Ichthyofauna of Paraíba State, Brazil: diversity patterns, knowledge gaps, and challenges for fish conservation and sustainable use

Viviana Márquez-Velásquez^1,2 , Ricardo S. Rosa³, Manoela Maria Ferreira Marinho⁴, João Paulo Capretz Batista da Silva³, Telton Pedro Anselmo Ramos⁵, Ana Carolina Figueiredo Lacerda⁶, Gabriel Beltrão³, André Castro³, Marcio Wanderson³, Danielle Cristina Gamba³, Laure Berti-Equille^1,7 and Rafael L. G. Raimundo^1,2,7

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Associate Editor: Osmar Luiz

Section Editor: Osmar Luiz

Editor-in-chief: José Birindelli

Abstract​

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Introduction​

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Aquatic ecosystems in the State of Paraíba, Brazil, hold over 560 fish species, a number resulting from the interplay of ecological gradients, biogeographical history, habitat heterogeneity, and anthropogenic processes (Márquez Velásquez et al., 2025). Freshwater fishes inhabit waters that drain two contrasting biomes: (i) the semiarid Caatinga, part of the Mid-Northeastern Caatinga Ecoregion (MNCE, sensu Rosa et al., 2003b), and (ii) the Atlantic Forest, being part of its highly threatened Pernambuco Endemism Centre (PEC; Silva, Casteleti, 2003), a global biodiversity hotspot (Myers et al., 2000). The Caatinga has a dense network of intermittent rivers that exhibit high fish endemism (Rosa et al., 2003b, 2004; Albert et al., 2011), and the Atlantic Forest includes PEC’s largest remaining forest fragments and small coastal streams flowing into the Atlantic. These biomes create unique conditions for faunal overlaps and transitions related to ecological and hydrological connectivity, which are intensified by anthropogenic processes.

Marine fishes of Paraíba occur in vast array of habitats, including tide pools, estuaries, shallow and deep reefs, and oceanic environments. Among these, coral reefs are particularly important, forming part of the richest coral reef system in the southwestern Atlantic (Leão et al., 2010), a biodiversity hotspot shaped by biogeographic and oceanographic processes. Locally, reefs are typically sandstone formations on the inner continental shelf (Leão et al., 2003), often forming long beachrock lines (Laborel, 1970). In these habitats, the ichthyofauna supports key socioeconomic activities, such as tourism and fishing (Costa et al., 2007; Paulo Júnior et al., 2012), with artisanal fishing comprising over 80% of the catches (IBAMA, 2008). The high fish diversity in this region is result of vicariance events, such as those associated with the final closure of the Isthmus of Panama (Coates, Obando, 1996); the presence of major dispersal barriers, such as the Amazon-Orinoco plume, which limits dispersal and contribute to the divergence between Brazilian and Caribbean assemblages (Rocha et al., 2005; Floeter et al., 2008). Additionally, the heavy sediment load in the Brazilian Northeastern Ecoregion of the Southwestern Atlantic (sensu Spalding et al., 2007) further isolates the northeastern and eastern portions of the coast (Leão et al., 2003).

Initial efforts to describe fish species in Northeastern Brazil began in the mid-17th century, with the publication of Historia Naturalis Brasiliae (Piso, Marcgrave, 1648), which documented the diversity of fishs from the São Francisco River to the state of Maranhão (Medeiros, Albuquerque, 2014; Alcantara-Rodriguez et al., 2019). This work presents detailed illustrations of marine and freshwater species (Willemsen, 2023). Later, at the end of the 19th century, the Geological Commission of the Empire conducted fish surveys in Paraíba, whose collected specimens are currently in the National Museum of Rio de Janeiro (Sanjad, 2004). Beaurepaire-Rohan (1911) compiled a list of Paraíba’s fishes by their common names and Miranda Ribeiro (1937, 1939) described several regional marine fish species in existing collections. Moreover, Starks (1913) surveyed the mouth of the Mamanguape River, while Fowler (1941, 1948, 1954) advanced systematic descriptions of freshwater species. Furthermore, the Technical Commission for Fish Farming in Northeast Brazil initiated its work in 1933, reinforcing pioneering studies of Paraíba fishes (Canan, 2011).

Over the last decades, ichthyology in the Paraíba State transitioned from a taxonomic focus to more multidisciplinary approaches, including ecology, reproduction, ethnobiology, parasitology, and fisheries management. This research has largely concentrated on major perennial and intermittent water courses, such as the Paraíba and Piranhas rivers, along with their respective coastal areas and basins (Rosa et al., 2003b; Honório et al., 2010; Xavier et al., 2012). However, other coastal rivers with ecological and socioeconomic relevance, such as the Jacu and Guaju rivers, remain poorly studied (Márquez Velásquez et al., 2025). In marine environments, research has focused mostly in coastal ecosystems, including estuaries and mangroves (e.g., Xavier et al., 2012; Medeiros et al., 2018, Ramos et al., 2019), intertidal zones (e.g., Rosa et al., 1997; Silva Nascimento, Ramos 2024), and coral reefs (e.g., Rocha et al., 1998; Souza et al., 2007).

Despite consistent growth of research in both freshwater (Lima et al., 2017; Oliveira-Silva et al., 2018, Ramos et al., 2018a,b, 2019; Marinho et al., 2023) and marine environments (Rosa et al., 1997; Honório et al., 2010; Silva et al., 2021; Marceniuk et al., 2023; Rosa et al., 2023), data remains uneven across time, space, habitats, and research fields, with most data available in grey literature. Scientometric methods can shed light on these imbalances and contribute with public policies and strategies that promote both basic and applied research. Although it is not biologically meaningful to use geopolitical borders to study biodiversity patterns, it is essential for policy-making to understand how knowledge trends vary across them (Beddington et al., 2007; Hortal et al., 2015). Data-driven strategies are critically lacking to guide policies promoting the conservation and sustainable use of aquatic biodiversity and ecosystems across spatial scales (Pullin, Stewart, 2006; Dallimer, Strange, 2015; Olawumi, Chan, 2018).

In this paper, we compiled data from published papers related to the ichthyofauna in the Paraíba State and data from specimens in relevant ichthyological collections. We combine these data with taxonomic and geographical analyses to suggest future directions for ichthyological research by (a) reviewing the development of ichthyological research over time in the Paraíba State, (b) exploring the temporal and disciplinary trends in research topics, (c) assessing gaps in current coverage of habitat types and geographical areas, and (d) describing trends and biases in the current coverage of taxonomic groups.

Material and methods

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Data sources. We performed a literature search in Web of Science (WoS) (https://www.webofscience.com) and Google Scholar (www.google.com) including publications up to September 2024, using the search strings: ((fish* OR Ichth* OR pisces OR Osteichthyes OR Chondrichthyes OR Elasmobranchii) AND (Coast* OR Estuar* OR Marine OR Reef OR Freshwater OR Oceanic OR mangrove OR Freshwater* OR lake*) AND (Paraíba AND Northeast* AND Brazil*)). We performed searches in both English and Portuguese, encompassing titles, abstracts, keywords, and methods. We focused on indexed journal articles and scientific or technical books and excluded grey literature to benefit from the transparency and replicability ensured by peer-reviewed sources (Tab. S1).

We employed a snowball method (Wohlin, 2014) to identify additional publications from the reference lists that met our search criteria. After manually screening titles, abstracts, and methodologies, and excluding articles that did not report data from Paraíba State, we found 198 publication records. We used the open-source software Zotero (www.zotero.org) to organize all bibliographical information. From each selected publication, we extracted the following data: (i) title; (ii) year of publication; (iii) authors (iv) journal; (v) type of article (primary research or review); (vi) sampled locations; (vii) habitat (marine, estuarine, freshwater); (viii) river basin, for continental fishes, according to the Brazilian National Water Agency (ANA, 2015); (ix) research topics; and (x) species or taxonomic group(s). Here, freshwater encompasses lakes, streams, creeks, ponds, wetlands, marshes, and reservoirs, while estuaries are considered a separate category. Additionally, we compiled data from five Brazilian ichthyological collections that holds relevant material from the Paraíba State: Universidade Federal da Paraíba, João Pessoa (UFPB), Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP), Universidade Federal de Rio Grande do Norte, Natal (UFRN), Museu Nacional, Rio de Janeiro (MNRJ), and Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre (PUCRS), focusing only on voucher specimens that were georeferenced or clearly linked to Paraíba State. This data compilation is available at the DATAPB repository (Márquez Velásquez et al., 2025, https://datapb.ccae.ufpb.br/).

Temporal trends. We analysed scientific production by year and habitat, categorising the habitats into three types: (a) freshwater, (b) marine, and (c) estuarine. When a study encompassed multiple habitats, we included it in each relevant habitat. We classified the papers into research topics (see Tab. S1) based on titles and keywords, but verifying also the abstract if the theme remained unclear. If a study covered multiple topics, we counted each one separately. Our classification followed a controlled vocabulary developed to cover the diversity of ecological, taxonomic, and applied themes (Tab. 1). We used these categories to describe temporal trends and thematic focus in research efforts.

