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Chondrichthyan systematics in Brazil depicted: historical overview, research trends and future perspectives

Abstract

Brazil comprises one of the world’s biodiversity hotspots of the chondrichthyan fauna, currently with 12 orders, 44 families, 90 genera and over 200 species of sharks, batoids, and chimaeras. These species inhabit marine, estuarine and freshwater realms of coastal and oceanic zones from the North equatorial central to the South-western Atlantic Ocean. Reporting on species occurrence in Brazil goes back to the pre-Linnaean Period. The golden era of the European Zoological studies put in place the knowledge on the local fauna with many Brazilian endemic and native species being officially described. The birth of Ichthyology in Brazil in the early 20th century, however, represents the milestone for the Chondrichthyan Systematics in the country when national scientists published the first list of species. This study provides an historical overview of the Chondrichthyan Systematics research in Brazil through a literature review of peer-reviewed publications in shark taxonomy, phylogenetics and morphology. Scientific trends of over 120-year period of dedicated investigations are herein pointed out regarding research scope and subject area, methodology, target study taxa, and gender diversity. Research recommendations and priorities are further given to assist researchers and interested stakeholders on future efforts in shark science and conservation in the country.

Keywords:
Biodiversity research; Brazilian waters; Elasmobranchii; Holocephali; Taxonomy

Resumo

O Brasil compreende um hotspot de biodiversidade da fauna de peixes Chondrichthyes, atualmente com 12 ordens, 44 famílias, 90 gêneros e mais de 200 espécies reconhecidas de tubarões, raias batóides e quimeras. Estas espécies habitam ambientes marinhos, estuarinos e dulcícolas de zonas costeiras e oceânicas do Oceano Norte Equatorial ao Atlântico Sudoeste. Registros da ocorrência de espécies no Brasil remonta do período Pré-Lineano. A era de ouro dos estudos zoológicos na Europa contribuiu para o conhecimento da fauna local com a descrição oficial de várias espécies nativas e endêmicas. O nascimento da Ictiologia no Brasil no início do século 20, contudo, representa o marco para a Sistemática de Chondrichthyes, quando cientistas nacionais publicaram a primeira lista de espécies registradas no país. Este estudo apresenta uma leitura histórica da pesquisa em Sistemática de Chondrichthyes no Brasil através de uma revisão da literatura de artigos revisados por pares sobre a taxonomia, filogenética e morfologia de peixes cartilaginosos. Tendências em pesquisa para o período de mais de 120 anos de dedicadas investigações científicas são aqui apontadas com relação ao escopo, área de estudo, metodologia, taxa-alvo estudado, e diversidade de gênero. Recomendações e prioridades em pesquisa são também indicadas para assistir cientistas e outras partes interessadas em esforços futuros da ciência e conservação de tubarões, raias e quimeras no país.

Palavras chave:
Águas brasileiras; Elasmobranchii; Holocephali; Pesquisa em biodiversidade; Taxonomia

INTRODUCTION

Brazil represents one of the world’s largest biodiversity hotspots of chondrichthyan fishes (Lucifora et al., 2011Lucifora LO, García VB, Worm B. Global diversity hotspots and conservation priorities for sharks. PLoS ONE. 2011; 6(4):e19356. https://doi.org/10.1371/journal.pone.0019356
https://doi.org/10.1371/journal.pone.001...
; Dulvy et al., 2014Dulvy NK, Fowler SL, Musick JA, Cavanagh RD, Kyne PM, Harrison LR et al. Extinction risk and conservation of the world’s sharks and rays. Elife. 2014; e00590(3):1–35. https://doi.org/10.7554/eLife.00590.001
https://doi.org/10.7554/eLife.00590.001...
). It is estimated that the country currently comprises 12 orders, 44 families, 90 genera (Fricke et al., 2023)Fricke R, Eschmeyer WN, van der Laan R. Eschmeyer’s catalog of fishes: genera, species, references [Internet]. San Francisco: California Academy of Science; 2023. Available from: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp.
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and over 200 species (SV, FFP, KS, work in progress). National sharks, batoids, and chimaeras are distributed along the equatorial tropical to temperate South-western Atlantic Ocean in a wide range of ecosystems, occurring in freshwater, brackish, and marine waters of inland river basins, and coastal and offshore zones as well as insular shelves and deep oceanic slopes. The gigantic profile of freshwater and marine realms under the Brazilian Federal jurisdiction have fostered a wide variability of habitats and phenotypic characters, species richness, high endemicity, and, possibly, revealing the exceptional evolutionary adaptability and the chondrichthyan diversity in the country.

Evidence of Chondrichthyes species in Brazil date back to the colonial time as it was mentioned in Pero Vaz de Caminha’s letters relating the Portuguese expeditions to the “New World” under Pedro Álvares Cabral leadership in the 16th century, as well as through the narratives of Frei Cristóvão de Lisboa (Rosa, 2009)Rosa RS. Elasmobranchii. In: da Rocha RM, Boeger WA, editors. Estado da arte e perspectivas para a zoologia no Brasil. Vol. 2. Curitiba: UFPR; 2009. p.56–62.. Pre-Linnaean sources that specifically accounted for the diversity of South American fishes such as Marcgrave (1648)Marcgrave G. Historiae rerum naturalium Brasiliae. In: Piso G, Marcgrave G, editors. Hist. Nat. Bras. auspicio Benef. Ilus. I. Mavritii Com. Nassav illius Prov. maris summi praefecti adornata. Leiden: F. H. Haack & L. Elzevier; 1648. and Willughby (1686)Willughby F. De historia piscium libri quatuor, jussu & sumptibus Societatis Regiæ Londinensis editi. Totum opus recognovit, coaptavit, supplevit, librum etiam primum & secundum integros adjecit Johannes Raius e Societate Regia. Oxonii (Oxford): Theatro Sheldoniano; 1686. included Brazilian records of sharks, batoids and chimaeras at that time. Progress in zoological studies and the Systematics foundation during the 18th century boosted the recognition of the first native nominal species (Rosa, 2009)Rosa RS. Elasmobranchii. In: da Rocha RM, Boeger WA, editors. Estado da arte e perspectivas para a zoologia no Brasil. Vol. 2. Curitiba: UFPR; 2009. p.56–62.. Ichthyological Systematics in the country commenced with the contributions of foreign naturalists and collection of specimens through scientific expeditions conducted during the 19th century. Nevertheless, its effective implementation arose much later in the beginning of the 20th century through the works of Alípio de Miranda-Ribeiro, the father of the Brazilian Ichthyology. Today, Brazil represents a solid Chondrichthyan research hub and accounts for one of the largest scientific communities, especially regarding taxonomic, phylogenetic, and biogeographic studies of marine and freshwater shark species. This scenario reflects 120 years of historical contributions to the Chondrichthyan biodiversity and evolutionary knowledge known to date. The efforts of the Sociedade Brasileira para o Estudo de Elasmobrânquios (SBEEL), which turned 39 years of foundation in 2023, also uplifted this scientific field through the dissemination of shark research, contribution to capacity building, and by strengthening conservation awareness and management countrywide and regionally.

Biodiversity knowledge and museum specimen data generated from Systematics research are an essential tool to inform conservationists and decision makers for better management of Chondrichthyan species, which are naturally vulnerable to threats such as overexploitation, habitat degradation, climate change and pollution (Stein et al., 2018)Stein RW, Mull CG, Kuhn TS, Aschliman NC, Davidson LNK, Joy JB et al. Global priorities for conserving the evolutionary history of sharks, rays and chimaeras. Nat Ecol Evol. 2018; 2(2):288–98. https://doi.org/10.1038/s41559-017-0448-4
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. It is thus imperative to pinpoint the progress in Chondrichthyan Systematics research so far to guide scientific policy and implementation, allocate scientific and research funding, improve local human capacity and strengthen knowledge-based information for conservation purposes. The present study thus aimed to chronologically retrieve the historical contributions to the Chondrichthyan Systematics in Brazil through a comprehensive research overview since 1903, reveal current research trends and provide recommendations for future research.

MATERIAL AND METHODS

Historical overview. A combination of data taken from literature review and personal communications of experts, curators, and collection managers were used to reconstruct the history of the Chondrichthyan Systematics in Brazil. Peer-reviewed scientific studies (e.g., articles, books, book chapters, guides, checklists) related to the taxonomy, phylogeny, and comparative anatomy of extant taxa from Brazil were incorporated into the analysis. We adopted the terminology ‘shark research’ to include all studies related to sharks, batoids and chimaeras. Fisheries reports or data from fishery statistics and abstracts of scientific meetings were excluded. Literature review included publications from 1903 to 2023 and were compiled from the “Portal de Periódicos da Capes” (an online scientific database of scientific publications available to Brazilian institutions), Google Scholar, and Web of Science. Additionally, personal libraries of experts were consulted to incorporate early literature that were absent in online repositories.

The following English keywords were used for searching these databases: “shark”, “rays”, “skates”, “stingrays”, “freshwater rays”, “elasmobranch”, “taxonomy”, “phylogeny”, “anatomy”, “Chondrichthyes”, “Southwestern Atlantic Ocean” and “Brazil”, besides their Portuguese translations: “tubarão”, “raias”, “raias com ferrão”, “raias de água doce”, “elasmobrânquios”, “taxonomia”, “filogenia”, “anatomia”, “Oceano Atlântico Sudoeste” and “Brasil”.

The historical overview was subdivided into different time periods from the Linnaean to the Modern eras. We also cited pre-Linnaean resources as these were relevant to trace the primary native Brazilian nominal species described in later taxonomic accounts. “A coleção de Peixes do Museu Nacional do Rio de Janeiro” (Schreiner, Miranda-Ribeiro, 1903)Schreiner C, Miranda-Ribeiro A. A coleção de peixes do Museu Nacional do Rio de Janeiro. Arq Mus Nac. 1903; 12:1–108., the first national publication in Brazilian Ichthyology, accounts for the origin of Ichthyological Systematics for historical purposes.

Analyses of research trends. We determined the year 1903 as a starting date to gather and retrieve data for analysis of research trends within the country. To reveal research trends in Shark research, detect knowledge gaps and identify research challenges, each study was classified as follows: A) research subject area: taxonomy, morphology, phylogeny; B) research scope: description, revision, checklist, identification guide, first record, new record, DNA barcoding, comparative anatomy, descriptive anatomy, molecular species delimitation, high- and low-level interrelationships; C) data source: morphological, molecular or integrative (more than one source of evidence) (definitions according to Mayr et al., 1953Mayr E, Linsley EG, Usinger RL. Methods and principles of systematics zoology. New York: McGraw Hill Book Company; 1953. ). Target study groups were classified by family and “multi-taxa” was applied to denote those studies whose target taxa comprised more than two families. We also ranked gender diversity in publications by examining the total number of female and male authors per publication either as first/last authors isolated or first and last authors combined. Gender definition was established on the author’s first names and followed a binary male/female score system. The complete list of papers analyzed is provided in Tab. S1.

RESULTS

Brazilian historical overview. Linnaean period.The earliest reporting of Brazilian Chondrichthyan species is available in the pre-Linnaean resources of Marcgrave (1648)Marcgrave G. Historiae rerum naturalium Brasiliae. In: Piso G, Marcgrave G, editors. Hist. Nat. Bras. auspicio Benef. Ilus. I. Mavritii Com. Nassav illius Prov. maris summi praefecti adornata. Leiden: F. H. Haack & L. Elzevier; 1648. and Willughby (1686)Willughby F. De historia piscium libri quatuor, jussu & sumptibus Societatis Regiæ Londinensis editi. Totum opus recognovit, coaptavit, supplevit, librum etiam primum & secundum integros adjecit Johannes Raius e Societate Regia. Oxonii (Oxford): Theatro Sheldoniano; 1686.. Eight species were listed as vernacular in Marcgrave’s study: “Puraquê”, “Araguagua”, “Cucuri”, “Aiereba”, “Jabebiretê”, “Raja sp. altera (jabebara)”, “Narinari”, and “Tiburonis sp. minor” (Miranda-Ribeiro, 1907)Miranda-Ribeiro A. Fauna Braziliense. Peixes. Tomo II, Desmobranchios. Arch Mus Nac. 1907; 14:137–71.. These species are currently assigned to the following valid nominal species: Pseudobatos percellens (Walbaum, 1792) for “Puraquê”, Pristis pristis (Linnaeus, 1758Linnaeus C. Systema Naturae. Regnum Animale. 10th ed. Cura Societatis Zoologicae Germanicae Iterum Edita; 1758. ) for “Araguagua”, Scyliorhinus cabofriensis Soares, Gomes & Carvalho, 2016 for “Cucuri”, Paratrygon aiereba (Walbaum, 1792) (not Müller, Henle, 1841Müller J, Henle FGJ. Systematische beschreibung der Plagiostomen. Berlin: Veit und Comp.; 1841. ) for “Aiereba”, Hypanus guttatus (Bloch & Schneider, 1801) for “Jabebiretê”, Aetobatus narinari (Euphrasen, 1790Euphrasen BA. Raja (Narinari). Kongliga Vetenskaps-Academiens Handlingar. 1790; 11:217–19.)for “Narinari”, and Sphyrna tiburo (Linnaeus, 1758Linnaeus C. Systema Naturae. Regnum Animale. 10th ed. Cura Societatis Zoologicae Germanicae Iterum Edita; 1758. ) for “Tiburonissp.minor”.

