Abstract:
Polymitarcyidae has a worldwide geographic distribution. Their nymphs are adapted to the fossorial habit, while the adults (imagos) have atrophied legs, except for the anterior legs of the males. This study represents the first survey of this family in the western region of Pará. For this purpose, we collected nymphs and adults. Nymphs were captured using aquatic entomological nets and sieves, while adults were obtained using light traps and flight interception. In addition, we used online sources, such as the Brazilian Fauna Taxonomic Catalog, the EPHEMBRAZIL platform and the Ephemeroptera da América do Sul website. The survey revealed the presence of three genera and 13 species of Polymitarcyidae in the region, including six new records for the state of Pará. These results highlight the importance of geographic distribution surveys to expand our knowledge about biodiversity.
Keywords: Aquatic insects; Freshwater; Survey; Amazon
Resumo:
Polymitarcyidae possui uma distribuição geográfica mundial. Suas ninfas são adaptadas ao hábito fossorial, enquanto os adultos (imagos) apresentam pernas atrofiadas, exceto pelas pernas anteriores dos machos. Este estudo representa o primeiro levantamento dessa família na região Oeste do Pará. Para isso, coletamos ninfas e adultos. As ninfas foram capturadas com redes entomológicas aquáticas e peneiras, enquanto os adultos foram obtidos por meio de armadilhas luminosas e de interceptação de voo. Além disso, utilizamos fontes online, como o Catálogo Taxonômico da Fauna do Brasil, a plataforma EPHEMBRAZIL e o website Ephemeroptera da América do Sul. O levantamento revelou a presença de três gêneros e 13 espécies de Polymitarcyidae na região, incluindo seis novos registros para o estado do Pará. Esses resultados destacam a importância dos levantamentos de distribuição geográfica para ampliar nosso conhecimento sobre a biodiversidade.
Palavras-chave: Insetos aquáticos; Ambiente dulcícola; Inventário; Amazônia
Introduction
The Polymitarcyidae Banks, 1900, a family of mayflies (Ephemeroptera: Insecta), exhibits a broad geographic distribution being found in the Holarctic, Palearctic, Eastern and Neotropical regions (McCafferty 2004) and inhabits both lentic and lotic environments (Molineri & Salles 2013). Nymphs of this family are notable for their behavior of excavating tunnels in wood, residing in freshwater porifers, or inhabiting inorganic sediments (Molineri et al. 2015). They are distinguished by sickle-shaped mandibular projections, fused anterior tibia and tarsus, and typically dorsal arrangement of gills (Da-Silva & Salles 2024). Adults of this group are characterized by the presence of atrophied legs, except for the anterior legs in males (Domínguez et al. 2006).
Numerous species actively participate in bioturbation of bottom sediments within watercourses (Fonseca & Assis, 2000; Figueiredo-Barros et al., 2007; Leal et al., 2007), playing crucial roles in both aquatic and terrestrial food chains. Species of Polymitarcyidae have emerged as significant indicators of water quality, exhibiting a preference for ecosystems with high substrate quality and availability (Hamada & Ferreira-Keppler, 2012; Hamada et al., 2019). This underscores the pivotal role of this group as a vital component within ecosystems (Kuhlmann et al., 2020).
The Polymitarcyidae family is divided into three subfamilies, two of which are found in South America: Asthenopodinae (comprising the genera AsthenopodesUlmer, 1924; Asthenopus Eaton, 1871; HubbardipesMolineri, Salles & Peters, 2015; and PriasthenopusMolineri, Salles & Peters, 2015) and Campsurinae (comprising the genera Campsurus Eaton, 1868; TortopusNeedham & Murphy, 1924; and TortopsisMolineri, 2010) (Molineri 2010, Molineri & Salles 2013, Molineri et al. 2015). In Brazil, there are 52 species distributed among seven genera (Salles et al. 2024), with Campsurus being the most representative genus, comprising 39 described species to date, and also one of the most specious genera on the American Continent (Molineri & Salles 2017).
