Taxonomic Catalog of the Brazilian Fauna: Bibionomorpha (Diptera) diversity and distribution

ABSTRACT

We provide an overview of the taxonomic history and systematics of Bibionomorpha families with occurrences in Brazil, focusing on Anisopodidae, Bibionidae, Cecidomyiidae, Diadocidiidae, Ditomyiidae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, Rangomaramidae, and Sciaridae. The study evaluated the Linnean and Wallacean shortfalls in the Brazilian fauna using the Taxonomic Catalog of the Fauna of Brazil (CTFB) database, until December, 2023. The analysis included species diversity and distribution patterns across states and biomes. Our results showed varied levels of taxonomic knowledge and distribution across families, highlighting the need for precise policies in terms of science funding and priorities. The study underscores the importance of ongoing projects, such as CTFB, to consolidate and disseminate taxonomic information for the next generations of researchers and biogeographers interested in the Brazilian and Neotropical fauna.

KEY WORDS:
Biodiversity; Brazilian biomes; distribution; Linnean shortfall; taxonomy; Wallacean shortfall

INTRODUCTION

Flies are insects with a pair of functional wings, with the hind wings modified into halters (e.g., de Carvalho et al. 2012, 2024). In terms of diversity, Diptera is a megadiverse clade. It has been considered either the second or third most speciose group of insects, after Coleoptera and Hymenoptera, until recently Hebert et al. (2016) suggested 1.8 million species of Cecidomyiidae alone. Indeed, in almost all recent studies (Hebert et al. 2016, Srivathsan et al. 2023) Cecidomyiidae, Sciaridae, and Mycetophilidae-all three being Bibionomorpha families-fit among the top 10 or top 20 most species-rich families of insects in traps in different parts of the planet. All three families fit into the definition of dark taxa by Hartop et al. (2022), corresponding to clades with more than 1,000 described species worldwide and with an undescribed fauna at least 10 times larger. D.S. Amorim et al. (unpublished data) and R. Meier et al. (unpublished data), for example, addressed the entire fauna of Mycetophilidae of Singapore alone, increasing the number of mycetophilid’ species of the Oriental region by 24 times. The number of extant species of Bibionomorpha has been estimated at 15,000 (Pape et al. 2009), but this number could be well over 1,000,000 (Hebert et al. 2016). The Brazilian fauna of Bibionomorpha has the same general pattern as seen in other parts of the world, with Mycetophilidae and Cecidomyiidae among the top 10 or 20 most species-rich families of Diptera in recent surveys in the country (Amorim et al. 2022, Riccardi et al. 2022, Lamas et al. 2023).

Bibionomorpha is one of the eight suborders of Diptera (in the sense of Amorim and Yeates 2006) and in a broader sense includes the extant families Anisopodidae, Bibionidae, Bolitophilidae, Cecidomyiidae, Diadocidiidae, Ditomyiidae, Keroplatidae, Lygistorrhinidae, Mycetophilidae, Pachyneuridae, Rangomaramidae, and Sciaridae. This set of families composes a clade (Amorim and Rindal 2007, Wiegmann and Yeates 2017, de Carvalho et al. 2024, Brazilian Zoology Group 2024), but there are issues with the position of some few groups. Pachyneura Zetterstedt, 1838, with forked R₂₊₃ and an unforked R₄₊₅, is sometimes included in the same family of Cramptonomyia Alexander, 1931 with an unforked R₂₊₃ and R₄₊₅, but belongs to a separate clade (Amorim 1993). All bibionomorph families have species known from Brazil, except Bolitophilidae and Pachyneuridae, being 944 species known so far (de Carvalho et al. 2024). There is also a question of the position of the Scatopsoidea: Wood and Borkent (1989), Amorim (1993), Amorim and Grimaldi (2006), and Amorim and Rindal (2007) keep the group (including Valeseguyidae, Canthyloscelidae, and Scatopsidae) close to the psychodomorph families based on features of the larval mandible and the fusion of the meron to the thoracic pleura, while Wiegmann et al. (2011), and Wiegmann and Yeates (2017), and Ševčík et al. (2016), using molecular data, suggest that the scatopsoids would belong to the Bibionomorpha.

There are some issues about the classification and internal relationships in the Bibionomorpha. In terms of rank, “Hesperinidae” is sometimes accepted as a family separate from the Bibionidae. More recently, the Lygistorrhinidae has been seen as a subclade of Keroplatidae (Ševčík et al. 2016), based on molecular data, in a study including four species of lygistorrhinids and six species of keroplatids. Lygistorrhinids indeed may represent a case of “morphology long branch attraction”, and their traditional position as the sister group to Mycetophilidae (Hennig 1973) could be misleading. However, morphological evidence to place lygistorrhinids as a subbranch of Keroplatidae remains scarce. Additional characters and wider taxon sampling would benefit reconstructions of the phylogeny of Bibionomorpha based respectively on morphological and molecular data. At this stage, we follow a more conservative position, keeping familial rank, as in Blagoderov and Ševčík (2017), Blagoderov (2017), and Söli (2017), for the Keroplatidae, Lygistorrhinidae, and Mycetophilidae, respectively.

