Introduction
The superorder Peracarida Calman, 1904 is considered one of the most widespread and adaptable groups among crustaceans. Characterized by direct development, where larvae transition directly into adults without free-swimming nauplius stages, peracarids currently encompass 11 recent orders (Lowry and Myers, Reference Lowry and Myers2013; WoRMS, 2024a), including Tanaidacea Dana, 1849. Tanaidaceans exhibit extensive morphological variety, with their body plan typically dorso-ventrally flattened, cylindrical, semi-cylindrical, or occasionally discoidal (Figure 1). Tanaidaceans have a global distribution, being found in all marine habitats and depths, including estuaries, albeit rarely in freshwater (Larsen, Reference Larsen2005; Błażewicz-Paszkowycz et al., Reference Błażewicz-Paszkowycz, Bamber and Anderson2012, Reference Błażewicz-Paszkowycz, Pabis and Jóźwiak2015; Pabis et al., Reference Pabis, Jóźwiak, Lörz, Schnabel and Błażewicz-Paszkowycz2015). Due to their limited dispersal capabilities, species with wide distribution are uncommon and questionable, but still registered in few groups (e.g., Chondrochelia dubia (Krøyer, Reference Krøyer1842), Neotanais spp., Sinelobus stanfordi Richardson, 1901, Agathotanais ingolfi Hansen, 1913) (Gardiner, Reference Gardiner1975; Sieg, Reference Sieg1986; Larsen, Reference Larsen2005 [Table IV, V, p. 291], Reference Larsen2013; Brandt et al., Reference Brandt, Błażewicz-Paszkowycz, Bamber, Mühlenhardt-Siegel, Malyutina, Kaiser, De Broyer and Havermans2012; Larsen et al., Reference Larsen, Guţu and Sieg2015).
According to Anderson (Reference Anderson2020) and the World Register of Marine Species website (2024b), there are currently 36 families, 316 genera, and 1575 species of recent Tanaidacea. In terms of biodiversity, is estimated that Tanaidacea could surpass 3000 species (Larsen, Reference Larsen2005) and the number of tanaids species might significantly increase in the coming years, mainly due to the expansion of material collected from numerous deep-sea (>200 m depth) expeditions, combined with the increasing interest in studying the group and advancements in taxonomy technology.
The deep-sea environment encompasses over 90% of the entire ocean, yet its extreme conditions make exploration significantly more expensive and challenging (Brandt et al., Reference Brandt, De Broyer, Gooday, Hilbig and Thomson2004) compared to shallower waters, requiring specialized sampling tools (e.g. box-corers, epibenthic sledges, grabs, multicorers). As a result, collecting abiotic and biotic data is often linked to deep-sea expedition efforts conducted by political incentives to survey economic exclusive zones (EEZ), or by oil companies seeking baseline studies for environmental impact assessments.
Brazil is a country of continental dimensions (8516,000 km2) (Machado et al., Reference Machado, Miranda, Salvador, Pimenta, Côrtes, Gomes, Miyahira, Agudo-Padrón, Oliveira, Caetano, Coelho, D’Ávila, Arruda, Almeida, Gomes, Alvim, Galvão-Filho, Ferreira-Júnior, Marques, Martins, Souza, Arruda, Cavallari, Santos, Pedro, Salles, Dornellas, Lima, Amaral, Silva, Passos, Thiengo, Leite and Simone2023), and has an extensive coastal zone, spanning the political regions North, Northeast, Southeast, and South. In the North Region littoral (Amapá and Pará), the coastline is characterized by the vast Amazon River delta, where the river meets the Atlantic Ocean, creating a blend of freshwater and saltwater habitats (Goes and Ferreira Jr, Reference Goes and Ferreira2017); the Northeast Region (Maranhão, Piauí, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, Sergipe, and Bahia) encompass an extensive coastline, with oligotrophic waters and a narrow and shallow continental shelf (40–80 m) (Knoppers et al., Reference Knoppers, Ekau and Figueiredo1999; Goes and Ferreira Jr, Reference Goes and Ferreira2017; Guimarães et al., Reference Guimarães, de Carvalho and Guimarães2020); the Southeast Region littoral (Espírito Santo, Rio de Janeiro and São Paulo) presents the upwelling occurring along the coast of the Rio de Janeiro State, where the cold, nutrient-rich deep water is brought to the surface, influencing the benthic assemblages diversity (Bernardino et al., Reference Bernardino, Berenguer and Ribeiro-Ferreira2016); and the South Region (Paraná, Santa Catarina and Rio Grande do Sul) has a series of coastal ranges with long stretches of beach and very few rocky shores, enclosed lagoons and lagoons connected to the sea (Goes and Ferreira Jr, Reference Goes and Ferreira2017).
