Not only for fruit flies: a study on richardiid flies (Diptera: Richardiidae) captured in Multilure traps in an anthropic environmentABSTRACT
This study is based on specimens of richardiid flies (Richardiidae) collected during surveys of fruit flies Anastrepha Schiner, 1868 (Diptera: Tephritidae) at the “Luiz de Queiroz” Campus, Piracicaba, São Paulo, Brazil. Sampling was conducted from April 2019 to April 2020 using Multilure traps baited with putrescine and ammonium acetate in four environments with different levels of anthropogenic activity: an agroforestry system, the central campus area, a riparian forest, and a preserved forest fragment. We collected 120 specimens belonging to two species of Richardiidae (Richardia teevani Curran, 1934 and Richardiodes trimaculata Hennig, 1937). Richardiodes trimaculata (n = 93) was more abundant in less disturbed areas, whereas R. teevani (n = 27) showed consistently low abundance across all sites. Both species exhibited population peaks during the dry season (winter), but in different months. While Ri. trimaculata has a peak in July and August, R. teevani has the highest abundance in September. Richardiodes trimaculata is recorded for the first time in the state of São Paulo. This is the first study on the population dynamics of richardiid flies using Multilure traps baited with the food attractants, putrescine and ammonium acetate.
Keywords:
Ecology; Population fluctuation; Richardia teevani; Richardiodes trimaculata; Sampling
1. Introduction
Richardiidae (Diptera), currently comprising 35 genera and approximately 200 described species (Steyskal, 1968; Hancock, 2010; Wendt and Ale-Rocha, 2014, 2015). The family is predominantly distributed in the Neotropics, with about 90% of its known species occurring in this region. In South America, richardiids have not been recorded in Chile or Patagonia (Aczél, 1950), while a few species are known from the Nearctic region (Steyskal, 1987). In Brazil, 67 species belonging to 15 genera have been documented (Steyskal, 1968; Ale-Rocha and Barros, 2026).
Larvae of some species are saprophagous and are commonly found in decomposing plant material on the ground (Perez-Gelabert and Thompson, 2006). Richardia Robineau-Desvoidy, 1830 species have also been reported as dominant in fallen fruits in La Selva, Costa Rica (Inouye, 1999). Other richardiid larvae are phytophagous, developing in the bracts or flowers of Heliconia L. (Heliconiaceae) (Seifert and Seifert, 1976), in Araceae species (Kitching, 2000; Hernández-Ortiz and Aguirre, 2015; Wendt et al., 2018; Amancio et al., 2019), and in diseased coconut palms (Steyskal, 1958). Notably, the larva of Melanoloma viatrix Hendel, 1911 is a pest of pineapple Ananas comosus (L.) Merr. (Bromeliaceae) in Peru (Bello-Amez et al., 2020) and Brazil (Adaime et al., 2024).
In Brazil, records and ecological studies of Richardiidae have been predominantly conducted in the Amazon biome, using suspended flight interception traps, Malaise traps (Alencar et al., 2013; Wendt and Ale-Rocha, 2015), and pineapple-based sampling (A.L Oliveira-Filho et al. 2025). In other biomes, ecological investigations are more specific: in the Cerrado, one study addressed mimicry (Pie and Del-Claro, 2002), while in the Atlantic Forest, only a single diversity survey has been reported (Wendt and Ale-Rocha, 2014).
Information regarding richardiid flies in anthropized areas is unknown, as well as the use of Multilure traps for collecting them. Therefore, the present study aimed to evaluate the abundance and population fluctuations of richardiid flies collected with Multilure traps across habitats with different anthropization levels.
2. Material and Methods
2.1. Study area
The study was conducted in an area of 914.5 ha, covered by vegetation of the Atlantic Forest biome, classified as Seasonal Semi-deciduous Forest. This forest is characterized by emergent trees forming an irregular canopy ranging from 15 to 20 m in height (Alexandrino et al., 2013). The climate in Piracicaba, São Paulo is transitional from Tropical to Mild Tropical (Novais and Machado, 2023), with a tropical Aw climate, with dry winters and an annual average precipitation of 1,100–1,300 mm (Alvares et al., 2022).