The book “Insetos do Brasil” (Rafael et al. 2012, 2024) was an effort to entirely compile the current knowledge about the Brazilian insect fauna, now a reference for students interested in the Brazilian entomofauna. A second ambitious project is the Taxonomic Catalog of the Brazilian Fauna (Catálogo Taxonômico da Fauna do Brasil - CTFB), which also had the Diptera led by José Albertino Rafael and involves many Brazilian taxonomists. The CTFB project started in 2015 and has the participation of over 500 experts in the taxonomy of different animal groups (Santos et al. 2020, see also Boeger et al. 2024). The platform is available online (http://fauna.jbrj.gov.br/) and provides information on the taxonomic status of the name, taxonomic hierarchy, whether the taxa are native to Brazil, its geographic distribution in Brazilian territory, and references documenting the taxon in Brazil (Santos et al. 2020).

Hortal et al. (2015) discussed seven key shortfalls of current biodiversity data and we highlighted the gaps in species taxonomy (Linnean shortfall), species distribution (Wallacean shortfall), and evolutionary patterns (Darwinian shortfall). Concerning Diptera in general and Bibionomorpha in particular, these knowledge gaps are evident. In this study, we used the CTFB Bibionomorpha database (Falaschi et al. 2024) to address the Linnean and Wallacean shortfalls in the Brazilian fauna. Specifically, we aimed to determine how many species are known from Brazil and to estimate the total number of Bibionomorpha species in the country, thus revealing the extent of the Linnean shortfall. Additionally, we aimed to provide a comprehensive view of the geographical distribution of the study groups, highlighting the Wallacean shortfall. This information will be valuable for the next generations of taxonomists and biogeographers studying the Brazilian and Neotropical fauna of these groups.

MATERIAL AND METHODS

We deal with the Bibionomorpha families known from Brazil individually, briefly describing their taxonomic improvements in the last decades, as well as perceptions of Linnean and Wallacean shortfalls.

The primary taxonomic data for Bibionomorpha species in the CTFB mostly come from the Neotropical catalogs of its families, until December of 2023. Additional sources include more recently published papers with revisions, checklists, and inventories. All searches on the CTFB were made considering the neotropical representants of the families Anisopodidae, Bibionidae, Cecidomyiidae, Diadocidiidae, Ditomyiidae, Keroplatidae, Mycetophilidae, Lygistorrhinidae, Rangomaramidae, and Sciaridae.

To understand both the Linnean and Wallacean shortfalls in Brazil, an overall analysis of the data was performed in two steps: 1) determining the number of species within each family in Brazil; 2) assessing the geographical distribution of each family.

For the first analysis, we entered the family name in the CTFB search area to understand the diversity of each family in Brazil. The second analysis aimed to compare Brazilian diversity with Neotropical diversity using catalogs of each family (see in Results, each family brief description). The third step involved understanding the geographical distributional data of these families in Brazil. Following Santos et al. (2020), we organize the data in two different datasets: political division in states, including the Federal District, and biomes as defined by the Instituto Brasileiro de Geografia e Estatística (IBGE 2004, 2020). To conduct this step, we accessed the CTFB page for each family and downloaded the distributional map of each family in Brazil. We then searched by biome for each family and compiled all the data to produce the maps presented herein.

Maps (Fig. 1A-J) with the number of species of Bibionomorpha and respective families (except Sciaridae) described or reported to occur are provided for the 26 states plus the Federal District. The software used for the maps is Quantum Gis, version 3.22.11 with data from the CTFB database and from IBGE (2004, 2020).

Species accumulation curves (Fig. 3A-K) were done for all Bibionomorpha and each family, by plotting the cumulative number of described species against the year of their descriptions.

RESULTS

Brief description of taxonomy history and systematics of Bibionomorpha families with occurrence in Brazil

Anisopodidae

This family encompasses eight extant genera worldwide, with over 160 described species (Hancock and Amorim 2009, Hancock 2017, González et al. 2019). The Neotropical fauna of Anisopodidae is composed of 65 described species within six genera: Carreraia Corrêa, 1947, Sylvicola Harris, 1780, Olbiogaster Osten Sacken, 1886, Lobogaster Philippi, 1865, and Mycetobia Meigen, 1818. Among these genera, Carreraia (one species), Olbiogaster (15 species), and Sylvicola (seven species) have been recorded for Brazil (Falaschi et al. 2016b, Falaschi and Oliveira 2024).