TABLE 1 | Research topic categories and their descriptions, used for classifying and assessing temporal trends of the compiled studies.

Topics	Description
Taxonomy, systematics, evolution	Species lists, inventories, descriptions of new species, updates on systematics and taxonomy, evolutionary history, phylogenetic systematics, and genetics.
Anatomy and morphology	Analysis and descriptions of anatomy and morphology.
Distribution and biogeography	Geographical distributions, occurrence, new records, biogeographic and phylogeographic patterns.
Trophic ecology	Diet, feeding habits, and trophic interactions.
Ecology	Species interactions, effects of abiotic variables on populations or communities, pollution, ecomorphology, and migration.
Ethnoecology	Local communities’ knowledge, management of natural resources, and cultural practices.
Physiology	Mechanical, physical, and biochemical functions; ecotoxicological analyses.
Reproduction	Reproductive biology, ontogeny, age and growth, weight-length and length-age relationships, demography.
Parasitology	Parasites, their hosts, and the interactions between them.
Fisheries	Landing data, description of catches, fishing effort, fishing gears, fish processing, marketing, and consumption.
Conservation and management	Vulnerability assessment, demographic analysis, population decline, design of closed areas, invasive species, and ecotourism.
Others	Papers not classified in any of the topics mentioned above, such as Palaeontology, and fish farming.

Geographical trends. We gathered the geographical coordinates of the sampling sites from each relevant publication. When studies provided coordinates, we use them directly. For studies that did not provide coordinates, we inferred the locations from suitably georeferenced maps using WebPlotDigitizer (https://automeris.io/WebPlotDigitizer/). We then created a distribution map in QGIS software v. 2.18. Furthermore, we analysed the number of published papers, categorising them by the main hydrographic basins of Paraíba State (http://www.aesa.pb.gov.br/). This basin-level spatial resolution enabled us to identify sampling imbalances and underrepresented river systems.

Taxonomic trends. We classified species by family, order, and class, and verified their taxonomic status in Eschmeyer’s Catalogue of Fishes (Fricke et al., 2025). Whenever possible, we verified and reported fish habits using sampling locations from collections (Camargo, Isaac, 2001; Morais, 2016; Vendel et al., 2022). We broadly defined estuarine species as those that use estuarine habitats at any life-stage, regardless of whether they primarily inhabit freshwater or marine environments. For each species, we considered human uses: commercial, local consumption, medicinal, aquaculture, and recreational fishing (Pimentel Rocha et al., 2008; Almeida et al., 2014; Marceniuk et al., 2023). We also reported global conservation status based on the IUCN Red List (IUCN, 2025) and national status according to the Sistema de Avaliação do Risco de Extinção da Biodiversidade – SALVE (ICMBio, 2026). We classified extinction risk following IUCN categories: Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT), Least Concern (LC), Data Deficient (DD), and Not Evaluated (NE).

We reported species endemic to the Northeastern Caatinga and Coastal Drainages ecoregion (NCCD) sensu Abell et al., 2008, and to the Northeastern Brazil Marine ecoregion (NBM) sensu Spalding et al., 2007. The NCCD encompasses all river basins located between the Parnaíba and São Francisco rivers. A large portion of this ecoregion is located within the Brazilian semi-arid region (Rosa et al., 2003b; Lima et al., 2017). Most of its tributaries, predominantly intermittent, drain areas of the Caatinga (Rosa et al., 2003b). In the easternmost portion of this ecoregion are small coastal rivers that lie entirely or largely under the influence of the Atlantic Forest and are distributed from southern Rio Grande do Norte to Alagoas (Rosa et al., 2003b; Lima et al., 2017). The NBM ecoregion, located between Piauí and Sergipe states, is nested within the Tropical South-western Atlantic Province (Northeastern and Eastern Brazil ecoregions) (Spalding et al., 2007). Additionally, we identified potentially undescribed or cryptic species in collections. For illustrative images of fish, we used figures from Phylopic.org (Keesey, 2024).

Results​

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Temporal trends. Our search identified 198 scientific publications (Tabs. S1, S2). Research articles comprised the majority (n = 184; 93%), while book or book sections accounted for only 14 (7%) of the total (Tabs. S1, S2). The first ichthyological study on Paraíba State documented the ichthyofauna of the estuarine waters in the Brazilian northeastern region during the 1970s (Oliveira, 1972). Advancements in taxonomy and ecology arose later, founded on investigations into shark fisheries biology (Lima, Ferreira, 1975) and preliminary surveys of marine fishes (Rosa, 1980a,b) (Fig. 1; Tab. S1).

FIGURE 1| Temporal distribution of scientific publications for fish species of Paraíba State, Brazil, between 1972 and 2024, based on their habitats.

A temporal increase in the number of publications across research topics was detected over the studied period, with ecological studies rising sharply over the last 20 years, followed by Taxonomy and Systematics, Reproduction, Ethnoecology, and Fisheries research (Fig. 2A). The most common research topic in the dataset was Ecology (n = 57, 29%), followed by Taxonomy (45, 23%) and Trophic Ecology (28, 14.14%). Together, these categories comprised more than 50% of the dataset. Other areas included Reproduction (21, 10.60%), Distribution/Biogeography (20, 10.10%), Ethnoecology (19, 9.60%), and Fisheries (17, 8.60%). Studies on Parasitology and Anatomy appear in the dataset, though to a lesser extent (6, 3% each). Only 2% (n = 4) of these studies focused on Conservation and Management (Fig. 2A).

FIGURE 2| A. A network summarizing temporal research trends in the scientific knowledge of the ichthyofauna of Paraíba State, Brazil, from 1972 to 2024. B. Number of research production during the same period, categorized by habitat.

Overall, a clear upward trend in publications was evident over the last twenty years, particularly since 2000, driven by increases in Ecology, Taxonomy, and Trophic ecology research (Fig. 2B). In contrast, when considering habitat types, the predominant research topics in marine and estuarine environments, the most frequent research topics were Ecology, followed by Taxonomy/Systematics/Evolution and Trophic Ecology, respectively. In freshwater habitats, Taxonomy/Systematics/Evolution was the most prevalent topic, followed by Ecology and Reproduction (Fig. 2B). For freshwater habitats, Taxonomy/Systematics/Evolution emerged as the most common topic, followed by Ecology and Reproduction (Fig. 2B; Tab. S1).

Representation of habitats and locations in scientific production. The Paraíba River basin had the highest number of studies (n = 61; 42.10%), followed by the Mamanguape basin (49; 33.80%) and the Piranhas River basin (11; 7.60%). The Curimatau (3, 2.10%), Camaratuba (2; 1.40%) and Miriri River basins (1; 0.70%) had much fewer studies (Figs. 3A, B). Notably, there were no publications for the interstate basins of the Jacu, Guaju and Trairi Rivers, all located in northern Paraíba State along the border with Rio Grande do Norte State. Although studies exist for the Trairi River basin, they report exclusively on species recorded in the portion of the basin situated within the State of Rio Grande do Norte. The Tracunhaém River, part of the Goiana River basin (Pernambuco), lies between the states of Paraíba and Pernambuco, with no records available for this drainage in Paraíba. Of the total studies, 70 included the estuarine area, with the Mamanguape River estuary having most of the studies (n = 41, 58.6%), followed by the Paraíba River estuary (20, 28.60%), Abiaí (5, 7.14%), and the Gramame and Camaratuba River estuaries (2, 2.90%).

FIGURE 3| Distribution pattern of sampling and scientific knowledge on the ichthyofauna of Paraíba State, Brazil, from 1972 to 2024. A. The purple points on the map represent the sampling locations across the state. B. The data is categorized by the main river basins, showing an uneven distribution of research efforts among them.

In marine regions, the coastal area accounted for nearly 40% of the published studies (Fig. 3A). While estuaries and shallow coastal zones are well studied, deep-sea and shelf-edge areas remain largely underexplored. The distribution of lots within hydrographic basins exhibits a similar pattern, with the Paraíba River basin having the largest number of lots (n = 2,161), followed by Mamanguape (n = 1,458) and Piranhas Rivers (n = 700). Abiaí (478), Gramame (369), Miriri (97), Curimataú (71), and Camaratuba Rivers (33) are less represented. There are no recorded lots for the Jacu, Guaju, and Trairi River basins in Paraíba territory. Marine records dominate the dataset, comprising a total of 2,909 lots, highlighting the research emphasis along the coast.

Scientific production by taxa. This assessment includes 565 valid species, comprising teleost and elasmobranch fishes found in freshwater and marine/estuarine environments. These species belong to 39 orders and 132 families (Tab. 2; Fig. 4). To ensure the accuracy of the diversity patterns presented, we excluded questionable records that may have resulted from misidentifications.