Despite Brazil ranks among the 17 most biodiverse countries on the planet, earning the distinction of ‘megadiversity’ (UNEP-WCMC, 2016) and Arthropods represents the group with the highest number of species (Machado et al., Reference Machado, Miranda, Salvador, Pimenta, Côrtes, Gomes, Miyahira, Agudo-Padrón, Oliveira, Caetano, Coelho, D’Ávila, Arruda, Almeida, Gomes, Alvim, Galvão-Filho, Ferreira-Júnior, Marques, Martins, Souza, Arruda, Cavallari, Santos, Pedro, Salles, Dornellas, Lima, Amaral, Silva, Passos, Thiengo, Leite and Simone2023), significant knowledge gaps remain for a substantial part of taxa (Guţu, Reference Guţu1996). Fox example, in the Brazilian EEZ, covering a vast area of 3.6 million km2, ranking 11th globally in terms of size (De Leo et al., Reference De Leo, Bernardino, Sumida, Sumida, Bernardino and De Leo2020), only 63 Tanaidacea species have been described to date (Segadilha, Reference Segadilha2024). Moreover, there are Brazilian areas where knowledge and sampling efforts are lacking, as well as unexplored marine ecosystems such as canyons and slope (Marques and Lamas, Reference Marques and Lamas2006).
This paper is a compilation resulting from the ‘Catálogo Taxonômico da Fauna do Brasil – CTFB’ (Taxonomic Catalogue of the Brazilian Fauna), an online catalogue exclusive on the Brazilian fauna, a project that started in 2015. The CTFB database is constantly updated by over 500 zoological experts in their respective group (Santos et al., Reference Santos, Dumas, Henriques-oliveira, Souza, Camargos, Calor and Pes2020; Machado and Martins, Reference Machado and Martins2022). There are over 130,000 valid animal species represented in the CTFB, and almost 3500 crustaceans (from marine, freshwater and terrestrial environments, excluding fossils) are recorded to Brazilian territory. Currently, a list containing all valid species of Brazilian tanaidacean is presented online. Detailed information about each taxon such as the hierarchy classification of species, year of publication, author's name, taxonomic status of the name, geographic distribution to Brazilian states and ecoregions, and bibliographic references can be freely accessed on the website http://fauna.jbrj.gov.br.
As for the present study, the authors give all information that will facilitate research on the taxonomic diversity of tanaidaceans in Brazil, presenting a comprehensive overview, and highlighting its historical taxonomic advancements. The database is also relevant since it shows Brazilian areas on which collection and research efforts should be concentrated.
Material and methods
Taxonomic information available in the CTFB website (http://fauna.jbrj.gov.br) was obtained from published papers with original descriptions, taxonomic revisions or checklists with species level identification provided. The CTFB database has been constantly updated (lastly updated in February 2024) and the higher-level classification for Tanaidacea adopted in the CTFB follows WoRMS (2024b). Data were extracted and compiled in an Excel spreadsheet, which was the basis for all analyses conducted in this study. Our dataset is organized first by family and then contains a list of species for each of these taxa. Detailed information was obtained for each tanaidacean species described or reported for Brazil: author's name, year of description, complete literature, year of first record in Brazil, if endemic to Brazil, and the distribution in biogeographic regions (ecoregions), and Brazilian states (political subdivision).