Anisopodids are often called ‘window gnats’ and are associated with humid habitats such as forests. Their immatures are saprophagous and are often found in decaying or fermenting organic matter, primarily of plant origin. However, some species may exploit animal carcasses or dung (Hancock 2017).

Bibionidae

A family with approximately 700 extant species distributed in eight genera (Pinto and Amorim 2000, Fitzgerald 2004, Fitzgerald et al. 2020). In the Neotropical region, 192 species distributed in six genera are known (Fitzgerald 1997, 2004, 2021a, 2021b). The South American and the worldwide fauna were inventoried in Hunter’s (1900) and Kertész’s (1902) catalogs, respectively. Hardy (1953) inventoried the fauna of Argentina and Hardy’s catalogs (1959, 1966) covered the Neotropical fauna. More recently, the fauna of Nicaragua, Mexico, Guatemala, Colombia, and Chile were addressed respectively by Maes (1990), Fitzgerald (2000, 2023), Falaschi et al. (2016a), and Fitzgerald et al. (2020). Brazil has 49 described species, and almost half of them (24) were described in the 1930s and 1940s (Falaschi and Schelesky-Prado 2024). Only nine species have been described since Hardy’s catalog (1966): one in the 1980s, five in the 1990s, and only three from the 2000s onwards (Falaschi and Schelesky-Prado 2024).

Bibionids are popularly known as march flies because of their emergence in the spring in the northern hemisphere. They are also referred to as lovebugs because of their habit of flying in copula, especially members of Plecia Wiedemann, 1828 (Fitzgerald 2004). The immature stages and biology of Neotropical Bibionidae are poorly known (Fitzgerald 2009). Immature stages of Plecia cf. collaris, P. nearctica Hardy, 1940, and P. plagiata (Wiedemann, 1824) are known, described (Kuitert 1975, Pinto and Amorim 1996). Additionally, the reproductive behavior of Dilophus sayi (Hardy, 1959) and P. nearctica were discussed (Thornhill 1976, Matthews and Matthews 1978). Lastly, the pollination behavior of D. espeletiae Sturm, 1990 was discussed (Sturm 1990). Bibionidae is considered a monophyletic group by most authors, but the inclusion of the Holarctic genus Hesperinus Walker, 1848 in the family is still controversial (Pinto and Amorim 2000, Fitzgerald 2004, Papp 2010).

Cecidomyiidae

Cecidomyiidae is one of the most speciose families of Diptera, with more than 6,600 species described in 832 genera (Gagné and Jaschhof 2021). Part of the cecidomyiids is popularly known as “gall midges”, with gall-inducer larvae. The family also includes mycetophagous, predaceous, and free-living phytophagous larvae. The family is monophyletic, and the most recent classification system divided it into six subfamilies (Gagné and Jaschhof 2021). The Nearctic and Palearctic faunas are known for more than two thousand species each, while the Neotropical fauna is still superficially known, comprising less than 10% of the species described worldwide. The first Neotropical cecidomyiid species were described at the end of the 19th century and the beginning of the 20th century. During this period, Jean Jacques Kieffer (1857-1925), Ewald Heinrich Rübsaamen (1857-1919), and Joachim da Silva Tavares (1866-1931) stood out for their pioneering and valuable studies. After them, taxonomic contributions there were scattered until 1959, when Edwin Möhn (1928-2008) began publishing on the fauna of El Salvador. A few years later, Raymond Gagné (1935-) published the first catalog of the Neotropical Cecidomyiidae. Gagné (1994) published the “green book” which is still an essential reference for the taxonomic study of the Neotropical Cecidomyiidae. In the 1990s, Brazilian taxonomists began to actively describe species and deal with the diversity of the family in the country. According to Maia (2021), the family is presently represented by 293 species of 105 genera in Brazil (Carmo-Neto et al. 2021, Maia 2022a, 2022b, 2022c, Garcia et al. 2023a, 2023b, 2023c, 2023d).

Diadocidiidae

This family includes two genera with 29 extant described species worldwide. Of these, 27 species belong to the extant genus Diadocidia Ruthe, 1831. There are only two species assigned to a fossil genus from the Cretaceous amber from Katchin, Myanmar, known as Docidiadia Blagoderov and Grimaldi, 2004 (Blagoderov and Grimaldi 2004, Amorim and Brown 2022), but the association of the genus to the Diadocidiidae still demands corroboration. In the Neotropical region, there are three described species of Diadocidia (Bechev and Chandler 2011), and only one known from Brazil (Falaschi 2024a). Papavero (1977a) mentioned a second undescribed species from Chile, Vockeroth (2009) referred to an undescribed species from Mexico, and Falaschi (2016a) reported the genus in Colombia.

