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Research Article: Taxonomic Catalog of the Brazilian Fauna • Zoologia (Curitiba) 41 • 2024 • https://doi.org/10.1590/S1984-4689.v41.e23095 linkcopy

Open-access The state of the art of the Brazilian Megaloptera (Insecta: Neuropterida)

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    ABSTRACT

    The present study provides a comprehensive overview of the Megaloptera fauna in Brazil. A total of 27 species-25 extant and two extinct-distributed into two families, Corydalidae and Sialidae, and six genera, are recorded from Brazil. The historical timeline of Megalopteran records in Brazil spans 180 years, from 1842 to 2022, and unfolds into three distinct periods: the 19th century and the first half of the 20th century dominated by European authors, the second half of the 20th century dominated by Mexican and US-born authors, and the 21st century dominated by Brazilian authors. Currently, about 75% of the type specimens of Brazilian Megaloptera are housed in foreign institutions. Among Brazilian states, Minas Gerais boasts the highest number of megalopteran records, with eight, while eight states have no records. The biome of Atlantic Forest exhibits the greatest richness of Megaloptera, with 15 species, whereas Pampas and Pantanal lack records of the order. The Amazon Basin leads with ten recorded species, although three Brazilian hydrographic basins remain with no records. Impressively, only ten Brazilian Megaloptera species have known immature stages.

    KEY WORDS:
    Alderfly; Brazilian Zoology Group; Corydalidae; dobsonfly; fishfly; Sialidae; taxonomy

    INTRODUCTION

    Megaloptera, a small order of holometabolous insects, encompasses 428 species sorted into 48 genera within two families, Corydalidae and Sialidae (Martins et al. 2022). The origin of the order has been recently elucidated through time-calibrated phylogenies, suggesting an origin predating the end of the Permian (ca 250 Ma) with its oldest known fossils dating back to the Early Jurassic of Europe (Winterton et al. 2017, Vasilikopoulos et al. 2020). This order, together with Neuroptera and Raphidioptera, belongs to the superorder Neuropterida. Recent studies support Raphidioptera as sister group to the clade Megaloptera + Neuroptera (Wang et al. 2017, Winterton et al. 2017, Vasilikopoulos et al. 2020).

    Despite their impressive size, megalopterans-commonly known in English as dobsonflies, fishflies, and alderflies-are often overlooked by non-biologists, save for their larvae, which are sought after as fishing bait. Megaloptera eggs are deposited in masses near freshwater containing few hundred eggs deposited in one layer and uncovered (Sialidae) or in multi-layers covered with a white or yellowish substance (Corydalidae) (New and Theischinger 1993, Ardila-Camacho and Contreras-Ramos 2018, Hamada and Azevêdo 2024). Megalopteran larvae are generalist predators in both lotic and, less commonly, lentic environments, inhabiting clear freshwater and being useful as indicators in environmental impact studies (Hamada and Azevêdo 2024). These larvae are characterized by a dorsoventrally flattened body, ranging in length from 0.8-12 mm (Sialidae) to 20-90 mm (Corydalidae), a well-developed head capsule with six stemmata per side, large prognathous mouthpart, short antennae, and an abdomen covered with small and large setae, featuring 7-8 pairs of lateral filaments (Ardila-Camacho and Contreras-Ramos 2018). Adults of Megaloptera exhibit a wide size range (7-100 mm forewing length), with a prognathous head and an enlarged hind wing anal area (New and Theischinger 1993, Ardila-Camacho and Contreras-Ramos 2018, Martins et al. 2022).

    Recently, Jiang et al. (2022) proposed an ancient Pangean distribution for early megalopterans. Presently, Megaloptera exhibits a global distribution across all major biogeographical regions, excluding near polar regions where they are notably absent. The substantial extant richness within the order is concentrated in Northern Hemisphere, with 283 extant species. The Oriental biogeographical area stands out as the most species-rich, housing 206 extant species, followed by the Neotropical region with 81 species (Martins et al. 2022). Regarding extinct diversity, Megaloptera comprises 25 species distributed across Europe (15 species), Asia (7 species), North America, and South America (two species each, Martins et al. 2022). Brazil emerges as richest country for Megalopteran species in the Neotropics, with 25 extant and two extinct (Martins 2024, Martins et al. 2022).

