Book chapter 9: Parasitoids of dipterous of forensic importance in Brazil

English version of chapter 9 Parasitóides de dípteros de importância forense no Brasil In: Entomologia Forense: novas tendências e tecnologias nas ciências criminais.1a. ed.Rio de Janeiro : T echnical Books Editora, 2010, v.1, p. 270-281. T he largest superfamily among parasitoid Hymenoptera is Ichneumonoidea that occurs in all habitats, being easily collected. Its hosts are mainly larvae and pupae of holometabolous insects, excluding Megaloptera and Siphonaptera, occurring rarely on individual eggs and are subdivided into Ichneumonidae and Braconidae. Many Braconidae parasitis hemimetabola nymphs (Homoptera, Heteroptera, Isoptera and Psocoptera) and some adult parasites of Coleoptera and Hymenoptera. Some species may be phytophages (Matthews, 1984; Wahl & Sharkey in Goulet & Huber, 1993). Askew & Shaw (1986) distinguished the Ichneumonoidea in idiobionts, those who permanently paralyze the host and in koinobionts, those who temporarily paralyze, allowing the development of the post-posture host. Ichneumonoidea are among the most diverse neotropical superfamilies with more than 40,000 described species distributed in 694 genera and together with Chalcidoidea correspond to 48% of all neotropical Hymenoptera (Grissell & Schauff, 1990). Among the Braconidae, the Subfamily Alysiinae is cosmopolitan with more than 1000 species described worldwide; of the 41 genera of the New World, 20 occur in the Neotropical region (Wharton et al. 1997). Alysiinae are koinobion andt endoparatoids of brachicerous dipterans; Alysiini are Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 1/14 associated with more than 20 Families of Diptera (Wharton, 1984). Many of its hosts occur associated with manure, fungi or decaying plant and animal matter; few parasites T ephritidae larvae. Pupation of the parasitoid occurs within the host's pupary.Most are solitary parasitoid; some species of Aphaereta sp. are gregarious. T he main function of exodont jaws seems to be to allow the adult to break the host's pupary to leave him (Griffiths, 1964). T hese jaws could also serve to break the substrate where the host's pupary is located. Some of the best studied Alysiinae are the paleatic species Alysia manducator and the nearctic species Aphaereta pallipes. T he Alysiinae of the genera Gnathopleura Wharton, Aphaereta and Alysia Latreille have been created and used for biological control of caliphorids and muscids, but without much success (Legner, 1978; Wharton, 1984). In Brazil, studies on Parasiinae alysides of muscoid dipterans refer mainly to species of the genera Aphaereta and Gnathopleura. Aphaereta sp. was one of the predominant parasitoids on muscoid dipterans collected on various substrates in the states of Goiás and Minas Gerais (Marchiori et al., 2002; Marchiori & Silva Filho, 2004; Silva, 2007). Gnathopleura quadridentate, Wharton 1986 was recorded in several studies conducted in Brazil, associated with species of Oxysarcodexia, Sarcodexia and Peckia (Diptera, Sarcophagidae) on various substrates, occurring in areas of mesophilic forest and urban environments (Marchiori et al. , 2004; Marchiori & Silva Filho, 2004; 2006; Bonani et al., 2006; Marchiori, 2007). Chalcididae are usually endoparasitoid, primary and solitary mainly of mature Diptera larvae (including aquatic as in Stratiomyidae) and Lepidoptera pupae (Grissell & Schauff, 1997; T avares, 2007), but are also registered as Coleoptera parasitoids (including the supposed primitive hosts whose larvae develop in wood), Neuroptera, Hymenoptera, Strepsiptera and as hyperparasitoids (Naumann, 1991). T here are also gregarious and ectoparasitoid species (Noyes, 2003). Species of the genus Brachymeria Westood are important primary parasitoids of couscoid dipterans, such as those of the family Sarcophagidae (Burks, 1960; Habu, 1960; Grissel & Schauff, 1990; Marchiori, 2001) and Calliphoridae (Bishop et al., 1996). Some species are of economic importance because they attack insect