TABLE 2 | Fish species found in Paraíba State, Northeastern Brazil, with non-native species indicated by asterisk. The symbol (?) denotes species whose occurrence is reported in biological collections but are possibly misidentifications. Endemic species to the Northeastern Caatinga and Coastal Drainagesecoregion (NCCD) are highlighted in bold. The IUCN status reflects the global assessment, and the BR status refers to the Sistema de Avaliação do Risco de Extinção da Biodiversidade – SALVE (ICMBio, 2026). NE: Not Evaluated, NA: Not Applicable. Species uses: Co: commercial, Or: ornamental in aquariums, F: food, Lc: local consumption, Me: medicinal, Aq: aquaculture, Rf: recreational fishing. Source: Li, Literature; ichthyological collections where the species is deposited (Tabs. S1, S2). ^a Endemic to the Mamanguape River, Paraíba, ^b Endemic to the Paraíba River, Paraíba, ^c Endemic to the Northeastern Brazil Marine ecoregion (NBM).

Class/Order	Family	Species	Habit	IUCN	BR	Use	Source
Elasmobranchii
Hexanchiformes	Hexanchidae	Hexanchus griseus (Bonnaterre, 1788)	Marine	NT	LC		Li
Orectolobiformes	Rhincodontidae	Rhincodon typus Smith, 1828	Marine	EN	VU		Li
Orectolobiformes	Ginglymostomatidae	Ginglymostoma cirratum (Bonnaterre, 1788)	Marine	VU	VU	Lc	CIUFPB, Li
Lamniformes	Lamnidae	Isurus oxyrinchus Rafinesque, 1810	Marine	EN	CR		Li
Carcharhiniformes	Scyliorhinidae	Scyliorhinus haeckelii (Miranda Ribeiro, 1907)	Marine	DD	LC		CIUFPB, Li
Carcharhiniformes	Triakidae	Mustelus higmani Springer & Lowe, 1963	Marine	EN	DD		CIUFPB, Li
Carcharhiniformes	Triakidae	Pseudotriakis microdon Capello, 1868	Marine	LC	LC		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus acronotus (Poey, 1860)	Marine	EN	VU		CIUFPB, Li
Carcharhiniformes	Carcharhinidae	Carcharhinus plumbeus (Nardo, 1827)	Marine	EN	CR		CIUFPB, Li
Carcharhiniformes	Carcharhinidae	Carcharhinus falciformis (Bibron, 1839)	Marine	VU	CR		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus leucas (Valenciennes, 1839)	Marine-Estuary	VU	VU		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus limbatus (Valenciennes, 1839)	Marine-Estuary	VU	NT		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus obscurus (Lesueur, 1818)	Marine-Estuary	EN	EN		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus perezi (Poey, 1876)	Marine	EN	VU		Li
Carcharhiniformes	Carcharhinidae	Carcharhinus porosus (Ranzani, 1839)	Marine-Estuary	CR	CR	Lc	CIUFPB, Li
Carcharhiniformes	Carcharhinidae	Prionace glauca (Linnaeus, 1758)	Marine	NT	NT		Li
Carcharhiniformes	Carcharhinidae	Rhizoprionodon lalandii (Valenciennes, 1839)	Marine	VU	DD	Co, Lc	CIUFPB, Li
Carcharhiniformes	Carcharhinidae	Rhizoprionodon porosus (Poey, 1861)	Marine-Estuary	VU	DD	Co	CIUFPB, Li
Carcharhiniformes	Galeocerdonidae	Galeocerdo cuvier (Péron & Lesueur, 1822)	Marine	NT	NT		CIUFPB, Li
Carcharhiniformes	Sphyrnidae	Sphyrna lewini (Griffith & Smith, 1834)	Marine-Estuary	CR	CR		CIUFPB, Li
Carcharhiniformes	Sphyrnidae	Sphyrna mokarran (Rüppell, 1837)	Marine-Estuary	CR	CR		Li
Carcharhiniformes	Sphyrnidae	Sphyrna aff. tiburo (Linnaeus, 1758)	Marine-Estuary	EN	CR		CIUFPB, Li
Carcharhiniformes	Sphyrnidae	Sphyrna zygaena (Linnaeus, 1758)	Marine-Estuary	VU	CR		CIUFPB, Li
Torpediniformes	Narcinidae	Narcine brasiliensis (Olfers, 1831)	Marine	NT	VU		CIUFPB, Li
Rhinopristiformes	Rhinobatidae	Pseudobatos horkelii (Müller & Henle, 1841)	Marine	CR	CR		CIUFPB, Li
Rhinopristiformes	Rhinobatidae	Pseudobatos percellens (Walbaum, 1792)	Marine	EN	VU	Co	CIUFPB, Li
Myliobatiformes	Dasyatidae	Hypanus americanus (Hildebrand & Schroeder, 1928)	Marine	NT	VU		CIUFPB, Li
Myliobatiformes	Dasyatidae	Hypanus berthalutzae Petean, Naylor & Lima, 2020	Marine	VU	VU		Li
Myliobatiformes	Dasyatidae	Hypanus guttatus (Bloch & Schneider, 1801)	Marine	NT	LC		CIUFPB, Li
Myliobatiformes	Dasyatidae	Hypanus marianae (Gomes, Rosa & Gadig, 2000)	Marine	EN	VU		CIUFPB, Li
Myliobatiformes	Urotrygonidae	Urotrygon microphthalmum Delsman, 1941	Marine	CR	VU	Lc	CIUFPB, Li
Myliobatiformes	Gymnuridae	Gymnura micrura (Bloch & Schneider, 1801)	Marine-Estuary	NT	DD		CIUFPB, Li
Myliobatiformes	Aetobatidae	Aetobatus narinari (Euphrasen, 1790)	Marine	EN	DD	Lc, Me	CIUFPB, Li
Myliobatiformes	Rhinopteridae	Rhinoptera bonasus (Mitchill, 1815)	Marine-Estuary	VU	DD		CIUFPB, Li
Myliobatiformes	Mobulidae	Mobula birostris (Walbaum, 1792)	Marine	EN	VU		Li
Actinopterygii
Elopiformes	Elopidae	Elops smithi McBride, Rocha, Ruiz-Carus & Bowen, 2010	Marine-Estuary	DD	LC		CIUFPB, Li
Elopiformes	Megalopidae	Megalops atlanticus Valenciennes, 1847	Marine-Estuary	VU	VU	Co, Lc, Me	CIUFPB, Li
Albuliformes	Albulidae	Albula nemoptera (Fowler, 1911)	Marine	DD	DD		Li
Albuliformes	Albulidae	Albula vulpes (Linnaeus, 1758)	Marine	NT	DD		CIUFPB, Li
Anguilliformes	Chlopsidae	Chilorhinus suensonii Lütken, 1852	Marine	LC	LC		CIUFPB, Li
Anguilliformes	Moringuidae	Moringua edwardsi (Jordan & Bollman, 1889)	Marine-Estuary	LC	LC		Li
Anguilliformes	Muraenesocidae	Cynoponticus savanna (Bancroft, 1831)	Marine	LC	LC		Li
Anguilliformes	Muraenidae	Echidna catenata (Bloch, 1795)	Marine	LC	LC		Li
Anguilliformes	Muraenidae	Enchelycore nigricans (Bonnaterre, 1788)	Marine	LC	LC		Li
Anguilliformes	Muraenidae	Gymnothorax funebris Ranzani, 1839	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Anguilliformes	Muraenidae	Gymnothorax miliaris (Kaup, 1856)	Marine	LC	LC		Li
Anguilliformes	Muraenidae	Gymnothorax moringa (Cuvier, 1829)	Marine	LC	DD		CIUFPB, Li
Anguilliformes	Muraenidae	Gymnothorax nigromarginatus (Girard, 1858)	Marine	LC	Ne		Li
Anguilliformes	Muraenidae	Gymnothorax ocellatus Agassiz, 1828	Marine	LC	DD		CIUFPB, Li
Anguilliformes	Muraenidae	Gymnothorax vicinus (Castelnau, 1855)	Marine	LC	DD		CIUFPB, MZUSP, Li
Anguilliformes	Muraenidae	Muraena pavonina Richardson, 1845	Marine	LC	LC		Li
Anguilliformes	Muraenidae	Uropterygius macularius (Lesueur, 1825)	Marine	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Ahlia egmontis (Jordan, 1884)	Marine	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Bascanichthys paulensis Storey, 1939	Marine	LC	DD		CIUFPB, Li
Anguilliformes	Ophichthidae	Callechelys bilinearis Kanazawa, 1952	Marine	LC	LC		Li
Anguilliformes	Ophichthidae	Myrichthys breviceps (Richardson, 1848)	Marine	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Myrichthys ocellatus (Lesueur, 1825)	Marine	LC	LC	Co(Or)	CIUFPB, Li
Anguilliformes	Ophichthidae	Myrophis platyrhynchus Breder, 1927	Marine-Estuary	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Myrophis punctatus Lütken, 1852	Marine-Estuary	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Ophichthus cylindroideus (Ranzani, 1839)	Marine-Estuary	LC	LC		CIUFPB, Li
Anguilliformes	Ophichthidae	Ophichthus ophis (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Anguilliformes	Congridae	Heteroconger camelopardalis (Lubbock, 1980)	Marine	LC	DD		Li
Anguilliformes	Congridae	Heteroconger longissimus Günther, 1870	Marine	LC	DD		Li