Accumulation curve was generated based on the year of descriptions or the year that the species was first reported from Brazil, if this occurred only in subsequent papers. Graphics were generated in Excel and maps in Quantum GIS v. 2.16.3, then edited in Adobe Photoshop 2024.
Results and discussion
Historical Tanaidacea data
The first reference of Tanaidacea from Brazil was Chondrochelia dubia, made by Kröyer as Tanais dubius, representative of the suborder Tanaidomorpha Sieg, Reference Sieg1980, collected on the coast of Bahia (Krøyer, Reference Krøyer1842; Brum, Reference Brum1973, Reference Brum1977). Guţu (Reference Guţu and Young Paulo1998), catalogued 29 species off Brazilian coast, comprising 17 of the suborder Apseudomorpha Sieg, Reference Sieg1980 and 12 of the suborder Tanaidomorpha. Larsen et al. (Reference Larsen, Araújo-Silva and Coelho2009) updated this list up to 41 species, with 26 of Apseudomorpha and 15 of Tanaidomorpha.
Currently, a total of 63 tanaidacean species are registered from Brazil (suppl. Table; Segadilha, Reference Segadilha2024): 30 Apseudomorpha species in 23 genera and six families and 33 Tanaidomorpha species in 23 genera and 12 families (Table 1). All three Tanaidomorphan superfamilies are represented: Neotanaoidea Sieg, Reference Sieg1980, Paratanaoidea Lang, 1949, and Tanaidoidea Nobili, 1906. Despite having nearly the same number of species described as the Apseudomorpha (30 spp.), the Tanaidomorpha (33 spp.) is the suborder with the highest number of families (12 compared to 6; Table 1).
Among the Brazilian apseudomorphan, Kalliapseudidae Lang, Reference Larsen1956 is the most diverse family with eight species (12.7%) divided into six genera, followed by Apseudidae Leach, 1814 and Parapseudidae Guţu, 1981 (six species each, 9.5%, divided into five genera each; Table 2). Among the tanaidomorphan, Typhlotanaidae Sieg, 1984 appears as the most diverse family with ten species (15.9%) divided into four genera, followed by Leptocheliidae Lang, 1973 with six species (9.5%) divided into four genera, and Tanaididae Nobili, 1906 with five species (7.9%) in five genera (Table 2).
Many authors, even foreigners, have been worked with Brazilian material describing new species. Until 2023, the author with more articles published of valid species of Tanaidacea described from Brazil was Juliana Segadilha (16), followed by Kim Larsen (9), Catarina Araújo-Silva, Modest Guţu and Cristiana Serejo (8 each), and Magdalena Błażewicz, Iva Nilce da Silva Brum and Kátia Christol dos Santos (5 each) (Figure 2).
Since the two first species described in Brazil in 1842 and later in 1849 by Dana (Leptochelia brasiliensis Dana, 1849), there was a gap of 100 years until the following species described by Mané-Garzón in 1949 (Monokalliapseudes schubarti Mané-Garzón, 1949) followed by Lang (Reference Lang and Wingstrand1956) (Psammokalliapseudes mirabilis Lang, Reference Lang and Wingstrand1956 and Teleotanais gerlachi Lang, Reference Lang and Wingstrand1956). During the second half of 19th century (1842–1900), six species were recorded from Brazil (Figure 3). Although Tanais dulongii (Audouin, 1826), Hexapleomera robusta (Moore, 1894), Paradoxapseudes intermedius (Hansen, 1895) and Leptochelia forresti (Stebbing, 1896) were first described offshore Brazilian waters, these species were reported to Brazilian coast in the second half of the following century (Brum, Reference Brum1969, Reference Brum1973; Sieg, Reference Sieg1980; Guţu, Reference Guţu and Young Paulo1998) (see supp. Table).
During the 20th century, a total of 33 species were recorded in Brazil. There was a slow growth on the cumulative number of tanaidacean species described from Brazil per year until the 1970s, with the Brazilian author Brum's works describing four species (Brum, Reference Brum1971, Reference Brum1973), among other authors (Figure 3; 1971–1975). After that, the second major increase occurred at the end of the 20th century, when Guţu (Reference Guţu1996) published the major work about Brazilian Tanaidacea describing eight new species (Figure 3; 1996–2000). The growth has reached the average of 3.0 species described per 5 years in the last 20 years.