In the Amazon biome, the state of Amazonas boasts the highest diversity of Polymitarcyidae, hosting a total of 28 species (Molineri et al. 2015, Salles et al. 2024). In contrast, the state of Pará harbors only two genera and seven species: Asthenopus curtus (Hagen, 1861); Campsurus albifilum (Walker, 1853); C. essequibo Traver, 1947; C. latipennis (Walker, 1853); C. lucidusNeedham & Murphy, 1924; C. segnisNeedham & Murphy, 1924; and C. violaceusNeedham & Murphy, 1924 (Hagen 1861, Walker 1853, Molineri & Salles 2017, Oliveira et al. 2023, Needham & Murphy 1924, Demoulin 1955).
Despite the increased focus on taxonomy, geographic distribution, and ecology of the order Ephemeroptera in the state of Pará in recent years (Oliveira et al. 2021, 2023), there remains a lack of knowledge regarding certain taxa, particularly the Polymitarcyidae, leading to limited information about their diversity. According to Cardoso et al. (2023), Pará is one of the North region states with the highest potential for Ephemeroptera species diversity. However, the overall diversity within the Amazon biome remains poorly understood due to various factors, including the scarcity of specialists in the region, insufficient investment in specialist training resources, and the challenges associated with accessing an extensive region (Cardoso et al. 2023).
Another significant challenge to understanding species diversity in Pará arises from human-induced degradation of ecosystems. Even within priority conservation areas like indigenous territories, there is a noticeable trend of progressive ecosystem degradation (Cardoso et al. 2023). This degradation is particularly concerning in areas lacking environmental protection measures, compounded by losses from deforestation. Therefore, there is an urgent need to prioritize undersampled taxa and localities to enhance our understanding of biodiversity. Additionally, it is crucial to emphasize the potential use of sensitive and threatened species in conservation efforts, thereby safeguarding both the species and their habitats (Cardoso et al. 2015, Principe et al. 2019).
Thus, we conducted the inaugural taxonomic survey of the Polymitarcyidae family in Western Pará. Our research delivers valuable insights into new records and geographic distribution within this region.
Material and Methods
1. Study area
The study was conducted from June 2019 to May 2024 across several municipalities in Western Pará, including Santarém, Alenquer, Oriximiná - Distrito de Porto Trombetas, Terra Santa, Monte Alegre, and Óbidos, covering a total of 11 locations (Table 1). The region features a tropical monsoon climate, characterized by an average temperature of 25.6°C, an average relative humidity of 80%, and annual rainfall of 2,000 mm (Fapespa 2015, De Andrade 2017).
The sampled locations encompassed the Tapajós and Lower Amazonas hydrographic regions, focusing on the main rivers such as the Amazon, Tapajós, and Trombetas. In addition to these major rivers, the study also included sampling from lakes and streams.
2. Collection, identification and illustration
The Polymitarcyidae nymphs were collected with the aid of an aquatic entomological net in D (rapiché) and sieves, both with a 1 mm mesh opening (Merrit & Cummins 2008). These tools were systematically dragged across various substrates, including marginal vegetation, sponges, and submerged logs.
The alates were collected at dusk using a white sheet illuminated with a 15W emergency lamp connected to a 12V battery, along with a Pennsylvania-type trap. After collection, the imagos were preserved in vials containing 80% ethyl alcohol (Edmunds et al. 1976), while the subimagos were maintained alive in plastic microtubes until imagos emerged.
Species identifications were conducted using dichotomous keys (e.g., Domínguez et al. 2006) and supplemented with updated bibliographic references on the group (e.g., Molineri & Emmerich 2010, Molineri & Salles 2013, Molineri et al. 2015, Molineri & Salles 2017). For nymph identification, some structures such as mouthparts, legs, and gills were examined under an optical microscope. Adult specimens were examined for features including wings, legs, and male genitalia. Wings were mounted between a slide and coverslip without mounting medium and sealed with adhesive tape. Genitalia were inserted between the slide and coverslip using Hoyer’s mounting medium and then dried in an oven at 60°C for two days to produce permanent slides (Edmunds et al. 1976).
For the photographs, a digital camera connected to a stereomicroscope was employed, alongside a computer equipped with the Leica Application Suite programs (Version 2019). Additionally, a Zeiss Axiocam ERc 5s photomicroscope contributed to image capture. The Combine ZP program was used to assemble the layer overlays. Subsequently, the photographs were edited into plates using the Adobe Photoshop CS6® program.