The biology of these gnats is poorly known; the limited knowledge available suggests that they are common in forest habitats (Bechev and Chandler 2011). The larvae of these gnats are associated with decaying wood and develop within a mucous tube under rotting logs. They primarily feed on the hymenium of higher fungi (Hutson et al. 1980, Yakovlev 1994, Zaitzev 1994).

Ditomyiidae

The ditomyiids constitute a relatively small family of fungus gnats, with over 100 described species in 10 genera in all regions, except from the Afrotropical Region (Vockeroth 2009). In the Neotropical Region, 35 species have been described in six genera (Falaschi 2016b). The known fauna from Brazil includes 11 described species in four genera: Australosymmerus Freeman, 1954-in the subgenus A. (Melosymmerus) Munroe, 1974 (seven species)-, Calliceratomyia Lane, 1946 (one species), Nervijuncta Marshall, 1896 (one species), and Rhipidita Edwards, 1940 (two species) (Falaschi 2024b), with a good number of undescribed species of the genera Rhipidita and Calliceratomyia available in collections, and an undescribed genus closely related to these two genera. These fungus gnats are typically associated with moist environments, such as decaying wood covered with moss and fungi. The immature stages live in galleries within rotting wood or on Polyporaceae fungi (Munroe 1974).

Keroplatidae

This family comprises about 1,000 species distributed in more than 100 extant and fossil genera (Mantič et al. 2020). The family has a wide biogeographic distribution, with most described species from the Palaearctic and Neotropical regions (Evenhuis 2006). In the Neotropical region, there are more than 30 genera and over 200 species, 90 of which are known from Brazil (Evenhuis 2006, Falaschi 2024c).

The knowledge about the biology of the group is still poor, and immature stages are unknown for most genera. The larval stage of some genera can be predatory, feeding on small invertebrates, while others are mycophagous, feeding on fungal spores (Skuse 1888, Matile 1997, Evenhuis 2006, Falaschi et al. 2019a). Adults are often found in damp places with low light, such as the entrance to caves and cavities of rotten trunks near fungi or rocks (Matile 1997, Evenhuis 2006, Falaschi 2014, 2016c). One species in Brazil, Neoceroplatus betaryensis Falaschi, Johnson & Stevani, 2019 is known to have blue bioluminescence (Falaschi et al. 2019a, 2019b).

Lygistorrhinidae

The Neotropical Lygistorrhinidae have 14 described species known in the genus Lygistorrhina Skuse, 1890 mainly from Brazil, French Guiana, and Mexico (Oliveira and Amorim 2012a, Grund 2012, Huerta et al. 2019, Blagoderov and Pollet 2020). Globally, there are 51 species of Lygistorrhinidae in 16 genera, of which 16 are known as fossils in nine extinct genera and 35 are extant species in eight genera (Blagoderov and Pollet 2020). The first Brazilian species were described by Edwards (L. brasiliensis Edwards, 1932) and Lane (L. barretoi Lane, 1947 and L. edwardsi Lane, 1947) from the states of Rio de Janeiro, Goiás, and São Paulo, respectively. Other Neotropical species are known from Mexico, Trinidad, St. Vincent, and French Guiana (Oliveira and Amorim 2012a, Grund 2012, Huerta et al. 2019, Blagoderov and Pollet 2020). The first version of the Neotropical catalog of the family (Papavero 1977b) had seven valid species, while the second version (Oliveira and Amorim 2012a) recorded nine species-with an increase of five species in the last decade (Grund 2012, Huerta et al. 2019, Blagoderov and Pollet 2020).

Although their biology is poorly known, the lygistorrhinids are flower visitors, with elongated mouthparts being used for feeding on nectar and pollen being recorded attached to their abdomen (Bertone 2018). The immature stages and larval habitat are unknown so far (Oliveira and Amorim 2012a, Blagoderov and Pollet 2020). Lygistorrhinidae is clearly monophyletic and its position found recent controversies in the literature (see above). A phylogenetic study of the genus Lygistorrhina is still pending, and the subgenus L. (Lygistorrhina) Skuse, 1890 may not represent a monophyletic group, L. (Probolaeus) Williston, 1896 probably being a small subclade within the genus (Oliveira and Amorim 2012a). For the time being, we abandon L. (Probolaeus) as a subgenus.