    Taxonomic studies on Brazilian Megaloptera started in the 19th century by European authors (e.g., Walker 1860). It was only in 2005 that a Brazilian author contributed to the original description of a Brazilian species (Contreras-Ramos et al. 2005). Noteworthy figures during this period include Atilano Contreras-Ramos (Universidad Nacional Autónoma de México, Mexico City, Mexico) and the Spanish Longino Navás (1858-1938), who described the highest number of Brazilian species per author of Megaloptera, five species each; the American Norman Dale Penny (1946-2016), for his contribution to faunistic of Megaloptera of the Amazon Forest during his presence in Manaus at Instituto Nacional de Pesquisas da Amazônia, Amazonas, Brazil-INPA in the late 1970s and 1980s; Neusa Hamada (INPA) and Carlos Augusto Silva de Azevêdo (Universidade Federal do Maranhão, Maranhão, Brazil-UEMA), who conducted studies on Brazilian Megaloptera, particularly focusing on their biology, taxonomy, and ecology (Penny 1982, Penny and Flint 1982, Contreras-Ramos 1998, 2000, Contreras-Ramos et al. 2005, Hamada and Azevêdo 2012, 2024, Mendes et al. 2022, 2024). The two extinct species of Megaloptera from Crato Formation in the Araripe Basin, Ceará, Brazil, were only described in the last decade by Jepson and Heads (2016).

    Data on the extant Brazilian Megaloptera fauna have been compiled in the Megaloptera section (Martins 2024) of the of the Taxonomic Catalog of the Brazilian Fauna. The section was coordinated by the first author of this study since the beginning of 2022, and prior to that, by the second and third authors. The Taxonomic Catalog of the Brazilian Fauna, in Portuguese: Catálogo Taxonômico da Fauna do Brasil (CTFB) is an online platform of the scientific valid names applied to the metazoan species occurring in Brazil. It is the product of the effort of around 800 qualified zoologists of all taxa that inhabit the Brazilian territory. Currently, CTFB encompasses around 125,000 valid extant animal nominal species, and includes information-e.g., year of publication, author’s name, synonymic list, hosts and parasites, distribution across Brazilian states, ecological biomes, and hydrographic basins-for an important part of these species (Boeger et al. 2024). The type locality, information on name-bearing types, and their museum repository have already started to be added to the Megaloptera section of the CTFB.

    This study draws inspiration from three recent papers: Santos et al. (2020), Duarte and Lecci (2023), and Machado and Martins (2023), which provided significant historical information into Brazilian species of Trichoptera, Plecoptera and Neuroptera, respectively. The objective of the current study is to conduct a similar comprehensive analysis of Megaloptera species of CTFB, along with an exploration of two extinct Megaloptera species from Brazil (not included in the CTFB). The focus is on highlighting locations of name-bearing type specimens, their distribution across states, biomes, and hydrographic basins, and presenting a historical narrative of how taxonomic studies on Brazilian Megaloptera have evolved over time. Additionally, the study aims to provide guidance on future areas of focus in this field.

    MATERIAL AND METHODS

    The extant species data analyzed in this study were sourced from the Megaloptera section of the CTFB website (http://fauna.jbrj.gov.br), last updated in November 2023. The continuous updating of Megaloptera data on the CTFB is grounded in specialized taxonomic literature and the Lacewing Digital Library - LDL (Oswald 2024). The classification for Megaloptera adopted in the CTFB follows Martins et al. (2022). Information on extinct species was extracted from Martins et al. (2023).

    All the data were extracted from the CTFB database and Martins et al. (2023) and adapted for presentation here (Supplementary Material 1), forming the basis for all analyses conducted in this study. The organizational structure follows Machado and Martins (2023), i.e. for each species, details such as the author’s name and nationality, year of description or first report from Brazil, location of deposit for name-bearing type specimens, endemism to Brazil, knowledge of immature stages, and distribution by states, biomes and hydrographic basins were provided (Supplementary Material 1). Author’s nationality was categorized into four categories: Brazil, Mexico, Europe, and USA, in order to organize the four main groups of authors who carry out studies on the taxonomy of Megaloptera in Brazil. The location of the deposit for name-bearing type specimens was divided into three categories: Brazil, Europe, and USA. The distributional data of extant species considered: (1) the 26 Brazilian states plus the Federal District and (2) the six Brazilian biomes: Amazônia (Amazon Forest), Mata Atlântica (Atlantic Forest), Cerrado (Brazilian savanna), Caatinga (arid shrubland), Pampas (subtropical grassland), and Pantanal (tropical wetlands), both as defined by the Instituto Brasileiro de Geografia e Estatistica (IBGE 2019); and (3) the 12 hydrographic basins: Amazon (Amazônica), East Atlantic (Atlântico Leste), Eastern Northeast Atlantic (Atlântico Nordeste Oriental), Paraguay (Paraguai), Paraná, Parnaíba, São Francisco, Southeast Atlantic (Atlântico Sudeste), South Atlantic (Atlântico Sul), Tocantins-Araguaia, Uruguay (Uruguai), Western Northeast Atlantic (Atlântico Nordeste Ocidental) as defined by the Agência Nacional de Águas e Saneamento Básico (ANA 2021). To extinct species were considered the Formation, Basin, and Chronostratigraphic ages that were found.