Linnaeus (1758)Linnaeus C. Systema Naturae. Regnum Animale. 10th ed. Cura Societatis Zoologicae Germanicae Iterum Edita; 1758. in Systema Naturae apparently did not examine Brazilian specimens at that time but he referred to Marcgrave and Willughby’s accounts to describe S. tiburo. Because syntypes of S. tiburo are unknown and its type-locality is regarded as “America” at its best this species is not considered to be the first nominal species described from Brazilian waters. Other species originally described from elsewhere in Linnaeus (1758)Linnaeus C. Systema Naturae. Regnum Animale. 10th ed. Cura Societatis Zoologicae Germanicae Iterum Edita; 1758. that are known to occur in Brazil today through subsequent works are: Gymnura altavela (Linnaeus, 1758),Squalus acanthias Linnaeus, 1758, Carcharodon carcharias (Linnaeus, 1758), Prionace glauca (Linnaeus, 1758), Galeorhinus galeus (Linnaeus, 1758), Sphyrna zygaena (Linnaeus, 1758), and Callorhinchus callorynchus (Linnaeus, 1758). The Spotted eagle ray or “raia-chita” A. narinari was described by Euphrasen (1790)Euphrasen BA. Raja (Narinari). Kongliga Vetenskaps-Academiens Handlingar. 1790; 11:217–19. based on ‘Narinari Brasiliensibus’ from Brazil of Marcgrave (1648:175)Marcgrave G. Historiae rerum naturalium Brasiliae. In: Piso G, Marcgrave G, editors. Hist. Nat. Bras. auspicio Benef. Ilus. I. Mavritii Com. Nassav illius Prov. maris summi praefecti adornata. Leiden: F. H. Haack & L. Elzevier; 1648. and later reproduced in Willughby (1686:66–67)Willughby F. De historia piscium libri quatuor, jussu & sumptibus Societatis Regiæ Londinensis editi. Totum opus recognovit, coaptavit, supplevit, librum etiam primum & secundum integros adjecit Johannes Raius e Societate Regia. Oxonii (Oxford): Theatro Sheldoniano; 1686., but also included specimens from St. Barthelémy in the Caribbean Sea. The type-locality of this species was undoubtedly reiterated in Euphrasen (1792)Euphrasen BA. Raja narinari. Der Königlich Schwedischen Akademie der Wissenschaften neue Abhandlungen aus der Naturlehre, Haushaltungskunst und Mechanik. 1792; 11:205–07., Kottelat (2013)Kottelat M. The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles B Zool. 2013; 27:1–663. https://doi.org/10.1080/02701367.2001.10608936
https://doi.org/10.1080/02701367.2001.10...
and again in Fricke et al. (2023)Fricke R, Eschmeyer WN, van der Laan R. Eschmeyer’s catalog of fishes: genera, species, references [Internet]. San Francisco: California Academy of Science; 2023. Available from: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp.
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. The lectotype (lost) for this nominal species, an illustration provided in Marcgrave’s account, was later designated in Kottelat (2013)Kottelat M. The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles B Zool. 2013; 27:1–663. https://doi.org/10.1080/02701367.2001.10608936
https://doi.org/10.1080/02701367.2001.10...
. Aetobatus narinari thus represents the first native nominal species described from Brazil according to this evidence and based on the Principle of Priority of the International Code of Zoological Nomenclature (ICZN, 1999)International Code of Zoological Nomenclature (ICZN). International Code of Zoological Nomenclature. 4th ed. London, UK: The International Trust for Zoological Nomenclature; 1999. . Pseudobatos percellens was described later by Walbaum (1792)Walbaum J. Petri Artedi sueci genera piscium. In quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Rose, Grypeswaldiae [Greifswald]: 1792. thus does not representing the earliest native nominal species described for the country as stated in Rosa (2009)Rosa RS. Elasmobranchii. In: da Rocha RM, Boeger WA, editors. Estado da arte e perspectivas para a zoologia no Brasil. Vol. 2. Curitiba: UFPR; 2009. p.56–62. and Gadig, Rosa (2014)Gadig OFB, Rosa RS. Diversity and conservation of the Brazilian marine Chondrichthyes: an overview. Program Poster Abstr Shark Int. Durban. 2014. . A third species, Myliobatis jussieui Cuvier, 1829, was also described based on pre-Linnaean sources as of Jussieu (1721)Jussieu A. Histoire de l’Académie royale des sciences. Paris: 1721. .

Bonnaterre (1788)Bonnaterre L. Tableau Encyclopédique et Méthodique des trois règnes de la nature, dédié et présenté a M. Necker, Ministre d’Etat & Directeur Général des Finances. Paris: 1788. in his “Tableau encyclopédique et méthodique des trois regnes de la nature Ichthyologie” also contributed to the knowledge of the local Chondrichthyan fauna when describing the following species: M. mobular (Bonnaterre, 1788Bonnaterre L. Tableau Encyclopédique et Méthodique des trois règnes de la nature, dédié et présenté a M. Necker, Ministre d’Etat & Directeur Général des Finances. Paris: 1788. ), Alopias vulpinus (Bonnaterre, 1788), Lamna nasus (Bonnaterre, 1788), Ginglymostoma cirratum (Bonnaterre, 1788), Heptranchias perlo (Bonnaterre, 1788), Hexanchus griseus (Bonnaterre, 1798), Echinorhinus brucus (Bonnaterre, 1788), and Dalatias licha (Bonnaterre, 1788). Other species described during this period included Cetorhinus maximus (Gunnerus, 1765), Pristis pectinata Latham, 1794 and Mobula birostris (Walbaum, 1792). However, these species were only recognized in Brazilian waters much later in the 19th century with the efforts of other fish taxonomists. A possible explanation for this is that scientific expeditions to the “New World” were extremely rare during that time due to the political exclusivity over the Brazilian waters and lands under the Portuguese Kingdom, and usually were associated to Portuguese, French and Dutch settlements’ expeditions (Vanzolini, 1996)Vanzolini PE. A contribuição zoológica dos primeiros naturalistas viajantes no Brasil. Rev USP. 1996; 30(30):190–238.. Two main political and societal events completely changed this scenario in the following century: the advent of the Portuguese Royal family to the country in 1808 in Rio de Janeiro; and in 1817 over the influence of the Austrian archduchess, D. Leopoldina, who officially implemented the zoological research in Brazil accompanied by a European scientific committee (Papavero, 1971Papavero N. Essays on the history of Neotropical dipterology: with special reference to collectors (1750-1905). São Paulo: Museu de Zoologia, Universidade de São Paulo; 1971.; Vanzolini, 1996Vanzolini PE. A contribuição zoológica dos primeiros naturalistas viajantes no Brasil. Rev USP. 1996; 30(30):190–238.).

Post-Linnaean period. Intensive naturalist expeditions in the South Atlantic Ocean and the South American rivers during the 19th century resulted with the description of the majority of the Brazilian natives and/or endemics, totalizing 39 nominal species of sharks and rays; for instance, Isistius brasiliensis (Quoy & Gaimard, 1824), Rhinoptera brasiliensis Müller, 1836, Rhizoprionodon lalandii (Valenciennes, 1839), Zapteryx brevirostris (Müller & Henle, 1841), Narcine nigra Duméril, 1852, Potamotrygon henlei (Castelnau, 1855Castelnau FL. Poissons. In: Bertrand P, editor. Animaux Nouveaux ou rares recueillis pendant l’expédition dans les parties centrales de l’Amérique du Sud, de Rio Janeiro a Lima, et de Lima au Para; exécutée par ordre du Gouvernement Français pendant les années 1843 a 1847. Part 7, Zoologie; Paris. 1855; 2:1–112.), and P. constellate (Vaillant, 1880). The diversity of Brazilian native/endemic freshwater batoids was intensively acknowledged with the studies of Müller, Henle (1841)Müller J, Henle FGJ. Systematische beschreibung der Plagiostomen. Berlin: Veit und Comp.; 1841. , Schomburgk (1843)Schomburgk RH. “The” Natural history of the fishes of Guiana (Vol. 1). WH Lizars; 1843., Castelnau (1855)Castelnau FL. Poissons. In: Bertrand P, editor. Animaux Nouveaux ou rares recueillis pendant l’expédition dans les parties centrales de l’Amérique du Sud, de Rio Janeiro a Lima, et de Lima au Para; exécutée par ordre du Gouvernement Français pendant les années 1843 a 1847. Part 7, Zoologie; Paris. 1855; 2:1–112. and Garman (1913)Garman S. The Plagiostomia (Sharks, Skates and Rays). Mem Mus Comp Zool Harvard Coll. 1913; 36:1–528. while marine species were more sporadically known after Müller (1836)Müller J. Vergleichende Anatomie der Myxinoiden, der Cyclostomen mit durchbohrtem Gaumen. Erster Theil. Osteologie und Myologie. Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin. 1836. p.65–340. , Olfers (1831)Olfers JFM von. Die Gattung Torpedo in ihren naturhistorischen und antiquarischen Beziehungen erläutert. Berlin: 1831., Ranzani (1839)Ranzani C. De novis speciebus piscium. Dissertatio prima. Novi Commentarii Academiae Scientiarum Instituti Bononiensis v. 4. 1839., and Regan (1903)Regan C. On a collection of fishes made by Dr. Goeldi at Rio Janeiro. Proc Zool Soc London. 1903; 2(pt. 1):59–68.. These efforts clearly evidenced from the upsurge of the Zoological research in Europe and North America that also had impacted the country.

Exploration of the Brazilian waters was undertaken by foreign naturalists from European and North American museums or universities, mainly as part of governmental and private efforts to stimulate Zoological research. PA Delalande who accompanied the botanist A de Saint-Hilaire from the Muséum National d’Histoire Naturelle onboard the L’Hermione (1816–1822) came to Brazil (Papavero, 1971)Papavero N. Essays on the history of Neotropical dipterology: with special reference to collectors (1750-1905). São Paulo: Museu de Zoologia, Universidade de São Paulo; 1971. and collected elasmobranchs (e.g., Rioraja agassizii (Müller & Henle, 1841), R. lalandii) in several Brazilian provinces, including the coastal areas of Rio de Janeiro, Espírito Santo, São Paulo, Paraná, Rio Grande do Sul and Santa Catarina. The L’Uranie (1817–1820) expedition around the world with Quoy and Gaimard describing the cookiecutter shark I. brasiliensis. The expedition “Les parties centrales de L’Amérique du Sud: Rio de Janeiro a Lima et de Lima au Para” (1843–1847) under leadership of FL de Castelnau and support from the French government resulted on the listing of nine nominal species of native elasmobranchs and description of four novel freshwater species collected in Rio de Janeiro, Bahia, Araguaia and Tocantins River basins. The Thayer Expedition to Brazil (1865–1866) under leadership of the Swiss ichthyologist L Agassiz from the Museum of Comparative Zoology – Harvard University, USA (MCZ), collected over 120 specimens in Bahia, Ceará, Piauí, Amazonas, Pará and Rio de Janeiro. Agassiz and his wife narrated their tropical experience in Agassiz, Agassiz (1868)Agassiz L, Agassiz EC. A Journey in Brazil. Boston: University Press: Welch, Bigelow & Co.; 1868. and described with passion their close relationship with D. Pedro II who fully supported this expedition at the point to collect a specimen of Sphyrna lewini (Griffith & Smith, 1834) (MCZ S–314, Rio Grande do Sul, Brazil). Agassiz published several fish species accounts but did not describe any elasmobranch species. His collection was later studied in Garman (1913)Garman S. The Plagiostomia (Sharks, Skates and Rays). Mem Mus Comp Zool Harvard Coll. 1913; 36:1–528., who made one of the most exhaustive scientific inputs to the Chondrichthyan Systematics until then. D Bourget (year 1863), CF Hartt (year 1867) and the Hassler Expedition (1872) also collected specimens in Brazil that are today part of the MCZ’s holdings.