3. Map construction
The map illustrating the geographic distribution of Polymitarcyidae species in the study area was constructed using QGIS (v. 3.30.2-’s-Hertogenbosch) with Shapefile files sourced from Brazil and the state of Pará (see Figure 1).
(A–C) Maps with distributional data. (A) Map of Brazil highlighting the Pará State. (B) Map of Pará with the western region of Pará in detail. (C) Distribution of Polymitarcyidae species in Western Pará.
4. Bibliographic survey and geographic distribution
The bibliographical survey and compilation of species’ geographic distribution were sourced from the Brazilian Fauna Taxonomic Catalog (http://fauna.jbrj.gov.br/), EPHEMBRAZIL (https://ephembrazil.netlify.app/grid_all), Ephemeroptera from South America (http://ephemeroptera.com.br/), as well as specific articles.
Results
The inventory resulted in the compilation of three genera and 13 species of Polymitarcyidae for the State of Pará, including six new records. Notably, this includes the first record of the genus Priasthenopus in the region. Table 1 provides geographic information for the described locations (S) in the examined material.
Checklist and new geographical records
Asthenopus curtus (Hagen, 1861)
Geographic distribution. North: Amazonas (Molineri et al. 2015), Pará (Hagen 1861).
Comment. Species previously recorded for the state of Pará by Hagen (1861), however without details of the type locality.
Priasthenopus gilliesi (Domínguez, 1988)
Male imagos of Polymitarcyidae (Ephemeroptera) recorded for the first time in Pará State. (A-B) Priasthenopus gilliesi, male imago. (A) Dorsal view. (B) Genitalia. (C–D) Campsurus demeni, male imago. (C) Dorsal view. (D) Genitalia. (E–F) Campsurus froehlichi, male imago. (E) Dorsal view. (F) Genitalia. (G–H) Campsurus truncatus, male imago. (G) Dorsal view. (H) Genitalia. (I-J) Campsurus povilla, male imago. (I) Dorsal view. (J) Genitalia. (K–L) Campsurus zunigae, male imago. (K) Dorsal view. (L) Genitalia.
Material Examined. One ♂ imago, S6, 04/vii/2023, L.A. Oliveira and S.E. Santos cols.
Geographic distribution. North: Amazonas (Domínguez 1988), Roraima (Salles et al. 2023). New record of the genus and species for the state of Pará.
Campsurus albifilum (Walker, 1853)
Geographic distribution. North: Amazonas (Molineri & Salles 2013), Pará (Walker 1853). Midwest: Mato Grosso do Sul (Molineri & Salles 2013). Southeast: Rio de Janeiro (Navás 1916). South: Paraná (Molineri & Salles 2013).
Comment. Species previously recorded for the state of Pará by Walker (1853), however without details of the type locality.
Campsurus demeni Molineri & Salles, 2017
Material Examined. Six ♂ imagos, S3, 09/ii/2024, L.A. Oliveira col.
Geographic distribution. North: Acre (Molineri & Salles 2017), Amazonas (Molineri & Salles 2017), Roraima (Molineri & Salles 2017). New record of the species for the state of Pará.
Campsurus essequibo Traver, 1947
Material Examined. 42 ♂ imagos, S1, 13/xii/2022, S.E. Santos and A.S. Corrêa cols.; same data as preceding except 33 ♂ imagos and 12 ♀ imagos, 21/ix/2023, L.A. Oliveira and A.S. Corrêa cols. 43 ♂ imagos, S3, 09/ii/2024, L.A. Oliveira col.
Geographic distribution. North: Amazonas (Molineri & Salles 2017), Pará (Molineri & Salles 2017), Roraima (Molineri & Salles 2017).
Campsurus froehlichi Cruz, Molineri & Hamada, 2022
Material Examined. 13 ♂ imagos and eight ♀ imagos, S11, 28-29/v/2023, L.A. Oliveira and S.E. Santos cols.
Geographic distribution. North: Amazonas (Cruz et al. 2022), Amapá (Cruz et al. 2022). New record of the species for the state of Pará.
Campsurus latipennis (Walker, 1853)
Geographic distribution. North: Pará (Walker 1853), Tocantins (Molineri & Salles 2017). Southeast: Espírito Santo (Salles et al. 2010).
