Expanding the menu: new host plants of fruit flies (Diptera: Tephritidae) in southern BrazilAbstract
Sampling fruit, especially those of native species that act as natural reservoirs for fruit flies (Diptera: Tephritidae), is crucial for fruit production. Therefore, this study aimed to identify fruit fly species, their hosts, and natural enemies in the Pelotas region (Rio Grande do Sul, Brazil) through fruit sampling. Samples were collected from 15 sites within the Pelotas microregion between October 2022 and March 2024. Following the identification of emerged fruit flies and parasitoids, infestation and parasitism rates were calculated. The following species were recovered from a total of 3,827 puparia: Anastrepha fraterculus (Wiedemann, 1830) (lato sensu) (91.33%), Anastrepha chiclayae Greene, 1934 (0.89%), and Ceratitis capitata (Wiedemann, 1824) (7.7%). Infestation occurred in 19 of the 25 sampled fruit species, with the highest incidence in Myrtaceae species. The highest infestation rates were observed in Acca sellowiana (O. Berg) Burret, Passiflora caerulea L., Psidium guajava L. and Psidium cattleyanum Sabine. Doryctobracon areolatus (Szépligeti, 1911) was the most frequent parasitoid and exhibited the highest parasitism rates. Four new hosts were recorded: three for A. fraterculus [Campomanesia aurea O. Berg, Passiflora caerulea L., and Psidium salutare (Kunth) O. Berg] and one for C. capitata [Citrus unshiu (Swingle) Marcow]. These records expand the understanding of associations between Anastrepha species and host plants. Moreover, they reinforce the importance of host surveys, especially of native species in woodlands adjacent to orchards, for the monitoring and management of A. fraterculus.
Keywords:
Anastrepha fraterculus (lato sensu); Ceratitis capitata; infestation indices; parasitoids
Resumo
A amostragem de frutos, principalmente de espécies nativas, que são reservatórios naturais para as moscas-das-frutas (Diptera: Tephritidae), é essencial para o manejo desses insetos. Assim, este estudo teve por objetivo conhecer as espécies de moscas-das-frutas, seus hospedeiros e inimigos naturais na região de Pelotas, RS, por meio da amostragem de frutos. Foram amostrados frutos em 15 áreas da microrregião de Pelotas, entre outubro de 2022 e março de 2024. Após a identificação das espécies de moscas-das-frutas e parasitoides emergidos, foram calculados os índices de infestação das moscas-das-frutas e os índices de parasitismo. De 3.827 pupários, emergiram Anastrepha fraterculus (Wiedemann, 1830) (lato sensu) (91,33%), Anastrepha chiclayae Greene, 1934 (0,89%) e Ceratitis capitata (Wiedemann, 1824) (7,7%). Houve infestação em 19 das 25 espécies de frutíferas amostradas, com maior incidência nas espécies de Myrtaceae e maiores índices de infestação em Acca sellowiana (O. Berg) Burret, Passiflora caerulea L., Psidium guajava L. e Psidium cattleyanum Sabine. Doryctobracon areolatus (Szépligeti, 1911) foi o parasitoide mais frequente e com maiores taxa de parasitismo. Foram descobertos quatro novos hospedeiros, sendo três para A. fraterculus [Campomanesia aurea O. Berg, Passiflora caerulea L., e Psidium salutare (Kunth) O. Berg] e um para C. capitata [Citrus unshiu (Swingle) Marcow]. Esses registros ampliam o conhecimento sobre a associação de espécies de Anastrepha com plantas hospedeiras e reforçam a importância dos levantamentos de hospedeiros, especialmente nas espécies nativas em matas vizinhas aos pomares, para o monitoramento e manejo de A. fraterculus.
Palavras-chave:
Anastrepha fraterculus (lato sensu); Ceratitis capitata; indices de infestação; parasitoides
1. Introduction
Fruit farming is a major economic activity in Rio Grande do Sul (RS), one of Brazil’s leading producers of temperate fruits. However, efforts to increase productivity and marketability face considerable challenges, particularly those associated with several species of fruit flies in the genus Anastrepha (Diptera: Tephritidae). These highly polyphagous pests sustain populations across seasons, compromising fruit production in multiple orchards within the same region (Schliserman et al., 2014). Anastrepha fraterculus (Wiedemann, 1830) prevails in RS, infesting both introduced commercial crops and wild hosts (Kovaleski et al., 1999). Currently, A. fraterculus comprises a complex of cryptic species (Norrbom et al., 2025) that has yet to be formally described.