    In constructing accumulation curves, the year of the species’ first report from Brazil was utilized. All the graphics were generated in MS Excel (Microsoft), while maps were created in QGIS (QGIS Development Team 2016), and both were subsequently edited in Adobe Illustrator 2020.

    RESULTS

    A total of 25 valid extant nominal species of Megaloptera (Fig. 1A-H), distributed into four genera, three subfamilies, and two families, are recorded from Brazil, with 11 species (44%) endemic to the country (Table 1). Corydalidae, with 20 species, dominates the Brazilian extant richness of the order, constituting 80%. This includes the richest megalopteran genus in the country, Corydalus Latreille, 1802 with 14 species, along with Chloronia Banks, 1908 (five species), both belonging to Corydalinae (dobsonflies), and the monotypic Puri Cardoso-Costa, Azevêdo & Ferreira Junior, 2013 from Chauliodinae (fishflies). Sialidae (alderflies) is represented in Brazil by five species of Ilyobius Enderlein, 1910. Concerning the extinct Megaloptera fauna, two monotypic genera of Corydalidae (subfamily incertae sedis)-Cratocorydalopsis Jepson & Heads, 2016 and Lithocorydalus Jepson & Heads, 2016-have been identified in the Crato Formation, Araripe Basin, Brazil (Fig. 1I-J). Presently, the recorded number of valid extant Megaloptera nominal species and genera in Brazil is 25 and four, respectively. However, the CTFB website registers 25 names for species and nine for genera, accounting junior synonyms and valid genera including these synonyms. Notably, the dobsonfly Chloronia corripiens (Walker, 1860) holds the record for the highest number of synonyms among Brazilian species, with a total of seven junior synonyms.

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    Figure 1
    Megaloptera from Brazil. (A) Corydalidae oviposition; (B) Larva of Chloronia sp. (Corydalidae: Corydalinae); (C) Larva of Corydalus sp. (Corydalidae: Corydalinae); (D) Larva of Ilyobius sp. (Sialidae); (E) Pupa of Corydalidae; (F) Chloronia corripiens (Walker, 1860) (Corydalidae: Corydalinae); (G) Female of Corydalus diasi Navás, 1936 (Corydalidae: Corydalinae); (H) Male of Puri aleca Cardoso-Costa, Azevêdo & Ferreira Junior, 2013 (Corydalidae: Chauliodinae); (I) Ilyobius flammatus (Penny, 1982); (J) Female of Ilyobius nubilus (Navás, 1933); (K) †Cratocorydalopsis brasiliensis Jepson & Heads, 2016; (L) †Lithocorydalus fuscata Jepson & Heads, 2016. Photos: A-C, E-G by Frederico Sales; D by Caleb Califre Martins; H after Cardoso-Costa et al. (2013); I by Guilherme Fischer (https://www.inaturalist.org/people/gafischer); J after Mendes et al. (2024); K-L after Jepson and Heads (2016).

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    Table 1
    Distribution and current knowledge of the immature stages and adult sexes of Brazilian species of Megaloptera.

    The first Megaloptera species recorded in Brazil was Corydalus cephalotes Rambur, 1842 from state of Rio de Janeiro, while the most recent is Ilyobius erebus Mendes et al., 2022 from Minas Gerais, both during their original description. Over the span of 180 years, 25 species have been documented in Brazil, resulting in an average rate of one new species recorded every 7.5 years. The 19th century witnessed the recording of five species in Brazil, averaging one new record every 4.8 years (1842-1866). The 20th century documented 14 species in Brazil (1914-2000), averaging one new record every 6.14 years. The 21st century (interval from 2005 to 2022) exhibited the highest average rate of Megaloptera species records for Brazil, with one new species documented every 2.83 years (see Fig. 2). The trend line in Fig. 1 indicates that there is much to discover about the extant Megaloptera fauna in Brazil. The two extinct Megaloptera species were recorded in Brazil in their original description by Jepson and Heads (2016).