Mycetophilidae

The Mycetophilidae at present have 1,219 Neotropical described species distributed in 56 genera (Oliveira and Amorim 2014b, 2014c, 2021, Amaral et al. 2023). A new Neotropical species was recently described by Afiune and Oliveira (2024), but not included in the database herein studied, totalizing 1,220 species in neotropics. Mycetophilids are widely distributed in the Neotropical Region. Recent inventories documented the presence of genera in different areas, widening their distribution in South America (e.g., Amorim et al. 2022, Riccardi et al. 2022, Lamas et al. 2023). The current number of known Neotropical species (Oliveira 2024b) in relation to the first version of Neotropical (Papavero 1978) shows an increase in the number of species from 749 to 1,219, 63% in 45 years.

Mycetophilids are part of the fungus gnats and their life cycle has mostly larvae feeding on fungi. The biology of the Neotropical species of the family began to be documented only recently (Oliveira et al. 2015, Passacq et al. 2017, Amaral et al. 2022a, 2022b, 2023). There is no question about the monophyly of the Mycetophilidae if we disregard a few mesosciophilid-like genera, such as Freemanomyia Jaschhof. The most recent classification system within the family was proposed by Oliveira and Amorim (2021).

Rangomaramidae

This family comprises 20 Neotropical species distributed in six genera (Amorim and Falaschi 2012). The Neotropical species fit in two of the four main rangomaramid subclades: the Chiletrichinae and the Ohakuneinae. The Chiletrichinae genus Chiletricha Chandler, 2002 is known from Chile, Argentina, and Chiletricha marginata (Edwards, 1940) from southern Brazil (Amorim and Falaschi 2012); Eratomyia Amorim & Rindal, 2007 is known from Ecuador and Colombia. The Ohakuneinae genus Ohakunea Tonnoir & Edwards, 1927 is known in South America from southern Chile and Argentina, but there is an undescribed species of the genus known from southern Brazil (D.S. Amorim unpublished data); the genus Colonomyia Colless, 1963 has species known from Argentina and southern Chile, with C. brasiliana Amorim and Rindal, 2007, known from the states of São Paulo (Amorim and Falaschi 2012) and Mato Grosso (unpublished data); the genus Rogambara Jaschhof, 2005 is known from Costa Rica and Cabamofa Jaschhof, 2005 is known from Costa Rica and Panama (Jaschhof 2004), but have not been recorded from Brazil.

The biology of rangomaramids remains largely unknown, and there is a lack of information regarding their immature stages (Amorim and Falaschi 2012). The evolutionary relationships among rangomaramids are still a topic of discussion. The connection of the Rangomarama Jaschhof & Didham, 2002, the Ohakuneinae, the Chiletrichinae and the Heterotrichinae in a clade was one of the most parsimonious solutions for the information available in Amorim and Rindal (2007), but a more extensive analysis of the relationships among Sciaroidea families is desirable. The monophyly of each of these four subfamilies of Rangomaramidae is well-founded.

Sciaridae

The family Sciaridae includes over 2,700 described species worldwide in more than 80 genera. The actual number of species known, however, is far below the actual diversity of the family. The BIN count for Sciaridae found by Hebert et al. (2016) in the Nearctic region is nearly 75 times the number of described species in the region; Srivathsan et al. (2023) studied the species composition of 39 Malaise trap samples in five biogeographic regions, eight countries, and numerous habitats, and found sciarids as the sixth most speciose in a set of 20 insect families that account for over half of the species diversity. There are almost 200 described Neotropical species of Sciaridae (Amorim 1992), of which 91 are known for Brazil. The concept of the many genera of sciarids has changed over the last 40 years (see, e.g., Mohrig and Menzel 2014) and most old keys for the genera of the family are not adequate for the Neotropical fauna. For the time being, the key in Mohrig and Menzel (2009) is the most adequate to identify the Brazilian fauna of Sciaridae.

Brief perceptions of Linnean and Wallacean shortfalls in the Brazilian fauna

An overall view of the species-richness of the Bibionomorpha families in Brazil is summarized in Table 1 and a comparison between the Brazilian known diversity and the Neotropical general diversity of these families is summarized in Table 2. The distributional data of these families resulted in maps based on a political division by states, including the Federal District (Fig. 1A-K) and on biomes for Bibionomorpha, in general, and Bibionidae, Cecidomyiidae, Lygistorrhinidae, Mycetophilidae, and Sciaridae (Fig. 2A-F).

Figure 1
Maps of Brazil in South America showing the 26 states plus the Federal District: (A) Bibionomorpha. (B) Anisopodidae, (C) Bibionidae, (D) Cecidomyiidae, (E) Diadocidiidae, (F) Ditomyiidae, (G) Keroplatidae, (H) Lygistorrhinidae, (I) Mycetophilidae, (J) Rangomaramidae, (K) Sciaridae. For each political region, the number of species described or reported to occur is provided for Bibionomorpha and its respective families. The north region is represented by the green color, the northeast region by the orange color, the southeast region by the blue color, the south region by the pink color, and the midwest region by the red color.