Monitoring is essential for the control of fruit fly populations. Traditionally, McPhail traps baited with attractants have been employed because of their ease in collecting and recording specimens (Bortoli et al., 2016). Data from these traps are valuable for correlating fruit fly occurrences with abiotic factors. However, this sampling method fails to provide information on biotic factors (e.g., tritrophic relationships among flies, hosts, and parasitoids) or fruit infestation levels (Nascimento et al., 2000). Comparative analyses of trap versus fruit sampling based on faunistic indices (abundance, diversity, and species richness) have demonstrated the importance of fruit sampling for monitoring Anastrepha species (Araujo et al., 2019).
In RS, A. fraterculus has been associated with 29 host plants, primarily within Myrtaceae, Rosaceae, and Rutaceae (Zucchi and Moraes, 2026). These include native species such as cattley guava (Psidium cattleianum Sabine), guava (Psidium guajava L.), and feijoa [Acca sellowiana (O. Berg) Burret], as well as introduced species such as citrus (Citrus spp.), apple (Malus spp.), and peach [Prunus persica (L.) Batsch]. However, given the abundance of native fruit-bearing plants in the state and the scarcity of fruit sampling studies, it is likely that there are unrecorded A. fraterculus hosts in the Pelotas (RS) microregion. When located near commercial orchards, these hosts can complicate fruit fly management efforts, as they provide suitable conditions for sustaining pest populations.
This study aimed to survey fruit flies, their hosts (cultivated and wild), and parasitoids in the Pelotas microregion, the main peach-producing area in RS, through fruit sampling.
2. Materials and Methods
Fruit sampling was conducted within the Pelotas microregion, covering rural areas in the municipalities of Canguçu, Morro Redondo, and Pelotas (including urban areas in the latter). Commercial and native fruits exhibiting signs of infestation (e.g., puncture marks and/or premature decay) were collected between October 2022 and March 2024 (Table 1). Sampling depended on fruit availability and seasonality. The goal was to sample 100 fruits per host plant whenever possible. Samples were gathered from the canopy, the ground, and the immediate vicinity of the plants, prioritizing fallen fruit. The scientific nomenclature for host plants follows the World Flora Online standards (WFO, 2026) standards (Table 2).
Fruit sampling sites in the municipalities of Canguçu (A), Morro Redondo (B), and Pelotas (C1, rural area; C2, urban area), Rio Grande do Sul, Brazil.
Fruit tree species sampled in the Pelotas microregion, Rio Grande do Sul, from October 2022 to March 2024.
The sampled fruits were isolated, weighed, and placed in 700 mL plastic cups over a 3-cm layer of fine vermiculite. The containers were covered with voile fabric secured by rubber bands. After 7 days, the vermiculite was sieved to collect puparia, which were then maintained in climate-controlled rooms at 27 ± 1 °C, 60 ± 20% relative humidity, and a 12-h photophase until the emergence of flies and/or parasitoids. The fruit flies and parasitoids were preserved in 70% ethanol and identified using the taxonomic keys of Zucchi (2000) and Marinho et al. (2018), respectively. Fruit fly parasitoid associations were determined following the method described by Leonel Junior et al. (1996). Fruit fly infestation rates were calculated based on the number of puparia per kilogram of fruit and the number of puparia per fruit. Parasitism was calculated using the following formula: (number of emerged parasitoids × 100) / (no. of emerged flies + no. of emerged parasitoids) (Matrangolo et al., 1998).
The collected fruit flies and parasitoids were preserved in 70% ethanol at Embrapa Clima Temperado in Pelotas, RS. Throughout the text, A. fraterculus is treated in the broad sense (lato sensu), as it comprises a complex of cryptic species.
3. Results
A total of 146.81 kg of fruit, representing 25 species, was sampled in the municipalities of Canguçu, Morro Redondo, and Pelotas. From this sample, 3,827 fruit fly puparia were collected, resulting in the emergence of 1,119 tephritids and 104 parasitoids (Braconidae and Figitidae). Infestation was observed in 19 of the sampled fruit species, including the first records in Brazil of A. fraterculus in Campomanesia aurea O. Berg (Myrtaceae), Passiflora caerulea L. (Passifloraceae), and Psidium salutare (Kunth) O. Berg (Myrtaceae), as well as the first record of C. capitata in Citrus unshiu (Swingle) Marcow (Figure 1). No fruit fly infestation was detected in Citrus limon (L.) Osbeck (Rutaceae), Duranta erecta L. (Verbenaceae), Inga marginata Willd. (Fabaceae), Pereskia aculeata Mill. (Cactaceae), Physalis peruviana Mill. (Solanaceae), or Solanum sp. (Solanaceae).