    The authors responsible for documenting extant Megaloptera species in Brazil can be categorized into three distinct intervals (Fig. 3). In the 19th century and the first half of 20th century, spanning between 1842 and 1936, European authors, such as Longinos Navás and Robert McLachlan (1837-1904) recorded ten species in Brazil, together. Moving to the second half of the 20th century (1977-2000), all nine megalopteran records in Brazil were attributed to authors from the USA such as Norman D. Penny and Oliver S. Flint Jr (1931-2019) or Mexico by Atilano Contreras-Ramos. It is important to highlight that Norman D. Penny lived in the Amazon Basin, Brazil Amazonas, Manaus for almost ten years (1976 to 1985). The third interval corresponds to the 21st century, where Brazilian authors actively participated in all six records of Megaloptera species in Brazil, with noteworthy contributions from Neusa Hamada and Carlos Augusto Silva de Azevêdo.

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    Figures 2-3
    Accumulation curve of Megaloptera extant species recorded from Brazil by year (1832-2022), based on the first record of each species to the country: (2) Trend line in red; (3) categorized by the citizenship of the first authors.

    Despite Brazil having ten authors involved in the descriptions of Megaloptera species from the country, each contributor is associated with the description of only one species. Additionally, nine Europeans, four US-Americans, and one Mexican participated in the description of Brazilian Megaloptera (Table 2). Atilano Contreras-Ramos from Mexico and the Spanish Longino Navás hold the record for describing the largest number of Brazilian Megaloptera species per author, five species each. While Gelson Luiz Fiorentin and Yuji Urakami contributed to the description of Ilyobius hauseri (Contreras-Ramos, Fiorentin & Urakami, 2005), it wasn’t until 2013 that Brazilian authors, namely Gil Cardoso-Costa, Carlos Augusto Silva de Azevêdo, and Nelson Ferreira-Jr, were the primary authors of a Megaloptera species, with the description of Puri aleca Cardoso-Costa, Azevêdo & Ferreira, 2013. The authors who described the extinct Megaloptera species and genera from Brazil are the European James E. Jepson and Sam W. Heads (Jepson and Heads 2016). Among the 24 authors who described the Brazilian Megaloptera species, only five are females: Gabriela C. Mendes, Jeane Marcelle C. do Nascimento, Lívia M. Fusari, Mireile R. dos Santos, and Neusa Hamada; all of them joined this list only in 2022 with the description of I. erebus.

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    Table 2
    Name of authors, and number of Brazilian species of Megaloptera they participated on the original description. († = described only extinct species).

    Approximately 75% of the type specimens of Brazilian Megaloptera (including fossils) are currently deposited in foreign institutions, notably in Europe (15 species, 55,5%) and the USA (three species, 11%). The remaining seven species have their name-bearing types deposited in four Brazilian collections: one species at the Coleção Entomológica Prof. José Alfredo Pinheiro Dutra, Universidade Federal do Rio de Janeiro, Rio de Janeiro (DZRJ), one species at Fundação Zoobotânica do Rio Grande do Sul (MCNZ), Rio Grande do Sul, two species at the Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP), and three species at the Instituto Nacional de Pesquisas da Amazônia, Amazonas (INPA).

    The overall knowledge of the Megaloptera in Brazil (Fig. 4) reveals the Southeastern region standing out with the highest richness of Megaloptera species, particularly in the state of Minas Gerais, which comprises the highest number of recorded species (eight). The richness of Megaloptera species in the state of Minas Gerais stems from studies conducted by researchers from various regions across the country and abroad. Most of the records of Megaloptera in the state were made by researchers from institutions outside of Minas Gerais. The two records of Chloronia were conducted by the US-Americans (Penny and Flint 1982), and the three of Corydalus by the Mexican Contreras-Ramos (Contreras-Ramos 1998). The record of Puri was made in its original description by Cardoso-Costa et al. (2013), with no authors from Minas Gerais. The only exceptions are the researchers from the Instituto Federal de Educação, Ciência e Tecnologia do Sul de Minas Gerais (IFSULDEMINAS): Mireile Reis dos Santos, who, along with other researchers, described the species I. erebus, and Gouvêa et al. (2020), who conducted a study on the distribution of Megaloptera in that state, leading to the recognition of Corydalus batesii (McLachlan, 1868) to this region.