The Natural History Museum (NHM, London) fish collection of Brazilian specimens was in majority acquired as a gift from the collections of the Portuguese helminthologist Wucherer who dedicated his academic life in Bahia, as well as through donation and/or purchase via von Ihering and his son. The former was a German naturalist that immigrated to the country in 1880 and worked for museums in Germany and the UK as well as the Museu Nacional da Universidade Federal do Rio de Janeiro collecting specimens especially from Rio Grande do Sul state. Later, H Ihering became the first director (1894–1916) of the Museu Paulista whose natural history collection became part of the Museu de Zoologia da Universidade de São Paulo (Nomura, 2012)Nomura H. Hermann von Ihering (1850–1930), the naturalist. Cad Hist Ciênc Inst Butantan. 2012; 8(1):9–60.. This author published several articles related to fish diversity and biology, (e.g., Ihering, 1893Ihering H von. Kustenfische von Rio Grande do Sul. Koseritz Deutscher Volkskalender für Brasilien. Porto Alegre: 1893; 89–119. ) even though he was an expert in malacology systematics and biology. Rodolpho von Ihering, however, played a major role in synthesizing vernacular names of fish species in tupi and Portuguese (e.g., Ihering, 1940Ihering R von. Dicionário de animais do Brasil. São Paulo: Diretoria de Produção Agrícola; 1940. ). E Goeldi, a Swiss ichthyologist and first director (1894–1916) of the Museu Paraense in Belém (Sanjad, Güntert, 2015)Sanjad N, Güntert M. Emil August Göldi (1859-1917): a life between Switzerland and Brazil. Mitt Der Naturforsch Ges Bern. 2015; 72:21–71. also sent out many specimens from the Amazonian Atlantic coast as a gift/exchange to the NHM. The Albatross expedition (1883–1900) further collected specimens of S. tiburo (in Bahia, year 1887) in the country while M. Ruth in 1878 collected a specimen of Pristis sp. from Pará.

Systematic studies particularly from French, German, British and American taxonomists during the golden era for Zoological Systematics recognized several wide-ranging species in Brazil including S. zygaena as Zygaena malleus (Valenciennes, 1822), Prionace glauca (Linnaeus, 1758) as Carcharias hirundinaceus Müller & Henle, 1839 and Rhinoptera bonasus (Mitchill, 1815) as R. lalandii (Valenciennes, 1841). Müller, Henle (1841) recognized the occurrence of R. porosus (Poey, 1861) but this was listed as R. terraenovae (Richardson, 1837). Castelnau (1855)Castelnau FL. Poissons. In: Bertrand P, editor. Animaux Nouveaux ou rares recueillis pendant l’expédition dans les parties centrales de l’Amérique du Sud, de Rio Janeiro a Lima, et de Lima au Para; exécutée par ordre du Gouvernement Français pendant les années 1843 a 1847. Part 7, Zoologie; Paris. 1855; 2:1–112. reported Hypanus say (Lesueur, 1817). Pteroplatea valenciennii Duméril, 1865 synonym of Gymnura altavela (Linnaeus, 1758)Linnaeus C. Systema Naturae. Regnum Animale. 10th ed. Cura Societatis Zoologicae Germanicae Iterum Edita; 1758. was described as a new species. Ihering (1893)Ihering H von. Kustenfische von Rio Grande do Sul. Koseritz Deutscher Volkskalender für Brasilien. Porto Alegre: 1893; 89–119. listed Mustelus canis (Mitchill, 1815), Squalus americanus Mitchill, 1815, and Myliobatis aquila (Linnaeus, 1758) in which the latter two species are synonyms of Carcharias taurus (Rafinesque, 1810) and Myliobatis ridens Ruocco, Lucifora, Díaz de Astarloa, Mabragaña & Delpiani, 2012, respectively.

Birth and rise in the 20tth century (Fig. 1). The early times: Miranda-Ribeiro (1903–1939). Chondrichthyan Systematics in Brazil developed later on in the beginning of the 20th century with the birth of the Brazilian Ichthyology through a series of publications of the “mineiro” naturalist Alípio de Miranda-Ribeiro from the MNRJ, between 1903 and 1939. Miranda-Ribeiro first co-authored the article “A coleção de Peixes do Museu Nacional do Rio de Janeiro” (Schreiner, Miranda-Ribeiro, 1903)Schreiner C, Miranda-Ribeiro A. A coleção de peixes do Museu Nacional do Rio de Janeiro. Arq Mus Nac. 1903; 12:1–108. which came to light as a result of the research efforts of Carlos Schreiner, former sub-director of the Zoology section of this museum, who died before its completion. Later, the first edition of “Fauna brasiliense (Peixes). Tomo II. Desmobranchios” (Miranda-Ribeiro, 1907) was published as an individual authorship. These studies represent pioneer scientific attempts of listing and describing national species that were conceptualized, written and published by a Brazilian specialist, and that provided in detail a global outline of the Systematics research of the group in Brazilian waters. A total of 18 and 31 nominal species of sharks and rays, respectively, were listed in these studies, including the first record of species (e.g., Mustelus canis (Mitchill, 1815), Squalus blainvillei (Risso, 1816), Squatina squatina Linnaeus, 1758, Mobula birostris (Walbaum, 1792Walbaum J. Petri Artedi sueci genera piscium. In quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Rose, Grypeswaldiae [Greifswald]: 1792.), Pristis pectinata Latham, 1794); some of which today are known to represent more than one valid species in Brazil), and the description of two native species (Atlantoraja castelnaui (Miranda-Ribeiro, 1907) and Scyliorhinus haeckelli Miranda-Ribeiro, 1907) were also provided. In the second edition of “Fauna brasiliense (Peixes)”, Miranda-Ribeiro (1923)Miranda-Ribeiro A. Fauna Braziliense. Peixes. (Volume II, 1a Parte). Fascículo 1. Imprensa Nac. 1923:1–52. described one more native species, Narcine blachypleura Miranda-Ribeiro, 1923 whose taxonomic status is still uncertain. Miranda-Ribeiro actively contributed to document, list and describe elasmobranch species having published works related to the ichthyological collections holdings in the MNRJ and the Museu de Zoologia da Universidade de São Paulo (MZUSP) (e.g., Miranda-Ribeiro, 1918Miranda-Ribeiro A. Lista dos peixes Brasileiros do Museu Paulista. Primeira parte e Terceira parte. Rev Mus Paul. 1918; 10:705–39; 759–83., 1928Miranda-Ribeiro A. Callorhynchus callorhynchus L. Bol Mus Nac Rio Janeiro. 1928; 4:47–48., 1937Miranda-Ribeiro A. Sobre una collecção de vertebrados do nordeste brasileiro. Primeira parte: peixes e batrachios. O Campo. 1937; 1937:54–56.) until his death in 1939.

FIGURE 1 |
Number of publications every 5–7 years regarding each era: orange, Miranda-Ribeiro (1903–1939); dark green, Hiatus (1940–1950); light green, Sadowsky (1951–1988); pink, Rosa-Gomes (1980–1999); purple, Modern (2000–2023). Scale to the left, absolute number of publications; scale to the right, cumulative percentage of published papers, as shown by the black line on top of graph.

The hiatus (1940–1950). After Miranda-Ribeiro, a few important studies or new data on Brazilian Chondrichthyes Systematics were added (e.g., Carvalho, Samaya, 1942Carvalho JP, Samaya P. Comentários sobre os peixes. In: Magalhães JP, editor. História Natural do Brasil (Jorge Marcgrave). São Paulo: Museu Paulista; 1942. p.61–65. ; Dalcina, 1943Dalcina J. Fauna marítima: tubarões e cações. Caça e Pesca. 1943; 2(21):9–12.; Batista, 1944Batista BV. Investigações anatômicas sobre os nervos cranianos ventrais in vertebrata. An Acad Bras Cienc. 1944; 16(1):79–110.). The main contributions to the knowledge of the Brazilian fauna during this time were achieved by non-native researchers who lived and worked in North America. Such contributions include the list of coastal Brazilian fishes of Fowler (1941)Fowler H. Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostarophysi obtained by the United States Bureau of Fisheries Steamer “Albatross” in 1907 to 1910, chie. Washington, DC: US Government Printing Office; 1941. and the regional systematic reviews of Bigelow, Schroeder (1948Bigelow HB, Schroeder WC. Sharks. In: Tee-Van J, Breder C, Hildebrand S, Parr A, Schroeder W, editors. Fishes West. North Atl. Part 1. Lancelets, Cyclostomes Sharks. New Haven: Memoirs of the Sears Foundation For Marine Research, Yale University; 1948. p.59–546. , 1953)Bigelow HB, Schroeder WC. Fishes of the western North Atlantic. Part two. Sawfishes, guitarfishes, skates and rays. New Haven: Memoirs of the Sears Foundation of Marine Research, Yale University; 1953. p.1–514. that reported, respectively, 53 and 49 chondrichthyan species. The limited financial resources imposed by the World War II and other political issues during this time possibly resulted in impediments of zoological studies in the country and worldwide. As an example, the Brazil-USA Scientific Cooperation (MNRJ and Stanford University) between 1943 and 1955 (Sá, Britto, 2018)Sá MR, Britto MR. A cooperação científica Brasil-Estados Unidos na Segunda Guerra Mundial: levantamento dos peixes marinhos brasileiros de importância comercial (1943-1955). Rio de Janeiro: 2018. did not provide significant inputs to the local fauna as expected.

An impressive number of refugees who were favored from multi-nations diplomatic treaties came to live and work in Brazil afterwards, having a positive impact on the future of shark research in the country. Victor Sadowsky, a 30-year-old Lithuanian marine biologist whose academic career was interrupted due the war in Europe, arrived in São Paulo coming from Poland and changed the course of studies in the upcoming years.

Starting over: Sadowsky (1951–1988). Financial and scientific research investments in Oceanography from off the Southwestern Atlantic Ocean between the 1950–1970’s decades expanded shark Systematics studies. The studies of Victor Sadowsky who was based at the Instituto Oceanográfico da Universidade de São Paulo(IOUSP) made long-term contributions to the field. His early research interests, however, were on inland fish farming till 1951 when Sadowsky coordinated the creation and building of this research station by invitation of the IOUSP former director, Wladimir Besnard, who became his close friend (Nomura, 1990)Nomura H. Necrológio: atividades científicas de Viktor Sadowsky(1909-1990). Rev Bras Zool. 1990; 7(4):645–49. https://doi.org/10.1590/S0101-81751990000400019
https://doi.org/10.1590/S0101-8175199000...
. Sadowsky dedicated his studies to the taxonomy, biology and anatomy of the local marine fauna, including fish and associated parasites caught at the Cananéia Biological Station, Southern Brazil. His first elasmobranch study (Sadowsky, 1958)Sadowsky V. Contribuição para o conhecimento da alimentação de jamantas jovens (Manta ehrenbergii Müller & Henle). Bol Inst Ocean. 1958; 9:37–38. refers to the feeding habits of Manta ehrenbergii (Müller & Henle, 1841) (= Mobula birostris). The description of Sphyrna nana Sadowsky, 1964 (= S. media) and the publication of a list of elasmobranch species occurring in the region in Sadowsky (1967)Sadowsky V. Selachier aus dem litoral von São Paulo, Bresilien. Beit Neotrop Fauna. 1967; 5(2):41–88. are examples of his contributions to the national Chondrichthyan taxonomy. Several accounts related to first records and biology of species caught by experimental gillnets were successively published between 1965 and 1973 (e.g., Sadowsky, 1965Sadowsky V. The hammerhead sharks of the littoral zone of São Paulo, Brazil, with the description of a new species. Bull Mar Sci. 1965; 15(1):1–12., 1967Sadowsky V. Selachier aus dem litoral von São Paulo, Bresilien. Beit Neotrop Fauna. 1967; 5(2):41–88., 1968Sadowsky V. On the measurement of the total length of sharks. Zool Anz. 1968; 181:197–99., 1970aSadowsky V. First record of broad-snouted seven-gilled shark from the Cananéia, Brazil. Bol Inst Ocean. 1970a; 18(1):33–35.,bSadowsky V. On the dentition of the sand shark, Odontaspis taurus, from Cananéia, coast of Brazil. Bol Inst Ocean. 1970b; 18(1):1–10., 1971aSadowsky V. Estudio económico sobre los elasmobranquios de la zona litoral paulista. Carpas. 1971a; 5(1):1–11.,bSadowsky V. Notes on the bull shark, Carcharhinus leucas, in the lagoon region of Cananéia, Brazil. Bol Inst Ocean. 1971b; 20(2):71–78.,cSadowsky V. First record of the occurrence of an adult hammerhead shark (Sphyrna mokarran) in southern Brazilian waters. Inst Ocean. 1971c; 24:1–03., 1973Sadowsky V. First record of a basking shark (Cetorhinus maximus (Gunnerus, 1765)) in the Brazilian Atlantic. Bol Inst Ocean. 1973; 22:01–10.). These studies resulted from numerous research expeditions coordinated by Sadowsky in the tropical and subtropical waters of the western South Atlantic Ocean onboard the IOUSP R/V “Prof. W. Besnard” during the 1970’s.

Undoubtedly, Sadowsky became the first national scientist to devote efforts to long-term studies on cartilaginous fishes, especially marine sharks and batoids, bringing up the Brazilian Chondrichthyan research into the spotlight and bridging the research gap of the earlier period. At total, Sadowsky published 16 articles related to the taxonomy of Chondrichthyes from Brazil. He established valuable contacts with renowned shark experts, including JA Garrick (New Zealand), S Springer (USA), J D’Aubrey (South Africa), and E Siccardi (Argentina) which resulted in important partnerships and collaborations such as the description of Scyliorhinus besnardi (Springer & Sadowsky, 1970) (= S. haeckelli) and the recognition of his works in relevant shark literature of that time (e.g., Compagno, 1984Compagno LJV. FAO species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of sharks species known to date. Part 1. Hexanchiformes to Lamniformes. Rome: Food and Agriculture Organization of the United Nations; 1984. ; Garrick, 1982Garrick JAF. Sharks of the genus Carcharhinus. NOAA (National Oceanic and Atmospheric Administration) Technical Report NMFS (National Marine Fisheries Service) Circular No. 445; 1982. ).

