First record of Zaprionus tuberculatus (Diptera: Drosophilidae) in the Serra do Espinhaço, Minas Gerais state, Brazil

The Serra do Espinhaço extends for more than 1,000 km across eastern Brazil and is divided into two major sections: the Minas Gerais portion in the south and the Bahia portion, known as Chapada Diamantina, in the north (Gontijo, 2008; Santos et al., 2020). Extensive high-altitude rocky outcrops, exceptional species richness, and high levels of endemism characterize the region. Consequently, it was designated as a UNESCO Biosphere Reserve (Brasil, 2018). In addition, sustainable land-use practices and the long-standing relationship between local communities and the landscape led the Food and Agriculture Organization (FAO) to recognize the Serra do Espinhaço as an Important Agricultural Heritage System (IAHS). Traditional communities in the region rely on the sustainable harvesting of native plant species and family farming, with fruit cultivation playing a central role in their livelihoods (Santos et al., 2020).

Among these cultivated crops, guava is one of the most economically important, both regionally and nationally. In 2023, guava was cultivated on 22,487 hectares in Brazil, yielding 582,832 tons, with an average productivity of 25,919 kg/ha. This production generated approximately 1.31 billion reais, highlighting the crop's importance to Brazilian agriculture (IBGE, 2023).

However, the profitability of guava production is threatened by several agricultural pests. Fruit flies are among the most important insect pests affecting fruit production and commercialization, posing major challenges for growers, regulatory agencies, and policymakers. Damage results from female oviposition, which punctures the fruit, and from larval feeding, which consumes the pulp and accelerates ripening by stimulating ethylene production in response to mechanical injury, frequently leading to premature fruit drop (Zart et al., 2011). In Brazil, several economically important fruit fly genera have already been reported (Zucchi, 2000).

The family Drosophilidae comprises approximately 4,000 species, with the genus Drosophila Fallén, 1823 being the most diverse, including more than 1,600 described species commonly known as fruit flies or vinegar flies (O'Grady and DeSalle, 2018). Understanding fruit fly diversity, host associations, and infestation rates is essential for developing effective management strategies (Silva et al., 2011). Among the fruit fly species associated with guava in Brazil are Anastrepha obliqua (Macquart, 1835), Anastrepha fraterculus (Wiedemann, 1830), Anastrepha serpentina (Wiedemann, 1830), Ceratitis capitata (Wiedemann, 1824), and Drosophila suzukii (Matsumura, 1931), all of which infest fruits and cause economically important losses (Benito et al., 2016; Trassato et al., 2017; Zucchi and Moraes, 2025).

Adding to this complex of guava pests, the Afrotropical genus Zaprionus Coquillett, 1902 gained attention in Brazil following the introduction of Zaprionus indianus Gupta, 1970 (Diptera: Drosophilidae) during the 1990s. In 2020, Zaprionus tuberculatus Malloch, 1932 (Diptera: Drosophilidae) was first recorded in the Federal District, Brazil. Until then, its known distribution included several African countries, such as Cameroon, Cape Verde, Egypt, Kenya, Mozambique, and South Africa, as well as regions surrounding the Mediterranean and western Asia, including Cyprus, Israel, the Canary Islands, and South Georgia (Oboňa et al., 2019; EPPO, 2020; Murvanidze et al., 2022).

Estimating the socioeconomic impacts of invasive fruit fly species is challenging because these impacts include reductions in agricultural productivity, high monitoring and containment costs, substantial investments in eradication programs despite their limited probability of success, and restrictions on domestic and international fruit trade. Furthermore, the inclusion of a new pest in export protocols substantially increases the costs associated with quarantine treatments, particularly when new disinfestation procedures are required (Papadopoulos et al., 2024).

Given the economic importance of guava production and the potential threat posed by invasive fruit flies to agricultural systems, this study reports the first occurrence of Z. tuberculatus in a cultivated guava area in the Serra do Espinhaço, Minas Gerais, Brazil. Additionally, a map showing the current distribution and host plant families of Z. tuberculatus in Brazil is provided.

Collection of study material. During the dry season, in August, five fallen guavas were collected from an orchard at Fazenda Mape Frutas Ltda., located in the municipality of Datas, Minas Gerais, Brazil (18°22'51.04" S, 43°45'49.11" W; 1,244 m.a.s.l) (Figure 1 A and B), a location with a predominantly tropical highland climate (Cwb).

Figure 1
Guava orchard in the municipality of Datas, Minas Gerais, Brazil (A); infested guava fruit collected in the area (B); adult specimen of Zaprionus tuberculatus Malloch, 1932 (C); and right prothoracic femur of Z. tuberculatus showing the characteristic diagnostic tubercle (D). Scale bars: 2 mm (C) and 1 mm (D).

Monitoring of the fruits. Each guava was placed individually in a 1 L plastic container, sealed with an organza-covered lid to allow air circulation and lined with paper towels to retain moisture. The fruits were kept under laboratory conditions in a climate-controlled room—with a temperature of 25 ± 2 °C, relative humidity of 70 ± 10%, and a 12-hour photoperiod—and monitored daily until adult emergence, which began approximately 48 hours after collection. The emerged drosophilids were immediately killed by freezing at -18 °C for 10 minutes and subsequently preserved in 70% ethanol.