    The North region ranks second in the number of Megaloptera species diversity in Brazil, particularly in the states of Amazonas and Roraima, which comprises an exceptional abundance of documented species. Furthermore, Rondônia, and Tocantins-also located in the north-emerge as states with some of the highest numbers of known Megaloptera species, contributing to the overall richness of this insect order in the region.

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    Figure 4
    Brazilian map showing the number of Megaloptera extant species recorded for each one of the states. States abbreviations: AC = Acre, AL = Alagoas, AM = Amazonas, AP = Amapá, BA = Bahia, CE = Ceará, DF = Federal District, ES = Espírito Santo, GO = Goiás, MA = Maranhão, MG = Minas Gerais, MS = Mato Grosso do Sul, MT = Mato Grosso, PA = Pará, PB = Paraíba, PE = Pernambuco, PI = Piauí, PR = Paraná, RJ = Rio de Janeiro, RN = Rio Grande do Norte, RO = Rondônia, RR = Roraima, RS = Rio Grande do Sul, SC = Santa Catarina, SE = Sergipe, SP = São Paulo, TO= Tocantins.

    In contrast, eight states-out of 26 and the Federal District-currently lack records of the order. Seven Brazilian states have endemic megalopteran species, with Minas Gerais leading with two species. The Southeastern region holds the highest number (12 species) of Megaloptera species, followed by the North (11 species), South (6 species), Midwest, and Northeast regions (4 species each). Despite the Northeast region having the largest number of states in the country, it exhibits the lowest known diversity of Megaloptera, with seven out of nine states lacking records of the order. Corydalus diasi Navás, 1936 has the widest distribution across Brazilian territory, occurring in nine states and Federal District and all five regions (North, Northeast, Midwest, Southeast, and South), followed by Corydalus affinis Burmeister, 1839, which is registered in nine states and three regions (North, Midwest, Southeast). Three species of Corydalus, one of Chloronia, four of Ilyobius and Puri have records in only one state (Table 1).

    Among Brazil’s six terrestrial biomes (Fig. 5), the Atlantic Forest stands out as the richest, harboring records of 15 extant megalopteran species, constituting more than 50% of the total Megaloptera extant fauna in the country. Following closely, the Amazon Forest is the second-highest richness with ten species, trailed by the Cerrado with six species, and Caatinga with one species. Conversely, there are no official records of Megaloptera species in the Pampa and Pantanal biomes. Given that Megaloptera larvae are aquatic, their distribution is intricately linked to the various hydrographic basins in Brazil (Fig. 6). Among the 12 basins present in the country, the Amazon Basin is the richest, hosting ten Megaloptera species. Paraná basin has the second-largest number of records, with eight recorded species. The basins of Western Northeast Atlantic, Parnaíba, and Paraguay currently lack records of Megaloptera. The first two are concentrated in the Northeastern region, where, unfortunately, there is a dearth of studies on this order. Similarly, the Paraguay basin is associated with the Pantanal biome, and like the Northeast, there is a notable absence of research on megalopterans. Corydalus diasi Navás, 1936 is the species occurring in the largest number of Brazilian hydrographic basins, spanning a total of eight throughout the country.

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    Figures 5-6
    Brazilian map showing the number of Megaloptera extant species recorded for each one of the Brazilian: (5) terrestrial biomes; (6) hydrographic basins.

    Despite their importance, especially in their larval stage, only ten Brazilian Megaloptera species-one Chloronia, six Corydalus, and three Ilyobius-have documented records of known immature stages. All these species have documented larvae, with information on eggs and pupae available for only two and three Brazilian megalopterans, respectively (Table 1).

    Valuable insights into the Brazilian Megaloptera fauna are scattered across various studies, particularly in recent years. Penny and Flint’s (1982) revision of Chloronia and Contreras-Ramos’s (2000) article provide important information and an identification key to the genus. The revision of Corydalus by Contreras-Ramos (1998) enables the identification of species within this genus. Hamada and Azevêdo have made substantial contributions to the understanding of Brazilian Megaloptera through several studies, encompassing insight into immature stages (Azevêdo 2003, 2009), key information about the fauna (Hamada and Azevêdo 2024), and an exploration of Amazonian fauna, including an excellent pictorial key for larvae (Azevêdo and Hamada 2019). Cardoso-Costa et al. (2013) described the singular Brazilian Chauliodinae, Puri aleca, providing comprehensive information on this unique species.

