropical Entomology, 32, 355 - 358. https: // doi. org / 10.1590 / S 1519 - 566 X 2003000200025","Baeza-Larios, G., Sivinski, J., Holler, T. & Aluja, M. (2002) The ability of Coptera haywardi (Ogloblin) (Hymenoptera: Diapriidae) to locate and attack the pupae of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), under seminatural conditions. Biological Control, 23, 213 - 218. https: // doi. org / 10.1006 / bcon. 2001.1010"]}

Family Eulophidae (Hymenoptera, Chalcidoidea) This is possibly the most diverse and common family of Chalcidoidea, with more than 4,000 species in 280 genera worldwide. In the Neotropics there are 500 species in 120 genera, but estimates are that Costa Rica alone has more than 2,000 species (LaSalle et al. 2006). The monophyly of the group is virtually undisputed (Heraty et al. 2013) and its position within Chalcidoidea seems to be basal, derived just after the two most basal egg-parasitoid families Mymaridae and Trichogrammatidae. Therefore, the ancestral Eulophidae would have been the first chalcidoids to attack hosts in stages other than eggs and to possess metallic coloration (Peters et al. 2018). The eulophids are extremely diverse in their natural histories, parasitism strategies and host associations. They attack hosts in many orders of insects and in any development stage from egg to adult, although the majority of the hosts are larvae of holometabolous insects. Exceptions include spiders and hemimetabolous insects (e.g., Odonata, Orthoptera and Hemiptera), in these cases usually parasitizing eggs (LaSalle et al. 2006). Tetrastichus giffardianus, a native of Africa, is the only species of Eulophidae that has been recorded in Brazil as parasitizing tephritids. It was introduced into Brazil in 1937 (Autuori 1938) for biological control of Ce. capitata. After its introduction, the species was believed to have failed to establish until it was rediscovered 60 years after the last report (Costa et al. 2005). Only one other eulophid parasitizes fruit-infesting tephritid flies in the New World, Aceratoneuromyia indica (Silvestri), which is also exotic, introduced from Southeast Asia, and has established successfully in several countries in the Americas (Ovruski et al. 2000)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 60, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["LaSalle, J., Schauff, M. E. & Hansson, C. (2006) Familia Eulophidae. In: Hanson, P. E. & Gauld, I. D. (Eds.), Hymenoptera de la Region Neotropical. Memoirs of the American Entomological Institute, 77, pp. 356 - 374.","Heraty, J. M., Burks, R. A., Cruaud, A., Gibson, G. A., Liljeblad, J., Munro, J., Rasplus, J. Y., Delvare, G., Jansta, P., Gumovsky, A. & Huber, J. (2013) A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics, 29, 466 - 542. https: // doi. org / 10.1111 / cla. 12006","Peters, R. S., Niehuis, O., Gunkel, S., Blasere, M., Mayer, C., Podsiadlowski, L., Kozlov, A., Donath, A., Noort, S., Liuj, X., Zhou, X., Misof, B., Heraty, J. & Krogmann, L. (2018) Transcriptome sequence-based phylogeny of chalcidoid wasps (Hymenoptera: Chalcidoidea) reveals a history of rapid radiations, convergence, and evolutionary success. Molecular Phylogenetics and Evolution, 120 (2018), 286 - 296. https: // doi. org / 10.1016 / j. ympev. 2017.12.005","Autuori, M. (1938) Notas sobre a introducao e multiplicacao do parasita Tetrastichus giffardianus Silv. no Brasil. O Biologico, 4 (4), 128 - 129.","Costa, V. A., Araujo, E. L., Guimaraes, J. A., Nascimento, A. S. & LaSalle, J. (2005) Redescoberta de Tetrastichus giffardianus (Hymenoptera: Eulophidae) apos 60 anos da sua introducao no Brasil. Arquivos do Instituto Biologico, 72, 539 - 541. http: // www. biologico. sp. gov. br / uploads / docs / arq / v 72 _ 4 / costa. PDF","Ovruski, S., Aluja, M., Sivinski, J. & Wharton, R. A. (2000) Hymenopteran parasitoids on fruit-infesting Tephritidae (Diptera) in Latin America and the southern United States: Diversity, distribution, taxonomic status and their use in fruit fly biological control. Integrated Pest Management Reviews, 5, 81 - 107. https: // doi. org / 10.1023 / A: 1009652431251"]}