Anguilliformes	Congridae	Paraconger caudilimbatus (Poey, 1867)	Marine	LC	Ne		CIUFPB, Li
Clupeiformes	Engraulidae	Anchoviella vaillanti (Steindachner, 1908)*	Freshwater	LC	LC		Li
Clupeiformes	Engraulidae	Anchoa filifera (Fowler, 1915)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Anchoa hepsetus (Linnaeus, 1758)	Marine-Estuary	LC	LC		Li
Clupeiformes	Engraulidae	Anchoa januaria (Steindachner, 1879)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Anchoa lyolepis (Evermann & Marsh, 1900)	Marine	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Anchoa spinifer (Valenciennes, 1848)	Marine-Estuary	LC	LC		CIUFPB, MNRJ, Li
Clupeiformes	Engraulidae	Anchoa tricolor (Spix & Agassiz, 1829)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Clupeiformes	Engraulidae	Anchovia clupeoides (Swainson, 1839)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Anchoviella brevirostris (Günther, 1868)	Marine-Estuary	LC	LC		Li
Clupeiformes	Engraulidae	Anchoviella lepidentostole (Fowler, 1911)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Cetengraulis edentulus (Cuvier, 1829)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Clupeiformes	Engraulidae	Engraulis anchoita Hubbs & Marini, 1935	Marine	LC	LC		CIUFPB, Li
Clupeiformes	Engraulidae	Lycengraulis grossidens (Spix & Agassiz, 1829)	Marine-Estuary	LC	LC	Lc	CIUFPB, Li
Clupeiformes	Pristigasteridae	Chirocentrodon bleekerianus (Poey, 1867)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Pristigasteridae	Odontognathus mucronatus Lacepède, 1800	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Clupeiformes	Pristigasteridae	Pellona harroweri (Fowler, 1917)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Dorosomatidae	Harengula clupeola (Cuvier, 1829)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Clupeiformes	Dorosomatidae	Lile piquitinga (Schreiner & Miranda Ribeiro, 1903)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Dorosomatidae	Opisthonema oglinum (Lesueur, 1818)	Marine-Estuary	LC	LC	Co, Lc	CIUFPB, UFRN, Li
Clupeiformes	Dorosomatidae	Rhinosardinia amazonica (Steindachner, 1879)	Marine-Estuary-Freshwater	LC	LC		CIUFPB
Clupeiformes	Dorosomatidae	Rhinosardinia bahiensis (Steindachner, 1879)	Marine-Estuary	LC	LC		CIUFPB, Li
Clupeiformes	Dorosomatidae	Sardinella aurita Valenciennes, 1847	Marine-Estuary	DD	DD	Co, Lc	CIUFPB, Li
Cypriniformes	Cyprinidae	Cyprinus carpio Wu, Yang, Yue & Huang, 1963*	Freshwater-Estuary	LC	Ne	Co(Aq)	UFRN, Li
Gymnotiformes	Gymnotidae	Gymnotus carapo Linnaeus, 1758	Freshwater	LC	LC		CIUFPB, UFRN, Li
Characiformes	Crenuchidae	Characidium bimaculatum Fowler, 1941	Freshwater	LC	LC		CIUFPB, MZUSP, PUCRS, UFRN , Li
Characiformes	Erythrinidae	Erythrinus erythrinus (Bloch & Schneider, 1801)	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Characiformes	Erythrinidae	Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829)	Freshwater	LC	LC		CIUFPB, Li
Characiformes	Erythrinidae	Hoplias malabaricus (Bloch, 1794)	Freshwater	LC	LC	Co, Lc, Me	CIUFPB, PUCRS, UFRN, Li
Characiformes	Parodontidae	Apareiodon davisi Fowler, 1941	Freshwater	LC	LC		CIUFPB, UFRN, Li
Characiformes	Serrasalmidae	Colossoma macropomum (Cuvier, 1816)*	Freshwater	NT	NT	Aq	CIUFPB, UFRN, Li
Characiformes	Serrasalmidae	Metynnis lippincottianus (Cope, 1870)	Freshwater	LC	LC		CIUFPB, Li
Characiformes	Serrasalmidae	Pygocentrus nattereri Kner, 1858	Freshwater	LC	LC		UFRN
Characiformes	Serrasalmidae	Serrasalmus cf. brandtii Lütken, 1875	Freshwater	LC	LC		CIUFPB
Characiformes	Serrasalmidae	Serrasalmus rhombeus (Linnaeus, 1766)	Freshwater	LC	LC		UFRN
Characiformes	Anostomidae	Leporinus piau Fowler, 1941	Freshwater	LC	LC	Co	CIUFPB, MZUSP, PUCRS, UFRN , Li
Characiformes	Anostomidae	Arhinolemur elongatus (Valenciennes, 1850)	Freshwater	LC	Ne		CIUFPB
Characiformes	Curimatidae	Curimatella lepidura (Eigenmann & Eigenmann, 1889)	Freshwater	LC	LC		UFRN
Characiformes	Curimatidae	Psectrogaster rhomboides Eigenmann & Eigenmann, 1889	Freshwater	LC	LC	Co, Lc	CIUFPB, PUCRS, UFRN, Li
Characiformes	Curimatidae	Steindachnerina notonota (Miranda Ribeiro, 1937)	Freshwater	LC	LC		CIUFPB, MNRJ, PUCRS, UFRN, Li
Characiformes	Prochilodontidae	Prochilodus brevis Steindachner, 1875	Freshwater	LC	LC	Co, Lc, Me	CIUFPB, MNRJ, PUCRS, UFRN, Li
Characiformes	Triportheidae	Triportheus signatus (Garman, 1890)	Freshwater	LC	LC	Co, Lc	CIUFPB, PUCRS, UFRN, Li
Characiformes	Characidae	Cheirodon jaguaribensis Fowler, 1941	Freshwater	DD	LC		CIUFPB, MZUSP, UFRN
Characiformes	Characidae	Compsura heterura Eigenmann, 1915	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Characiformes	Characidae	Phenacogaster calverti (Fowler, 1941)	Freshwater	LC	LC		PUCRS, UFRN, MZUSP
Characiformes	Characidae	Serrapinnus heterodon (Eigenmann, 1915)	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Characiformes	Characidae	Serrapinnus piaba (Lütken, 1875)	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Characiformes	Acestrorhamphidae	Astyanax bimaculatus (Linnaeus, 1758)	Freshwater	LC	LC	Lc	CIUFPB, MZUSP, PUCRS, UFRN , Li
Characiformes	Acestrorhamphidae	Ctenobrycon kennedyi (Eigenmann, 1903)	Freshwater	LC	LC		CIUFPB, PUCRS, MZUSP
Characiformes	Acestrorhamphidae	Ctenobrycon spilurus (Valenciennes, 1850)	Freshwater	LC	LC		UFRN
Characiformes	Acestrorhamphidae	Hemigrammus marginatus Ellis, 1911	Freshwater	LC	LC		CIUFPB, MZUSP, PUCRS, UFRN , Li
Characiformes	Acestrorhamphidae	Hemigrammus rodwayi Durbin, 1909	Freshwater	LC	Ne		CIUFPB, PUCRS, UFRN, Li
Characiformes	Acestrorhamphidae	Hemigrammus unilineatus (Gill, 1858)	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Characiformes	Acestrorhamphidae	Hyphessobrycon piabinhas Fowler, 1941	Freshwater	DD	LC		CIUFPB
Characiformes	Acestrorhamphidae	Moenkhausia costae (Steindachner, 1907)*	Freshwater	LC	LC		CIUFPB, UFRN, Li
Characiformes	Acestrorhamphidae	Psalidodon fasciatus (Cuvier, 1819)	Freshwater	LC	LC		CIUFPB, UFRN, MZUSP, Li
Siluriformes	Callichthyidae	Brochis cf. garbei (Ihering, 1911)	Freshwater	LC	LC		To be cataloged
Siluriformes	Callichthyidae	Callichthys callichthys (Linnaeus, 1758)	Freshwater	LC	LC		CIUFPB, PUCRS
Siluriformes	Callichthyidae	Hoplosternum littorale (Hancock, 1828)	Freshwater	LC	LC		To be cataloged
Siluriformes	Callichthyidae	Megalechis thoracata (Valenciennes, 1840)	Freshwater	LC	LC		CIUFPB, Li
Siluriformes	Loricariidae	Hypostomus pusarum Starks, 1913	Freshwater	LC	LC	Co, Lc	CIUFPB, PUCRS, UFRN, Li
Siluriformes	Loricariidae	Parotocinclus cearensis Garavello, 1977	Freshwater	LC	LC		UFRN, PUCRS
Siluriformes	Loricariidae	Parotocinclus jacksoni Ramos, Lustosa-Costa, Barros-Neto & Barbosa, 2021a	Freshwater	Ne	Ne		CIUFPB, MNRJ, Li
Siluriformes	Loricariidae	Parotocinclus jumbo Britski & Garavello, 2002	Freshwater	LC	LC		CIUFPB, MNRJ, MZUSP, PUCRS, UFRN, Li
Siluriformes	Loricariidae	Parotocinclus seridoensis Ramos, Barros-Neto, Britski & Lima, 2013	Freshwater	LC	EN		CIUFPB, PUCRS, UFRN, Li