Comment. Species previously recorded for the state of Pará by Walker (1853), however without details of the type locality.
Campsurus lucidus Needham & Murphy, 1924
Geographic distribution. North: Pará (Oliveira et al. 2023), Roraima (Molineri & Salles 2017). Midwest: Mato Grosso do Sul (Molineri & Salles 2017). South: Santa Catarina (Ulmer 1942).
Comment. Species previously recorded for the state of Pará by Oliveira et al. (2023) for the metropolitan region of Santarém.
Campsurus segnis Needham & Murphy, 1924
Geographic distribution. North: Amapá (Cruz et al. 2022), Pará (Needham & Murphy 1924).
Comment. Species previously recorded for the state of Pará by Needham & Murphy (1924) for the municipality of Igarapé-Açu.
Campsurus truncatus Ulmer, 1920
Material Examined. 15 ♂ imagos, S6, 06-07/vii/2023, L.A. Oliveira col. Five ♂ imagos, S8, 04/v/2023, S.E. Santos col.
Geographic distribution. North: Amazonas (Molineri et al. 2015). North East: Bahia (Lima et al. 2016), Pernambuco (Lima et al. 2015), Piauí (Takiya et al. 2016). Midwest: Goiás (Raimundi 2019). Southeast: Espírito Santo (Ulmer 1920), Minas Gerais (Molineri et al. 2015), Rio de Janeiro (Pereira & Da Silva 1991), São Paulo (Molineri et al. 2015). New record of the species for the state of Pará.
Campsurus violaceus Needham & Murphy, 1924
Material Examined. 41 ♂ imagos, S2, 14/vii/2023, S.E. Santos col. 213 ♂ imagos, S5, 04-05/vii/2023, S.E. Santos and L.A. Oliveira cols. 77 ♂ imagos and 15 ♀ imagos, S6, 04-07/vii/2023, S.E. Santos and L.A. Oliveira cols. 19 ♂ imagos and 10 ♀ imagos, S7, 28/v/2023, S.E. Santos col.
Geographic distribution. North: Amazonas (Molineri et al. 2015), Pará (Demoulin 1955). North East: Piauí (Takiya et al. 2016). Midwest: Goiás (Raimundi 2019), Mato Grosso do Sul (Molineri et al. 2015), Mato Grosso (Molineri et al. 2015).
Campsurus povilla Molineri & Salles, 2017
Material Examined. Eight ♂ imagos, S9, 13/v/2024, L.A. Oliveira and S.E. Santos cols. Three ♂ imagos, S10, 14/v/2024, L.A. Oliveira and S.E. Santos cols.
Geographic distribution. North: Amazonas (Molineri & Salles 2017). Midwest: Goiás (Molineri & Salles 2017). New record of the species for the state of Pará.
Campsurus zunigae Molineri & Salles, 2017
Material Examined. 19 ♂ subimagos, S4, 14-15/v/2024, L.A. Oliveira and S.E. Santos cols.
Geographic distribution. North: Amazonas (Molineri & Salles 2017), Roraima (Molineri & Salles 2017). North East: Bahia (Molineri & Salles 2017). Midwest: Mato Grosso do Sul (Molineri & Salles 2017). New record of the species for the state of Pará.
Discussion
The bibliographic survey initially identified only seven species, distributed across two genera in Pará. Our study has expanded the state’s representation to 13 species spanning three genera. Among the collections, Campusurus violaceus emerged as the most abundant species (n = 375), while Priasthenopus gilliesi was the least abundant, with only one individual recorded. In terms of geographic distribution, only C. froehlichi, C. demeni, C. povilla, C. zunigae and P. gilliesi were found to be restricted to a single municipality.
Currently, Pará ranks as the third state with the highest number of Polymitarcyidae records in the North region, trailing behind Amazonas and Roraima (Salles et al., 2024). Following our study, there has been 86% increase in the number of Polymitarcidae species documented for the state. This increase elevates Pará to the second-best sampled state in the region.
Priasthenopus gilliesi, Campsurus demeni, Campsurus froehlichi, Campsurus truncatus, Campsurus povilla and Campsurus zunigae were the species highlighted as new records for Pará. These species exhibit limited geographical distribution, with some restricted to the northern region, except for C. truncatus, which has a broader distribution throughout the country but had not been previously recorded in the state (Salles et al. 2024). This underscores the importance of regular sampling efforts and the expansion of survey areas, as it helps to reduce the Wallacean shortfall (Cardoso et al. 2015).