Key to species of Diapriidae, Eulophidae and Pteromalidae (Hymenoptera) parasitoids of fruit flies (Diptera, Tephritidae) in Brazil 1 Fore wing with submarginal, marginal and stigmal veins present (Figs 5, 8, 15); antennae inserted close to or below lower margin of eyes, not on a projecting ledge (Figs 3, 6) (Chalcidoidea)............................................. 2 - Fore wing at most with a short submarginal vein (Figs 16, 17); antennae inserted on a projecting ledge near upper margin of eyes (Figs 1, 2) (Diaprioidea)............................................................................ 6 2 Tarsi 4-segmented; scutellum with a pair of submedian grooves and a pair of sublateral grooves (Fig. 4) (Eulophidae)...................................................................................... Tetrastichus giffardianus - Tarsi 5-segmented; scutellum without longitudinal grooves (Fig. 7) (Pteromalidae)................................. 3 3 Antennae separated from clypeus, inserted nearly in the middle of face, slightly above lower margin of eyes (Fig. 6); marginal vein distinctly and uniformly thickened (Fig. 8)...................................... Pachycrepoideus vindemmiae - Antennae inserted just above oral margin, far below lower margin of eyes (Fig. 3); marginal vein not thickened (Fig. 15) (Spalangia).......................................................................................... 4 4 Pronotum smooth, with setae on tiny bumps (Fig. 14); median lobe of mesoscutum with a single median pit (Fig. 14)............................................................................................ Spalangia simplex - Pronotum punctate or reticulate-rugose (Figs 12, 13); medial lobe of mesoscutum with different sculpturing............. 5 5 Pronotum with circular punctures distinctly separated by smooth interstices (Fig. 12); malar sulcus present; propodeum with paramedian crenulate furrows sub parallel, all cells with similar size................................ Spalangia endius - Pronotum mostly reticulate-rugose (Fig. 13); malar sulcus absent; propodeum with paramedian crenulate furrows widened anteriorly, forming a Y, with anterior cells larger than posterior cells................................ Spalangia gemina 6 Fore wing lacking tubular veins, with a mid-longitudinal fold line, apex with apical notch (Fig. 17)....... Coptera haywardi - Fore wing with a tubular vein (submarginal), without fold-line, apex rounded (Fig. 15)...... Trichopria sp. aff. anastrephae, Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 65, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551

Spalangia simplex Perkins, 1910 (Figs 3, 11, 14, 15) Diagnosis. Head and pronotal collar smooth, with scattered setae originating from pinprick-like punctures on head and tiny bumps on pronotal collar; gena with malar sulcus present; midlobe of mesoscutal median lobe smooth and shiny, with only a transverse row of setiferous punctures and a single median puncture posteriorly males with ventral setae of scape distinctly longer than dorsal setae (Bouček 1963; Gibson 2009). Taxonomy. See S. endius. Biology. This species has fewer host records than the other two species of Spalangia discussed here, most of them associated with species of Tephritidae and Drosophilidae (Gibson 2009; Sureshan & Farsana 2014). Tephritid hosts in Brazil are A. serpentina (Fernandes et al. 2013), Ce. capitata (Silva et al. 2020). Biological control. There is insufficient information on the host range to address the potential of this species for biological control programs. Distribution. Mainly tropical, but reaching temperate zones of the New and Old World, and Australasia. Distribution in Brazil (associated with Tephritidae). AP (Silva et al. 2007), CE and RN (Silva et al. 2020) and SP (Fernandes et al. 2013)., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 64, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Boucek, Z. (1963) A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Sbornik Entomologickeho Oddeleni Narodniho Musea v Praze, 35, 429 - 512.","Gibson, G. A. P. (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae). Zootaxa, 2259 (1), 1 - 159. https: // doi. org / 10.11646 / zootaxa. 