By the beginning of the 21st century (2001–present), 30 species have already been described, totalling 63 species reported to Brazil. The third major increase in the cumulative number of tanaidacean species was due mainly to two Brazilian authors: Santos, with four new species described (Santos, Reference Santos2007; Santos & Hansknecht, Reference Santos and Hansknecht2007) and Araújo-Silva, with three new species described (Larsen et al., Reference Larsen, Araújo-Silva and Coelho2009; Araújo-Silva and Larsen, Reference Araújo-Silva and Larsen2010) (Figure 3; 2006–2010). At this point until nowadays, the growth has reached the higher average, with 7.25 species described per 5 years: six new species described until 2015 (five by Araújo-Silva and Larsen, Reference Araújo-Silva and Larsen2012b; and Araújo-Silva et al., Reference Araújo-Silva, Coelho and Larsen2013) (Figure 3; 2011–2015), seven described until 2020 (Segadilha et al., Reference Segadilha, Gellert and Błażewicz2018, Reference Segadilha, Serejo and Błażewicz2019; Segadilha and Serejo, Reference Segadilha and Serejo2020) (Figure 3; 2016–2020), and nine described until the present moment (Iwasa-Arai et al., Reference Iwasa-Arai, Siqueira, Segadilha and Leite2021; Segadilha and Serejo, Reference Segadilha and Serejo2022; Segadilha et al., Reference Segadilha, Bird and Tavares2023) (Figure 3; 2021–2023). This steep increase suggests that the total number of Tanaidacea species in Brazil is much greater than what was recorded in previous centuries, with less than a quarter of the 21st century almost 50% of the total species (30 species from 63) have been described.
It is relevant to emphasize that among these six points of great increase in the accumulation curve, five were due to four Brazilian authors, all of them women who were without permanent positions (such as professors in institutions), meaning that they were early-career researchers who received only temporary research scholarships, but who were essential for the tanaidaceans studies in Brazil. This highlights the importance of government investment in science, promoting equal opportunities between genders in academic careers and the need to encourage women in the field of taxonomy.
Tanaidacea distribution across Brazil
From the 63 species currently registered in Brazilian waters, only one species occurred in freshwater (Halmyrapseudes spaansi Băcescu & Guţu, 1975; 1.6%, suppl. Table) and four species were recorded from brackish waters (estuary and mangroves; 6.3%, suppl. Table). According to Larsen & Hansknecht (Reference Larsen and Hansknecht2004) and Błażewicz-Paszkowycz et al. (Reference Błażewicz-Paszkowycz, Bamber and Anderson2012), while 188 Tanaidaceans are predominantly found in the marine habitats, there are also some records from 189 brackish and freshwater habitats. The limited presence of brackish water species and the 190 specific conditions regarding the freshwater species indicate that tanaidaceans have not been 191 successful in colonizing inland waters (Larsen, Reference Larsen2005; Larsen et al., Reference Larsen, Guţu and Sieg2015).
Regarding the marine environment, 38 species were recorded to shallow waters (coastal waters) and continental shelf (suppl. Table), whereas about 1/3 was registered to the deep-sea (24 species; suppl. Table), with five occurring in both shallow and deep waters: Intermedichelia gracilis Guţu, Reference Guţu1996; I. jesseri Araújo-Silva & Larsen, 2012; Hamatipeda prolata Segadilha et al., Reference Segadilha, Serejo and Błażewicz2019; Makraleptochelia potiguara Araújo-Silva & Larsen, 2012 and Typhlotanais ischnochela Segadilha and Serejo, Reference Segadilha and Serejo2022. However, these results may be subject to bias as deep-sea studies have only begun to increase more recently. The first paper on tanaidacean found on the Brazilian slope was published by Guţu (Reference Guţu1996), followed by samples collected from abyssal plains 13 years later by Larsen et al. (Reference Larsen, Araújo-Silva and Coelho2009). This suggests that in the near future this difference may reduce or even the number of Brazilian deep-sea tanaidaceans may be greater than those from shallow areas.