New hosts for Anastrepha fraterculus and Ceratitis capitata. (A) Campomanesia aurea (Myrtaceae); (B) Psidium salutare (Myrtaceae); (C) Citrus unshiu (Rutaceae)*; (D) Passiflora caerulea (Passifloraceae). *Photography by Paulo Lanzetta.
Most fruit flies collected were identified as A. fraterculus (91.33%), which infested nearly all native and exotic hosts, especially those in the Myrtaceae (Table 3). This species was found in all municipalities, including urban areas. Mediterranean fruit fly, Ceratitis capitata (Wiedemann, 1824) accounted for 7.7% of the collected flies, predominantly infesting introduced hosts such as Diospyros kaki L.f. (Ebenaceae) in Canguçu and Citrus unshiu (Swingle) Marcow. (Rutaceae) in Pelotas (first record). Infestation by the Mediterranean fruit fly in Psidium guajava L. (Myrtaceae) was observed in both urban and rural environments. Anastrepha chiclayae Greene, 1934 was collected exclusively from P. caerulea L. in the rural area of Pelotas.
Infestation rates of fruit flies in different host plants in peach-producing municipalities of the Pelotas microregion, Rio Grande do Sul, Brazil.
Anastrepha fraterculus was parasitized by four species belonging to the families Braconidae (three species) and Figitidae (one species) (Figure 2A). Doryctobracon areolatus (Szépligeti, 1911) exhibited the highest parasitism rates across hosts in both rural and urban environments in Pelotas. Opius bellus (Gahan, 1930) parasitized A. fraterculus in P. caerulea (urban area) and E. uniflora (rural area) in Pelotas (Figure 2A). A single specimen of Doryctobracon brasiliensis (Szépligeti, 1911) was recovered from A. fraterculus in Eriobotrya japonica (Thunb.) Lindl. in the rural area of Morro Redondo (Figure 2A). Aganaspis pelleranoi (Brèthes, 1924) was recorded in P. guajava (rural and urban areas) and P. cattleianum (rural area) (Figure 2A). The highest parasitoid incidence (Figure 2A) and parasitism rates (Figure 2B) were associated with wild fruit species, such as E. uniflora, P. cattleianum, and P. guajava. However, these values were also notably high in P. guajava (native host) (Figure 2A) and pear (Pyrus communis) (introduced host) (Figure 2B), respectively.
Number of emerged parasitoids (A) and % of parasitism (B) of Anastrepha fraterculus on different host fruits.
4. Discussion
Fruit sampling confirmed the prevalence of A. fraterculus in the Pelotas microregion. Its presence in 17 of the 25 host plants underscores the need for continued monitoring in fruit crops, particularly those of commercial importance. The novel records of A. fraterculus in new hosts within Brazil (Figure 1) highlight the need to extend sampling to other fruit-bearing species. These plants may serve as alternative hosts, sustaining or increasing populations of A. fraterculus that will eventually migrate to commercial orchards. Furthermore, understanding the seasonality of wild fruits is essential, as they act as reservoirs for fruit flies that may subsequently infest commercial crops.
The detection of A. fraterculus in C. aurea and P. salutare represents the first record of an Anastrepha species in these hosts (see Norrbom, 2022). However, this is the first record of A. fraterculus in P. caerulea in Brazil, despite previous records in Argentina (Putruele, 1996).
Campomanesia aurea (commonly known locally as guabirobinha-do-campo) is a subshrub native to the Pampa biome of RS that is also distributed across the states of Paraná and Santa Catarina (Stumpf, 2009; Lima et al., 2011). Its range also extends into Argentina, Paraguay, and Uruguay (Lima et al., 2011). Growing up to 1 m in height, C. aurea is considered an ornamental species characterized by woody stems and white flowers, with fruiting occurring between October and February (Stumpf, 2009). Psidium salutare is another native subshrub found in RS that bears a strong morphological resemblance to C. aurea. It typically inhabits open environments dominated by Poaceae and Fabaceae species and, in many instances, including the site of this study, coexists with other Myrtaceae, such as those in the genus Campomanesia (see Landrum, 2003). Because both species develop during the hot, dry season (February), they may serve as potential reservoirs for A. fraterculus during periods of resource scarcity.