pests and others Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 2/14 of forensic importance (Gauld & Bolton, 1988; Santana et al.2008). T he species Brachymeria podagrica (Fabricus) was found parasitising the following species of dipterans of forensic importance: Chrysomya sp., Chrysomya albiceps (Wiedemann), Chrysomya megacephala (Fabriicus) (Calliphoridae), Hemilucilia flavifacies (Calliphoridae) Erderlein, Ophyra aenescens (Wiedemann) (Muscidae), Oxysarcodexia thornax (Walker) (Sarcophagidae), Peckia chrysostoma (Wiedemann) (Sarcophagidae), Peckia intermutnas (Walker) (Diptera: Sarcophagidae), Sarcodexia lambens (Wiedemann) and Squamatoids trivittatus (Curran, 1927) (Diptera: Sarcohagidae) (Silva, 1999; Marchiori et al., 2001a; Marchiori et al., 2002a; Marchiori et al., 2003a; b; c; Silva et al., 2005). Granato & Oliveira Costa (2008) collected from immature specimens of dipterans of forensic importance Brachymeria sp. and Conura sp. T he family Diapriidae (Hymenoptera: Proctotrupoidea) has many common species, little is known about the biology of these parasitoids. Oviposition can occur in the host larva or pupa. Most Diapriidae are parasitoids of Calliphoridae, Muscidae, Sarcophagidae, Sphaercoeridae and T achinidae (Silva, 1991). Species of the genus T richopria are usually of immature stages of Diptera (Legner et al., 1976). T richopria sp. is cited in the literature as parasitoid of some Sarcophagidae, Sepsidae, Muscidae and Calliphoridae (Figg et al., 1983; Blume, 1984; Silva, 1991). T richopria sp. was found parasitizing the following species of dipterans of forensic importance: C. albiceps, Chrysomya putoria (Wiedemann), Fannia canicularis (L.), H. flavifacies, Ophyra sp., Palaeosepsis spp., Phaenicia eximia (Wiedemann), P. chrysosotma , P. intermutnas, P. eximia and Synthesiomyia nudiseta (Wulp) (Muscidae) (Silva, 1991; Monteiro & Prado, 2000; Marchiori et al., 2000a; Marchiori et al., 2001; Bonani et al., 2004; Marchiori et al., 2002; Marchiori et al., 2004) T he Family Encyrtidae and a general form are endoparasitoids of eggs, immature and adults or egg predators mainly of cochineals (Hemiptera: Coccidae and Pseudococcidae), although several hosts have already been registered in several groups of Arthropoda (T achikawa, 1981), as well as hyperparasitoids and polyembryonic parasitoids (Noyes, 1995). Most egg and pupe parasitoids have idiobiont strategy, while larval parasitoids have koinobiont strategy. Several particularities are noted in the biology of Encyrtidae. T he Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 3/14 phenomenon of polyembryonic consists of the development of multiple larvae from a single egg placed in the host (Noyes, 1985; 1987; 1988; 1990a; b; 1996; 2000; Noyes & Valentini, 1989; T rjapitzin, 1973a; b). T he species T achinaephagus zealandicus Ashmead was found parasitating the following species of dipterans of forensic importance: C. megacephala, C. putoria, Musca domestica (L.) (Muscidae) and S. lambens (Monteiro, 1995; Sereno & Neves, 2003; Costa et al., 2004; Morretti & Ribeiro, 2006) and the species Hemencrytus herbertii parasitoids the species: C. albiceps, Euboettcheria colusor (Curran et Walley) (Sarcophagidae), Hemilucilia flavifacies (Calliphoridae), M. domestica, Ophrya sp., O. thornax, P. eximia, P. chrysosotma, P. intermutnas and S. nudiseta (Silva, 1991; Noyes & Wodley, 1994; Marchiori et al., 2003b; d; Marchiori et al., 2004). T he Eulophidae family, as you would expect in such a large group, various habits of life occur as parasitoids of eggs, larvae and pupae of more than 10 orders of insects can be solitary, gregarious, idiobionts, koinobionts, ectoparasitoids and endoparastoids, including aquatic, predators of Arachnida oothetes and even Nematoda (LaSalle & Suffcha, 1995). However, there is a pattern among hosts, predominantly those whose immature larvae are housed in gallhouses, mines or stems, especially of Lepidoptera, Coleoptera, Diptera and Hymenoptera orders. T achinobia species behave like gregarious