Siluriformes	Loricariidae	Parotocinclus spilosoma (Fowler, 1941)b	Freshwater	LC	EN		CIUFPB, PUCRS, UFRN, Li
Siluriformes	Loricariidae	Pseudancistrus papariae Fowler, 1941	Freshwater	Ne	Ne		UFRN
Siluriformes	Loricariidae	Pseudancistrus genisetiger Fowler, 1941	Freshwater	LC	LC		CIUFPB, PUCRS
Siluriformes	Auchenipteridae	Trachelyopterus galeatus (Linnaeus, 1766)	Freshwater	LC	LC	Co	ANSP-Ichthyology, Li, UFRN
Siluriformes	Heptapteridae	Pimelodella enochi Fowler, 1941	Freshwater	LC	LC		ANSP-Ichthyology, UFRN, Li
Siluriformes	Heptapteridae	Rhamdia quelen (Quoy & Gaimard, 1824)	Freshwater	LC	LC		CIUFPB, UFRN, Li
Siluriformes	Ariidae	Bagre bagre (Linnaeus, 1766)	Marine-Estuary	LC	NT	Co	CIUFPB, Li
Siluriformes	Ariidae	Bagre filamentosus (Swainson, 1839)	Marine	Ne	LC	Co	CIUFPB
Siluriformes	Ariidae	Cathorops agassizii (Eigenmann & Eigenmann, 1888)	Marine-Estuary	LC	LC		CIUFPB, Li
Siluriformes	Ariidae	Cathorops arenatus (Valenciennes, 1840)	Marine-Estuary	LC	LC		Li
Siluriformes	Ariidae	Cathorops spixii (Agassiz, 1829)	Marine-Estuary	Ne	LC		CIUFPB, Li
Siluriformes	Ariidae	Notarius luniscutis (Valenciennes, 1840)	Marine-Estuary	LC	LC		CIUFPB
Siluriformes	Ariidae	Notarius parmocassis (Valenciennes, 1840)	Marine-Estuary	Ne	DD		Li
Siluriformes	Ariidae	Sciades herzbergii Bloch, 1794	Marine-Estuary	LC	LC		CIUFPB, Li
Siluriformes	Ariidae	Sciades proops (Valenciennes, 1840)	Marine-Estuary	Ne	DD		CIUFPB, Li
Stomiiformes	Gonostomatidae	Diplophos taenia Günther, 1873	Marine	LC	LC		Li
Stomiiformes	Gonostomatidae	Gonostoma atlanticum Norman, 1930	Marine	LC	LC		Li
Stomiiformes	Gonostomatidae	Sigmops elongatus (Günther, 1878)	Marine	LC	LC		Li
Stomiiformes	Gonostomatidae	Zaphotias pedaliotus (Goode & Bean, 1896)	Marine	LC	Ne		Li
Stomiiformes	Sternoptychidae	Argyropelecus aculeatus Valenciennes, 1850	Marine	LC	LC		Li
Stomiiformes	Sternoptychidae	Argyropelecus affinis Garman, 1899	Marine	LC	LC		Li
Stomiiformes	Sternoptychidae	Sternoptyx diaphana Hermann, 1781	Marine	LC	LC		Li
Stomiiformes	Sternoptychidae	Sternoptyx pseudobscura Baird, 1971	Marine	LC	LC		Li
Stomiiformes	Phosichthyidae	Ichthyococcus polli Blache, 1964	Marine	LC	Ne		Li
Stomiiformes	Phosichthyidae	Vinciguerria nimbaria (Jordan & Williams, 1895)	Marine	LC	LC		Li
Stomiiformes	Stomiidae	Aristostomias tittmanni Welsh, 1923	Marine	LC	Ne		Li
Stomiiformes	Stomiidae	Astronesthes similus Parr, 1927	Marine	LC	Ne		Li
Stomiiformes	Stomiidae	Chauliodus sloani Bloch & Schneider, 1801	Marine	LC	LC		Li
Stomiiformes	Stomiidae	Melanostomias bartonbeani Parr, 1927	Marine	LC	Ne		Li
Stomiiformes	Stomiidae	Melanostomias tentaculatus (Regan & Trewavas, 1930)	Marine	LC	LC		Li
Aulopiformes	Giganturidae	Gigantura indica Brauer, 1901	Marine	LC	Ne		Li
Stomiiformes	Stomiidae	Photonectes achirus Regan & Trewavas, 1930	Marine	LC	Ne		Li
Stomiiformes	Stomiidae	Stomias danae Ege, 1933	Marine	LC	LC		Li
Aulopiformes	Scopelarchidae	Rosenblattichthys hubbsi Johnson, 1974	Marine	LC	LC		Li
Aulopiformes	Scopelarchidae	Scopelarchus guentheri Alcock, 1896	Marine	LC	LC		Li
Aulopiformes	Synodontidae	Synodus bondi Fowler, 1939	Marine	LC	LC		Li
Aulopiformes	Synodontidae	Synodus foetens (Linnaeus, 1766)	Marine-Estuary	LC	LC		CIUFPB, Li
Aulopiformes	Synodontidae	Synodus intermedius (Agassiz, 1829)	Marine	LC	LC		CIUFPB, Li
Aulopiformes	Synodontidae	Synodus synodus (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Aulopiformes	Synodontidae	Trachinocephalus myops (Forster, 1801)	Marine	LC	LC		CIUFPB, Li
Gadiformes	Stylephoridae	Stylephorus chordatus Shaw, 1791	Marine	LC	Ne		Li
Beryciformes	Holocentridae	Holocentrus adscensionis (Osbeck, 1765)	Marine	LC	LC	Lc	CIUFPB, MZUSP, Li
Beryciformes	Holocentridae	Holocentrus rufus (Walbaum, 1792)	Marine	LC	LC		CIUFPB, Li
Beryciformes	Holocentridae	Myripristis jacobus Cuvier, 1829	Marine	LC	LC	Lc	CIUFPB, Li
Beryciformes	Holocentridae	Plectrypops retrospinis (Guichenot, 1853)	Marine	LC	LC		CIUFPB, Li
Ophidiiformes	Ophidiidae	Lepophidium brevibarbe (Cuvier, 1829)	Marine	LC	DD		Li
Batrachoidiformes	Batrachoididae	Amphichthys cryptocentrus (Valenciennes, 1837)	Marine-Estuary	LC	LC		OBIS_BR, Li
Batrachoidiformes	Batrachoididae	Batrachoides surinamensis (Bloch & Schneider, 1801)	Marine-Estuary	LC	LC		Li
Batrachoidiformes	Batrachoididae	Porichthys kymosemeum Gilbert, 1968	Marine	LC	LC		CIUFPB, Li
Batrachoidiformes	Batrachoididae	Porichthys plectrodon Jordan & Gilbert, 1882	Marine	LC	LC		Li
Batrachoidiformes	Batrachoididae	Thalassophryne nattereri Steindachner, 1876	Marine-Estuary	LC	LC		CIUFPB, Li
Batrachoidiformes	Batrachoididae	Thalassophryne punctata Steindachner, 1936	Marine-Estuary	LC	LC		CIUFPB, Li
Gobiiformes	Apogonidae	Apogon americanus Castelnau, 1855	Marine	LC	LC		CIUFPB, MZUSP, Li
Gobiiformes	Apogonidae	Apogon pseudomaculatus Longley, 1932	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Apogonidae	Apogon quadrisquamatus Longley, 1934	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Apogonidae	Apogon robbyi Gilbert & Tyler, 1997	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Apogonidae	Astrapogon puncticulatus (Poey, 1867)	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Apogonidae	Astrapogon stellatus (Cope, 1867)	Marine	DD	Ne		CIUFPB, Li
Gobiiformes	Apogonidae	Phaeoptyx pigmentaria (Poey, 1860)	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Eleotridae	Eleotris pisonis (Gmelin, 1789)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, MNRJ, PUCRS, UFRN, Li
Gobiiformes	Eleotridae	Erotelis smaragdus (Valenciennes, 1837)	Marine-Estuary	LC	LC		CIUFPB, Li
Gobiiformes	Eleotridae	Dormitator maculatus (Bloch, 1792)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Eleotridae	Guavina guavina (Valenciennes, 1837)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Awaous tajasica (Lichtenstein, 1822)	Freshwater-Estuary-Marine	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Ctenogobius apogonus Pezold, 2022	Marine	Ne	Ne		Li
Gobiiformes	Oxudercidae	Ctenogobius boleosoma (Jordan & Gilbert, 1882)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Ctenogobius saepepallens (Gilbert & Randall, 1968)	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Ctenogobius shufeldti (Jordan & Eigenmann, 1887)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Ctenogobius smaragdus (Valenciennes, 1837)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Ctenogobius stigmaticus (Poey, 1860)	Marine-Estuary	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Evorthodus lyricus (Girard, 1858)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Gnatholepis thompsoni Jordan, 1904	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Gobioides broussonnetii Lacepède, 1800	Marine-Estuary-Freshwater	LC	DD		CIUFPB, Li
Gobiiformes	Oxudercidae	Gobionellus oceanicus (Pallas, 1770)	Marine-Estuary	LC	LC		CIUFPB, Li