Apseudomorphan species are predominantly shallow-waters taxa, with few groups extending their distribution to slope depths (Błażewicz-Paszkowycz et al., Reference Błażewicz-Paszkowycz, Bamber and Anderson2012). In Brazil, from the 30 species registered, most occurred between 0 and 100 m depth (24 spp.; Figure 4), being from family Kalliapseudidae the most representative with nine species, followed by Metapseudidae with five species, Apseudidae and Parapseudidae with four species each and Pagurapseudidae with two. Only six species occurred below 200 m depth: three species of Sphyrapodidae Guţu, 1980 (Kudinopasternakia amazonica; K. brasiliensis and K. trispinosa), two Apseudidae (Carpoapseudes prospectnes and Atlantapseudes brasiliensis) and one Parapseudidae (Leptolicoa thokozele) (Figure 4) - for taxon authority see suppl. Table.
According to Błażewicz-Paszkowycz et al. (Reference Błażewicz-Paszkowycz, Bamber and Anderson2012), within Tanaidomorpha, four families are predominantly shallow-water: Tanaididae, Leptocheliidae, Mirandotanaidae Błażewicz-Paszkowycz and Bamber, 2009 and Paratanaidae Lang, 1949. In the Brazilian coast, 22 species were registered until 100 m depth, being Leptocheliidae the most representative (6 spp.), followed by Tanaididae (5 spp.), and Paratanaidae (2 spp.). Families represented in deep-sea in Brazil are: Typhlotanaidae, Tanaellidae, Neotanaidae, Akanthophoreidae, and Hamatipedidae (Figure 5). The species Typhlotanais ischnochela and Hamatipeda prolata had a wider range of depth occurring from continental shelf to lower slope (46–1898.7 m and 77–1310 m, respectively).
With this compilate, it was possible to carry out a regional analysis of the diversity of Brazilian tanaids and track the under-sampling in some Brazilian areas. According to the present literature revision, the Southeast Region of Brazil holds the majority of Tanaidacea records with 46 occurrences (with 7 species occurring concomitantly in the South and 6 in the Northeast Region), followed by the Northeast Region with 19 occurrences. Poorly known Brazilian regions are the South with nine and the North with only two registers so far (Figure 6), though no region is considered adequately known. Moreover, the number of tanaidacean specialists seems to be directly related to the number of species records per region. The highest numbers of records are mainly due to localities in Brazil where taxonomists are based: Santos, Segadilha and Brum for Southeast Region, and Araújo-Silva for Northeast Region of Brazil (e.g. Brum, Reference Brum1971, Reference Brum1978; Santos, Reference Santos2007, Reference Santos2014; Araújo-Silva and Larsen, Reference Araújo-Silva and Larsen2010, Reference Araújo-Silva and Larsen2012a, Reference Araújo-Silva and Larsen2012b; Araújo-Silva et al., Reference Araújo-Silva, Coelho and Larsen2013; Segadilha et al., Reference Segadilha, Serejo and Błażewicz2019, Reference Segadilha, Bird and Tavares2023; Segadilha and Serejo, Reference Segadilha and Serejo2020, Reference Segadilha and Serejo2022).
The locations of oil prospecting sites where energy companies worked in Brazilian offshore certainly influenced the distribution of species records within the country. The states of Espírito Santo (12 spp.), Rio de Janeiro (35 spp.) and São Paulo (13 spp.) were the three most speciose states in Brazil (Figure 7), corresponding to the areas where the Petrobras Company oil conducted big deep-sea fauna projects in the last two decades (Falcão et al., Reference Falcão, Curbelo-Fernandez, Borges, Filgueiras, Kowsmann, Martins, Curbelo-Fernandez and Braga2017; Segadilha and Serejo, Reference Segadilha and Serejo2022). Among the Brazilian coastline states, only Amapá, Maranhão, and Piauí have no records of Tanaidacea until now. In general, the North Region in Brazil is the least explored area in the country, with only two species registered to Pará state (Figure 7). This disparity is certainly the result of a lack of studies targeting local biodiversity, but it is also due to the Amazon river plume, which might have influence on the local fauna, changing the composition of species (Sumida et al., Reference Sumida, Bernardino and De Leo2020; Targino and Gomes, Reference Targino and Gomes2020). Nevertheless, further studies need to be conducted in these areas to confirm this influence on the number and the composition of tanaidacean species in the Brazilian North Region.