2259.1.1","Sureshan, P. N. & Farsana, V. K. R. (2014) Two new distributional and host records for Spalangia Latreille parasitizing Dro- sophila sp. on putrefied tender jack fruit (Artocarpus heterophyllus) from Kerala, India. Journal of Biological Control, 28 (2), 57 - 61.","Fernandes, D. R. R., Vacari, A. M., Araujo, E. L., Guimaraes, J. A., Bortoli, A. S. & Perioto, N. W. (2013) Frugivorous larvae (Diptera, Tephritidae and Lonchaeidae) and native parasitoids (Hymenoptera) associated with Pouteria caimito (Sapotaceae) in Brazil. Florida Entomologist, 96, 255 - 257. https: // doi. org / 10.1653 / 024.096.0139","Silva, B. K. A., Silva, H. M., Fernandes, C. E., Costa, V. A. & Araujo, E. L. (2020) Pupal parasitoids associated with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) in a semiarid environment in Brazil. Revista Brasileira de Entomologia, 64 (2), e 20190002. https: // doi. org / 10.1590 / 1806 - 9665 - rbent- 2019 - 0002","Silva, R. A., Silva, W. R. & Jesus, C. R. (2007) Diversidade de parasitoides de Tephritidae em goiabeiras no Estado do Amap. In: Embrapa Recursos Geneticos e Biotecnologia (Org) Anais do Simposio de Controle Biologico, Brasilia, D. F., 2007, CD-ROM."]}

Family Diapriidae (Hymenoptera, Diaprioidea) This family comprises about 2,300 species in 197 genera (Johnson 1992; Arias 2003). Most of them are pupal parasitoids on dipterans and are frequently associated with fungi (Masner 2006). Diapriidae is subdivided into four subfamilies. The two species (Co. haywardi and Tr. anastrephae) that are parasitoids of fruit-infesting tephritids in Brazil belong to the subfamily Diapriinae. An illustrated key to genera of the New World Diapriinae, based mostly on Masner & Garc��a (2002), is available online (Yoder et al. 2007). Both the genera Coptera and Trichopria are highly speciose, and a revision of the Neotropical fauna is necessary (Ovruski et al. 2000). Within the parasitoid species treated here, the Diapriinae are exceptional in their parasitism strategy, because, as opposed to most pupal parasitoids, they are endophagous and, in the few studies observing oviposition stage, allow further development of their hosts (i.e. koinobionts) (Sivinski et al. 1998, Masner 2006). These parasitoids prefer young pupae and are subject to most of the same physiological constraints as other koinobionts, because the pupa is not permanently paralyzed but continues to develop for some time before the parasitoid finally kills it (Sivinski et al. 1998). Most species of Diapriinae recorded on fruit-flies are stenophagous, specialists on species of a single family of Diptera (Ovrurski et al. 2000; Sivinski & Aluja 2012). There is evidence that some species of Coptera could be oligophagous, or even monophagous, using one or a very few species of Rhagoletis as hosts (Forbes et al. 2012). The foraging strategy of pupal parasitoids (i.e., searching and ovipositing in puparia in the soil) is complementary to that of the larval parasitoids, and an interesting trait for biological control when allied with a narrow host range (e.g. Wang et al. 2016), especially when the prey larvae in the fruit are nearly inaccessible to larval parasitoids., Published as part of Shimbori, Eduardo Mitio, Costa, Valmir Antonio & Zucchi, Roberto Antonio, 2020, Annotated checklist and illustrated key to parasitoids (Hymenoptera: Diapriidae Eulophidae and Pteromalidae) of fruit flies (Diptera, Tephritidae) in Brazil, pp. 53-70 in Zootaxa 4858 (1) on page 57, DOI: 10.11646/zootaxa.4858.1.3, http://zenodo.org/record/4411551, {"references":["Johnson, N. (1992) Catalog of world Proctotrupoidea excluding Platygastridae. Memoirs of the American Entomological Institute, 51, 1 - 825.","Arias, T. M. (2003) Lista de los generos y especies de la superfamilia Proctotrupoidea (Hymenoptera) de la region Neotropical. Biota Colombiana, 4 (1), 3 - 32.","Masner, L. (2006) Superfamilia \" Proctotrupoidea. In: Hanson, P. E. & Gauld, I. D. (Eds.), Hymenoptera de la Region Neotropical. Memoirs of the American Entomological Institute, 77, pp. 236 - 253.","Yoder, M. J., Dole, K. & Deans, A. R. (2007) Updated New World genera of Diapriidae. Available from: http: // www. diapriid. org (accessed 11 August 2019)","Ovruski, S., Aluja, M., Sivinski, J. & Wharton, R. A. (2000) Hymenopteran parasitoids on fruit-infesting Tephritidae (Diptera) in Latin America and the southern United States: Diversity, distribution, taxonomic status and their use in fruit fly biological