Figure 2
Maps of Brazil in South America showing the six biomes: (A) Bibionomorpha, (B) Bibionidae, (C) Cecidomyiidae, (D) Lygistorrhinidae, (E) Mycetophilidae, (F) Sciaridae. For each biome, the respective number of species described or reported to occur is provided. The green color represents the Amazonic Forest (tropical rainforest), the red color the Caatinga (semi-arid tropical vegetation), the purple color the Atlantic Forest (tropical forest), the orange color the Pampa (low grasslands), the yellow color the Cerrado (the Brazilian savanna), and the blue color the Pantanal (tropical flooded grasslands).

Table 1
Family diversity in Brazil according to CTFB.

Family	Species recorded for Brazil	Endemic species*	Genera recorded for Brazil	CTFB authorship
Anisopodidae	23	20	3	Falaschi and Oliveira 2024
Bibionidae	49	33	6	Falaschi and Schelesky-Prado 2024
Cecidomyiidae	293	80	105	Maia 2023
Diadocidiidae	1	1	1	Falaschi 2024a
Ditomyiidae	11	10	4	Falaschi 2024b
Keroplatidae	90	56	24	Falaschi 2024c
Lygistorrhinidae	5	4	1	Oliveira 2024a
Mycetophilidae	431	306	41	Oliveira 2024b
Rangomaramidae	2	2	2	Falaschi 2024d
Sciaridae	91	60	17	Amorim 2024

• * “Endemic species” corresponds to the nomenclature used in the CTFB platform indicating species so far only recorded for Brazil.

Table 2
Diversity of Bibionomorpha families in Brazil in comparison with the known Neotropical fauna.

Family	Species known in Brazil	Neotropical species	Percentage of species in Brazil compared with Neotropical diversity (%)
Anisopodidae	23	65	35.0
Bibionidae	49	192	25.5
Cecidomyiidae	293	630	46.5
Diadocidiidae	1	3	33.0
Ditomyiidae	11	35	31.0
Keroplatidae	90	200	45.0
Lygistorrhinidae	5	9	55.0
Mycetophilidae	431	1219	35.0
Rangomaramidae	2	20	10.0
Sciaridae	91	200	45.5

DISCUSSION

The Brazilian diversity of Bibionomorpha was mainly described by John Lane (1905-1963), Frederick Wallace Edwards (1888-1940), and José Pedro Duret (1913-2007), with a relevant contribution also from Edward Irving Coher (1920-2023) and Loïc Matile (1938-2000), added by the recent taxonomic works done, mainly, by Dalton de Souza Amorim (1958-), Valéria Cid Maia (1965-), Maria Virginia Urso-Guimarães (1970-), Rafaela Lopes Falaschi (1982-), and Sarah Siqueira de Oliveira (1984-).

Most of the species described by Edwards are from the district of Nova Teutônia, municipality of Seara, in the state of Santa Catarina (e.g., Edwards 1940a). These were collected by Fritz Plaumann in temperate areas of the Atlantic Forest, in southern Brazil (for more details, refer to Amaral et al. 2022a). Edwards also described species from the Pantanal (e.g., Edwards 1941) and Cerrado biomes (e.g., Edwards 1933, 1940b). John Lane mostly described species from the Atlantic Forest biome. He also described species from the Cerrado (e.g., Lane 1950), and from the Amazon Forest (e.g., Lane 1958, 1959).

Brazilian taxonomists have been involved in describing species from Brazil and other South American countries (e.g., Peru, Chile, Colombia), playing a role in enhancing the knowledge of the fauna of the region, particularly on Mycetophilidae and Cecidomyiidae. Oliveira and Amorim have so far 74 described mycetophilid species from the Neotropics (e.g., Amorim and Oliveira 2008, Oliveira and Amorim 2010a, 2010b, 2011, 2012b, Amorim et al. 2018), Maia described 126 cecidomyiid species, mostly from Brazil, but also from Peru and Chile (e.g., Maia and Vasquez 2011, Maia and Villagra 2017). They also took part in other Diptera inventories across South America (Amorim et al. 2002, Falaschi and Amorim 2014a, 2014b, Amorim and Falaschi 2014, Oliveira and Amorim 2014a, 2014b, 2014c, Amorim et al. 2016, Falaschi 2016a, 2016b, 2016c, Falaschi and Amorim 2016, Falaschi et al. 2016a, 2016b, Oliveira and Amorim 2016a, 2016b, Oliveira et al. 2017, González et al. 2019, Amorim et al. 2022, Riccardi et al. 2022, Lamas et al. 2023). This includes organizing the taxonomic information for various families (Amorim and Falaschi 2012, Oliveira and Amorim 2012a, 2014c, Oliveira et al. 2017), in continuation of the effort by Dr. Nelson Papavero began in the 1960s.