Identification of the specimens and deposit of biological material in a reference collection. Morphological identification of the specimens was performed using a Leica EZ4 binocular stereomicroscope (Leica Microsystems). Identification followed the terminology proposed by Cavalcanti et al. (2022), Moreira et al. (2023), and Mendes and Lamas (2025). As three drosophilid species emerged from the collected fruits, the literature was consulted to distinguish Z. tuberculatus from Z. indianus and Drosophila melanogaster (Meigen, 1830) (Diptera: Drosophilidae), with particular focus on the morphology of the prothoracic leg (Figure 1C). Zaprionus tuberculatus is characterized, in both sexes, by the presence of a prominent tubercle on the outer surface of the medioventral margin of the forefemur, associated with a long seta adjacent to a short cuticular expansion on the inner surface of the same margin (Figures 1C and D) (Cavalcanti et al., 2022). Specimens of Z. tuberculatus were deposited in the entomological collection of the Universidade Federal dos Vales do Jequitinhonha e Mucuri (UFVJM), JK Campus, where accession numbers are pending.

Photodocumentation. Images of the study area were obtained using a Canon EOS Rebel 850D camera with an EF-S 18-55 mm (IS/STM) lens. Images of Z. tuberculatus were obtained using the same camera mounted on an AXOC NDPL 1 lens on the stereomicroscope at the Cell Biology Laboratory of UFVJM, Diamantina, Minas Gerais, Brazil.

Distribution map. The distribution map of Z. tuberculatus was created in QGIS, version 3.44, using information from the following sources: Cavalcanti et al. (2022); Mateus and Machado (2022); Montes and Vilela (2022); Jobim et al. (2023); Moreira et al. (2023); Santos et al. (2023); Ribeiro et al. (2024); Santos et al. (2024); Jobim et al. (2025); and Mendes and Lamas (2025).

Among the drosophilid species that emerged from guava fruits collected in a commercial orchard, Z. tuberculatus was recorded for the first time in a cultivated area of the Serra do Espinhaço, Minas Gerais, Brazil (Figure 1C).

Three drosophilid species emerged from the collected guava fruits: Z. indianus (105 individuals, 47%), Z. tuberculatus (87 individuals, 39%), and D. melanogaster (30 individuals, 14%).

Native to the Afrotropical region, Z. tuberculatus has expanded its distribution to several regions worldwide, likely due to its polyphagous feeding habits and ecological plasticity, which allow it to exploit fruits from a wide range of botanical families (Ebejer, 2001; Araujo et al., 2019; EPPO, 2020). Reflecting this high adaptability, Z. tuberculatus has rapidly expanded throughout Brazil and is currently confirmed in all five geographic regions of the country, encompassing eight states and the Federal District (Figure 2A) (Cavalcanti et al., 2022; Mateus and Machado, 2022).

Figure 2
Brazilian states (A) with confirmed records and botanical families (B) that include host plant species of Zaprionus tuberculatus Malloch, 1932 reported in Brazil. Legend: Distrito Federal (DF), Minas Gerais (MG), Pará (PA), Paraíba (PB), Paraná (PR), Rio de Janeiro (RJ), Rio Grande do Sul (RS), Santa Catarina (SC), and São Paulo (SP); Orange coloration indicates states with confirmed occurrence of Z. tuberculatus in Brazil.

Current Brazilian records indicate that Z. tuberculatus has been associated with 19 fruit species belonging to 10 botanical families. Among these, Myrtaceae and Moraceae account for 53% of the reported host species (Figure 2B) (Jobim et al., 2023; Moreira et al., 2023; Santos et al., 2023; Ribeiro et al., 2024; Santos et al., 2024; Jobim et al., 2025; Mendes and Lamas, 2025).

The establishment and rapid expansion of Z. tuberculatus in Brazil are of concern because the species occupies ecological niches similar to those of Z. indianus. The coexistence of these drosophilid species in guava orchards, together with D. melanogaster, may increase fruit infestation and damage. As generalists, these species exploit both native and cultivated hosts, increasing the potential for negative impacts on fruit production (Valadão et al., 2019; Vieira et al., 2019).

This study represents the second report of Z. tuberculatus infesting guava fruits in Brazilian orchards and the first record of the species in the Serra do Espinhaço, the only Globally Important Agricultural Heritage System (GIAHS) recognized by the FAO in Brazil. This record extends the known distribution of Z. tuberculatus in Minas Gerais and reinforces the rapid geographic expansion of this invasive species across the country.

Acknowledgements

To the Brazilian funding institutions: Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) [financial code 001], Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG) [project APQ-04955-23], for their support of the research. We also thank Mape Frutas Ltda. and its employees for their assistance in this study. The authors also thank Lucas Fernandes Silva, master of Animal Biology (UFVJM), for creating the map included in this manuscript.

Data Availability Statement

Data or information may be sent, upon prior request, to the corresponding author.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  21 Aug 2026
• Date of issue
  2026

History

• Received
  08 Dec 2025
• Accepted
  26 June 2026