control. Integrated Pest Management Reviews, 5, 81 - 107. https: // doi. org / 10.1023 / A: 1009652431251","Sivinski, J., Vulinec, K., Meneses, E. & Aluja, M. (1998) The bionomics of Coptera haywardi (Ogloblin) (Diapriidae: Hymenoptera) and other pupal parasitoids of tephritid fruit flies (Diptera). Biological Control, 11, 193 - 202. https: // doi. org / 10.1006 / bcon. 1997.0597","Sivinski, J. & Aluja, M. (2012) The Roles of Parasitoid Foraging for Hosts, Food and Mates in the Augmentative Control of Tephritidae. Insects, 3 (3), 668 - 691. https: // doi. org / 10.3390 / insects 3030668","Forbes, A. A., Satar, S., Hamerlink, G., Nelson, A. E. & Smith, J. J. (2012) DNA Barcodes and targeted sampling methods identify a new species and cryptic patterns of host specialization among North American Coptera (Hymenoptera: Diapriidae). Conservation Biology and Biodiversity, 105 (4), 608 - 612. https: // doi. org / 10.1603 / AN 12012","Wang, X. G., Kacar, G., Biondi, A. & Daane, K. M. (2016) Foraging efficiency and outcomes of interactions of two pupal parasitoids attacking the invasive spotted wing drosophila. Biological Control, 96, 64 - 71. https: // doi. org / 10.1016 / j. biocontrol. 2016.02.004"]}

Family Eulophidae (Hymenoptera, Chalcidoidea) This is possibly the most diverse and common family of Chalcidoidea, with more than 4,000 species in 280 genera worldwide. In the Neotropics there are 500 species in 120 genera, but estimates are that Costa Rica alone has more than 2,000 species (LaSalle et al. 2006). The monophyly of the group is virtually undisputed (Heraty et al. 2013) and its position within Chalcidoidea seems to be basal, derived just after the two most basal egg-parasitoid families Mymaridae and Trichogrammatidae. Therefore, the ancestral Eulophidae would have been the first chalcidoids to attack hosts in stages other than eggs and to possess metallic coloration (Peters et al. 2018). The eulophids are extremely diverse in their natural histories, parasitism strategies and host associations. They attack hosts in many orders of insects and in any development stage from egg to adult, although the majority of the hosts are larvae of holometabolous insects. Exceptions include spiders and hemimetabolous insects (e.g., Odonata, Orthoptera and Hemiptera), in these cases usually parasitizing eggs (LaSalle et al. 2006). Tetrastichus giffardianus, a native of Africa, is the only species of Eulophidae that has been recorded in Brazil as parasitizing tephritids. It was introduced into Brazil in 1937 (Autuori 1938) for biological control of Ce. capitata. After its introduction, the species was believed to have failed to establish until it was rediscovered 60 years after the last report (Costa et al. 2005). Only one other eulophid parasitizes fruit-infesting tephritid flies in the New World, Aceratoneuromyia indica (Silvestri), which is also exotic, introduced from Southeast Asia, and has established successfully in several countries in the Americas (Ovruski et al. 2000).

The hymenopteran parasitoids of pest species of Tephritidae in Brazil are best known by their most prominent species, members of Braconidae and Figitidae. Species in the less-studied families Diapriidae, Eulophidae and Pteromalidae, which are mostly pupal parasitoids, have been largely neglected and the literature on these groups is sparse and scattered. Therefore, their importance as natural enemies of fruit flies is likely underestimated. Here, we present a parasitoid-host-plant checklist of all diapriids, eulophids and pteromalids that parasitize fruit flies of economic importance in Brazil, namely Anastrepha species and Ceratitis capitata. A compilation of information of the seven species of these parasitoids occurring in Brazil—Coptera haywardi Loiácono and Trichopria anastrephae Lima (Diapriidae), Tetrastichus giffardianus Silvestri (Eulophidae), Pachycrepoideus vindemmiae (Rondani), Spalangia endius Walker, S. gemina Bouček and S. simplex Perkins (Pteromalidae)—including their taxonomic status, general biology, and potential as biocontrol agents, is presented. Additionally, we provide an illustrated key to species, aiming to highlight key morphological features and facilitate identification at species level, stimulating future research on these groups.