Sadowsky’ legacy had a major impact on subsequent research in Systematics of elasmobranch fishes from Brazil even after his retirement in 1979 as he inspired a variety of eminent experts in the country. Figueiredo (1977)Figueiredo JL. Manual de peixes marinhos do Sudeste do Brasil. I. Introdução. Cações, raias e quimeras. São Paulo: Museu de Zoologia, Universidade de São Paulo; 1977. , for instance, in the first edition of “Manual de peixes marinhos do Sudeste do Brasil” examined numerous specimens collected by Sadowsky. This study represents one of the most comprehensive accomplishments in the Brazilian Chondrichthyan Systematics in which 72 species were listed (for more detail about the contributions of J. L. Figueiredo, see Rosa, Gadig, 2014Rosa RS, Gadig OBF. Conhecimento da diversidade dos Chondrichthyes marinhos no Brasil: a contribuição de José Lima De Figueiredo. Arq Zool Mus Zool Univ São Paulo. 2014; 45:89–104. https://doi.org/10.11606/issn.2176-7793.v45iespp89-104
https://doi.org/10.11606/issn.2176-7793....
). Lucena, Lucena (1981)Lucena CAS, Lucena ZMS. Catálogo dos peixes marinhos do Museu de Ciências da Pontifícia Universidade Católica do Rio Grande do Sul. Elasmobranchii. Teleostomi (1 parte). Comun Mus Cienc PUCRS. 1981; 21:1–66. also included Sawdosky’s specimens when listing the marine fishes from the Ichthyological collection of the Museu de Ciências da Pontifícia Universidade Católica do Rio Grande do Sul (MCP-PUCRS). After retirement, Sadowsky continued to contribute towards shark taxonomy with the collaboration of A. F. de Amorim and C. A. Arfelli from the Instituto de Pesca de Santos (São Paulo) through reporting of first occurrences from off Southeast-South Brazil. This includes Odontaspis noronhai (Maul, 1955), Lamna nasus (Bonnaterre, 1788), Etmopterus gracilispinis Krefft, 1968 and Isistius plutodus Garrick & Springer, 1964 in Sadowsky et al. (1988Sadowsky V, Arfelli CA, Amorim AF. Primeiro registro de ocorrência de Isistius plutodus (Chondrichthyes) no hemisfério sul. Supl Ciência e Cult Resumos, Soc Bras Para Prog Ciência. 1988; 4(7):919., 1986Sadowsky V, Arfelli CA, Amorim AF. First record of broadbanded lanternshark, Etmopterus gracilispinis (Krefft, 1968) (Squalidae), in the Brazilian waters. Bol Inst Pesca. 1986; 13(2):1–04., 1985Sadowsky V, Arfelli CA, Amorim AF. First record of porbeagle, Lamna nasus (Bonnaterre, 1788), in the Brazilian Atlantic. Bol Inst Pesca. 1985; 12(2):49–53., 1984Sadowsky V, Amorim AF, Arfelli CA. Second occurrence of Odontaspis noronhai (Maul, 1955). Bol Inst Pesca. 1984; 11:45–50., respectively).

Contemporary to Sadowsky, some individual initiatives were published in Sasso, Santos (1961)Sasso WS, Santos HS. Electron microscopy of enamel and dentin of teeth of the Odontaspis (Selachii). J Dent Res. 1961; 40(1):49–57. https://doi.org/10.1177/00220345610400012301
https://doi.org/10.1177/0022034561040001...
who investigated the dentition of Carcharias taurus (as Odontaspis sp.) as well as Miranda-Ribeiro (1961)Miranda-Ribeiro A. Catálogo dos peixes do Museu Nacional VI. 1961. and Barcellos (1963)Barcellos BN. Ictiofauna marinha do Rio Grande do Sul. Torpedo Nobiliana. 1963; 1963:1–15. who provided taxonomic listings, biological aspects of species and pinpointed the economic importance of certain taxa to local artisanal fisheries. Penna (1967)Penna I. Tubarões do Brasil. Rio de Janeiro: Sociedade Amigos do Museu Nacional; 1967. published the first comprehensive list of Brazilian sharks, as an initiative from a collaborative consortium of the MNRJ.

Rosa-Gomes (1980–1999). Because of investigations on comparative anatomy and phylogeny of elasmobranch fishes came to light with assiduity during the 80–90’s decades, it depicts the Rosa-Gomes era. Substantial changes in the classification using Cladistics and novel discussions on the inner and outer phylogenetic relationships of higher taxa were first introduced here at this stage. Starting with the culmination of the PhD thesis of R. S. Rosa in 1985Rosa RS. A systematic revision of the South American freshwater stingrays (Chondrichthyes: Potamotrygonidae). [PhD Thesis]. Gloucester Point, VA: College of William and Mary; 1985. Available from: https://dx.doi.org/doi:10.25773/v5-6ts0-6v68.
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on the family Potamotrygonidae whilst at the Virginia Institute of Marine Science, College of William and Mary, USA. The first native genus of freshwater rays, Plesiotrygon Rosa, Castello & Thorson, 1987, was then supported in Rosa et al. (1987)Rosa RS, Castello HP, Thorson TB. Plesiotrygon iwamae, a new genus and species of Neotropical freshwater stingray (Chondrichthyes: Potamotrygonidae). Copeia. 1987; 1987(2):447–58. https://doi.org/10.2307/1445783
https://doi.org/10.2307/1445783...
. Rosa has worked for over 40 years since 1977 when he became Professor at the Universidade Federal da Paraíba(UFPB), Northeastern Brazil. His performance has played a role in disseminating Chondrichthyan Systematics and conservation in Brazil and South America as recently highlighted in Viana et al. (2020)Viana STFL, Lima D, Viana K, Felinto A, Rosa RS. Cartilaginous fishes (Class Chondrichthyes) from the Ichthyological Collection at the Federal University of Paraíba, Brazil. Rev Nord Biol. 2020; 27(1):25–58. https://doi.org/10.22478/ufpb.2236-1480.2019v27n1.46558
https://doi.org/10.22478/ufpb.2236-1480....
and has published over 22 papers on the subject.

Concomitantly, U. L. Gomes, former professor at the Universidade do Estado do Rio de Janeiro (UERJ), initiated investigations on the dentition, anatomy of the vertebral column, clasper skeleton, pectoral and pelvic girdles, and neurocranium for descriptive purposes. A series of studies came to light targeting the sharpnose sharks of the genus Rhizoprionodon Whitley, 1929 in Gomes, Reis (1991)Gomes UL, Reis MAF. Considerações sobre a dentição de Rhizoprionodon lalandii (Valenciennes, 1839) e Rhizoprionodon porosus (Poey, 1861) (Elasmobranchii, Carcharhinidae). Parte I. Morfologia, Padrão Dentário e Heterodontia. An Acad Bras Cienc. 1991; 63(3):285–94., and species of Rajiformes, Myliobatiformes and Carcharhiniformes in Carvalho, Gomes (1992)Carvalho MR, Gomes UL. Reinterpretation of the clasper morphology of Prionace glauca (Chondrichthyes, Carcharhiniformes) with notes on clasper terminology. An Acad Bras Cienc. 1992; 64(2):199–206., da Cruz Lima et al. (1997)da Cruz Lima M, Leite Gomes U, Souza-Lima W, Paragó C. Anatomical study on the pre branchial region of Sphyrna lewini (Griffith & Smith) and Rhizoprionodon lalandii (Valenciennes) (Elasmobranchii, Carcharhiniformes) related with the cephalofoil in Sphyrna Rafinesque. Rev Bras Zool. 1997; 14(2):347–70. https://doi.org/10.1590/S0101-81751997000200009
https://doi.org/10.1590/S0101-8175199700...
, Gomes et al. (1997b)Gomes UL, Winkelstein C, Souza-Lima W. Estudo da cartilagem sinarcual cervico-toracica em Rajideos (Batomorphii, Rajiformes) da região sudeste do Brasil. An Acad Bras Cienc. 1997b; 69(1):95–107., Souza et al. (1999)Souza AM, Kamimoto JE, Intelizano W. Anatomia descritiva dos cingulos peitoral e pelvino, e respectivos pterigioforos de Rhizoprionodon lalandii (Valenciennes, 1839) (Carcharhinidae, Elasmobranchii). Publ Avulsas Inst Pau Brasil Hist Nat. 1999; 2:13–33.. Gomes’ contributions date back to the 1980’s though when providing data on new species occurrences in Brazilian waters such as for Rhincodon typus Smith, 1829 in Santos et al. (1988)Santos CAA, Andreata JV, Gomes UL. Sobre o ocorrencia de Rhincodon typus Smith, 1829 (Elasmobranchii: Rhincodontidae) em águas brasileiras e aspectos sobre sua taxonomia. Acta Biol Leopoldinensia. 1988; 10(2):175–82. and Cetorhinus maximus (Gunnerus, 1765) in Tomás, Gomes (1989)Tomás ARG, Gomes UL. Observações sobre a presença de Cetorhinus maximus (Gunnerus, 1765) (Elasmobranchii, Cetorhinidae) no sudeste e sul do Brasil. Bol Inst Pesca. 1989; 16(1):111–16.. Later, Gomes and collaborators released the book entitled “Catálogo das coleções ictiológicas do Departamento de Biologia Animal e Vegetal, Instituto de Biologia” in Gomes et al. (1997a)Gomes UL, Lima MC, Paragó CLD, Quintans AP. Catálogo das coleções ictiológicas do Departamento de Biologia Animal e Vegetal. Rio de Janeiro: Instituto de Biologia. Rio de Janeiro, Universidade do Estado do Rio de Janeiro; 1997a. , representing the largest species account published at that time with over 100 nominal species of sharks, batoids and chimeras recognized. Since then, Gomes published over 55 articles in Systematics (Tab. S1), becoming one of the most productive experts in the country (for more details on his biography, see Soares et al. (2022)Soares KDA, Gadig O, Santos HRS. O Professor Ulisses Leite Gomes e sua contribuição para o estudo da anatomia e taxonomia de elasmobrânquios (Chondrichthyes: Elasmobranchii). Arq Zool. 2022; 53:1–10. https://doi.org/10.11606/2176-7793/2022.53.01
https://doi.org/10.11606/2176-7793/2022....
.

Rosa-Gomes era coincides with the SBEEL foundation in Rio Grande, Rio Grande do Sul State in 1995. Before becoming a society, it was known as “Grupo de Trabalho sobre Pesca e Pesquisa de Tubarões e Raias no Brasil (GTPPTR)” and founded in July 1985 through the joint efforts of Amorim, Arfelli and Rosa under Sadowsky’s support. In that same year, the first Brazilian national meeting in Santos (São Paulo) was held with a handful of early career researchers having the chance to build up a new network of experts and discuss ideas of practical research on biology and natural history. The society thus has been determinant to promote national research and contribute towards scientific and community engagement. SBEEL furthermore has dedicated to strengthening conservation actions in Brazil and has been responsible for directly planning, preparing and delivering of the Brazilian National Plan of Action for the Conservation and Stock Management of Elasmobranch fishes whose first edition was released in 2005. SBEEL initiative has definitely influenced decision makers on protecting endangered sharks and batoids nationwide to avoid the collapse of populations and species.

Many regional checklists and identification guides also became evident in this period after national scientific incentives in marine explorations. The Brazilian Program for Evaluating the Living Marine Resources in the Exclusive Economic Zone (REVIZEE) between 1995–1998 comprised the first national initiative of concatenated multidisciplinary scientific surveys for this purpose and emphasized research dealing with potential fishing stocks, species diversity, marine topography, and ocean chemistry and biology of coastal and deep-water marine organisms, including cartilaginous fishes. REVIZEE oceanographic expeditions undergone within four major areas subdivided according to regional boundaries: North, Northeast, Central, and South. Lessa et al. (1999)Lessa R, Santana FM, Rincón G, Gadig OBF, El-Deir ACA. Biodiversidade de elasmobrânquios do Brasil. Ministério do Meio Ambiente (MMA). Projeto de Conservação e Utilização Sustentável da Diversidade Biológica Brasileira (PROBIO). Recife, PE: 1999. summarized the data available from these areas, providing information regarding the diversity, abundance, distribution, and ecology of elasmobranchs at a national level. Rosângela Lessa, professor at the Universidade Federal Rural de Pernambuco(UFRPE) and female pioneer in shark research in Brazil since the 1980’s, conducted extensive surveys off the Western Equatorial Atlantic region.