Sampling was carried out in four areas of the “Luiz de Queiroz” Campus (Figure 1), characterized as follows (Ribeiro et al., 2021; Soares et al., 2023): (i) Areão Farm (Agroforestry Systems – SAF) (22°41′41.90″S; 47°38′25.22″W, 556 m a.s.l.) – This area is located within an industrial hub and surrounded by highways. It features coffee cultivation under a canopy of native and exotic plant species, along with various fruit trees (Figure 1a); (ii) Central Area (22°42′40.56″S; 47°37′55.71″W, 560 m a.s.l.) – An open field with small fragments of native and exotic plants, situated near administrative buildings with intense vehicle and pedestrian traffic (Figure 1b); (iii) Piracicamirim Riverbank (22°42′22.58″S; 47°37′39.05″W, 531 m a.s.l.) – A native riparian forest with some surrounding exotic plants and a few dirt roads with low vehicle and pedestrian traffic (Figure 1c); (iv) Monte Olimpo (22°43′9.41″S; 47°36′43.37″W, 585 m a.s.l.) – A remnant native forest surrounded by pasture and agricultural areas (Figure 1d).
Study area: (a) Areão Farm; (b) Central area; (c) Piracicamirim Riverbank; (d) Monte Olimpo.
2.2. Sampling and identification
Thirty-two Multilure traps were installed between April 2019 and April 2020 in four areas of the “Luiz de Queiroz” Campus (Figure 1a–d). In each area, traps were organized into four stations, with eight traps per block: four placed at 2 m and four at 10 m above ground, resulting in four sampling sites per block, each separated by at least 100 m. Traps were installed at two heights (2 and 10 m), attached to the same tree using an extension cable at 2 m and suspended from branches with nylon ropes at 10 m (Ribeiro et al., 2021; Soares et al., 2023, 2024a).
Traps were baited with putrescine and ammonium acetate, food-based attractants commonly used to capture fruit flies (Thomas et al., 2001; Soares et al., 2023, 2024a; M.C. Oliveira-Filho et al. 2025;2026), as the sampling targeted fruit flies of Anastrepha Schiner, 1868 (Diptera: Tephritidae). Besides the food attractant, which was replaced every three months, each trap contained 250 mL of propylene glycol to preserve the specimens. During inspections, traps were lowered, and all sampled material was transferred to labeled plastic containers with the respective trap information. Specimens were collected every 15 days, following the annual week calendar adapted from Silveira Neto et al. (1976).
The material was taken to the Insect Taxonomy Laboratory at the “Luiz de Queiroz” College of Agriculture (ESALQ/USP) for sorting. Flies were transferred to 5 mL tubes, with a maximum of five specimens per tube, containing 99% ethanol for subsequent taxonomic identification. Ethanol was replaced after three days to prevent dilution by body fluids (Savaris and Lampert, 2023). Each tube was labeled with collection data.
Richardiids were identified using the key by Wendt and Ale-Rocha (2016) and original species descriptions (Curran, 1934; Hennig, 1937), based on external morphological characters and the genitalia of males and females. Twenty specimens (8♂,12♀) were deposited in the Museu de Entomologia “Luiz de Queiroz” – MELQ (ESALQENT0002337-57), Department of Entomology and Acarology, “Luiz de Queiroz” Campus, ESALQ/USP, Piracicaba, São Paulo, Brazil.
2.3. Statistics
To compare the abundance of Richardiidae species across different sites on the campus, generalized linear models (GLMs) assuming a Poisson distribution were used. The same approach was applied to evaluate whether species abundance varied significantly throughout the sampling period (April 2019 to April 2020). When necessary, models were adjusted for overdispersion using quasi-Poisson distributions. The significance of the effects of area and sampling period on richardiid abundance was assessed by analysis of deviance with F- or Chi-square tests (α = 0.05). Model fit was evaluated using probability envelopes based on half-normal plots generated with the hnp package (Moral et al., 2017). Pairwise comparisons among sampling areas were performed with Tukey’s test, applying a Bonferroni adjustment. All analyses were conducted in R (R Core Team, 2025).