Nelson Papavero (1942-) stands out as the most influential Brazilian Dipterist. His contributions encompass species descriptions, organizing the Neotropical catalog of flies, catalogs for numerous particular Diptera families, curation of the insect collection in the Museu de Zoologia in São Paulo, and a vast body of work covering the history of dipterology, systematics, biogeography, and zoology. According to Klassa and Santos (2012, 2014), Papavero made groundbreaking contributions to science by publishing across multiple domains, concurrently with his involvement in graduate courses and mentorship of teachers and students. The present status of Brazilian Zoology worldwide (see e.g., Rafael et al. 2009 and https://www.scimagojr.com/countryrank.php?area=1100&category=1103) is largely the consequence of Papavero’s enduring contributions over the last decades.

As mentioned earlier, Mycetophilidae has been the most prolific family in the last years in Neotropics in terms of species descriptions, particularly in the genus Manota Williston, 1896. In the Neotropical catalog of the family (Papavero 1978), three species of Manota were recorded; in 2023, 92 species were recorded (Kurina et al. 2018), representing a 3,000% increase in the fauna. Regarding the Brazilian fauna, only two Manota species were previously known (Papavero 1978, Oliveira and Amorim 2014c) while in 2023 there are records of 34 additional species in Brazil (Kurina et al. 2017, 2018), an increase of 1,800% in the known fauna. There are additional unidentified Manota specimens from Roraima (Riccardi et al. 2022) and Manaus (Amorim et al. 2022), indicating that the Linnean gap in the genus (actually in the family) is far away from being healed in the country, in particular, and in the Neotropics in general, as well as the Wallacean shortfall.

A detailed view of the Bibionomorpha species accumulation curves (Fig. 3A-K) shows that bibionomorphs (Fig. 3A), in general, and Bibionidae (Fig. 3C), Cecidomyiidae (Fig. 3D), Keroplatidae (Fig. 3G), and Mycetophilidae (Fig. 3I), in particular, still not achieve the total of taxonomic knowledge in Brazil. Indeed, we expect thousands of species of each of these families to be described in the future. The species accumulation curves of Anisopodidae (Fig. 3B), Diadocidiidae (Fig. 3E), Ditomyiidae (Fig. 3F), Lygistorrhinidae (Fig. 3H), Rangomaramidae (Fig. 3J), and Sciaridae (Fig. 3K), shows some stability, which is evidence of lack of specialists and studies, not lack of undescribed species.

An analysis of the distributional maps (Fig. 1A-K) shows that bibionomorphans are known from all the Brazilian states, with major records from the states of São Paulo (397), Santa Catarina (293), and Rio de Janeiro (235) (Fig. 1A), primarily under the Atlantic Forest biome (Fig. 2A-F). This is the bias caused by the first taxonomists describing species from Brazil, working with material from these states. The only Bibionomorpha family known from all Brazilian states and biomes is Cecidomyiidae (Figs 1D, 2C). Regarding other dark taxa in Bibionomorpha, Mycetophilidae is not recorded from the states: Rio Grande do Sul (in the Pampa biome), in southern Brazil; Tocantins, in northern Brazil (in the Cerrado biome), and Rio Grande do Norte, Ceará, Piauí, and Maranhão, in northeastern Brazil (in the Caatinga biome) (Figs 1I, 2E). Sciaridae is not recorded from the states of Espírito Santo (under the Atlantic Forest biome) in the southeast region of Brazil, Acre, Rondônia, Amapá, Roraima, and Tocantins in the north region (under the Amazonia and Cerrado biomes), and in the northeast region (under Caatinga and Atlantic Forest biomes), except by the state of Bahia (Figs 1K, 2F). For the remaining bibionomorph families, the Wallacean shortfalls are more evident, with distributional gaps, especially in the north, central-west, and northeast of the country, mainly under Caatinga and Cerrado biomes (Figs 1B, C, G, H, 2B, D). The families Diadocidiidae (Fig. 1E) and Rangomaramidae (Fig. 1J) are known only from the state of Santa Catarina in the south region of Brazil, and the family Ditomyiidae only from the states of Santa Catarina and Rio de Janeiro (Fig. 1F), under the Atlantic Forest biome. The diadocidiids are widespread, so they are expected in tropical areas of the country, but not very speciose. The rangomaramid genera are largely temperate, so they are not expected to be present in most of the country, except for the genus Colonomyia. The ditomyiids in Brazil include relatively species of temperate clades-especially Nervijuncta and Australosymmerus (Melosymmerus)-, but have a considerable number of undescribed species of Rhipidita and Calliceratomyia.