The prevalence of A. fraterculus across fruit species aligns with reported findings for the southern region of the state (Araujo et al., 2025; Nunes et al., 2012; Salles, 1995), as well as for the rest of the state (Gattelli et al., 2008; Garcia and Corseuil, 1998). The highest A. fraterculus infestation rates were observed in feijoa (A. sellowiana), blue passionflower (P. caerulea), guava (P. guajava), and cattley guava (P. cattleianum), respectively (Table 3). However, infestation in blue passionflower is likely influenced by anthropogenic factors within the urban environment, given that this host was not attacked by A. fraterculus in rural areas. Fruiting among the Myrtaceae species exhibiting the highest infestation rates occurred between late summer and early autumn under milder climatic conditions. This finding explains the increased abundance of A. fraterculus during this period, as fruit fly population dynamics mirror the phenological development of native fruit hosts (Souza-Filho et al., 2009). Nevertheless, the flies also persist through winter in alternative hosts that serve as reservoirs. Consequently, it was observed that A. fraterculus is present year-round, facilitated by its ability to develop in a diverse range of commercial and native hosts.
Ceratitis capitata infestation levels were highest in persimmon (D. kaki) and loquat (E. japonica), both of which are common introduced species to the Pelotas microregion. In 2023, however, infestation by A. fraterculus was observed in loquat (an exotic host), although not by C. capitata. The presence of A. fraterculus in loquat within the Pelotas microregion had previously been noted by Nunes et al. (2012). Conversely, along the western border of RS, the Mediterranean fruit fly was found to be prevalent in loquat (Dias et al., 2013), indicating regional variation in the species' behavior within the Pampa biome. The detection of C. capitata in C. unshiu marks the first record of this association in Brazil, despite previous records in Argentina (Putruele, 1996).
Anastrepha chiclayae, previously misidentified in Brazil as A. dissimilis Stone, 1942 (see Araujo et al., 2023), infested P. caerulea. This occurrence had already been reported (as A. dissimilis) in the Middle Plateau region of RS, in the municipalities of Passo Fundo and Quatro Irmãos (Marsaro Júnior, 2014), and recently in Pelotas (Araujo et al., 2023).
Regarding fruit species not infested (C. limon, D. erecta, I. marginata, P. aculeata, P. peruviana, and Solanum sp.), most of them are poorly associated with fruit flies. These hosts appear to have been little explored by Anastrepha species or only minimally sampled in surveys conducted across Brazil. According to Zucchi and Moraes (2026), there is only a single record of infestation for these species (except for I. emarginata for which no records are available) in the states of Mato Grosso do Sul, Minas Gerais and São Paulo. These patterns may be explained by a combination of factors, including: (i) physicochemical properties of the fruits, (ii) structural barriers to oviposition, (iii) fruit size and resource limitation, and (iv) species-specific host preferences. The low interaction between these plants and fruit flies may be related to their physicochemical and structural characteristics. For instance, high pulp acidity in C. limon and the presence of potentially toxic compounds in Solanum sp. may impair larval development. In addition, small fruit size, as in D. erecta, may limit resource availability, while mechanical barriers to oviposition, such as protective fruit structures in I. marginata and P. peruviana, may reduce successful infestation. The absence of fruit flies recorded on P. aculeata was due to the lack of collection of Anastrepha barbiellini, (Lima, 1938) the only species that develops on this host (Almeida et al., 2019; Garcia et al., 2021). Future studies should experimentally evaluate these hypotheses to confirm host suitability.
Although the levels of pupal mortality, fruit fly emergence, and parasitism observed are generally consistent with the regional literature (Nunes et al., 2012; Araujo et al., 2025), particularly for native host species such as E. uniflora, it is plausible to acknowledge the existence of methodological limitations that may have influenced these results. Fruit decay after collection and isolation may lead to larval mortality or promote microbial contamination of pupae, especially in fruits with succulent pulp, which increase moisture within the container and may release liquids onto the pupae and substrate. Additionally, thermal fluctuations during sample handling, particularly when samples are transferred between different environments, may cause stress to immature stages. Biological factors may also be considered important contributors to pupal mortality, including fruit hyperinfestation and high larval density, which intensify intraspecific competition for resources. This may have been the case for A. sellowiana, which in this study produced 1,039 pupae from only 50 fruits. Furthermore, parasitism may contribute to pupal mortality even in the absence of adult parasitoid emergence, due to developmental failure or early mortality of parasitoids. In this context, future studies incorporating pupal dissection or complementary approaches may help detect non-emergent parasitism and improve understanding of these processes.