Pará stands out among states requiring priority surveys for Ephemeroptera, given its areas characterized by low sampling effort and high potential for discovering new species (Cardoso et al. 2023). Furthermore, many areas are unprotected by the arc of deforestation, which accelerates habitat degradation and puts biodiversity at risk (Veríssimo & Pereira 2014, Jacobus et al. 2019). According to Cardoso et al. (2015), despite certain well-studied groups, there remains a pressing need to revise estimates of taxon diversity due to the significant potential for new discoveries, the model predicts that 11,155 species of Ephemeroptera may occur in Brazil, of which 10,864 are not yet known. This is particularly true for Polymitarcyidae, which remain understudied across various Brazilian regions (Ottoni et al. 2009).
Thus, geographic distribution surveys play a crucial role in advancing our understanding of species, particularly those threatened or vulnerable to anthropogenic effects. This is especially pertinent for the species highlighted in this study, as many of them currently have limited distribution across the country.
Acknowledgments
This study was carried out with the support of the Municipal Environment Secretaries of the municipalities of Santarém, Óbidos, Terra Santa, Alenquer and ICMBio- Porto Trombetas, as well as the financial support provided by CNPq through notice 02/2023/PROPPIT/UFOPA. With the support of the Universidade Federal do Oeste Pará and the Bionorte Postgraduate Program. We also thank Suzane Evaristo dos Santos, Iandra Braz Sousa and Carlos Souza for their contributions to field collection, Hugo from the Multifunctional Multiuser Analysis Laboratory for helping with the photographs and Diego Sousa for creating the map.
Data Availability
The data used in our analysis is available at Biota Neotropica: https://doi.org/10.48331/scielodata.OBROHU
References
-
CARDOSO, M. N., SHIMANO, Y., NABOUT, J. C. & JUEN, L. 2015. An estimate of the potential number of mayfly species (Ephemeroptera, Insecta) still to be described in Brazil. Revista Brasileira de Entomologia, 59, 147. http://dx.doi.org/10.1016/j.rbe.2015.03.014.
» https://doi.org/10.1016/j.rbe.2015.03.014 -
CRUZ, P.V., MOLINERI, C. & HAMADA, N. 2022. Two new species and new records of Campsurus Eaton (Ephemeroptera: Polymitarcyidae) in Amazon. Zootaxa, 5100 (3), 435–444. https://dx.doi.org/10.11646/zootaxa.5100.3.7.
» https://doi.org/10.11646/zootaxa.5100.3.7 -
DA-SILVA, E.R. & SALLES, F.F. 2024. Cap 14, Ephemeroptera Hyatt & Arms, 1891, pp. 174–186. In: RAFAEL, J.A., MELO, G.A.R., CARVALHO, C.J.B., CASARI, S. & CONSTANTINO, R. (eds). Insetos do Brasil: Diversidade e Taxonomia. 2°ed. Instituto Nacional de Pesquisas da Amazônia, Manaus. 880 pp. https://doi.org/10.61818/56330464c14.
» https://doi.org/10.61818/56330464c14 - DE ANDRADE, V.M.S, CORDEIRO, I.M.C.C., SCHWARTZ, G., VASCONCELOS, L.G.T.R. & OLIVEIRA, F.D.A. 2017. Considerações sobre clima e aspectos edafoclimáticos da mesorregião Nordeste paraense. In: CORDEIRO, I.M.C.C., RANGEL-VASCONCELOS, L.G., SCHWARTZ, G. & OLIVEIRA, F.D.A. 2017. Belém: EDUFRA. pp. 59–96.
- DEMOULIN, G. 1955. Une mission biologique belge au Bresil. Ephéméroptères. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 31(20), 1–32.
-
DE SOUZA, M. R. & MOLINERI, C. 2012. The adults and nymphs of Asthenopus angelae new species (Ephemeroptera: Polymitarcyidae) from Argentina, Bolivia, Brazil and Colombia. Zootaxa, 3399, 45–52. https://doi.org/10.11646/zootaxa.3399.1.4.