    Information about the Brazilian Sialidae fauna is mainly found in the original descriptions of the species (Navás 1933, Penny 1982, Contreras-Ramos et al. 2005), and in the article by Liu et al. (2015) about taxonomic notes of Ilyobius. Additionally, in the same article featuring the original description of I. erebus (Mendes et al 2022), the authors provided an important discussion about the Brazilian fauna of Sialidae. Martins et al. (2022) curated a comprehensive global checklist of extant and extinct species within the order, offering identification keys, distribution details, and historical data for all genera, including Brazilian fauna. The Megaloptera section on the “Planeta Invertebrados Brasil” website (PIB 2023) shares important information about Brazilian fauna, featuring images of most species. Concerning the fossil fauna, primary information is present in original descriptions (Jepson and Heads 2016) and in a recent article on the Neuropterida fauna from Brazil (Martins et al. 2023).

    Based on the available information regarding the Brazilian Megaloptera fauna, it is evident that many species are known from only a few specimens. Chloronia pennyi Contreras-Ramos, 2000, is officially documented only by its type series, comprising the holotype and four paratypes. Penny (1977) recorded Chloronia hieratica Navás, 1928-a junior synonym of Chloronia mirifica (Navás, 1925)-for Brazil, however, he did not provide further locality or material examined, casting doubt on this record. Corydalus armatus Hagen, 1861, was mentioned for Brazil without further details by Navás (1929). Contreras-Ramos (1998) confirmed this record with a Brazilian specimen, but without additional locality details. Corydalus arpi Navás, 1936, is represented by several specimens from Venezuela; however, only one specimen from Rondônia is officially known for Brazil (Contreras-Ramos 1998). Corydalus amazonas Contreras-Ramos, 1998, is known only from the holotype and two paratypes, all females; the male, hitherto unknown, is being described by the first author of this study. Corydalus cephalotes Rambur, 1842, is known from a small series (holotype and three additional specimens), all from the state of Rio de Janeiro (Contreras-Ramos 1998). Both Corydalus flavicornis Stitz, 1914, and Corydalus ignotus Contreras-Ramos, 1998, are known from Brazil based solely on a series of specimens studied by Azevêdo (2009). Corydalus peruvianus Davis, 1903, is known only from three male specimens from Roraima (Gama-Neto and Passos 2018) and one specimen from Mato Grosso, identified as Corydalus primitivus Van der Weele, 1909, by Glorioso (1981) (see Felix et al. 2024 for further details). Corydalus tridentatus Stitz, 1914, is also known from only a few Brazilian specimens (Contreras-Ramos 1998). The only Brazilian Chauliodinae, Puri aleca, is represented by three males from the type series; additional specimens are currently unknown. All five Brazilian species of Ilyobius are known from only a few specimens collected in specific regions of Brazil.

    DISCUSSION

    Species richness in Brazil and the distribution of subfamilies align with the global pattern for Megaloptera, with Corydalidae having the highest number of species (Martins et al. 2022). Considering the species of Megaloptera found in neighboring countries (Martins 2019, Oswald 2024) and the vast unexplored areas within Brazil, we anticipate that taxonomic knowledge of this group will expand (Hamada and Azevêdo 2024).

    Our dataset reveals that the Atlantic Forest, Amazon Forest, and Cerrado are well-studied biomes, while the diversity of Megaloptera of Caatinga, Pampa, and Pantanal are virtually unknown. The understanding of insect diversity in Brazil is significantly hindered by sampling bias, impacting knowledge of species composition, distribution, and ecological niches (Oliveira et al. 2016). This is closely linked to Brazil’s historical socioeconomic development, which began in the southeastern coastal area and spread to the southern and northern coast and inland regions (Marques et al. 2016). Consequently, the earliest Brazilian universities and biodiversity research groups were established in coastal states and the southeastern countryside, including Minas Gerais state, an area with rich fragments of Atlantic Forest and Cerrado.

    Prominent biodiversity research centers such as the Instituto Nacional de Pesquisas da Amazônia in Manaus and the Museu Paraense Emílio Goeldi in Belém have significantly contributed to the Amazônia status as a highly studied biome. Research in this biome is concentrated around these centers. Although there has been some biodiversity research in the Caatinga, Pampa, and Pantanal, dedicated Megaloptera research groups are relatively new compared to those in the southeastern and major northern regions, explaining the lower number of studies in these areas. Including studies from neighboring South American countries that share the Pampa and Pantanal biomes could increase the number of papers, but it would also artificially inflate the study count for already well-studied biomes like the Amazon Forest, Atlantic Forest, and Cerrado.