3. Results
A total of 120 richardiids were collected across four areas of the “Luiz de Queiroz” Campus, identified as Richardia teevani Curran, 1934 (n = 27) (Figure 2a–b) and Richardiodes trimaculata Hennig, 1937 (n = 93) (Figure 2 c–d). Only these two species of Richardiidae were collected, with Ri. trimaculata representing the first record for the state of São Paulo. The areas with the highest specimen abundance were Piracicamirim Riverbank and Monte Olimpo, with a total of 103 individuals collected (Table 1). The abundance of Ri. trimaculata differed significantly among areas (F = 11.67, df = 3, p < 0.001), with the highest number of individuals at the Piracicamirim Riverbank, followed by Monte Olimpo (Table 1). In contrast, R. teevani showed similar abundance across all studied areas (χ2 = 2.36, df = 3, p = 0.49) (Table 1).
Lateral habitus: (a) Richardia teevani Curran, 1934; (c) Richardiodes trimaculata Hennig, 1937. Wing pattern: (b) R. teevani; (d) Ri. trimaculata. (Scale bars = 1 mm).
Richardiidae species captured in four areas on the "Luiz de Queiroz" Campus, Piracicaba, São Paulo, Brazil, from April 2019 to April 2020. Different letters within the same row indicate significant differences (p<0.05).
In terms of population fluctuations, both species showed significant variation over the collection period (R. teevani: χ2 = 31.71, df = 12, p = 0.001; Ri. trimaculata: F = 5.83, df = 12, p<0.001). Richardiodes trimaculata exhibited a remarkable abundance peak in July and August dry season (winter) followed by a decline from September, with abundance remaining low from October 2019 to April 2020 (Figure 3a). Richardia teevani also showed increases in abundance during the dry season, in May, July, August, reaching its highest level in September, low abundance was observed from November to April 2020 (Figure 3b).
Population fluctuation at four sites on the "Luiz de Queiroz". Campus, Piracicaba, São Paulo, Brazil, from April 2019 to April 2020. (a) Richardiodes trimaculata Hennig, 1937; (b) Richardia teevani Curran, 1934.
4. Discussion
Multilure traps baited with putrescine and ammonium acetate captured two Richardiidae species, indicating low diversity of these richardiid flies in the anthropized areas of the “Luiz de Queiroz” Campus, Piracicaba, São Paulo. A higher number of Ri. trimaculata individuals was collected in less disturbed areas, such as Monte Olimpo and the Piracicamirim Riverbank. In contrast, R. teevani was recorded in low numbers across all studied areas and did not differ significantly among anthropized sites. These findings suggest that Ri. trimaculata is more generalist in its resource use and more tolerant of variation in microclimatic conditions and habitat quality, with less disturbed environments being more favorable to this species. However, there is no supporting evidence in the literature regarding its ecological aspects. In contrast, the low abundance of R. teevani across the studied fragments may reflect naturally low population densities, limited attraction to the sampling method, or ecological factors not evaluated in this study, rather than a clear effect of habitat fragmentation.
The higher abundance of Ri. trimaculata in less anthropized areas (Piracicamirim Riverbank and Monte Olimpo) compared to more disturbed sites (Areão Farm and Central Area) likely reflects the impacts of structural simplification in fragmented habitats, which can reduce habitat quality and resource availability (Ribeiro et al., 2009). Such patterns are consistent with functional declines reported in fragmented landscapes, where populations may persist at lower densities and face constraints on maintaining ecological interactions (Tabarelli et al., 2010). Conversely, R. teevani was consistently recorded at low abundance across all areas, and no clear spatial pattern related to environmental disturbance was detected for this species. The influence of trap height could not be evaluated due to the low number of specimens collected for both species.
Our results are consistent with patterns reported for Anastrepha species in the same areas, where some specialist species were more abundant in less anthropized environments (Soares et al., 2024a). However, generalist species, such as Anastrepha fraterculus (Wiedemann, 1830) and Anastrepha obliqua (Macquart, 1835) persisted at high abundance even in more disturbed environments on the “Luiz de Queiroz” Campus. It is important to highlight that both fruit fly species are highly polyphagous and exploit a wide range of host plants (Zucchi and Moraes, 2026). As fruit fly pests, they may exhibit considerable phenotypic plasticity, potentially enhancing their adaptability to environmental disturbances (Silva and Elliot, 2016).