» https://doi.org/10.11646/zootaxa.3399.1.4 - DOMÍNGUEZ, E. 1988. Asthenopus gilliesi sp. n. y su importancia en la taxonomia de la subfamilia Asthenopodinae (Ephemeroptera: Polymitarcyidae). Anales del Museo de Historia Natural de Valparaiso, 19, 21–26.
- DOMÍNGUEZ, E., MOLINERI, C., PESCADOR, M.L., HUBBARD, M.D. & C. NIETO. 2006. Ephemeroptera of South America. In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.): Aquatic Biodiversity in Latin America (ABLA). Vol. 2. Moscow: Pensoft. pp. 646.
- EDMUNDS, G.F., JENSEN, S.L. & BERNER, L. 1976. Mayflies of North and Central America. Minneapolis: Minnesota Archive Editions, 330.
-
FAPESPA - Fundação Amazônia de Amparo a Estudos e Pesquisas do Pará. 2015. Estatísticas Municipais Paraenses: Santarém. Diretoria de Estatística e de Tecnologia e Gestão da Informação, Belém, pp. 58. https://www.fapespa.pa.gov.br/ last access on 07/07/2023.
» https://www.fapespa.pa.gov.br/ - FIGUEIREDO-BARROS, M. P., SAVASSI, F., PIRES, A. P. F., DANTAS, M., LEAL, J. J. F., ESTEVES, F. A. & BOZELLI, R. L. 2007. Efeito da bioturbação por Campsurus notatus (Ephemeroptera: Polymitarcyidae) sobre a concentração de sólidos totais em suspensão e nitrogênio do séston. In Anais do VIII Congresso de Ecologia do Brasil. Caxambú: Sociedade de Ecologia do Brasil.
-
FONSECA LEAL, J. J. & ASSIS ESTEVES, F. 2000. Life cycle and production of Campsurus notatus (Ephemeroptera, Polymitarcyidae) in an Amazonian lake impacted by bauxite tailings (Pará, Brazil). Hydrobiologia, 437, 91–99. http://dx.doi.org/10.1023/A:1026526101039.
» https://doi.org/10.1023/A:1026526101039 - HAGEN, H.A. 1861. Synopsis of the Neuroptera of North America. With a list of the South American species. Smithosnian Miscellaneous Collections, Washingon. pp 1-347.
- HAMADA, N. & FERREIRA-KEPPLER, R. L. M. 2012. Guia ilustrado de insetos aquáticos e semiaquáticos da Reserva Florestal Ducke. Manaus: Editora da Universidade Federal do Amazonas. IBSN 978-85-7401-460-9.
- HAMADA, N., NESSIMIAN, J. L. & QUERINO, R. B. 2019. Insetos aquáticos na Amazônia brasileira: taxonomia, biologia e ecologia (2nd ed.). Manaus: Editora do INPA. ISBN 978-85-211-0194-9.
-
JACOBUS, L.M., MACADAM, C.R. & SARTORI, M. 2019. Mayflies (Ephemeroptera) and their contributions to ecosystem services. Insects, 10(6), 170. https://doi.org/10.3390/insects10060170
» https://doi.org/10.3390/insects10060170 -
KUHLMANN, M.L., IMBIMBO, H.R.V. & WATANABE, H.M. 2020. Campsurus truncatus Ulmer, 1920 (Polymitarcyidae): an Ephemeroptera in eutrophic waters. Acta Limnologica Brasiliensia, 32, e17. https://doi.org/10.1590/S2179-975X10019.
» https://doi.org/10.1590/S2179-975X10019 -
LEAL, J. J. F., DOS SANTOS FURTADO, A. L., DE ASSIS ESTEVES, F., BOZELLI, R. L. & FIGUEIREDO-BARROS, M. P. 2007. The role of Campsurus notatus (Ephemeroptera: Polymitarcytidae) bioturbation and sediment quality on potential gas fluxes in a tropical lake. Hydrobiologia, 586, 143-154. http://dx.doi.org/10.1007/s10750-006-0570-9.
» https://doi.org/10.1007/s10750-006-0570-9 -
LIMA, L. R., SALLES, F. F. & PINHEIRO, U. 2015. New records of mayflies (Ephemeroptera: Insecta) from Pernambuco state, northeastern Brazil. Check List, 11(3), 1652-1652. https://doi.org/10.15560/11.3.1652.