Gobiiformes	Oxudercidae	Gobionellus stomatus Starks, 1913	Marine-Estuary	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Barbulifer ceuthoecus (Jordan & Gilbert, 1884)	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Barbulifer cf. enigmaticus Joyeux, Van Tassell & Macieira, 2009	Marine	DD	LC		CIUFPB
Gobiiformes	Gobiidae	Bathygobius geminatus Tornabene, Baldwin & Pezold, 2010	Marine	DD	LC		Li
Gobiiformes	Gobiidae	Bathygobius soporator (Valenciennes, 1837)	Marine-Estuary-Freshwater	LC	LC	Lc	CIUFPB, MNRJ, MZUSP, Li
Gobiiformes	Gobiidae	Coryphopterus dicrus Böhlke & Robins, 1960	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Coryphopterus aff. eidolon Böhlke & Robins, 1960	Marine	VU	Ne		CIUFPB
Gobiiformes	Gobiidae	Coryphopterus glaucofraenum Gill, 1863	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Coryphopterus thrix Böhlke & Robins, 1960	Marine	VU	LC		CIUFPB
Gobiiformes	Gobiidae	Elacatinus figaro Sazima, Moura & Rosa, 1997	Marine	VU	EN		CIUFPB, Li
Gobiiformes	Gobiidae	Gobiosoma alfiei Joyeux & Macieira, 2015	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Gobiosoma spilotum? (Ginsburg, 1939)	Marine	EN	Ne		CIUFPB
Gobiiformes	Gobiidae	Gobulus myersi Ginsburg, 1939	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Lythrypnus brasiliensis? Greenfield, 1988	Marine	Ne	LC		CIUFPB
Gobiiformes	Gobiidae	Microdesmus bahianus Dawson, 1973	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Microdesmus longipinnis (Weymouth, 1910)	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Microgobius carri Fowler, 1945	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Microgobius meeki Evermann & Marsh, 1899	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Priolepis dawsoni Greenfield, 1989	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Psilotris cf. alepis Ginsburg, 1953	Marine	LC	Ne		CIUFPB
Gobiiformes	Gobiidae	Ptereleotris randalli Gasparini, Rocha & Floeter, 2001	Marine	LC	LC		CIUFPB, Li
Gobiiformes	Gobiidae	Risor ruber (Rosén, 1911)	Marine	LC	LC		CIUFPB, Li
Syngnathiformes	Dactylopteridae	Dactylopterus volitans (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Syngnathiformes	Mullidae	Mulloidichthys martinicus (Cuvier, 1829)	Marine	LC	LC	Co	Li
Syngnathiformes	Mullidae	Pseudupeneus maculatus (Bloch, 1793)	Marine	LC	LC	Co	CIUFPB, Li
Syngnathiformes	Mullidae	Upeneus parvus (Cuvier, 1829)	Marine	LC	LC	Co	Li
Syngnathiformes	Callionymidae	Callionymus bairdi Jordan, 1888	Marine	LC	LC		CIUFPB, Li
Syngnathiformes	Aulostomidae	Aulostomus strigosus Wheeler, 1955	Marine	LC	LC	Co	Li
Syngnathiformes	Fistulariidae	Fistularia petimba Lacépède, 1803	Marine	LC	LC	Co	CIUFPB, Li
Syngnathiformes	Fistulariidae	Fistularia tabacaria Linnaeus, 1758	Marine	LC	LC	Co	CIUFPB, Li
Syngnathiformes	Syngnathidae	Bryx dunckeri (Metzelaar, 1919)	Marine	LC	LC		CIUFPB, Li
Syngnathiformes	Syngnathidae	Cosmocampus elucens (Poey, 1868)	Marine	LC	LC		CIUFPB, Li
Syngnathiformes	Syngnathidae	Halicampus crinitus (Jenyns, 1842)	Marine	Ne	LC		CIUFPB, Li
Syngnathiformes	Syngnathidae	Hippocampus aff. erectus Perry, 1810	Marine	VU	VU	Co(Or)	OBIS_BR
Syngnathiformes	Syngnathidae	Hippocampus reidi Ginsburg, 1933	Marine	NT	VU	Co(Or), Me	CIUFPB, Li
Syngnathiformes	Syngnathidae	Microphis lineatus (Kaup, 1856)	Marine-Estuary-Freshwater	Ne	Ne		CIUFPB, Li
Syngnathiformes	Syngnathidae	Syngnathus folletti Herald, 1942	Marine-Estuary	LC	LC		Li
Syngnathiformes	Syngnathidae	Syngnathus pelagicus Linnaeus, 1758	Marine	LC	LC		CIUFPB, Li
Scombriformes	Stromateidae	Peprilus crenulatus Cuvier, 1829	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Scombriformes	Pomatomidae	Pomatomus saltatrix (Linnaeus, 1766)	Marine-Estuary	VU	LC		Li
Scombriformes	Nomeidae	Nomeus gronovii (Gmelin, 1789)	Marine	LC	LC		Li
Scombriformes	Scombridae	Acanthocybium solandri (Cuvier, 1832)	Marine	LC	LC	Co	Li
Scombriformes	Scombridae	Auxis thazard (Lacepède, 1800)	Marine	LC	LC	Co	Li
Scombriformes	Scombridae	Euthynnus alletteratus (Rafinesque, 1810)	Marine	LC	LC	Co	Li
Scombriformes	Scombridae	Scomberomorus brasiliensis Collette, Russo & Zavala-Camin, 1978	Marine	LC	NT	Co	CIUFPB, Li
Scombriformes	Scombridae	Scomberomorus cavalla (Cuvier, 1829)	Marine	LC	NT	Co	CIUFPB, Li
Scombriformes	Scombridae	Scomberomorus regalis (Bloch, 1793)	Marine	LC	LC	Co	CIUFPB, Li
Scombriformes	Scombridae	Thunnus albacares (Bonnaterre, 1788)	Marine	LC	LC	Co	Li
Scombriformes	Scombridae	Thunnus atlanticus (Lesson, 1831)	Marine	LC	LC	Co	Li
Scombriformes	Scombridae	Thunnus obesus (Lowe, 1839)	Marine	VU	LC	Co	CIUFPB, Li
Scombriformes	Trichiuridae	Trichiurus lepturus Linnaeus, 1758	Marine	LC	LC	Co	CIUFPB, Li
Synbranchiformes	Synbranchidae	Synbranchus aff. marmoratus Bloch, 1795	Freshwater	LC	LC		CIUFPB, UFRN, Li
Carangiformes	Centropomidae	Centropomus ensiferus Poey, 1860	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Centropomidae	Centropomus parallelus Poey, 1860	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Centropomidae	Centropomus pectinatus Poey, 1860	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Centropomidae	Centropomus undecimalis (Bloch, 1792)	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Sphyraenidae	Sphyraena barracuda (Edwards, 1771)	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Sphyraenidae	Sphyraena borealis DeKay, 1842	Marine	LC	Ne	Co	CIUFPB, Li
Carangiformes	Sphyraenidae	Sphyraena guachancho Cuvier, 1829	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Carangiformes	Polynemidae	Polydactylus oligodon (Günther, 1860)	Marine	LC	LC	Co	CIUFPB
Carangiformes	Polynemidae	Polydactylus virginicus (Linnaeus, 1758)	Marine-Estuary	LC	LC	Lc	CIUFPB, Li
Carangiformes	Cyclopsettidae	Citharichthys arenaceus Evermann & Marsh, 1900	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Cyclopsettidae	Citharichthys cornutus (Günther, 1880)	Marine	LC	LC		Li
Carangiformes	Cyclopsettidae	Citharichthys macrops Dresel, 1885	Marine	LC	LC		CIUFPB, Li
Carangiformes	Cyclopsettidae	Citharichthys spilopterus Günther, 1862	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Carangiformes	Cyclopsettidae	Cyclopsetta fimbriata (Goode & Bean, 1885)	Marine	LC	LC		Li
Carangiformes	Cyclopsettidae	Etropus crossotus Jordan & Gilbert, 1882	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Cyclopsettidae	Syacium micrurum Ranzani, 1842	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Cyclopsettidae	Syacium papillosum (Linnaeus, 1758)	Marine	LC	LC		Li
Carangiformes	Bothidae	Bothus lunatus (Linnaeus, 1758)	Marine	LC	LC		Li
Carangiformes	Bothidae	Bothus ocellatus (Agassiz, 1831)	Marine	LC	LC		CIUFPB, Li
Carangiformes	Bothidae	Bothus robinsi Topp & Hoff, 1972	Marine	LC	LC		Li
Carangiformes	Paralichthyidae	Paralichthys brasiliensis (Ranzani, 1842)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Carangiformes	Achiridae	Achirus achirus (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Achiridae	Achirus declivis Chabanaud, 1940	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Carangiformes	Achiridae	Achirus lineatus (Linnaeus, 1758)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Carangiformes	Achiridae	Gymnachirus nudus Kaup, 1858	Marine-Estuary	LC	LC		Li