Doryctobracon areolatus was the most frequently collected parasitoid, consistent with observations across nearly the entire Brazilian territory (Marinho et al., 2018; Marinho and Zucchi, 2023). The highest parasitism rates for frugivorous larvae were recorded in Surinam cherry (E. uniflora), pear (P. communis), and cattley guava (P. cattleianum). The association between D. areolatus and soft-skinned, thin-fleshed myrtaceous fruits, such as those sampled in this study, is well documented (Marinho et al., 2009). However, parasitism events in sweet orange (C. sinensis), guava (P. guajava), and pear (P. communis) demonstrate that the long ovipositor of D. areolatus enables it to reach larvae in fruits with thick pulp and tougher skins, particularly when these hosts are located near myrtaceous areas infested by fruit flies. In the Pelotas microregion, Araujo et al. (2025) had previously associated D. areolatus only with A. sellowiana and P. cattleianum. In the present study, in addition to P. cattleianum, D. areolatus was found parasitizing A. fraterculus larvae in six other host plants. This expands the number of known host plants for this parasitoid in the Pelotas microregion to seven, as D. areolatus was not recovered from fruit fly larvae in A. sellowiana in our samples.
The second most frequent species was O. bellus, found in blue passionflower in the urban area and Surinam cherry in the rural area. Although this species represents the first known record of parasitism in an urban environment, rural parasitism had previously been observed in the region, as had the occurrence of A. pelleranoi in cattley guava and guava (Salles, 1996; Nunes et al., 2012; Araujo et al., 2025). This parasitoid is primarily recovered from fruit samples collected from the ground (Nunes et al., 2012; Canejo et al., 2023; Monteiro et al., 2023), a pattern confirmed by this study. Parasitism of larvae in E. japonica by D. brasiliensis was low, consistent with previous regional records (Salles, 1996; Araujo et al., 2025).
In the Pelotas microregion, A. fraterculus is the most notable fruit fly pest for fruit production, as it infests a wide range of hosts, including peaches, the region’s most economically important fruit, throughout almost the entire year. Based on Zucchi and Moraes (2026) database and our findings, the number of recorded hosts for A. fraterculus has increased to 32 in RS and 170 across Brazil.
Carvalho (1940) first reported C. capitata in RS, infesting pears and plums. Today, 85 years later, the Mediterranean fruit fly continues to broaden its host range within the state. With this new record on C. unshiu, there are now 14 known hosts in RS and 118 in Brazil. This figure demonstrates that, 125 years after its introduction to the country, the Mediterranean fruit fly is still expanding its invasive potential by adapting to new host species.
5. Conclusion
Anastrepha fraterculus infests a wide range of fruits throughout the year in the peach-producing microregion of Pelotas, including both commercial and wild hosts. The occurrence of three new host records for the species in Brazil, all of them native, highlights the need to expand knowledge on interactions among native hosts, potential population reservoirs, and fruit flies. Doryctobracon areolatus, found parasitizing A. fraterculus in a wide variety of fruits, constitutes an important ecological agent in the biological control of the species. Taken together, these findings provide valuable information for the improvement of monitoring and integrated management strategies for fruit flies in southern Brazil.
Acknowledgements
This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001, and by the National Council for Scientific and Technological Development (CNPq), which provided financial support to the last two authors. We are grateful to our colleague Ângelo da Silva Lopes for his field assistance, which was essential for the sample collection. We are grateful Paulo Lanzetta by one picture shared with us.
Data Availability Statement
All datasets supporting the results of this study are included in the article.
References
-
ALMEIDA, L.B.M., COELHO, J.B., UCHOA, M.A. and GISLOTI, L.J., 2019. Diversity of fruit flies (Diptera: Tephritoidea) and their host plants in a conservation unit from Midwestern Brazil. The Florida Entomologist, vol. 102, no. 3, pp. 562-570. https://doi.org/10.1653/024.102.0333
» https://doi.org/10.1653/024.102.0333 -
ARAÚJO, A.S., ZUCCHI, R.A., NORRBOM, A.L., NANINI, F., CORREA, A.S., ALVARENGA, C.D., SOUZA-FILHO, M.F., NAVA, D.E. and SAVARIS, M., 2023. Integrative approach reveals the identity of Brazilian specimens previously recognized as Anastrepha dissimilis Stone, 1942 (Diptera: tephritidae). Zootaxa, vol. 5228, no. 3, pp. 317-336. https://doi.org/10.11646/zootaxa.5228.3.5 PMid:37044649.