» https://doi.org/10.15560/11.3.1652 - LIMA, L. R. C., KNAPP, W. & DOCIO, L. 2016. New records of mayflies (Insecta: Ephemeroptera) from Bahia state, northeastern Brazil. Entomotropica, 31, 212-220.
- MCCAFFERTY, W.P. 2004. Higher classification of the burrowing mayflies (Ephemeroptera : Scap phodonta). Entomological News 115: 84–92.
- MERRIT, R. W., CUMMINS, K. W. & BERG, M. B. 2008. An Introduction to the Aquatic Insects Of North America. 4aed. Iowa.
-
MOLINERI, C. 2010. A cladistic revision of Tortopus Needham & Murphy with description of the new genus Tortopsis (Ephemeroptera: Polymitarcyidae). Zootaxa, 2481, 1–36. http://dx.doi.org/10.5281/zenodo.195431.
» https://doi.org/10.5281/zenodo.195431 -
MOLINERI, C. & EMMERICH, D. 2010. New species and new stage descriptions of Campsurus major species group (Polymitarcyidae: Campsurinae), with first report of silk-case construction in mayfly nymphs. Aquatic Insects, 32(4), 265–280. http://dx.doi.org/10.1080/01650424.2010.533131.
» https://doi.org/10.1080/01650424.2010.533131 -
MOLINERI, C. & Salles, F.F. 2013. Phylogeny and biogeography of the ephemeral Campsurus Eaton (Ephemeroptera, Polymitarcyidae). Systematic Entomology, 38(2), 265–277. https://doi.org/10.1111/j.1365-3113.2012.00656.x.
» https://doi.org/10.1111/j.1365-3113.2012.00656.x -
MOLINERI, C., SALLES, F.F. & PETERS, J.G. 2015. Phylogeny and biogeography of Asthenopodinae with a revision of Asthenopus, reinstatement of Asthenopodes, and the description of the new genera Hubbardipes and Priasthenopus (Ephemeroptera, Polymitarcyidae). ZooKeys, 478, 45–128. https://doi.org/10.3897/zookeys.478.8057.
» https://doi.org/10.3897/zookeys.478.8057 -
MOLINERI, C. & SALLES, F.F. 2017. Review of selected species of Campsurus Eaton 1868 (Ephemeroptera: Polymitarcyidae), with description of eleven new species and a key to male imagos of the genus. Zootaxa, 4300 (3), 301–354. http://dx.doi.org/10.11646/zootaxa.4300.3.1.
» https://doi.org/10.11646/zootaxa.4300.3.1 - NAVÁS, L. 1916. Neurópteros Sudamericanos. Tercera serie. Neurópteros del Brasil recogidos por el R. P. Joaquín da Silva Tavares S. J. Brotéria (Serie Zoológica) 14:14–35.
- NEEDHAM, J.G. & MURPHY, H.E. 1924. Neotropical mayflies. Bulletin of the Lloyd Library, 24, Entomological Series, 4, 1–79.
-
OTTONI, B.M.P., PINHEIRO, M.P.G., DE ALMEIDA ANDRADE, H.T. & ANDREAZZE, R. 2009. Ocorrência do gênero Campsurus (Ephemeroptera, Polymitarcyidae) no Semi-Árido do Estado do Rio Grande do Norte, Brasil. EntomoBrasilis , 2 (3), 90–91. https://doi.org/10.12741/ebrasilis.v2i3.55.
» https://doi.org/10.12741/ebrasilis.v2i3.55 -
OLIVEIRA, L.A., NASCIMENTO, J.M.C. & COUCEIRO, S.R.M. 2021. A new species of Tricorythopsis Traver, 1958 (Ephemeroptera: Leptohyphidae) from Pará state, Brazil. Zootaxa, 5023(1), 131–141. https://doi.org/10.11646/zootaxa.5023.1.8.
» https://doi.org/10.11646/zootaxa.5023.1.8 -
OLIVEIRA, L.A.D., COUCEIRO, S.R.M. & NASCIMENTO, J.M.C.D. 2023. Ephemeroptera (Insecta) from the metropolitan region of Santarém, Pará, Brazil. Biota Neotropica, 23(1). https://doi.org/10.1590/1676-0611-BN-2022-1437.