parasitoids of Lepidoptera and Diptera pupae (Boucek, 1977; La Salle, 1994). T achinobia spp. parasitised the following species of dipterans of forensic importance: C. albiceps, P. eximia, Megaselia scalaris (Loew) (Phoridae) O. aenescens, O. thornax, parasitou C. albiceps, P. eximia, Oxysarcodexia sp. and P. intermutnas and S. nudiseta (Silva, 1991; Marchiori et al., 2005a; Marchiori et al., 2006a). T he Eurytomide family are mostly parasitoids of usually endophytic immature insects tenants or phytophages. T enants can feed on the galley material and also from the gallant. Phytophages can be galleys, blowers or feed on seeds from a wide variety of plants. T here are several types of idiobiont parasitoids, koinobionts, solitary or gregarious, kleptoparasitoids, facultative hyperparasitoids or not (Noyes, 2003) and these occur in several orders, such as Diptera, Coleoptera, Hymenoptera, Lepidoptera, Heteroptera eggs and Orthoptera. Eurytoma sp. parasitized the following species of dipterans of forensic importance: Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 4/14 Fannia pusio (Wiedemann) (Fanniidae) (Marchiori, 2007a) and M. domestica (Bruno et al., 1992; Costa, et al., 2004). T he Figitidae family are parasitoids of Neuroptera and Diptera (Gauld & Bolton, 1988). T hey behave like primary parasitoids of dipteran larvae (Díaz & Gallardo, 1995; 1996). Eucoilinae are koinobionts and endoparatoids that ovipose in the larval stage of Diptera Cyclorrapha and emerge as adults of their host's pupary (Wharton et al., 1998), which remains alive until the pupa stage (Ovruski, 1994). T hey are the largest group of Cynipoidea presenting a wide geographical distribution (Gauld & Bolton, 1988). Several species of eucoilines have been reported as important natural enemies of different groups of dipterans (Wharton et al., 1998). T he species Acantheucoela sp. was found parasitating Oxysarcodexia sp. Kleidotoma nigra (Hartig) Palaeosepsis spp: Paraganapis egeria Díaz, Gallardo Walsh parasitating Cyrtoneurine pararescita Couri (Muscidae), Palaeosepsis spp., and F. pusio; T riplasta artocoxalis Ashmead C. pararescita, Palaeosepsis spp., and Sphaeroceridae spp.; T riplasta coxalis Ashmead parasitating Palaeosepsis spp.; T rybliographa sp. parasitating O. thornax (Silva, 1991; Marchiori et al., 2006) (Marchiori & Linhares, 1999; Marchiori et al., 1999; Marchiori et al., 200b;c;d; Marchiori et al., 2001b; Marchiori et al., 2001c; Marchiori et al., 2002b; Marchiori et al., 2005;b;c;d;e; Marchiori et al., 2007b) and Neralsia esplendens (Borgmeier) parasitating O. thornax (Marchiori et al., 2006b). T he Pteromalidae family (Hymenoptera, Chalcidoidea) includes a large number of parasitoid species, many of which are important in biological control of synanthropic muscoids (Cardoso & Milward-De-Azevedo, 1996). T hey can be solitary or gregarious, ectoparasitoids or endoparasitoids, primary or secondary parasitoids, koinobionts or idiobionts. Most are idiobiont and many develop as ectoparasitoids in larvae or puparies of Diptera, Coleoptera, Hymenoptera, Lepidoptera and Siphonaptera (Ioriatti, 1995). T he main species of parasitoids and their hosts are: Muscidifurax raptor Girault & Sanders: M. domestica and Palaeosepsis spp. Muscidifurax raptoroides (Kogan & Legner): M. domestica and C. putoria. Nasonia vitripennis (Walker) Cochliomyia hominivorax (Coquerel) (Diptera: Calliphoridae), C. albiceps, C. megacephala, C. putoria, H. flavifacies, M. domestica, O. thornax, P. chrysostoma, P. intermutans e Synthesiomyia nudiseta (Diptera: Muscidae). Pachycrepoideus. vindemmiae (Rondani): C. albiceps, C. megacephala, C. putoria, F. pusio, H. flavifacies, M. scalaris, M domestica, O. thornax, Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 5/14 Ophyra sp. (Diptera: Muscidae), Poeciolosomella sp. (Diptera: Sphaeroceridae), O. thornax, P. chrysostoma and Sarcodexia lambens (Diptera: Sarcophagidae). Spalangia cameroni Perkins: C. putoria, C. parescita, M. domestica, Palaeosepsis spp., R. belforti and Stomoxys calcitrans