Afterwards, studies on sharks, batoids and chimeras were a product of localized initiatives of oceanographic institutions in the Northeast, South and Southeast Brazil as well as occasional surveys on landings of artisanal fisheries and fish markets, including Gadig, Moreira Junior (1992)Gadig OBF, Moreira Junior W. Tubarões da costa brasileira. Leopoldianum. 1992; 18(52):111–19., Amorim et al. (1998)Amorim A, Arfelli C, Fagundes L. Pelagic elasmobranchs caught by longliners off southern Brazil during 1974-97: an overview. Mar Freshw Res. 1998; 49:621–32. https://doi.org/10.1071/MF97111
https://doi.org/10.1071/MF97111...
, and Menni, Lessa (1998)Menni RC, Lessa RP. The chondrichthyan community off Maranhão (northeastern Brazil). II. Biology of species. Acta Zool Lilloana. 1998; 44(1):69–89.. Taxonomic accounts that report first records and new species occurrences prevailed in this eraand only a handful number of taxa was targeted in more comprehensive studies such as Vooren, Silva (1991)Vooren CM, Silva KG. On the taxonomy of the angel sharks from southern Brazil, with the description of Squatina occulta sp. n. Rev Bras Biol. 1991; 51(3):589–602. for angelsharks and Gallo-da-Silva et al. (1997)Gallo-da-Silva V, Afonso AF, Gomes UL. Estudo taxonômico das espécies de Rhinoptera Cuvier, 1829 ocorrentes no litoral brasileiro. Parte 1: condroneurocranio de exemplares jovens de Rhinoptera brasiliensis Muller & Henle, 1841 e Rhinoptera bonasus (Mitchill, 1815) (Elasmobranchii, Myliobatiformes, Rhinopteridae). An Acad Bras Cienc. 1997; 69(4):585–600. for cownose rays. Phylogenetic studies were not yet solid here except for Carvalho (1996)Carvalho MR. Higher-level elasmobranch phylogeny, basal squaleans, and paraphyly. In: Stiassny M, Parenti L, Johnson G, editors. Interrelationships of fishes. California: Academic Press; 1996. p.35–62. , who brought the country into the spotlight of the elasmobranch research community after reiterating the Hypnosqualean hypothesis on the evolutionary relationships between sharks and batoids.

Both Rosa and Gomes’ inputs in the subject area surpass the turn of the century in the history of Chondrichthyan Systematics in Brazil having a thorough influence over successive generations of shark experts. Publications related to the recognition of novel species, taxonomic accounts and revisions have been provided till recently in partnership with current and former students, colleagues and collaborators. The first generation of researchers that were under their influence and that has been active today include Getúlio Rincón (Universidade Federal do Maranhão), Patrícia Charvet (Universidade Federal do Ceará) and Marcelo Carvalho (former Universidade de São Paulo).

Modern systematics in the 21st century (2000–today). Scientific productivity over the last 23 years has been recurrent and strong, which thus set Brazil as one of the world’s leading shark research hubs. The quality and scope of publications has also stood out with more comprehensive studies of taxa currently prevailing. Phylogenetic studies of native and non-native taxa using morphological characters have been slowly implemented. Descriptive anatomical studies of species and comparative anatomy for evolutionary inferences have been widely employed, especially for Squatiniformes, Squaliformes, Carcharhiniformes (e.g., Mello, 2009Mello W. The electrosensorial pore system of the cephalofoil in the four most common species of hammerhead shark (Elasmobranchii: Sphyrnidae) from the Southwestern Atlantic. CR Biol. 2009; 332(4):404–12.; Carvalho et al., 2012Carvalho MR, Faro C, Gomes UL. Comparative neurocranial morphology of angelsharks from the south-western Atlantic Ocean (Chondrichthyes, Elasmobranchii, Squatinidae): Implications for taxonomy and phylogeny. Acta Zool. 2012; 93(2):171–83. https://doi.org/10.1111/j.1463-6395.2010.00495.x
https://doi.org/10.1111/j.1463-6395.2010...
; Moreira et al., 2018Moreira RA, Loboda TS, Carvalho MR. Comparative anatomy of the clasper of the subfamily Potamotrygoninae (Chondrichthyes: Myliobatiformes). J Morphol. 2018; 279(5):598–608. https://doi.org/10.1002/jmor.20795
https://doi.org/10.1002/jmor.20795...
; Silva et al., 2018Silva JPCB, Vaz DFB, Carvalho MR. Phylogenetic inferences on the systematics of squaliform sharks based on elasmobranch scapular morphology (Chondrichthyes: Elasmobranchii). Zool J Linn Soc. 2018; 182(3):614–30. https://doi.org/10.1093/zoolinnean/zlx051
https://doi.org/10.1093/zoolinnean/zlx05...
), and Myliobatiformes and Rajiformes in Afonso, Gallo (2001)Afonso AF, Gallo V. Study of the scapulocoracoid and cervico-thoracic synarcual cartilage of Rhinoptera brasiliensis Mueller & Henle and Rhinoptera bonasus (Mitchill) (Elasmobranchii, Rhinopteridae). Rev Bras Zool. 2001; 18(2):319–31. https://doi.org/10.1590/S0101-81752001000200002
https://doi.org/10.1590/S0101-8175200100...
, Oddone, Vooren (2008)Oddone MC, Vooren CM. Comparative morphology and identification of egg capsules of skate species of the genera Atlantoraja Menni, 1972, Rioraja Whitley, 1939 and Sympterygia Müller & Henle, 1837. Arq Ciênc Mar. 2008; 41(2):5–13. https://labomar.ufc.br/wp-content/uploads/2017/01/acm-2008-41-2-01.pdf
https://labomar.ufc.br/wp-content/upload...
, Shibuya et al. (2010)Shibuya A, Zuanon JAS, Araújo MLG, Tanaka S. Morphology of lateral line canals in Neotropical freshwater stingrays (Chondrichthyes: Potamotrygonidae) from Negro River, Brazilian Amazon. Neotrop Ichthyol. 2010; 8(4):867–76. https://doi.org/10.1590/S1679-62252010000400017
https://doi.org/10.1590/S1679-6225201000...
, Bini et al. (2015)Bini IRJ, Santos CMH, Fadig OBF. Anatomical description of scapulocoracoid and gill arches of Benthobatis kreffti. Univ Sci. 2015; 20(3):305–12. https://doi.org/10.11144/Javeriana.SC20-3.adsg.
https://doi.org/10.11144/Javeriana.SC20-...
, and Cunha et al. (2016)Cunha CM, Oliveira LE, Kfouri Jr. J.R. Comparative anatomy of the extraocular muscles in four Myliobatoidei rays (Batoidea, Myliobatiformes). J Anat. 2016; 228(5):877–86. https://doi.org/10.1111/joa.12438
https://doi.org/10.1111/joa.12438...
. Regional taxonomic revisions have been implemented to provide morphological separation between species and elucidate regional synonyms using comprehensive examination of specimens (e.g., Silva, Carvalho, 2011Silva JPCB, Carvalho MR. A taxonomic and morphological redescription of Potamotrygon falkneri Castex & Maciel, 1963 (Chondrichthyes: Myliobatiformes: Potamotrygonidae). Neotrop Ichthyol. 2011; 9(1):209–32. https://doi.org/10.1590/S1679-62252011000100021
https://doi.org/10.1590/S1679-6225201100...
; Loboda, Carvalho, 2013Loboda TS, Carvalho MR. Systematic revision of the Potamotrygon motoro (Müller & Henle, 1841) species complex in the Paraná-Paraguay basin, with description of two new ocellated species (Chondrichthyes: Myliobatiformes: Potamotrygonidae). Neotrop Ichthyol. 2013; 11(4):693–737. https://doi.org/10.1590/S1679-62252013000400001
https://doi.org/10.1590/S1679-6225201300...
; Vaz, Carvalho, 2013Vaz DFB, Carvalho MR. Morphological and taxonomic revision of species of Squatina from the Southwestern Atlantic Ocean (Chondrichthyes: Squatiniformes: Squatinidae). Zootaxa. 2013; 3695(1):1–81. https://doi.org/10.11646/zootaxa.3695.1.1
https://doi.org/10.11646/zootaxa.3695.1....
; Fontenelle, Carvalho, 2017Fontenelle JP, Carvalho MR. Systematic revision of the Potamotrygon scobina Garman, 1913 species-complex (Chondrichthyes: Myliobatiformes: Potamotrygonidae), with the description of three new freshwater stingray species from Brazil and comments on their distribution and biogeography. Zootaxa. 2017; 4310(1):1–63. https://doi.org/10.11646/zootaxa.4310.1.1
https://doi.org/10.11646/zootaxa.4310.1....
; Petean Carvalho, 2018Petean FF, Carvalho MR. Comparative morphology and systematics of the cookiecutter sharks, genus Isistius Gill (1864) (Chondrichthyes: Squaliformes: Dalatiidae). PLoS ONE. 2018; 13(8):e0201913. https://doi.org/10.1371/journal.pone.0201913
https://doi.org/10.1371/journal.pone.020...
; Soares, Carvalho, 2019Soares KDA, Carvalho MR. The catshark genus Scyliorhinus (Chondrichthyes: Carcharhiniformes: Scyliorhinidae): taxonomy, morphology and distribution. Zootaxa. 2019; 4601(1):1–147. https://doi.org/10.11646/ZOOTAXA.4601.1.1
https://doi.org/10.11646/ZOOTAXA.4601.1....
). Meanwhile, checklists and species listings have still been provided as seen in Gadig, Gomes (2003)Gadig OFB, Gomes UL. Various families. In: Menezes NA, Buckup PA, Figueiredo JL, Moura RL, editors. Catálogo das espécies peixes Marinhos do Brasil. São Paulo: Museu de Zoologia, Universidade de São Paulo; 2003. , Soto, Mincarone (2004)Soto JMR, Mincarone MM. Collections of the Museu Oceanográfico do Vale do Itajaí. I. Catalog of cartilaginous fishes (Myxini, Cephalaspidomorphi, Elasmobranchii, Holocephali). Mare Magnum. 2004; 2(1–2):1–125., Nunan, Senna (2007)Nunan GW, Senna MLV. Tubarões (Selachii) coletados pelo navio oceanográfico Thalassa sobre a plataforma externa e talude continental do Brasil entre 118 e 228. In: Costa PAS, Olavo G, Martins AS, editors. Biodiversidade da Fauna Mar. Profunda na Costa Central Brasileira. Rio de Janeiro: Museu Nacional; 2007. p.184. , Gomes et al. (2010Gomes UL, Signori CN, Gadig OBF, Santos HRS. Guia para identificação de tubarões e raias do Rio de Janeiro. 1a ed. Rio de Janeiro: Technical Books Editora; 2010. , 2019Gomes UL, Santos HRS, Gadig OFB, Signori CN, Vicente MM. Guia para identificação dos tubarões, raias e quimeras do Rio de Janeiro (Chondrichthyes: Elasmobranchii e Holocephali). Rev Nord Biol. 2019; 27(1):171–368. https://doi.org/10.22478/ufpb.2236-1480.2019v27n1.47122
https://doi.org/10.22478/ufpb.2236-1480....
), and thus constantly updating the number of species of sharks, batoids and chimeras recognized in the country. Novel species were also widely recognized in this period within a variety of taxa, including Dasyatidae stingrays and Rajidae skates in Gomes et al. (2000)Gomes UL, Rosa RS, Gadig OBF. Dasyatis macropththalma sp. n.: a new species of stingray (Chondrichthyes: Dasyatidae) from the Southwestern Atlantic. Copeia. 2000; 2000(2):510–15. https://doi.org/10.1643/0045-8511(2000)000[0510:DMSNAN]2.0.CO;2
https://doi.org/10.1643/0045-8511(2000)0...
, Gomes, Paragó (2001)Gomes UL, Paragó CLD. Espécie nova de rajídeo (Chondrichthyes, Rajiformes) do Atlântico Sul Ocidental. Bol Mus Nac Zool. 2001; 448:1–10., Gomes, Picado (2001)Gomes UL, Picado S. Distribution of the species of Dipturus Rafinesque. Distribution. 2001; 18(1):171–85., Santos, Carvalho (2004)Santos HRS, Carvalho MR. Description of a new species of whiptailed stingray from the Southwestern Atlantic Ocean (Chondrichthyes, Myliobatiformes, Dasyatidae). Bol Mus Nac. 2004; 516:1–24., Carvalho et al. (2005)Carvalho MR, Gomes UL, Gadig OBF. Description of a new species of skate of the genus Malacoraja Stehmann, 1970: the first species from the southwestern Atlantic Ocean, with notes on generic monophyly and composition (Chondrichthyes: Rajidae). Neotrop Ichthyol. 2005; 3(2):239–58. https://doi.org/10.1590/S1679-62252005000200002
https://doi.org/10.1590/S1679-6225200500...
, Petean et al. (2020)Petean FF, Naylor GJP, Lima SMQ. Integrative taxonomy identifies a new stingray species of the genus Hypanus Rafinesque, 1818 (Dasyatidae, Myliobatiformes) from the Tropical Southwestern Atlantic. J Fish Biol. 2020; 97(4):1120–42. https://doi.org/10.1111/jfb.14483
https://doi.org/10.1111/jfb.14483...
deep-water species such as deep-water catsharks in Soto (2001aSoto JMR. Galeus mincaronei sp. nov. (Carcharhiniformes, Scyliorhinidae), a new species of sawtail catshark from southern Brazil. Mare Magnum. 2001a; 1(1):11–18. https://www.univali.br/institucional/museu-oceanografico-univali/mare-magnum/volume-1-numero-1/Documents/maremagnum3.pdf
https://www.univali.br/institucional/mus...
,b)Soto JMR. Schroederchthys saurisqualus sp. nov. (Carcharhiniformes, Scyliorhinidae), a new species of catshark from southern Brazil, with further data on Schroederichthys species. Mare Magnum. 2001b; 1(1):37–50. https://www.univali.br/institucional/museu-oceanografico-univali/mare-magnum/volume-1-numero-1/Documents/maremagnum7.pdf
https://www.univali.br/institucional/mus...
and Soares et al. (2019)Soares KDA, Carvalho MR, Schwingel PR, Gadig OBF. A new species of Parmaturus (Chondrichthyes: Carcharhiniformes: Scyliorhinidae) from Brazil, Southwestern Atlantic. Copeia. 2019; 107(2):314–22. https://doi.org/10.1643/CI-18-152
https://doi.org/10.1643/CI-18-152...
, dogfish sharks in Viana et al. (2016)Viana STF, Carvalho MR, Gomes UL. Taxonomy and morphology of species of the genus Squalus Linnaeus, 1758 from the Southwestern Atlantic Ocean (Chondrichthyes: Squaliformes: Squalidae). Zootaxa. 2016; 4133(1):1–89. https://doi.org/10.11646/zootaxa.4133.1.1
https://doi.org/10.11646/zootaxa.4133.1....
, electric rays in Rincon et al. (2001)Rincon G, Stehmann MFW, Vooren CM. Results of the research cruises of FRV “Walter Herwig” to South America. LXXIV. Benthobatis kreffti n. sp. (Chondrichthyes, Torpediniformes, Narcinidae), a new deep-water electric ray from off South Brazil and the third species of the genus. Arch Fish Mar Res. 2001; 49:45–60., rabbitfishes in Soto, Vooren (2004)Soto JMR, Vooren CM. Hydrolagus matallanasi sp. nov. (Holocephali, Chimaeridae) a new species of rabbitfish from southern Brazil. Zootaxa. 2004; 687(1):1–10. https://doi.org/10.11646/zootaxa.687.1.1, freshwater rays in Carvalho, Ragno (2011)Carvalho MR, Ragno M. An unusual, dwarf new species of Neotropical freshwater stingray, Plesiotrygon nana sp. nov., from the upper and mid Amazon basin: the second species of Plesiotrygon (Chondrichthyes: Potamotrygonidae). Pap Avulsos Zool. 2011; 51(7):101–38. https://doi.org/10.1590/S0031-10492011000700001
https://doi.org/10.1590/S0031-1049201100...
, and Fontenelle et al. (2014)Fontenelle JP, Lovejoy NR, Kolmann MA, Marques FPL. Molecular phylogeny for the Neotropical freshwater stingrays (Myliobatiformes: Potamotrygoninae) reveals limitations of traditional taxonomy. Biol J Linn Soc. 2021; 134(2):381–401. https://doi.org/10.1093/biolinnean/blab090
https://doi.org/10.1093/biolinnean/blab0...
.