» https://doi.org/10.1590/1676-0611-BN-2022-1437 - PEREIRA, S. M. & DA-SILVA, E. R. 1991. Descrição de uma nova espécie de Campsurus Eaton, 1868 do sudeste do Brasil, com notas biológicas (Ephemeroptera: Polymitarcyidae: Campsurinae). Revista Brasileira de Biologia, 51(2), 321–326.
-
PRINCIPE, R. E., MARQUEZ, J. A. & CIBILS-MARTINA, L. 2019. Distribution and habitat preference of Ephemeroptera and Trichoptera in subtropical mountain streams: implications for monitoring and conservation. Anais da Academia Brasileira de Ciências, 91, e20180692. https://doi.org/10.1590/0001-3765201920180692.
» https://doi.org/10.1590/0001-3765201920180692 -
RAIMUNDI, E. A. 2019. Contribution to the knowledge of Ephemeroptera (Insecta) from Goiás State, Brazil. Journal of Insect Biodiversity, 12(2), 33–47. https://doi.org/10.12976/jib/2019.12.2.1.
» https://doi.org/10.12976/jib/2019.12.2.1 -
SALLES, F. F., NASCIMENTO, J. M. C. D., MASSARIOL, F. C., ANGELI, K. B., SILVA, P. B., RÚDIO, J. A. & BOLDRINI, R. 2010. First survey of mayflies (Ephemeroptera, Insecta) from Espírito Santo state, southeastern Brazil. Volume 10, Número 1, Pags. 294–307. http://dx.doi.org/10.1590/S1676-06032010000100025.
» https://doi.org/10.1590/S1676-06032010000100025 -
SALLES, F.F., BRANT, A., MASSARIOL, F.C., GONÇALVES, I.C., LIMA, L.R.C., OLIVEIRA, L.H.M., SOUZA, M.R., FERNANDES, O.L., CRUZ, P.V., BOLDRINI, R., SILVA, R.M.L. & FEITOZA, Y.S. 2023. Priasthenopus gilliesi. Sistema de Avaliação do Risco de Extinção da Biodiversidade - SALVE. Disponível em: https://salve.icmbio.gov.br Digital Object Identifier (DOI): https://doi.org/10.37002/salve.ficha.32715 - Acesso em: 23 de nov. de 2023.
» https://salve.icmbio.gov.br -
SALLES, F.F., BOLDRINI, R. & LIMA, L.R.C. 2024. Polymitarcyidae in Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/1951 Last access on 17/03/2024.
» http://fauna.jbrj.gov.br/fauna/faunadobrasil/1951 -
TAKIYA, D. M., SANTOS, A. P. M., PINTO, Â. P., HENRIQUES-OLIVEIRA, A. L., do Lago Carvalho, A., Sampaio, B. H. L. & RAFAEL, J. A. 2016. Aquatic insects from the caatinga: checklists and diversity assessments of Ubajara (Ceará State) and Sete Cidades (Piauí State) national parks, Northeastern Brazil. Biodiversity data journal, (4). https://doi.org/10.3897%2FBDJ.4.e8354.
» https://doi.org/10.3897%2FBDJ.4.e8354 - ULMER, G. 1920. Übersicht über die Gattungen der Ephemeropteren, nebst Bemerkungen über einzelne Arten. Stettiner Entomologische Zeitung, 81(1–2), 97–144.
- ULMER, G. 1942. Alte und neue Eintagsfliegen (Ephemeropteren) aus Süd- und Mittelamerika. Stettiner Entomologische Zeitung 103:98–128.
- VERÍSSIMO, A. & PEREIRA, D. 2014. Produção na Amazônia Florestal: características, desafios e oportunidades. Parcerias Estratégicas, 19(38), 13–44.
- WALKER, F. 1853. Ephemerinae. List of the specimens of neuropterous insects in the collection of the British Museum. Part III - (Termitidae- Ephemeridae). pp. 533–585.
Publication Dates
-
Publication in this collection
10 Jan 2025 -
Date of issue
2024
History
-
Received
03 June 2024 -
Accepted
25 Nov 2024