Carangiformes	Achiridae	Trinectes inscriptus (Gosse, 1851)	Marine-Estuary	LC	Ne		CIUFPB
Carangiformes	Achiridae	Trinectes microphthalmus (Chabanaud, 1928)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Carangiformes	Achiridae	Trinectes paulistanus (Miranda Ribeiro, 1915)	Marine-Estuary-Freshwater	LC	LC		CIUFPB, Li
Carangiformes	Cynoglossidae	Symphurus plagusia (Bloch & Schneider, 1801)	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Cynoglossidae	Symphurus rhytisma? Böhlke, 1961	Marine	LC	LC		CIUFPB
Carangiformes	Cynoglossidae	Symphurus tessellatus (Quoy & Gaimard, 1824)	Marine-Estuary	LC	LC		CIUFPB, Li
Carangiformes	Carangidae	Alectis ciliaris (Bloch, 1787)	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Caranx bartholomaei (Cuvier, 1833)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Caranx crysos (Mitchill, 1815)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Caranx hippos (Linnaeus, 1766)	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Caranx latus Agassiz, 1831	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Caranx lugubris Poey, 1860	Marine	LC	LC	Co	MNRJ, Li
Carangiformes	Carangidae	Caranx ruber (Bloch, 1793)	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Chloroscombrus chrysurus (Linnaeus, 1766)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Decapterus macarellus (Cuvier, 1833)	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Decapterus tabl Berry, 1968	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Elagatis bipinnulata (Quoy & Gaimard, 1825)	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Hemicaranx amblyrhynchus (Cuvier, 1833)	Marine	LC	LC		CIUFPB
Carangiformes	Carangidae	Naucrates ductor (Linnaeus, 1758)	Marine	LC	LC		Li
Carangiformes	Carangidae	Oligoplites palometa (Cuvier, 1832)	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Oligoplites saliens (Bloch, 1793)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Oligoplites saurus (Bloch & Schneider, 1801)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Selar crumenophthalmus (Bloch, 1793)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Selene brownii (Cuvier, 1816)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Selene setapinnis (Mitchill, 1815)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Selene vomer (Linnaeus, 1758)	Marine-Estuary	LC	LC	Co, Lc	CIUFPB, Li
Carangiformes	Carangidae	Seriola dumerili (Risso, 1810)	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Seriola rivoliana Valenciennes, 1833	Marine	LC	LC	Co	Li
Carangiformes	Carangidae	Trachinotus carolinus (Linnaeus, 1766)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Trachinotus cayennensis Cuvier, 1832	Marine	LC	LC		Li
Carangiformes	Carangidae	Trachinotus falcatus (Linnaeus, 1758)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Trachinotus goodei Jordan & Evermann, 1896	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Carangidae	Trachurus lathami? Nichols, 1920	Marine	LC	LC		CIUFPB
Carangiformes	Carangidae	Uraspis secunda (Poey, 1860)	Marine	LC	LC		Li
Carangiformes	Echeneidae	Echeneis naucrates Linnaeus, 1758	Marine	LC	LC		CIUFPB, Li
Carangiformes	Echeneidae	Remora remora (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Carangiformes	Rachycentridae	Rachycentron canadum (Linnaeus, 1766)	Marine	LC	LC	Co	CIUFPB, Li
Carangiformes	Coryphaenidae	Coryphaena hippurus Linnaeus, 1758	Marine	LC	LC	Co	Li
Atheriniformes	Atherinopsidae	Atherinella blackburni (Schultz, 1949)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Atheriniformes	Atherinopsidae	Atherinella brasiliensis (Quoy & Gaimard, 1825)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Atheriniformes	Atherinopsidae	Membras dissimilis? (Carvalho, 1956)	Marine	Ne	NT		CIUFPB
Beloniformes	Belonidae	Ablennes hians (Valenciennes, 1846)	Marine-Estuary-Freshwater	LC	LC		Li
Beloniformes	Belonidae	Strongylura marina (Walbaum, 1792)	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, Li
Beloniformes	Belonidae	Strongylura timucu (Walbaum, 1792)	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, Li
Beloniformes	Belonidae	Tylosurus crocodilus (Péron & Lesueur, 1821)	Marine	LC	LC	Co	Li
Beloniformes	Hemiramphidae	Hemiramphus brasiliensis (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, Li
Beloniformes	Hemiramphidae	Hemiramphus balao Lesueur, 1821	Marine	LC	DD	Co	Li
Beloniformes	Hemiramphidae	Hyporhamphus roberti (Valenciennes, 1847)	Marine-Estuary	LC	LC		CIUFPB, Li
Beloniformes	Hemiramphidae	Hyporhamphus unifasciatus (Ranzani, 1841)	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Beloniformes	Exocoetidae	Cheilopogon cyanopterus (Valenciennes, 1847)	Marine	LC	LC		Li
Beloniformes	Exocoetidae	Cheilopogon melanurus (Valenciennes, 1847)	Marine	LC	LC		Li
Beloniformes	Exocoetidae	Exocoetus volitans Linnaeus, 1758	Marine	LC	LC		Li
Beloniformes	Exocoetidae	Hirundichthys affinis (Günther, 1866)	Marine	LC	LC		Li
Cyprinodontiformes	Rivulidae	Hypsolebias sp.	Freshwater	Na	Na		To be cataloged
Cyprinodontiformes	Poeciliidae	Poecilia reticulata Peters, 1859*	Freshwater	LC	Ne		CIUFPB, PUCRS, UFRN, Li
Cyprinodontiformes	Poeciliidae	Poecilia vivipara Bloch & Schneider, 1801	Freshwater-Estuary	LC	LC	Co, Lc	CIUFPB, MNRJ, PUCRS, UFRN, Li
Cyprinodontiformes	Poeciliidae	Xiphophorus maculatus (Günther, 1866)*	Freshwater	DD	Ne		CIUFPB, Li
Cichliformes	Cichlidae	Astronotus ocellatus (Agassiz, 1831)*	Freshwater	LC	LC	Co, Lc	CIUFPB, Li
Cichliformes	Cichlidae	Cichla ocellaris Bloch & Schneider, 1801*	Freshwater	LC	LC	Co, Lc	CIUFPB, Li
Cichliformes	Cichlidae	Cichla monoculus Spix & Agassiz, 1831*	Freshwater	LC	LC	Co, Lc	CIUFPB, UFRN, Li
Cichliformes	Cichlidae	Cichlasoma orientale Kullander, 1983	Freshwater	LC	LC		CIUFPB, PUCRS, UFRN, Li
Cichliformes	Cichlidae	Cichlasoma sanctifranciscense Kullander, 1983	Freshwater	LC	LC		CIUFPB
Cichliformes	Cichlidae	Coptodon rendalli (Boulenger, 1897)*	Freshwater	LC	Ne	Co(Aq), Lc	CIUFPB, PUCRS, Li
Cichliformes	Cichlidae	Geophagus brasiliensis (Quoy & Gaimard, 1824)	Freshwater-Estuary	LC	LC		CIUFPB, PUCRS, UFRN, Li
Cichliformes	Cichlidae	Oreochromis niloticus (Linnaeus, 1758)*	Freshwater	LC	Ne	Co(Aq), Me, Lc	CIUFPB, UFRN, Li
Cichliformes	Cichlidae	Parachromis managuensis (Günther, 1867)*	Freshwater	LC	Ne		CIUFPB, UFRN, Li
Cichliformes	Cichlidae	Saxatilia brasiliensis (Bloch, 1792)	Freshwater	LC	LC		CIUFPB, UFRN, PUCRS, Li
Mugiliformes	Mugilidae	Mugil brevirostris Miranda Ribeiro, 1915	Marine-Estuary	Ne	DD	Co	Li
Mugiliformes	Mugilidae	Mugil curema Valenciennes, 1836	Marine-Estuary	LC	DD	Co, Lc	CIUFPB, Li
Mugiliformes	Mugilidae	Mugil curvidens Valenciennes, 1836	Marine-Estuary	Ne	DD	Co	CIUFPB, Li
Mugiliformes	Mugilidae	Mugil incilis Hancock, 1830	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Mugiliformes	Mugilidae	Mugil liza Valenciennes, 1836	Marine-Estuary	DD	NT	Co, Lc	CIUFPB, Li