Application of innovative methodologies for data sampling also helped to evolve the Modern Systematics in the country. The pioneer study of Solé-Cava et al. (1983)Solé-Cava AM, Vooren CM, Levy JA. Isozymic differentiation of two sibling species of Squatina (Chondrichthyes) in south Brazil. Comp Biochem Physiol. 1983; 75B(2):355–58. http://www.intranet.biologia.ufrj.br/lbdm/pdfs/Sole-Cavaetal1983Squatina.pdf
http://www.intranet.biologia.ufrj.br/lbd...
provided data for genetic frequencies of allozymes to identify species of Squatina occurring in Southern Brazil. Subsequent molecular data related studies focusing on DNA barcoding and molecular species delimitation have been published for freshwater rays, lamnid and carcharhinid sharks, guitarfishes, sawfishes, and stingrays.

Research trends in shark systematics within the 120-year period. Over the period 1903–2023 a total of 354 scientific publications related to Systematics of Chondrichthyes from Brazil were produced (Fig. 1). Miranda-Ribeiro era has a total of six publications, followed by three publications during the Hiatus era. Thirty-two studies were published in the Sadowsky era, and 54 in Rosa-Gomes era. The Modern era, which comprises the last 23 years, exhibited a total of 259 publications with an average of 11 studies per year (Fig. 1).

Research subject area. Taxonomic studies have been the major focus (70.1%) followed by morphological (22.3%) and phylogenetic ones (7.6%) (Fig. 2A).

Research scope. Taxonomic scopes comprise almost all categories mentioned in Material in Methods, except high-level interrelationships. Within the taxonomy subject area, checklists, new record, description, and revision together represent 72.6% of total studies published between 1903–2023 while first record and DNA barcoding exhibit 10.9% and 8.9%, respectively. The other five categories were represented by less than 8% of taxonomic papers (Tab. 1).

FIGURE 2 |
Percentages of papers within (A) three distinct subject areas (taxonomy (green), morphology (orange), and phylogeny (purple) on the inner circle, and data sources (morphological, molecular, integrative) on the outer circle. Percentage of papers under one of the 12 research scopes (checklist, comparative anatomy, description, descriptive anatomy, DNA barcoding, first record, high-level interrelationships, identification guide, low-level interrelationships, molecular species delimitations, new record, and revision) using each of the data sources: (B) integrative; (C) molecular; (D) morphological.
TABLE 1 |
Percentage of studies under distinct research scopes within each one of the subject areas.

Within Morphology, only four research scopes were identified and the two most expressive were comparative anatomy (53.2%) and descriptive anatomy (43.0%). Finally, within phylogenetic studies, low- and high-level interrelationships were the most common scopes with 40.7% and 33.3% of studies, respectively (Fig. 3B).

When analyzing the Research Scopes regarding the Data Source used, within integrative data, revision was the most common (42.9%) (Fig. 2B); within molecular, DNA barcoding (46.8%) (Fig. 2C), and within morphological, checklist (23.0%), followed by new records (16.3%) and comparative anatomy (16.7%) (Fig. 2D).

Data source. Morphological-based methodology stands out, representing 84.7% of the total studies on Systematics of Chondrichthyes, while those based exclusively on molecular data are 13.3%, and research combining two or more data sources (integrative) correspond to only 2.0% (Fig. 2A). Within phylogeny subject area, 48.1% of studies were based on molecular data, 33.3% morphological, and 18.5% integrative; while within taxonomy, 85.5% on morphological, 13.7% molecular, and 0.8% integrative. Phylogenetic research is the only area herein considered in which molecular-based data is larger than morphological (Fig. 2A).

Targeted study taxa. Most manuscripts focused on single species, genus, or family-level (68.9%), followed by those focused on two or more taxa (multi-taxa, 27.7%). Some studies focused on Batoidea as a whole, and 10 studies on the orders Carcharhiniformes (4), Squaliformes (3), Lamniformes (1), Rajiformes (1), and Rhinopristiformes (1). Among those working on family-level or lower taxa, 17.6% regarded Potamotrygonidae and 13.5%, Carcharhinidae, whereas 29 families have been mentioned in less than 10 manuscripts in a total of 37. Excluding those multi-taxa papers, the majority comprised research on Batoidea (46.7%), followed by Galeomorphi (36.5%), Squalomorphi (15.6%), and Holocephali (1.2%). Potamotrygonidae was the most studied group within the subject areas “taxonomy” (12.1%) and “phylogeny” (18.5%), while within “morphology”, Carcharhinidae (19.0%) was the most studied family (Fig. 3).

Of the few manuscripts that integrate different data sources (7), two regarded Potamotrygonidae and the other five were on distinct families. Among the 47 manuscripts using molecular data, 17% concerned Carcharhinidae and 36.2%, multi-taxa. Finally, among the 300 manuscripts using morphological data, 12.3% concerned Potamotrygonidae and 27.0%, multi-taxa.

FIGURE 3 |
Percentage of studies focused on each family or on two or more (multi-taxa) within the subject areas (A) taxonomy, (B) morphology, and (C) phylogeny.

Taking into account the eight most-cited families (Potamotrygonidae, Carcharhinidae, Dasyatidae, Scyliorhinidae, Arhynchobatidae, Rajidae, Sphyrnidae, and Squatinidae) as those that were studied in more than 10 papers (Figs. 45), morphological data used in taxonomical and morphological studies were the only ubiquitous data source and subject area among them. Phylogenetic studies based exclusively on morphological data are lacking for Potamotrygonidae, Carcharhinidae, Dasyatidae and Squatinidae. Research focused on Rajidae (Fig. 5) was only based on morphological data within taxonomy and morphology, without any studies performing phylogenetic analyses, or using integrative and molecular data, demonstrating a clear gap in systematic studies. Besides, Scyliorhinidae and Arhynchobatidae, when the only target taxa of a study, also lack studies based exclusively on molecular data.

FIGURE 4 |
Percentage of studies within the shark families Carcharhinidae, Scyliorhinidae, Sphyrnidade, and Squatinidae, and within subject areas taxonomy, morphology, and phylogeny.
FIGURE 5 |
Percentage of studies within the batoid families Arhynchobatidae, Dasyatidae, Potamotrygonidae, and Rajidae, and within subject areas taxonomy, morphology, and phylogeny.

Authorship and gender diversity. There is an average of 3.23 authors by study, ranging from 1 (14.7%) to 15 (0.3%), with two authors being the most common (30.5%) (Fig. 6A). Of the 52 studies authored by only 1 person, 5.8% were women and 94.2% were men. Of all 354 studies, 20.3% were first-authored and 15.8% last-authored by a woman, with only 6.5% with first and last authors being women. Conversely, 55.6% were first- and last-authored by men (Fig. 6B). The average percentage of women as co-authors was 17.7% in each study; however, the most common scenario was the absence of women in authorships (60.7% of studies, 215 out of 354).

FIGURE 6 |
A. Number of papers written by 1 to 15 authors; B. Percentage of publications authored by men and women as first and last authors.

DISCUSSION

Overall research trends in Brazil. Literature review of the 120-years period (1903–2023) revealed that scientific outputs within the Chondrichthyan Systematics in Brazil have constantly evolved. The number of publications has increased considerably since the Miranda-Ribeiro era. Least impactful studies were conducted during this first period, in which general species listings related to national diversity of fishes were the main study scope along with new species descriptions, with exception to the pioneer work of Batista (1944)Batista BV. Investigações anatômicas sobre os nervos cranianos ventrais in vertebrata. An Acad Bras Cienc. 1944; 16(1):79–110. concerning cranial anatomy. Sadowsky era mostly accompanied occurrence reports of wide-ranging species and remarkably the first taxonomic revisions of native species raised exponentially to over five times the total number of publications. Rosa-Gomes era kept the total number of publications rising notwithstanding comprising the shortest era within the Chondrichthyan Systematic history of the country and exhibited a publication average above the previous eras.

Substantial contributions to the field were noticed in the last 23 years, with the total number of studies and the publication average superseding earlier periods of research in Brazil altogether (Fig. 1). Brazil thus has contributed to exceptional inputs to the local and international knowledge, a protagonism that has never been experienced before in more than 120 years of Brazilian research on cartilaginous fishes. Heavy investments in research and higher education during this period were possibly the main determinant in the application of rapid taxonomic approaches such as molecular genetic data (see detailed explanation further below).

Modalities of research in Brazil have been disparate over time. Taxonomy stands out with 70.0% of the scientific productivity related to Chondrichthyes followed by morphology and phylogeny. Studies of taxonomic scope that involve revisions, species descriptions, checklists and first/new records have been constantly implemented since the Miranda-Ribeiro era, remaining the main study field to date. Comparative anatomical investigations have been carried out since early 1990 throughout the Rosa-Gomes era till recently although the first descriptive morphological studies were conducted earlier during the 1940 and 1960’s decades. Phylogenetic analyses, on the other hand, have been sporadically applied and thus it is underrepresented in the overall shark research trend.

Research knowledge gaps and challenges. Traditional Systematics dominates in studies related to the diversity and evolution of Brazilian sharks, batoids and chimeras. Most morphological-based studies have constantly provided high quality data of characters for species diagnosis and evolutionary support of higher clades within Chondrichthyes. Methods in traditional Systematics such as dissection, radiography and CT-scan, and clearing and staining are usually more time consuming for data gathering and analyses. Not surprisingly, the fields of phylogeny and morphology, that require detailed morphological characters, are less commonly employed than taxonomic studies. Within the latter field, low research effort is also applied for studies with scope on taxonomic revisions and descriptions of species. Thus, generic taxonomic studies such as checklists/lists and first records/new occurrences of species have superseded the scientific production in the country because it usually relies on rapid methodology for data acquisition such as literature compilation and/or superficial morphological investigations.