The population fluctuations of R. teevani and Ri. trimaculata showed similar seasonal patterns, with peaks occurring between July and September, during the dry season (winter). This result is noteworthy, as it contradicts the expectation that saprophagous organisms, such as Richardiidae species (Hancock, 2010), are more prevalent during rainy periods. Based on Talora and Morellato (2000), fruiting phenology on the “Luiz de Queiroz” Campus is likely more evenly distributed throughout the year, including the winter season. Thus, even during the dry season, richardiid larvae may feed on decomposing fruits on the ground, ensuring a continuous supply of resources. Additionally, the areas where more specimens were collected are located near moist environments, such as the Piracicamirim Riverbank and a lake at Monte Olimpo, which may support the development of immature stages and adult activity by providing resources even during periods of low rainfall in the region, usually from May to September.
Richardia species are dominant in fallen fruits in La Selva, Costa Rica (Inouye, 1999), suggesting a strong dependence on decomposing substrates for oviposition and larval development. This pattern suggests that local dynamics of fruit phenology and subsequent decomposition may be key factors regulating populations of richardiid flies. However, the occurrence of population peaks during the dry season may also reflect reduced competition or predation, or the possibility that lower relative humidity limits the activity of decomposer fungi (Logan et al., 2021), thereby increasing the persistence of fallen fruits and potentially enhancing their suitability for the development of these two richardiid species.
Although Multilure traps are widely used to capture fruit flies (Tephritidae) (Thomas et al., 2001; Soares et al., 2023, 2024a; M.C. Oliveira-Filho et al. 2025; 2026), their application to other insect groups remains poorly documented. However, Multilure traps baited with food attractants have been successfully used to collect green lacewings (Neuroptera: Chrysopidae) on the “Luiz de Queiroz” Campus (Soares et al., 2024b). Our results further demonstrate that this method is also effective for sampling richardiid species across different habitats with different anthropization levels.
This study expands the known geographic distribution of Ri. trimaculata in Brazil. Previously, this species had been recorded in the Brazilian Amazon (Amazonas) and in Peru (Rio Urubamba – type locality) (Hennig, 1937; Wendt and Ale-Rocha, 2016). In contrast, the distribution of R. teevani remains poorly documented, being known only from its original description, which reports its occurrence in Brazil (unknown locality), Guyana, Panama, Suriname, and Trinidad (Curran, 1934; Ale-Rocha and Barros, 2026). Therefore, the present study provides new distributional data for Ri. trimaculata, representing the first record for the state of São Paulo.
The low number of R. teevani specimens may reflect limitations of the trapping method or lower tolerance of disturbed areas. Alternatively, R. teevani may occur at a low population density on the campus; nevertheless, its capture indicates that Multilure traps baited with putrescine and ammonium acetate are suitable for sampling richardiids, even at low population levels. This pattern is consistent with surveys of Anastrepha fruit flies, in which a few species are collected in large quantities, whereas most are represented by only a few specimens (Soares et al., 2024a).
Future studies could combine additional approaches, such as direct collection from natural substrates, to better understand the diversity and behavior of these insects, as well as test other trap types for collecting richardiid flies.
The ecological aspects of Richardiidae species remain poorly known, largely due to the limited number of ecological studies conducted in both forest fragments and well-preserved areas. In this study, we present data on two Richardiidae species sampled across four sites with different levels of anthropization, showing that their abundance patterns differ. Richardiodes trimaculata occurred in less disturbed areas, whereas R. teevani showed a relatively uniform abundance across all sites. Additionally, our results indicate that Multilure traps baited with synthetic food-based attractants can be useful for sampling richardiid flies. Therefore, these traps could be used in combination with other sampling methods for collecting richardiids.
Acknowledgements
The authors thank the Graduate Program in Entomology of the “Luiz de Queiroz” College of Agriculture, University of São Paulo (ESALQ/USP). The senior author was supported by a scholarship from the Coordination for the Improvement of Higher Education Personnel Brazil (CAPES – Finance Code - 001). The authors thank José Gabriel Gonçalves Ribeiro for the help during the collections. MS acknowledges financial support from the National Council for Scientific and Technological Development (CNPq). RAZ is a research fellow of CNPq. The specimens collected during this project were obtained through a collection license by the "Sistema de Autorização e Informação em Biodiversidade – SISBIO", number: 61549-7.
Data statement
The entire dataset supporting the results of this study was published in the article itself.
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