Figure 3
Species accumulation curves of Bibionomorpha, and respective families, of species described for Brazil by year (from 1805 to 2023): (A) Bibionomorpha, (B) Anisopodidae, (C) Bibionidae, (D) Cecidomyiidae, (E) Diadocidiidae, (F) Ditomyiidae, (G) Keroplatidae, (H) Lygistorrhinidae, (I) Mycetophilidae, (J) Rangomaramidae, (K) Sciaridae.

The community of Brazilian dipterists is working diligently to address the Linnean and Wallacean shortfalls regarding the Bibionomorpha fauna. Considering the large area of the country and the highly diverse families, the number of taxonomists is not enough to fully accomplish the challenge of describing the fauna before the wide devastation of the natural areas. Recent inventories, particularly in the Cerrado and Amazon biomes, have been conducted, along with efforts to employ new technologies (as R. Meier et al. unpublished data, and D.S. Amorim et al. unpublished data) to know the Brazilian fauna. The number of known species is expected to improve rapidly in the coming years (e.g., Afiune and Oliveira 2024).

ACKNOWLEDGMENTS

Many thanks to the CTFB Coordinators, Hussam Zaher (MZUSP), José A. Rafael (INPA), Michel P. Valim (MZUSP), and Walter A.P. Boerger (UFPR), for their dedication to managing this ambitious project. Special thanks to José A. Rafael for the coordination of the insect groups, especially Diptera. SSO was funded by CNPq 308298/2021-7 and FAPEG 202310267000883; GPSA was funded by CAPES 88887.841455/2023-00; DCSP was funded by CNPq 131837/2022-2; DSA is supported by a FAPESP grant #14.092-0; RLF was funded by FAPEG 202410267000126.

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ADDITIONAL NOTES

• ZooBank register
  https://zoobank.org/B1E9AAB7-B87A-4BF9-92D4-19AB56484B3A
• How to cite this article
  Oliveira SS, Afiune GPS, Schelesky-Prado DC, Maia VC, Amorim DS, Falaschi RL (2024) Taxonomic Catalog of the Brazilian Fauna: Bibionomorpha (Diptera) diversity and distribution. Zoologia 41: e23103. https://doi.org/10.1590/S1984-4689.v41.e23103
• Published by
  Sociedade Brasileira de Zoologia at Scientific Electronic Library Online - https://www.scielo.br/zool

• CNPq 308298/2021-7 and FAPEG 202310267000883; GPSA was funded by CAPES 88887.841455/2023-00; DCSP was funded by CNPq 131837/2022-2; DSA is supported by a FAPESP grant #14.092-0; RLF was funded by FAPEG 202410267000126. SSO was funded by CNPq 308298/2021-7 and FAPEG 202310267000883; GPSA was funded by CAPES 88887.841455/2023-00; DCSP was funded by CNPq 131837/2022-2; DSA is supported by a FAPESP grant #14.092-0; RLF was funded by FAPEG 202410267000126.

Edited by

• Editorial responsibility
  Sionei R. Bonatto

Data availability

Data citations

Amorim DS (2024) Sciaridae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1556 [Accessed: 05/08/2024]

Brazilian Zoology Group (2024) Catálogo Taxonômico da Fauna do Brasil. Available at http://fauna.jbrj.gov.br/fauna

Falaschi RL (2024a) Diadocidiidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1610 [Accessed: 05/08/2024]

Falaschi RL (2024b) Ditomyiidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1855 [Accessed: 05/08/2024]

Falaschi RL (2024c) Keroplatidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1632 [Accessed: 05/08/2024]

Falaschi RL (2024d) Rangomaramidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/2358 [Accessed: 05/08/2024]

Falaschi RL, Oliveira SS (2024) Anisopodidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/2616 [Accessed: 05/08/2024]

Falaschi RL, Schelesky-Prado DC (2024) Bibionidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1843 [Accessed: 05/08/2024]

Falaschi RL, Oliveira SS, Schelesky-Prado DC (2024) Bibionomorpha. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/453 [Accessed: 05/08/2024]

Maia VC (2023) Cecidomyiidae. Catálogo Taxonômico da Fauna do Brasil. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/2608 [Accessed: 25/11/2023]

Oliveira SS (2024a) Lygistorrhinidae. Catálogo Taxonômico da Fauna do Brasil . PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/1250 [Accessed: 05/08/2024]

Oliveira SS (2024b) Mycetophilidae. Catálogo Taxonômico da Fauna do Brasil . PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/2709 [Accessed: 05/08/2024]

Publication Dates

• Publication in this collection
  20 Dec 2024
• Date of issue
  2024

History

• Received
  14 Dec 2023
• Accepted
  06 Sept 2024