    Megaloptera sampling in all Brazilian biomes is highly concentrated near major biodiversity research centers, similar to other groups. This high aggregation of samples is primarily due to historical financial limitations, particularly in the Amazon Forest, where accessing remote areas is costly and time-consuming (Magnusson et al. 2013). Consequently, many sampling sites are near Manaus and Belém, where researchers can conduct studies more economically. In the Atlantic Forest and Cerrado, besides cost limitations, habitat loss, fragmentation, and degradation further restrict sampling to geographically limited and permanent sites.

    Our data visually highlight the gaps in Megaloptera diversity studies across Brazilian biomes and identify under-sampled or unknown areas for future research. Therefore, models that incorporate both sampling density and habitat loss are essential for pinpointing high-priority areas for new surveys (Divieso et al. 2020). Furthermore, it is essential to expand the coverage of Megaloptera diversity studies in Brazil due to the pervasive habitat loss, fragmentation, and degradation affecting all biomes, not just the Atlantic Forest and Cerrado (Divieso et al. 2020).

    Despite the presence of various gaps that contribute to biodiversity shortcomings, particularly highlighted by Hortal et al. (2015) and Faria et al. (2020), in the Midwest and South regions, it is worth noting that the Megaloptera fauna in the Northeast region is virtually unknown. This pattern, seen in other insect groups like Neuroptera (Machado and Martins 2023), results from insufficient studies on local diversity, with seven of nine states lacking Megaloptera records. Preliminary findings from a project led by the first author indicate that this knowledge gap in the Northeast is expected to narrow in the coming years, addressing the Brazilian Linnean and Wallacean shortfalls.

    Future biodiversity projects should prioritize regions, biomes, and hydrographic basins without Megaloptera records to fill critical knowledge gaps. Even in biomes with the highest Megaloptera diversity, achieving comprehensive understanding remains challenging. The scarcity of information also extends to Brazil’s extinct Megaloptera fauna. Despite various Brazilian formations, including those from the Cretaceous period, only two species have been documented from the Crato Formation due to the lack of comprehensive collections (Martins et al. 2023).

    As evidenced by our dataset, the documentation of Megaloptera species in Brazil during the 19th century and the first half of 20th century was dominated by European authors, a pattern consistent with observations in other insect groups, such as lacewings (Machado and Martins 2023) and caddisflies (Santos et al. 2020). This trend is likely linked to historical events shaping scientific exploration. The notable surge in records by Mexican and U.S.A. authors in the latter half of 20th century is primarily attributed mainly to the comprehensive revisions of the Chloronia and Corydalus genera by Penny and Flint (1982) and Contreras-Ramos (1998), respectively. Furthermore, the extensive impact of Norman Penny, an American researcher, cannot be overstated. He dedicated his time to collecting and studying a diverse array of insect specimens, including those from the Megaloptera group. The increased average rate of Megaloptera species records in Brazil during the 21st century aligns with a shift in authorship dominance by Brazilian researchers, emphasizing the pivotal role of local researchers in advancing our understanding of native faunas. Despite this progress, the order still requires heightened attention in diversity studies. It is conceivable that the number of Megaloptera could potentially double in the coming years (Hamada and Azevêdo 2024, Rafael et al. 2024). For instance, ongoing descriptions of a new Chloronia species from the Atlantic Forest (Paraná state) and a new Corydalus species from the Parnaíba basin (Maranhão state) are indicative of this trend.

    The deposition of name-bearing types of Brazilian Megaloptera in foreign institutions is closely tied to the nationalities of the reaches studying this fauna. Despite efforts in digitizing insect collections in prominent international institutions, local researchers often face challenges accessing these crucial specimens, presenting a significant taxonomic impediment for Brazil. This challenge extends to extinct species, as their holotypes are currently housed in the Stuttgart State Museum of Natural History, Germany (SMNS).

    Beyond the aforementioned issues, several other aspects merit prioritization in future studies. Ongoing initiatives are training Brazilian students to delve into the research of extant Megaloptera across diverse institutions and regions in the country (e.g., Instituto Nacional de Pesquisas da Amazônia in the North; Universidade Estadual do Maranhão and Universidade Federal da Bahia in the Northeast; Museu Nacional, Universidade Federal do Rio de Janeiro in the Southeast; and Universidade Federal do Paraná in the South). However, the emphasis on this training needs to be sustained and expanded in the future, with a crucial priority being the inclusion of studies on extinct fauna. since, as highlighted by Martins et al. (2023), Brazil has an important number of geological formations with great potential for new materials of this order.