» https://doi.org/10.11646/zootaxa.5228.3.5 -
ARAUJO, E.S., KRÜGER, A.P., CALVO, M.V., TELLES, M.H., NEUMANN, A.M., SCATONI, I.B., COSTA, V.A., NAVA, D.E., MIRÁS-AVALOS, J.M. and GARCIA, F.R.M., 2025. Tritrophic interactions among fruit flies (Diptera: Tephritidae), its parasitoids and cultivated and wild hosts in the Pampa Biome, Rio Grande do Sul, Brazil. Agriculture, vol. 15, no. 19, pp. 1993. https://doi.org/10.3390/agriculture15191993
» https://doi.org/10.3390/agriculture15191993 -
ARAUJO, M.R., URAMOTO, K., FERREIRA, E.N.L., MESQUITA-FILHO, W., WALDER, J.M.M., SAVARIS, M. and ZUCCHI, R.A., 2019. Fruit fly (Diptera: Tephritidae) diversity and host relationships in diverse environments estimated with two sampling methods. Environmental Entomology, vol. 48, no. 1, pp. 227-233. https://doi.org/10.1093/ee/nvy177 PMid:30544259.
» https://doi.org/10.1093/ee/nvy177 -
BORTOLI, L.C., MACHOTA JUNIOR, R., GARCIA, F.R.M. and BOTTON, M., 2016. Evaluation of food lures for fruit flies (Diptera: Tephritidae) captured in a citrus orchard of the Serra Gaúcha. The Florida Entomologist, vol. 99, no. 3, pp. 381-384. https://doi.org/10.1653/024.099.0307
» https://doi.org/10.1653/024.099.0307 -
CANEJO, F.W.G., CANEJO, R.P.R., DAL MOLIN, A. and SOUZA, J.M.G.A., 2023. New native host plant records for frugivorous flies (Diptera: Tephritidae, Lonchaeidae) in Northeastern Brazil. Entomological Communications, vol. 5, ec05040. https://doi.org/10.37486/2675-1305.ec05040
» https://doi.org/10.37486/2675-1305.ec05040 - CARVALHO, J.H., 1940. Existe a môsca do Mediterrâneo (Ceratitis capitata) [sic] no Rio Grande do Sul. Revista Agronomica, vol. 4, pp. 411-412.
-
DIAS, N.P., SILVA, F.F., ABREU, J.A., PAZINI, J.B. and BOTTA, R.A., 2013. Nível de infestação de moscas-das-frutas em faixa de fronteira, no Rio Grande do Sul. Revista Ceres, vol. 60, no. 4, pp. 589-593. https://doi.org/10.1590/S0034-737X2013000400020
» https://doi.org/10.1590/S0034-737X2013000400020 -
GARCIA, F.R.M. and CORSEUIL, E., 1998. Flutuação populacional de Anastrepha fraterculus (Wiedemann) e Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) em pomares de pessegueiro em Porto Alegre, Rio Grande do Sul. Revista Brasileira de Zoologia, vol. 15, no. 1, pp. 153-158. https://doi.org/10.1590/S0101-81751998000100013
» https://doi.org/10.1590/S0101-81751998000100013 -
GARCIA, F.R.M., ARAUJO, E.S., KRÜGER, A.P., TELLES, M.H.F. and COSTA, V.A., 2021. First record of parasitoids (Hymenoptera: Braconidae) associated with Anastrepha barbiellinii Lima, 1938 (Diptera: Tephritidae). Brazilian Journal of Biology, vol. 81, no. 4, pp. 1125-1126. https://doi.org/10.1590/1519-6984.234183 PMid:33053136.
» https://doi.org/10.1590/1519-6984.234183 -
GATTELLI, T., SILVA, F.F., MEIRELLES, R.N., REDAELLI, L.R. and DAL SOGLIO, F.K., 2008. Moscas frugívoras associadas a mirtáceas e laranjeira “Céu” na região do Vale do Rio Caí, Rio Grande do Sul, Brasil. Ciência Rural, vol. 38, no. 1, pp. 236-239. https://doi.org/10.1590/S0103-84782008000100038
» https://doi.org/10.1590/S0103-84782008000100038 -
KOVALESKI, A., SUGAYAMA, R.L. and MALAVASI, A., 1999. Movement of Anastrepha fraterculus from native breeding sites into apple orchards in Southern Brazil. Entomologia Experimentalis et Applicata, vol. 91, no. 3, pp. 459-463. https://doi.org/10.1046/j.1570-7458.1999.00514.x
» https://doi.org/10.1046/j.1570-7458.1999.00514.x - LANDRUM, L.R., 2003. A revision of the Psidium salutare complex (Myrtaceae). SIDA, Contributions to Botany, vol. 20, no. 4, pp. 1449-1469.