Modern Systematics is still evolving in Brazil, which is in contrast with the current global scenario related to Chondrichthyan research but in accordance with the South American trend as noticed in Awruch et al. (2019)Awruch CA, Somoza GM, Baldock C. Chondrichthyan research in South America: Endocrinology overview and research trends over 50 years (1967–2016) compared to the rest of the world. Gen Comp Endocrinol. 2019; 273:118–33. https://doi.org/10.1016/j.ygcen.2018.06.005
https://doi.org/10.1016/j.ygcen.2018.06....
. Molecular taxonomy for species identification might undermine ichthyofaunistic listings (e.g., Brandão et al., 2016Brandão JHSG, Bitencourt JA, Santos FB, Watanabe LA, Schneider H, Sampaio I et al. DNA barcoding of coastal ichthyofauna from Bahia, northeastern Brazil, South Atlantic: High efficiency for systematics and identification of cryptic diversity. Biochem Syst Ecol. 2016; 65:214–24. https://doi.org/10.1016/j.bse.2016.02.012
https://doi.org/10.1016/j.bse.2016.02.01...
; Pereira et al., 2011Pereira LHG, Maia GMG, Hanner R, Foresti F, Oliveira C. DNA barcodes discriminate freshwater fishes from the Paraíba do Sul River Basin, São Paulo, Brazil. Mitochondrial DNA. 2011; 22:71–79. https://doi.org/10.3109/19401736.2010.532213
https://doi.org/10.3109/19401736.2010.53...
), focus on forensics of target species in the fisheries industry (e.g., Bunholi et al., 2018Bunholi IV, Ferrette BLS, Biasi JB, Magalhães COM, Rotundo MM, Oliveira C et al. The fishing and illegal trade of the angelshark: DNA barcoding against misleading identifications. Fish Res. 2018; 206:193–97. https://doi.org/10.1016/j.fishres.2018.05.018
https://doi.org/10.1016/j.fishres.2018.0...
) and specific taxa, such as sharpnose sharks in Mendonça et al. (2011)Mendonça FF, Oliveira C, Burgess G, Coelho R, Piercy A, Gadig OBF et al. Species delimitation in sharpnose sharks (genus Rhizoprionodon) in the western Atlantic Ocean using mitochondrial DNA. Conserv Genet. 2011; 12(1):193–200. https://doi.org/10.1007/s10592-010-0132-6
https://doi.org/10.1007/s10592-010-0132-...
and Pinhal et al. (2011)Pinhal D, Shivji MS, Vallinoto M, Chapman DD, Gadig OBF, Martins C. Cryptic hammerhead shark lineage occurrence in the western South Atlantic revealed by DNA analysis. Mar Biol. 2011; 159:829–36. https://doi.org/10.1007/s00227-011-1858-5
https://doi.org/10.1007/s00227-011-1858-...
, guitarfishes in Mariguela et al. (2009)Mariguela TC, De-Franco B, Almeida TV, Mendonça FF, Gadig OB, Foresti F et al. Identification of guitarfish species Rhinobatos percellens, R. horkelli, and Zapteryx brevirostris (Chondrichthyes) using mitochondrial genes and RFLP technique. Conserv Genet Resour. 2009; 1:393–96. https://doi.org/10.1007/s12686-009-9091-y
https://doi.org/10.1007/s12686-009-9091-...
and De-Franco et al. (2010)De-Franco B, Mendonça FF, Hashimoto DT, Porto-Foresti F, Oliveira C, Foresti F. Forensic identification of the guitarfish species Rhinobatos horkelli, R. percellens and Zapteryx brevirostris using multiplex‐PCR. Mol Ecol Resour. 2010; 10(1):197–99. https://doi.org/10.1111/j.1755-0998.2009.02728.x
https://doi.org/10.1111/j.1755-0998.2009...
, angelsharks in Falcão et al. (2014)Falcão LHO, Furtado-Neto MAA, Maggioni R, Faria VV. Prospective molecular markers for the identification of illegally traded angelsharks (Squatina) and dolphin (Sotalia guianensis). Genet Mol Res. 2014; 13(4):9710–17. https://doi.org/10.4238/2014.November.24.2
https://doi.org/10.4238/2014.November.24...
, requiem sharks in Domingues et al. (2013)Domingues RD, Amorim AF, Hilsdorf AW. Genetic identification of Carcharhinus sharks from the southwest Atlantic Ocean (Chondrichthyes: Carcharhiniformes). J Appl Ichthyol. 2013; 29(4):738–42. https://doi.org/10.1111/jai.12154
https://doi.org/10.1111/jai.12154...
, sawfishes in (Faria et al. (2013)Faria VV, McDavitt MT, Charvet P, Wiley TR, Simpfendorfer CA, Naylor GJP. Species delineation and global population structure of Critically Endangered sawfishes (Pristidae). Zool J Linn Soc. 2013; 167(1):136–64. https://doi.org/10.1111/j.1096-3642.2012.00872.x
https://doi.org/10.1111/j.1096-3642.2012...
, guitarfishes in Cruz et al. (2023)Cruz VP, Rotundo MM, Charvet P, Boza BR, Souza BC, Cerqueira NN et al. Investigating an unknown biodiversity: evidence of distinct lineages of the endemic chola guitarfish Pseudobatos percellens Walbaum, 1792 in the Western Atlantic Ocean. Diversity. 2023; 15(3):344. https://doi.org/10.3390/d15030344
https://doi.org/10.3390/d15030344...
, and daggernose sharks in Nachtigall et al. (2017)Nachtigall PG, Rodrigues Filho LF, Sodré DC, Vallinoto M, Pinhal D. A multiplex PCR approach for the molecular identification and conservation of the Critically Endangered daggernose shark. Endanger Species Res. 2017; 32:169–75. https://doi.org/10.3354/esr00798
https://doi.org/10.3354/esr00798...
, to mention a few. Dasyatids have been well documented in this aspect whilst freshwater stingrays have been less investigated than marine species (e.g., Valentim et al., 2006Valentim FC, Falcão JD, Porto JI, Feldberg E. Chromosomes of three freshwater stingrays (Rajiformes Potamotrygonidae) from the Rio Negro basin, Amazon, Brazil. Genetica. 2006; 128(1–3):33–39. https://doi.org/10.1007/s10709-005-3816-y
https://doi.org/10.1007/s10709-005-3816-...
; Toffoli et al., 2008Toffoli D, Hrbek T, Araújo MLG, Almeida MP, Charvet-Almeida P, Farias IP. A test of the utility of DNA barcoding in the radiation of the freshwater stingray genus Potamotrygon (Potamotrygonidae, Myliobatiformes). Genet Mol Biol. 2008; 31:324–36. https://doi.org/10.1590/S1415-47572008000200028
https://doi.org/10.1590/S1415-4757200800...
; Cruz et al., 2015Cruz VP, Vera M, Mendonça FF, Pardo BG, Martinez P, Oliveira C et al. First identification of interspecies hybridization in the freshwater stingrays Potamotrygon motoro and P. falkneri (Myliobatiformes, Potamotrygonidae). Conserv Genet. 2015; 16:241–45. https://doi.org/10.1007/s10592-014-0642-8
https://doi.org/10.1007/s10592-014-0642-...
; Torres et al., 2022Torres Y, Faria VV, Charvet P. Current status and future perspectives of Neotropical freshwater stingrays (Potamotrygoninae, Myliobatiformes) genetics. Environ Biol Fishes. 2022; 105:1111–27. https://doi.org/10.1007/s10641-022-01320-x
https://doi.org/10.1007/s10641-022-01320...
). Unexpected trends in research like these are unjustified as freshwater stingrays are abundant and commonly found in freshwater rivers of the country but still unexplored by the scientific community, as already shown by Soares, Petean (2023)Soares KDA, Petean FF. Three decades of Chondrichthyan research in Brazil assessed from conferences’ abstracts: patterns, gaps, and expectations. Neotrop Ichthyol. 2023; 21(2):e230027. https://doi.org/10.1590/1982-0224-2023-0027
https://doi.org/10.1590/1982-0224-2023-0...
. However, disparity in research efforts of certain taxa may be related to other reasons such as shortage of permanent human resources (e.g., with shark molecular taxonomy qualifications), lack of equipment and facilities (e.g., molecular laboratories and supplies), and, more precisely, direct research funding to conduct costly genetic analyses.

Molecular phylogenetic studies are also in shortage and have been applied to a handful taxa that include only six families occurring in Brazil (out of 42 families): Aetobatidae (Sales et al., 2019)Sales JBL, Oliveira CN, Santos WCR, Rotundo MM, Ferreira Y, Ready J et al. Phylogeography of eagle rays of the genus Aetobatus: Aetobatus narinari is restricted to the continental western Atlantic Ocean. Hydrobiologia. 2019; 836(1):169–83. https://doi.org/10.1007/s10750-019-3949-0
https://doi.org/10.1007/s10750-019-3949-...
, Arhynchobatidae (Coelho et al., 2020Coelho JFR, Lima SMQ, Petean FF. Phylogenetic conservatism of abiotic niche in sympatric Southwestern Atlantic skates. Mar Biol Res. 2020; 16(6–7):458–73. https://doi.org/10.1080/17451000.2020.1837883
https://doi.org/10.1080/17451000.2020.18...
), Carcharhinidae (Mendonça et al., 2011Mendonça FF, Oliveira C, Burgess G, Coelho R, Piercy A, Gadig OBF et al. Species delimitation in sharpnose sharks (genus Rhizoprionodon) in the western Atlantic Ocean using mitochondrial DNA. Conserv Genet. 2011; 12(1):193–200. https://doi.org/10.1007/s10592-010-0132-6
https://doi.org/10.1007/s10592-010-0132-...
; Rodrigues Filho et al., 2023Rodrigues Filho LFS, Nogueira PC, Sodré D, Leal JRS, Nunes JL, Rincon G et al. Evolutionary history and taxonomic reclassification of the Critically Endangered daggernose shark, a species endemic to the Western Atlantic. J Zool Syst Evol Res. 2023; 2023. https://doi.org/10.1155/2023/4798805
https://doi.org/10.1155/2023/4798805...
), Dasyatidae (Vaz et al., 2006Vaz LAL, Carreiro CR, Goulart Filho LR, Furtado Neto MA. Relações filogenéticas em raias (Dasyatis, Elasmobranchii) do estado do Ceará, Brasil. Arq Ciênc Mar Fortaleza. 2006; 39(1):86–88.; Petean et al., 2020Petean FF, Naylor GJP, Lima SMQ. Integrative taxonomy identifies a new stingray species of the genus Hypanus Rafinesque, 1818 (Dasyatidae, Myliobatiformes) from the Tropical Southwestern Atlantic. J Fish Biol. 2020; 97(4):1120–42. https://doi.org/10.1111/jfb.14483
https://doi.org/10.1111/jfb.14483...
), Potamotrygonidae (Carvalho, Lovejoy, 2011Carvalho MR, Lovejoy NR. Morphology and phylogenetic relationships of a remarkable new genus and two new species of Neotropical freshwater stingrays from the Amazon basin (Chondrichthyes: Potamotrygonidae). Zootaxa. 2011; 2776(1):13–48. https://doi.org/10.11646/ZOOTAXA.2776.1.2
https://doi.org/10.11646/ZOOTAXA.2776.1....
; Fontenelle et al., 2021Fontenelle JP, Lovejoy NR, Kolmann MA, Marques FPL. Molecular phylogeny for the Neotropical freshwater stingrays (Myliobatiformes: Potamotrygoninae) reveals limitations of traditional taxonomy. Biol J Linn Soc. 2021; 134(2):381–401. https://doi.org/10.1093/biolinnean/blab090
https://doi.org/10.1093/biolinnean/blab0...
; Kolmann et al., 2022Kolmann MA, Marques FP, Weaver JC, Dean MN, Fontenelle JP, Lovejoy NR. Ecological and phenotypic diversification after a continental invasion in Neotropical freshwater stingrays. Integr Comp Biol. 2022; 62(2):424–40. https://doi.org/10.1093/icb/icac019
https://doi.org/10.1093/icb/icac019...
), and Sphyrnidae hammerheads in Cavalcanti (2007)Cavalcanti MJ. A phylogenetic supertree of the hammerhead sharks (Carcharhiniformes, Sphyrnidae). Zool Stud. 2007; 46(1):6–11. and Gonzalez et al. (2021)Gonzalez C, Postaire B, Domingues RR, Feldheim KA, Caballero S, Chapman D. Phylogeography and population genetics of the cryptic bonnethead shark Sphyrna aff. tiburo in Brazil and the Caribbean inferred from mtDNA markers. J Fish Biol. 2021; 99(6):1899–911. https://doi.org/10.1111/jfb.14896
https://doi.org/10.1111/jfb.14896...
. More rarely phylogenetic studies that applied combined morphological and molecular data have been represented to a minimum as seen in Carvalho, Lovejoy (2011)Carvalho MR, Lovejoy NR. Morphology and phylogenetic relationships of a remarkable new genus and two new species of Neotropical freshwater stingrays from the Amazon basin (Chondrichthyes: Potamotrygonidae). Zootaxa. 2011; 2776(1):13–48. https://doi.org/10.11646/ZOOTAXA.2776.1.2
https://doi.org/10.11646/ZOOTAXA.2776.1....
and Soares, Mathubara (2022)Soares KDA, Mathubara K. Combined phylogeny and new classification of catsharks (Chondrichthyes: Elasmobranchii: Carcharhiniformes). Zool J Linn Soc. 2022; 195(3):761–814. https://doi.org/10.1093/zoolinnean/zlab108
https://doi.org/10.1093/zoolinnean/zlab1...
. Genomic studies of phylogenetic inference are still exceptional (e.g., Amaral et al., 2018Amaral CRL, Pereira F, Silva DA, Amorim A, Carvalho EF. The mitogenomic phylogeny of the Elasmobranchii (Chondrichthyes). Mitochondrial DNA Part A DNA Mapping, Seq Anal. 2018; 29(6):867–78. https://doi.org/10.1080/24701394.2017.1376052
https://doi.org/10.1080/24701394.2017.13...
). Slow-growing trends of molecular Systematics in Brazil are also evident through examination of the online repositories of genetic sequences in which the availability of genetic information related to national and endemic species are scant.