    Despite the larvae of Brazilian Megaloptera playing a pivotal role in potential biomonitoring studies for aquatic environment degradation, their knowledge remains largely obscure. Prioritizing studies on the immature stages of this order is essential to address the Haeckelian shortfall (see Faria et al. 2020). Field research often struggles to locate adult Megaloptera, necessitating the refinement of laboratory rearing methods for larvae-a task already undertaken by the research group led by Neusa Hamada at Instituto Nacional de Pesquisas da Amazônia. Lastly, a noteworthy gender gap is evident in Megaloptera studies, with a disproportionately low percentage of female authors. Addressing this disparity is crucial to encourage future actions aimed at reducing the significant gender imbalance in Brazilian science and should be a focal point in future research endeavors.

    ACKNOWLEDGMENTS

    We thank the CTFB administrators: Walter A.P. Boeger (UFPR), Hussam Zaher (MZUSP) and Michel P. Valim (MZUSP); and John D. Oswald (TAMU), coordinator of the LDL. We are very grateful to Ministério da Ciência, Tecnologia e Inovação, Governo Federal, Brazil) and Ministério do Meio Ambiente, Governo Federal, Brazil) for the financial support at the beginning of the CTFB project; to all of the coordinators of the CTFB taxa for the management of this enormous project; to the botanist Rafaela Campostrini Forzza for the support and accommodation of the Botanical Garden of Rio de Janeiro, and to all zoologists for participating in this enormous project. Our special thanks to the website INaturalist, and Frederico Falcão Sales and Guilherme Fischer Roger Dias for making available and allowing the use of their images of Brazilian Megaloptera. We are grateful to Alexandre de Aguiar Cavalcante for the help with the map’s figures. We offer our sincere acknowledgments to Atilano Contreras-Ramos, Frederico Sales, and Ângelo P. Pinto for corrections and suggestions. CCM is grateful to the FAPEMA/SECTI of the Maranhão State government for the visiting professor/research grant (BPV-06012/21). JAR: to Conselho Nacional de Desenvolvimento Científico e Tecnológico, by the financial support, research fellow, processes 300997/2016-7 and 306661/2021-7; PROTAX proc 440423/2015-5 and 443641/2020-0 and recent grants, process 404225/2021-7 (Universal) and 406833/2022-2 (INCT Biodossel). JTC: to Conselho Nacional de Desenvolvimento Científico e Tecnológico, for the research grant, process 433694/2018-1 (Universal).

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    ADDITIONAL NOTES

    • FAPEMA/SECTI BPV-06012/21. Conselho Nacional de Desenvolvimento Científico e Tecnológico 300997/2016-7, 306661/2021-7; 440423/2015-5, 443641/2020-0, 404225/2021-7, 406833/2022-2, 433694/2018-1. CCM is grateful to the FAPEMA/SECTI of the Maranhão State government for the visiting professor/research grant (BPV-06012/21). JAR: to Conselho Nacional de Desenvolvimento Científico e Tecnológico, by the financial support, research fellow, processes 300997/2016-7 and 306661/2021-7; PROTAX proc 440423/2015-5 and 443641/2020-0 and recent grants, process 404225/2021-7 (Universal) and 406833/2022-2 (INCT Biodossel). JTC: to Conselho Nacional de Desenvolvimento Científico e Tecnológico, for the research grant, process 433694/2018-1 (Universal).

    Supplementary material 1

    Supplementary S1. General data on Brazilian species of Megaloptera.

    Authors: Caleb Califre Martins.

    Data type: Distribution, taxonomy and knowledge about different stages.

    Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

    Link: https://doi.org/10.1590/S1984-4689.v41.e23095

    Edited by

    • Editorial responsibility
      Ângelo Parise Pinto

    Data availability

    Data citations

    Martins CC (2024) Megaloptera. In: Taxonomic Catalog of the Brazilian Fauna. PNUD, PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/249 [Acessed: 24/05/2024]

    Publication Dates

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

    History

    • Received
      05 Dec 2023
    • Accepted
      06 Sept 2024
    Creative Common - by 4.0
    This is an open-access article distributed under the terms of the Creative Commons Attribution License
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