-
LEONEL JUNIOR, F.L., ZUCCHI, R.A. and DAZA, N.A.C., 1996. Parasitismo de moscas-das-frutas (Diptera: Tephritidae) por Braconidae (Hymenoptera) em duas localidades do Estado de São Paulo. Anais da Sociedade Entomológica do Brasil, vol. 25, no. 2, pp. 199-206. https://doi.org/10.37486/0301-8059.v25i2.1116
» https://doi.org/10.37486/0301-8059.v25i2.1116 -
LIMA, D.F., GOLDENBERG, R. and SOBRAL, M., 2011. O gênero Campomanesia (Myrtaceae) no estado do Paraná, Brasil. Rodriguésia, vol. 62, no. 3, pp. 683-693. https://doi.org/10.1590/2175-7860201162313
» https://doi.org/10.1590/2175-7860201162313 - MARINHO, C.F. and ZUCCHI, R.A., 2023. Taxonomia de parasitoides das moscas-das-frutas—Braconidae (Alysiinae e Opiinae). In: R.A. ZUCCHI, D.E. NAVA, R. ADAIME and A. MALAVASI, eds. Moscas-das-frutas no Brasil: conhecimento básico e aplicado. Piracicaba: Editora FEALQ, no. 2, pp. 293-332.
-
MARINHO, C.F., COSTA, V. and ZUCCHI, R.A., 2018. Annotated checklist and illustrated key to braconid parasitoids (Hymenoptera, Braconidae) of economically important fruit flies (Diptera, Tephritidae) in Brazil. Zootaxa, vol. 4527, no. 1, pp. 21-36. https://doi.org/10.11646/zootaxa.4527.1.2 PMid:30651473.
» https://doi.org/10.11646/zootaxa.4527.1.2 -
MARINHO, C.F., SOUZA-FILHO, M.F., RAGA, A. and ZUCCHI, R.A., 2009. Parasitóides (Hymenoptera: Braconidae) de moscas-das-frutas (Diptera: Tephritidae) no Estado de São Paulo: plantas associadas e parasitismo. Neotropical Entomology, vol. 38, no. 3, pp. 321-326. https://doi.org/10.1590/S1519-566X2009000300004 PMid:19618046.
» https://doi.org/10.1590/S1519-566X2009000300004 -
MARSARO JÚNIOR, A.L., 2014. Novos registros de hospedeiros de moscas-das-frutas (Diptera: Tephritidae) para o Rio grande do Sul. Revista de Agricultura (Piracicaba), vol. 89, no. 1, pp. 65-71. https://doi.org/10.37856/bja.v89i1.116
» https://doi.org/10.37856/bja.v89i1.116 -
MATRANGOLO, W.J.R., NASCIMENTO, A.S., CARVALHO, R.S., MELO, E.D. and JESUS, M., 1998. Parasitoides de moscas-das-frutas (Diptera: Tephritidae) associados a fruteiras tropicais. Anais da Sociedade Entomológica do Brasil, vol. 27, no. 4, pp. 593-603. https://doi.org/10.1590/S0301-80591998000400012
» https://doi.org/10.1590/S0301-80591998000400012 -
MONTEIRO, L.B., NISHIMURA, G. and MONTEIRO, R.S., 2023. Natural parasitism in fruit fly (Diptera: Tephritidae) and interaction with wild hosts surrounding apple orchards adjacent to Atlantic Forest fragments in Paraná State, Brazil. Brazilian Journal of Biology, vol. 83, e250505. https://doi.org/10.1590/1519-6984.250505 PMid:34669805.
» https://doi.org/10.1590/1519-6984.250505 - NASCIMENTO, A.S., CARVALHO, R.S. and MALAVASI, A., 2000. Monitoramento populacional. In: A. MALAVASI and R.A. ZUCCHI, eds. moscas-das-frutas importância econômica no Brasil conhecimento básico e aplicado Ribeirão Preto: Holos, pp. 109-112.