L. (Diptera: Muscidae). Spalangia drosophilae (Ashmead): F. pusio, O. thornax, Palaeosepsis spp. and P. chrysostoma. Spalangia endius Walker: C. putoria, C. parescita, F. pusio, M. domestica, O. thornax, Palaeosepsis spp., P. chrysostoma, S. lambens. and S. calcitrans. Spalangia gemina Boucek: M domestica, C. putoria and S. calcitrans. Spalangia nigra Latrielle: C. pararescita, F. pusio, M. domestica, O. thornax, Palaeosepsis spp. and R. belforti. Spalangia nigroaenae Curtis: C. parescita, M. domestica, O. thornax, Palaeosepsis spp., R. belforti and S. calcitrans. Spalangia sp.: O. thornax and P. intermutans (Silveira et al., 1980; Silva, 1991; Sereno & Neves, 1993a; 1993b; Monteiro, 1995; Almeida, 1996; Marchiori & Linhares, 1999; Marchiori et al., 1999; Marchiori et al., 200b;c;d; Marchiori et al., 2001b; Marchiori et al., 2001c; Marchiori et al., 2002b; Carvalho et al., 2003; Carvalho et al., 2003b; Carvalho et al., 2004; Marchiori et al., 2005;b;c;d;e; Marchiori et al., 2007b) (Marchiori et al., 2006a; Marchiori et al., 2006b, Rodrigues-Guimarães, 2006). References ALMEIDA, M.A.F. Abundância relativa e sazonal de Musca domestica L., 1758 (Diptera: Muscidae) e de seus parasitóides em microhabitats de um curral de gado bovino, em Pirassununga (SP). Campinas: 1996. 79p. [Dissertação (Mestrado) Instituto de Biologia UNICAMP]. ASKEW, R.R. & SHAW, M.R. Parasitoid communities: their size, structure and development. 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T riplasta coxalis Ashmead (Hymenoptera: Figitidae: Eucoilinae) como parasitóide de Palaeosepsis spp. (Diptera: Sepsidae) em fezes de búfalos em Itumbiara, Goiás, Brasil. Biotemas. , v.18, p.217 221, 2005c. MARCHIORI C. H.; SILVA FILHO, O. M;, PAULA, B. M. & ARANT ES, V. Ocorrência de T rybliographa sp. (Hymenoptera: Figitidae: Eucolinae) como parasitóide de Oxysarcodexia thornax (Walker, 1849) (Diptera: Sarcophagidae) no Brasil. Arquivos do Instituto Biológico. , v.72, p.265 266, 2005d. MARCHIORI, C. H.; SILVA FILHO, O. M.; FORT ES, F. C. Al.; BRUMES, R. R.; BORGES, R. F.; GONÇALVES, P. L. P. & LAURINDO, J. F. Parasitóides de Fannia pusio (Wiedemann, 1830) (Diptera: Fanniidae) coletados em Caldas Novas, Goiás, Brasil.. Ciência e Agrotecnologia. , v.29, p.1288 1291, 2005e. MARCHIORI, C.H.; SILVA FILHO, O. M.; PEREIRA, L. A.;, RIBEIRO, L. C. S.& BORGES, V. R. Hemencyrtus herbertii Ashmead (Hymenoptera: Encyrtidae) como parasitóides de dípteros muscóides coletados em Itumbiara, Goiás. Arquivo Brasileiro de Medicina Veterinária e Zootecnia. , v.56, p.691 693, 2004. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.; RIBEIRO, L. C. S. & BORGES, V. R. Parasitoids of Ophyra aenescens (Wiedemann) (diptera: Muscidae): fly of medicalsanitary importance collected in State of Goiás, Brazil. Revista da Sociedade Brasileira de Medicina T ropical. , v.36, p.629 631, 2003a. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.; RIBEIRO, L. C. S. & BORGES, V. R. Ocorrência de Brachymeria podagrica (Fabricius) (Hymenoptera: Chalcididae) como parasitóide de dípteros coletados em diferentes substratos em Itumbiara, Sul de Goiás, Brasil. Arquivo Brasileiro de Medicina Veterinária e Zootecnia. , v.55, p.246 248, 2003b. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.; RIBEIRO, L. C. S.; BORGES, V. R. & ARANT ES, S. B. Microhimenópteros parasitóides de dípteros muscóide coletados em diferentes substratos em área de mata em Itumbiara, Goiás, Brasil. Arquivo Brasileiro de Medicina Veterinária e Zootecnia. , v.55, p.510 513, 2003c. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.; RIBEIRO, L. C. S.; BORGES, V. R. First occurrence of parasitoid Hemencyrtus herbertii Ashmead (Hymenoptera: Encyrtidae) in pupae of Hemilucilia flavifacies Enderlein (Diptera: Calliphoridae) in Brazil. Revista Brasileira de Ciência Veterinária. , v.10, p.124 125, 2003d. MARCHIORI, C.H. & SILVA FILHO, O. M. Peckia chrysostoma (Wiedemann) (Diptera: Sarcophagidae) e seus parasitóides coletados no sul do Estado de Goiás. Revista Qeios, CC-BY 4.0 · Article, November 23, 2019 Qeios ID: 249386 · https://doi.org/10.32388/249386 10/14 Brasileira de Parasitologia Veterinária 13, n. 4, p. 165-168. 