For instance, in Brazil, there is a significant amount of material from both deep-sea and shallow waters in zoological collections that remain unpublished (e.g. MNRJ, MZUSP, UFPE; C. Serejo, M. Tavares and J. Souza-Filho – pers. comm., respectively). This highlights that the lack of specialists remains an obstacle to the knowledge of the group, failures in sampling and/or treatment, and, most critical, funding for the development of pure science (Błażewicz-Paszkowycz et al., Reference Błażewicz-Paszkowycz, Bamber and Anderson2012).
Conclusions
Our results highlight the increasing number of publications and new Brazilian species of Tanaidacea in the past 15 years, and this is directly correlated to where the specialists in alpha-taxonomy, systematics/phylogeny, and ecology are based. There is a vast sum of deep-sea and shallow-water material in the Brazilian zoological collections, however, remains unpublished.
Knowing about the current status of Brazilian taxonomy is a required step to establish future strategies to develop unfilled areas and overlooked groups (Marques and Lamas, Reference Marques and Lamas2006). An important key to understanding the biodiversity of a country is the ability to form taxonomists, as well as investment in science and technology. However, the taxonomy is currently maintained in a weak appreciation (Giangrande, Reference Giangrande2003), reflecting the worlds lack of interest in the area. As indicated by our current dataset, one important aspect that must be addressed in future studies is the lack of taxonomic experts of Tanaidacea, especially in Brazil.
None of the Brazilian specialists in the order Tanaidacea had permanent positions. All previous studies were carried out by students and researchers whose publications were funded by their own scholarships, one of the greatest problems faced by taxonomists in Brazil. And that is the Brazilian biodiversity challenge: an impasse between the lack of taxonomists and the urgent need for knowledge of our biodiversity, especially because many wild habitats are rapidly disappearing.
We conclude that the lack of specialists is still an obstacle to the future knowledge of the order, as well as failures in sampling and/or treatment and, most critical, funding for the development of alpha-taxonomy and pure science in Brazil. Currently, there is no research group in Brazil for order Tanaidacea and this must be changed soon. Authors hope that this paper will stimulate further studies on this group in the country, which is in urgent need of experts.
Finally, the overall knowledge on the Brazilian tanaidaceans still contains gaps that should be addressed in future studies. One great issue is the under sampled regions in the country. Future research efforts should focus on areas such as the North and South regions, although Southeast and Northeast regions should not be left out, as it is unlikely that the local fauna is not even close to being cataloged.
Supplementary material
The supplementary material for this article can be found at https://doi.org/10.1017/S0025315424000808.
Acknowledgements
The authors would like to thank Dr Kátia Christol dos Santos (MZUSP) for supporting our idea and constructive discussions. We also thank the contributions of the anonymous referees for their highly valuable comments on the manuscript.
Data
Data supporting the conclusions of this article are available in full from the references listed below.
Author contributions
JLS concept of the manuscript, data analysis, manuscript writing, and discussion. CLAS concept of the manuscript, data analysis, manuscript writing, and discussion. All authors contributed to the draft provided critical feedback and helped shape the research. All authors read and approved the final manuscript.
Financial support
This work was supported by the funding agency CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) which provided the fellowship for the senior author – JL Segadilha during the CTFB database compilation. Junior author – CL Araújo-Silva thanks to her post-doctoral research funded by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) (Grant n◦ APQ-0755-1.08/16) and Fundação de Amparo a Ciência e Tecnologia do Estado de Pernambuco (FACEPE) (Grant n◦ BCT-0234-1.08/17).
Competing interests
None.
Ethical standards
Not applicable.