-
NORRBOM, A.L., 2022 [viewed 27 May 2025]. Tephritidae databases. In: N.J. LIQUIDO, A.L. NORRBOM, G.T. MCQUATE, K.A. SUITER, W.L. YEE and C.L. CHANG, eds. The USDA compendium of fruit fly host information (CoFFHI). Edition 5.0 [online]. Available from: https://coffhi.cphst.org
» https://coffhi.cphst.org -
NORRBOM, A.L., RODRIGUEZ, E.J., STECK, G.J., CASSEL, B.K., RUIZ-ARCE, R., MULLER, A., GANGADIN, A., SAVARIS, M., NOLAZCO, N., TROYA, H., RODRIGUEZ, P., PEÑALOZA BARRIA, Y., MADRIZ PICADO, J., RODRIGUEZ CLAVIJO, P.A., MOORE, M.R., BRANHAM, M., SUTTON, B.D., RASZICK, T., CATTLE, E. and WIEGMANN, B.M., 2025. A new phylogeny of Anastrepha (Diptera: Tephritidae) based on nuclear loci obtained by phylogenomic methods. Systematic Entomology, vol. 51, no. 1, e70003. https://doi.org/10.1111/syen.70003
» https://doi.org/10.1111/syen.70003 -
NUNES, A.M., MÜLLER, F.A., GONÇALVES, R.S., GARCIA, M.S., COSTA, V.A. and NAVA, D.E., 2012. Moscas frugívoras e seus parasitoides nos municípios de Pelotas e Capão do Leão, Rio Grande do Sul, Brasil. Ciência Rural, vol. 42, no. 1, pp. 6-12. https://doi.org/10.1590/S0103-84782012000100002
» https://doi.org/10.1590/S0103-84782012000100002 - PUTRUELE, M.T.G., 1996. Hosts for Ceratitis capitata and Anastrepha fraterculus in the northeastern Province of Entre Ríos, Argentina. In: B.A. MCPHERON and G.J. STECK, eds. Fruit fly pests: a world assessment of their biology and management Boca Raton: CRC Press, 586 p.
- SALLES, L.A.B., 1995. Bioecologia e controle da mosca-das-frutas sul-americana Pelotas: Embrapa Clima Temperado, 58 p.
- SALLES, L.A.B., 1996. Parasitismo de Anastrepha fraterculus (Wied.) (Diptera: Tephritidae) por Hymenoptera, na região de Pelotas, RS. Pesquisa Agropecuária Brasileira, vol. 31, no. 11, pp. 769-774.
-
SCHLISERMAN, P., ALUJA, M., RULL, J. and OVRUSKI, S.M., 2014. Habitat degradation and introduction of exotic plants favor persistence of invasive species and population growth of native polyphagous fruit fly pests in a Northwestern Argentinean mosaic. Biological Invasions, vol. 16, no. 12, pp. 2599-2613. https://doi.org/10.1007/s10530-014-0690-5
» https://doi.org/10.1007/s10530-014-0690-5 -
SOUZA-FILHO, M.F., RAGA, A., AZEVEDO-FILHO, J.A., STRIKIS, P.C., GUIMARÃES, J.A. and ZUCCHI, R.A., 2009. Diversity and seasonality of fruit flies (Diptera: Tephritidae and Lonchaeidae) and their parasitoids (Hymenoptera: Braconidae and Figitidae) in orchards of guava, loquat and peach. Brazilian Journal of Biology, vol. 69, no. 1, pp. 31-40. https://doi.org/10.1590/S1519-69842009000100004 PMid:19347143.
» https://doi.org/10.1590/S1519-69842009000100004 - STUMPF, E.R.T., 2009. Cores e formas no Bioma Pampa: plantas ornamentais nativas Pelotas: Embrapa Clima Temperado, 272 p.
-
WORLD FLORA ONLINE – WFO, 2026 [viewed 5 May 2025]. World flora online plant list [online]. Available from: http://www.worldfloraonline.org
» http://www.worldfloraonline.org - ZUCCHI, R.A., 2000. Taxonomia. In: A. MALAVASI and R.A. ZUCCHI, eds. Moscas-das-frutas de importância econômica no Brasil: conhecimento básico e aplicado Ribeirão Preto: Holos Editora, 327 p.
-
ZUCCHI, R.A. and MORAES, R.C.B., 2026. [viewed 12 March 2025]. Fruit flies in Brazil – Anastrepha species their host plants and parasitoids [online]. Available from: http://www.lea.esalq.usp.br/anastrepha/
» http://www.lea.esalq.usp.br/anastrepha/
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Editor:
Takako Matsumura Tundisi