2008. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.; RIBEIRO, L. C. S. & BORGES, V. R. Brachymeria podagrica (Fabricius) (Hymenoptera: Chalcididae) as parasitoid of Sarcodexia lambens (wiedemann) (Diptera: Sarcophagidae) in Brazil. Arquivos do Instituto Biológico. , v.69, p.121 122, 2002a. MARCHIORI, C.H.; PEREIRA, L. A.;, SILVA FILHO, O. M.& RIBEIRO, L. C. S.; Paraganaspis egeria Díaz, Gallardo & Walsh (Hymenoptera: Figitidae: Eucoilinae) as potential agent in the biocontrol of muscoid dipterous collected in several substracts in Itumbiara,. Goias, Brazil. Arquivo Brasileiro de Medicina Veterinária e Zootecnia, v.54, p.662-664, 2002b. MARCHIORI, C. H. Ocorrência de Brachymeria podagrica (Fabricius) (Hymenoptera: Chalcididae) como parasitóide de Peckia chrysostoma (Wiedemann) (Diptera: Sarcophagidae) no Brasil. Entomología Y Vectores, v.8, n.4, p.513-517, 2001a. MARCHIORI, C. H.; OLIVIEIRA, A. T . & LINHARES, A. X. Artrópodes associados a massas fecais bovinas no Sul do Estado de Goiás. Neotropical Entomology. , v.30, p.19 24, 2001b. MARCHIORI, C. H.; CALDAS, E. R. & ALMEIDA, K. G. S. Primeira ocorrência de T riplasta coxalis Ashmead em pupas de Palaeosepsis sp. e nova ocorrência de Kleidotoma nigra (Hartig) (Hymenoptera: Figitidae: Eucoilinae) no Brasil. Arquivo Brasileiro de Medicina Veterinária e Zootecnia. , v.53, p.449 450, 2001c. MARCHIORI, C.H.; PEREIRA, L. A.; FILHO, O.M.S. & RIBEIRO, L.C.S. parasitóides gregários de dípteros muscoideos coletados em vários substratos em Itumbiara, Goiás. Biotemas 15, n. 1, p. 111-118 2002. MARCHIORI, C. H.; OLIVIEIRA, A. T . & LINHARES, A. X. T richopria sp. (Hymenoptera: Diapriidae) parasitóides de Diptera muscóidea. Arquivos do Instituto Biológico. , v.67, p.131 133, 2000a. MARCHIORI, C. H.; CALDAS, E. R.; SILVA, C. G.; T EIXEIRA, F. F.& LINHARES, A. X. Dípteros muscóides associados com fezes bovinas e seus parasitóides em Goiás. Arquivo Brasileiro de Medicina Veterinária e Zootecnia. , v.52, p.354 356, 2000b. MARCHIORI, C. H.; OLIVEIRA, A. T .; DÍAZ, N. B.; GALLARDO, F. PENT EADO-DIAS, A. M. & LINHARES, A.X. 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associated with more than 20 Families of Diptera (Wharton, 1984). Many of its hosts occur associated with manure, fungi or decaying plant and animal matter; few parasites T ephritidae larvae. Pupation of the parasitoid occurs within the host's pupary.Most are solitary parasitoid; some species of Aphaereta sp. are gregarious.
T he main function of exodont jaws seems to be to allow the adult to break the host's pupary to leave him (Griffiths, 1964). T hese jaws could also serve to break the substrate where the host's pupary is located.
Some of the best studied Alysiinae are the paleatic species Alysia manducator and the nearctic species Aphaereta pallipes. T he Alysiinae of the genera Gnathopleura Wharton, Aphaereta and Alysia Latreille have been created and used for biological control of caliphorids and muscids, but without much success (Legner, 1978;Wharton, 1984).
T he family Diapriidae (Hymenoptera: Proctotrupoidea) has many common species, little is known about the biology of these parasitoids. Oviposition can occur in the host larva or pupa. Most Diapriidae are parasitoids of Calliphoridae, Muscidae, Sarcophagidae, Sphaercoeridae and T achinidae (Silva, 1991).
parasitised the following species of dipterans of forensic importance: C. albiceps, P.
T hey behave like primary parasitoids of dipteran larvae (Díaz & Gallardo, 1995;. Eucoilinae are koinobionts and endoparatoids that ovipose in the larval stage of Diptera Cyclorrapha and emerge as adults of their host's pupary (Wharton et al., 1998), which remains alive until the pupa stage (Ovruski, 1994).
T hey are the largest group of Cynipoidea presenting a wide geographical distribution (Gauld & Bolton, 1988 T riplasta coxalis Ashmead parasitating Palaeosepsis spp.; T rybliographa sp.