Being a mega diverse country, it is expected that studies related to the taxonomy, phylogeny and morphology of Chondrichthyes in Brazil exhibit diversity of target taxa as well. Shark studies are more frequently conducted than batoids and, more rarely, chimeras, representing, respectively, 52.0%, 46.7% and 1.2% of the total number of studies produced during over 120-year period. Potamotrygonidae and Carcharhinidae consist of the two main study groups, representing over 10% each of the total number of studies. Dasyatids, hammerhead sharks, skates, catsharks and angelsharks represent, each, approximately 5.0% of the total studies. Availability and abundance of material in the wild, fish markets and scientific collections possibly have influenced the research outputs related to these taxa as freshwater stingrays, requiem sharks, dasyatids, skates, catsharks and hammerhead sharks are more easily spotted.

Remaining 26 families that occur in Brazil represent less than 30.0% of the total studies. Most of these taxa are pelagic species and demersal dwellers that inhabit deep-water realms such as cownose rays, thresher sharks, deep-water skates, lantern sharks which makes us infer that the lack of scientific expeditions and research funding in the Brazilian open ocean serve as drivers to the taxonomic impediment. Surprisingly, guitarfishes, electric rays, butterfly rays and smooth-hound sharks that are easily found in shallow coastlines and in artisanal fish markets have been scarcely investigated probably due its non-charismatic features even though these taxa represent one of the most problematic groups with guitarfishes being one of the most vulnerable taxa to extinction (Moore, 2017)Moore ABM. Are guitarfishes the next sawfishes? Extinction risk and an urgent call for conservation action. Endanger Species Res. 2017; 34:75–88. https://doi.org/10.3354/esr00830
https://doi.org/10.3354/esr00830...
. Other taxa that have been listed to the country, including Ginglymostomatidae, Pseudocarcharhidae, Pseudotriakidae, Torpedinidae, and Rhinochimaeridae have not yet received any direct attention to Systematics research.

Future research perspectives and priorities. High susceptibility to environmental impacts such as overfishing and climate changes arises from the combining exposure to fishing mortality, life history and ecological traits of sharks, rays and chimeras (Dulvy et al., 2014)Dulvy NK, Fowler SL, Musick JA, Cavanagh RD, Kyne PM, Harrison LR et al. Extinction risk and conservation of the world’s sharks and rays. Elife. 2014; e00590(3):1–35. https://doi.org/10.7554/eLife.00590.001
https://doi.org/10.7554/eLife.00590.001...
. Many species of sharks, batoids and chimeras are naturally vulnerable due to its k-selected (e.g., slow growth rate, late maturity, long gestation periods, low fecundity) and ecological features (e.g., habitat-specific, philopatry, limited dispersal) (Moore, 2017)Moore ABM. Are guitarfishes the next sawfishes? Extinction risk and an urgent call for conservation action. Endanger Species Res. 2017; 34:75–88. https://doi.org/10.3354/esr00830
https://doi.org/10.3354/esr00830...
. It is estimated that 32% of the elasmobranch marine species are under extinction risk in Brazil (Rosa, Gadig, 2014Rosa RS, Gadig OBF. Conhecimento da diversidade dos Chondrichthyes marinhos no Brasil: a contribuição de José Lima De Figueiredo. Arq Zool Mus Zool Univ São Paulo. 2014; 45:89–104. https://doi.org/10.11606/issn.2176-7793.v45iespp89-104
https://doi.org/10.11606/issn.2176-7793....
; Reis et al., 2016Reis RE, Albert JS, Di Dario F, Mincarone MM, Petry P, Rocha LA. Fish biodiversity and conservation in South America. J Fish Biol. 2016; 89(1):12–47. https://doi.org/10.1111/jfb.13016
https://doi.org/10.1111/jfb.13016...
; ICMBio, 2018Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio). Livro vermelho da fauna brasileira ameaçada de extinção. Volume 1. Brasília: ICMBio/MMA; 2018. ). Industrial and artisanal fisheries play a major influence over the resiliency of local populations and species together with habitat loss, degradation and coastal development as additional threats. Currently, Brazil represents the second in place for the global ranking of shark meat producer and fin exporter, especially of highly migratory and endangered species (Falcão et al., 2014Falcão LHO, Furtado-Neto MAA, Maggioni R, Faria VV. Prospective molecular markers for the identification of illegally traded angelsharks (Squatina) and dolphin (Sotalia guianensis). Genet Mol Res. 2014; 13(4):9710–17. https://doi.org/10.4238/2014.November.24.2
https://doi.org/10.4238/2014.November.24...
; Barreto et al., 2017Barreto RR, Bornatowski H, Motta FS, Santander-Neto J, Vianna GMS, Lessa RP. Rethinking use and trade of pelagic sharks from Brazil. Mar Policy. 2017; 85:114–22. https://doi.org/10.1016/j.marpol.2017.08.016
https://doi.org/10.1016/j.marpol.2017.08...
). Forensic investigations in the fishing industry have evidenced that shark meat is widely effective for consumption under mislabeled products in supermarkets and fish markets (e.g., Bornatowski et al., 2013Bornatowski H, Braga RR, Vitule JRS. Shark mislabeling threatens biodiversity. Science. 2013; 340(6135):923. https://doi.org/10.1126/science.340.6135.923-a
https://doi.org/10.1126/science.340.6135...
; Feitosa et al., 2018Feitosa LM, Martins APB, Giarrizzo T, MacEdo W, Monteiro IL, Gemaque R et al. DNA-based identification reveals illegal trade of threatened shark species in a global elasmobranch conservation hotspot. Sci Rep. 2018; 8(1):1–11. https://doi.org/10.1038/s41598-018-21683-5
https://doi.org/10.1038/s41598-018-21683...
; Rodrigues Filho et al., 2020Rodrigues Filho LFS, Feitosa LM, Nunes JLS, Palmeira ARO, Martins APB, Giarrizzo T et al. Molecular identification of ray species traded along the Brazilian Amazon coast. Fish Res. 2020; 223:105407. https://doi.org/10.1016/j.fishres.2019.105407
https://doi.org/10.1016/j.fishres.2019.1...
). This scenario thus set the country in peril regarding shark sustainable fisheries as the fisheries sector has faced the worst management crises over the last decade (Di Dario et al., 2015Di Dario F, Alves C, Boos H, F’redou FL, Lessa RPT, Mincarone MM et al. A better way forward for Brazil’s fisheries. Science. 2015; 347:1079. https://doi.org/10.1126/science.347.6226.1079-a
https://doi.org/10.1126/science.347.6226...
; Martins et al., 2018Martins APB, Feitosa LM, Lessa RP, Almeida ZS, Heupel M, Silva WM et al. Analysis of the supply chain and conservation status of sharks (Elasmobranchii: Superorder Selachimorpha) based on fisher knowledge. PLoS ONE. 2018; 13(3):e0193969. https://doi.org/10.1371/journal.pone.0193969
https://doi.org/10.1371/journal.pone.019...
).

Target or by-catch species are usually reported on fishing logbooks lacking accurate taxonomic identifications (e.g., Sphyrna and Carcharhinus) and/or are barely assigned to general vernacular names such as “cação” or “raia”. Fisheries statistics available for Brazil commonly list five species that are easily identifiable (P. glauca, I. oxyrinchus, C. longimanus, C. falciformis,and A. superciliosus; Barreto et al., 2017Barreto RR, Bornatowski H, Motta FS, Santander-Neto J, Vianna GMS, Lessa RP. Rethinking use and trade of pelagic sharks from Brazil. Mar Policy. 2017; 85:114–22. https://doi.org/10.1016/j.marpol.2017.08.016
https://doi.org/10.1016/j.marpol.2017.08...
) while less charismatic and small-sized species that are still under fishing pressures are unreported. Inability to identify and apply species-name of sharks, rays and chimeras is a global tendency and not less locally Bornatowski et al. (2014)Bornatowski H, Braga RR, Vitule JRS. Threats to sharks in a developing country: the need for effective and simple conservation measures. Nat Conservacao. 2014; 12(1):11–18. https://doi.org/10.4322/natcon.2014.003
https://doi.org/10.4322/natcon.2014.003...
which hampers species-specific fisheries jurisdictions and conservation discharges. Knowledge gaps persist for a variety of taxa from Brazil that require taxonomic clarification regarding morphological and genetic characterization, application of synonyms, and type designation such as for guitarfishes Pseudobatos, stingrays Hypanus, skates Rajella, smooth-hound sharks Mustelus, and requiem sharks Carcharhinus.

Taxonomic impediment in Brazil obscures research development in Ichthyology. Absence of substantiate funding in STEM research that includes Systematics, limited research efforts through scientific expeditions in open ocean and remote freshwater regions, scarcity of permanent and full-time taxonomists in museums and universities, and reduced investment in education constitute its major drivers. Deficiency of scientific awareness on the sharks, rays and chimaeras diversity in the country also urges the impediment as it does not attract new students, research interests and public engagement. To gather Brazil within the worldwide framework in the modern Chondrichthyan Systematics it is imperative the implementation of strategic initiatives in research, education and infrastructure.

Modernization of zoological collections, such as digitalizing inventories of available specimens and tissues, will facilitate access to biodiversity data for the academic and general public. Currently, to know some collections’ holdings, the manager should be inquired to send a list, since it is not freely available on an online database, hindering a fast access to data and progress of studies. Often, lists of elasmobranchs deposited in national museums carry misidentifications and/or outdated identifications, hampering the knowledge on taxa in need of review. For taxonomic and systematic research, it is of utmost importance for this transition toward digitalized collections to happen soon, so scientists can acknowledge taxa with distributional gaps that need higher sampling efforts to accomplish comprehensive studies. In this way, offering scholarships aimed at updating and digitalizing museum collection catalogs could represent a viable solution. In prevailing scenario, only those taxa easily accessed are studied, generating the bias we have noticed throughout this research.

Novel methodologies in Systematics research allied to evolutionary ecology and genomics which exemplify the so-called Integrative Systematics (e.g., population genomics; eDNA; phylogenomics) will indeed uncover species delimitations and boost taxonomic and phylogenetic resolutions. Strengthening collaborative network among national and international shark experts, especially in Latin America, will further joint multi-lateral forces to minimize taxonomic confusions of taxa. Not less importantly, public outreach involvement in universities, institutions and museums by promoting educative programs and activities as well as publication of easy-to-learn identification guides will definitely raise the scientific awareness regarding the conservation and diversity of the group in Brazil.

As observed in our study, and in Soares, Petean (2023)Soares KDA, Petean FF. Three decades of Chondrichthyan research in Brazil assessed from conferences’ abstracts: patterns, gaps, and expectations. Neotrop Ichthyol. 2023; 21(2):e230027. https://doi.org/10.1590/1982-0224-2023-0027
https://doi.org/10.1590/1982-0224-2023-0...
, Brazilian chondrichthyan research is very male dominated research field. More than half (60.7%) of analyzed papers were authored only by men, revealing a huge gender bias that still hampers women to participate as lead authors. It is mandatory the gender inclusion in future studies, research expeditions, regional book editions and other types of scientific outputs.

ACKNOWLEDGEMENTS

Authors thank Otto B. F. Gadig and Ricardo S. Rosa for their inputs and suggestions in the initial stages of this study. Hugo Santos (UERJ) and James Maclaine (BMNH) are thanked for assisting with information in the biodiversity database of the respective ichthyological collections. SV thanks Mark Lisher, Dylan Clarke and Wayne Florence (Iziko Museums of South Africa) for their institutional support. SV thanks III Sharks International Conference 2018 through the Sharks International Travel Grant in support to the oral presentation of the preliminary results of this study.

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ADDITIONAL NOTES

  • HOW TO CITE THIS ARTICLE

    Viana S, Petean FF, Soares K. Chondrichthyan systematics in Brazil depicted: historical overview, research trends and future perspectives. Neotrop Ichthyol. 2024; 22(3):e240011. https://doi.org/10.1590/1982-0224-2024-0011

Edited-by

Osmar Luiz

Publication Dates

  • Publication in this collection
    23 Sept 2024
  • Date of issue
    2024

History

  • Received
    19 Feb 2024
  • Accepted
    10 June 2024
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E-mail: neoichth@nupelia.uem.br