Mugiliformes	Mugilidae	Mugil rubrioculus Harrison, Nirchio, Oliveira, Ron & Gaviria, 2007	Marine-Estuary	LC	DD	Co	Li
Blenniiformes	Pomacentridae	Abudefduf saxatilis (Linnaeus, 1758)	Marine	LC	LC	LC	CIUFPB, Li
Blenniiformes	Pomacentridae	Azurina multilineata (Guichenot, 1853)	Marine	LC	LC	Co(Or)	CIUFPB, Li
Blenniiformes	Pomacentridae	Chromis flavicauda (Günther, 1880)	Marine	DD	LC	Co(Or)	CIUFPB, Li
Blenniiformes	Pomacentridae	Chromis jubauna Moura, 1995	Marine	LC	LC	Co(Or)	CIUFPB
Blenniiformes	Pomacentridae	Microspathodon chrysurus (Cuvier, 1830)	Marine	LC	VU	Co(Or)	Li
Blenniiformes	Pomacentridae	Stegastes fuscus (Cuvier, 1830)	Marine	LC	LC	Co(Or)	CIUFPB, MZUSP, Li
Blenniiformes	Pomacentridae	Stegastes pictus (Castelnau, 1855)	Marine	LC	LC	Co(Or)	CIUFPB, MZUSP, Li
Blenniiformes	Pomacentridae	Stegastes variabilis (Castelnau, 1855)	Marine	LC	LC	Co(Or)	CIUFPB, MZUSP, Li
Blenniiformes	Grammatidae	Gramma brasiliensis Sazima, Gasparini & Moura, 1998	Marine	LC	DD	Co(Or)	CIUFPB, Li
Blenniiformes	Opistognathidae	Opistognathus aurifrons (Jordan & Thompson, 1905)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Opistognathidae	Opistognathus cuvierii? Valenciennes, 1836	Marine	LC	LC		CIUFPB
Blenniiformes	Gobiesocidae	Gobiesox barbatulus Starks, 1913	Marine	LC	LC		Li
Blenniiformes	Gobiesocidae	Gobiesox strumosus? Cope, 1870	Marine	LC	Ne		CIUFPB
Blenniiformes	Gobiesocidae	Tomicodon fasciatus (Peters, 1859)	Marine	LC	Ne		CIUFPB, Li
Blenniiformes	Tripterygiidae	Enneanectes altivelis Rosenblatt, 1960	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Blenniidae	Entomacrodus vomerinus (Valenciennes, 1836)	Marine	LC	LC		CIUFPB, MZUSP, Li
Blenniiformes	Blenniidae	Hypleurochilus fissicornis (Quoy & Gaimard, 1824)	Marine	LC	LC		Li
Blenniiformes	Blenniidae	Hypleurochilus pseudoaequipinnis Bath, 1994	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Blenniidae	Ophioblennius atlanticus? (Valenciennes, 1836)	Marine	LC	Ne		CIUFPB
Blenniiformes	Blenniidae	Ophioblennius trinitatis Miranda Ribeiro, 1919	Marine	LC	LC		Li
Blenniiformes	Blenniidae	Parablennius marmoreus (Poey, 1876)	Marine	LC	LC	LC	Li
Blenniiformes	Blenniidae	Parablennius pilicornis (Cuvier, 1829)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Blenniidae	Scartella cristata (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, MZUSP, Li
Blenniiformes	Labrisomidae	Gobioclinus kalisherae (Jordan, 1904)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Labrisomidae	Labrisomus cricota Sazima, Gasparini & Moura, 2002	Marine	LC	LC		Li
Blenniiformes	Labrisomidae	Labrisomus nuchipinnis (Quoy & Gaimard, 1824)	Marine	LC	LC		CIUFPB, MNRJ, MZUSP, Li
Blenniiformes	Labrisomidae	Malacoctenus delalandii (Valenciennes, 1836)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Labrisomidae	Malacoctenus triangulatus? Springer, 1959	Marine	LC	LC		CIUFPB
Blenniiformes	Labrisomidae	Malacoctenus zaluari Carvalho-Filho, Gasparini & Sazima, 2020	Marine	LC	Ne		Li
Blenniiformes	Labrisomidae	Paraclinus arcanus Guimarães & Bacellar, 2002	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Labrisomidae	Paraclinus rubicundus (Starks, 1913)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Labrisomidae	Starksia brasiliensis (Gilbert, 1900)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Chaenopsidae	Emblemaria australis Ramos, Rocha & Rocha, 2003c	Marine	LC	DD		CIUFPB, Li
Blenniiformes	Chaenopsidae	Emblemariopsis signifer (Ginsburg, 1942)	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Dactyloscopidae	Dactyloscopus crossotus Kanazawa, 1952	Marine	LC	LC		CIUFPB, Li
Blenniiformes	Dactyloscopidae	Dactyloscopus tridigitatus Böhlke, 1968	Marine	LC	LC		CIUFPB, Li
Perciformes	Serranidae	Diplectrum formosum (Linnaeus, 1766)	Marine	LC	LC	Co	CIUFPB, Li
Perciformes	Serranidae	Diplectrum radiale (Quoy & Gaimard, 1824)	Marine	LC	LC	Co	Li
Perciformes	Serranidae	Serranus annularis (Günther, 1880)	Marine	LC	LC		CIUFPB, Li
Perciformes	Serranidae	Serranus baldwini (Evermann & Marsh, 1899)	Marine	LC	LC		CIUFPB, Li
Perciformes	Serranidae	Serranus flaviventris (Cuvier, 1829)	Marine	LC	LC		CIUFPB, Li
Perciformes	Serranidae	Serranus phoebe Poey, 1851	Marine	LC	LC		CIUFPB, Li
Perciformes	Epinephelidae	Alphestes afer (Bloch, 1793)	Marine	LC	DD	Co	CIUFPB, Li
Perciformes	Epinephelidae	Cephalopholis fulva (Linnaeus, 1758)	Marine	LC	NT	Co	CIUFPB, MZUSP, Li
Perciformes	Epinephelidae	Dermatolepis inermis (Valenciennes, 1833)	Marine	DD	DD	Co	Li
Perciformes	Epinephelidae	Epinephelus adscensionis (Osbeck, 1765)	Marine	LC	DD	Co	CIUFPB, Li
Perciformes	Epinephelidae	Epinephelus guttatus (Linnaeus, 1758)	Marine	LC	Ne		Li
Perciformes	Epinephelidae	Epinephelus itajara (Lichtenstein, 1822)	marine	VU	CR	Co	CIUFPB, Li
Perciformes	Epinephelidae	Epinephelus morio (Valenciennes, 1828)	Marine	VU	EN	Co	Li
Perciformes	Epinephelidae	Hyporthodus niveatus (Valenciennes, 1828)	Marine	VU	VU	Co	Li
Perciformes	Epinephelidae	Mycteroperca bonaci (Poey, 1860)	Marine	NT	EN	Co	CIUFPB, Li
Perciformes	Epinephelidae	Mycteroperca interstitialis (Poey, 1860)	Marine	VU	VU	Co	Li
Perciformes	Epinephelidae	Paranthias furcifer (Valenciennes, 1828)	Marine	LC	Ne	Co	CIUFPB, Li
Perciformes	Grammistidae	Rypticus bistrispinus (Mitchill, 1818)	Marine	LC	LC		CIUFPB, Li
Perciformes	Grammistidae	Rypticus randalli Courtenay, 1967	Marine	LC	LC		CIUFPB, Li
Perciformes	Grammistidae	Rypticus saponaceus (Bloch & Schneider, 1801)	Marine	LC	LC		CIUFPB, Li
Perciformes	Grammistidae	Rypticus subbifrenatus Gill, 1861	Marine	LC	LC		CIUFPB
Perciformes	Triglidae	Prionotus cf. nudigula Ginsburg, 1950	Marine	LC	LC		CIUFPB, Li
Perciformes	Triglidae	Prionotus punctatus (Bloch, 1793)	Marine	LC	LC		CIUFPB, Li
Perciformes	Liopropomatidae	Liopropoma carmabi (Randall, 1963)	Marine	LC	LC		Li
Perciformes	Scorpaenidae	Pterois volitans (Linnaeus, 1758)*	Marine	LC	Ne		CIUFPB, Li
Perciformes	Scorpaenidae	Scorpaena bergii Evermann & Marsh, 1900	Marine	LC	LC		CIUFPB, Li
Perciformes	Scorpaenidae	Scorpaena brasiliensis Cuvier, 1829	Marine	LC	LC		CIUFPB, Li
Perciformes	Scorpaenidae	Scorpaena inermis Cuvier, 1829	Marine	LC	LC		Li
Perciformes	Scorpaenidae	Scorpaena isthmensis Meek & Hildebrand, 1928	Marine	LC	LC		CIUFPB, Li
Perciformes	Scorpaenidae	Scorpaena melasma Eschmeyer, 1965	Marine	DD	LC		CIUFPB
Perciformes	Scorpaenidae	Scorpaena plumieri Bloch, 1789	Marine-Estuary	LC	LC		CIUFPB, Li
Perciformes	Scorpaenidae	Scorpaenodes caribbaeus Meek & Hildebrand, 1928	Marine	LC	LC		Li
Perciformes	Scorpaenidae	Scorpaenodes tredecimspinosus (Metzelaar, 1919)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Bodianus brasiliensis (Heiser, Moura & Robertson, 2000)	Marine	LC	Ne		Li
Labriformes	Labridae	Bodianus pulchellus (Poey, 1860)	Marine	LC	LC	Co(Or)	Li
Labriformes	Labridae	Bodianus rufus (Linnaeus, 1758)	Marine	LC	LC	Co(Or)	CIUFPB, Li
Labriformes	Labridae	Clepticus brasiliensis Heiser, Moura & Robertson, 2000	Marine	LC	LC		Li
Labriformes	Labridae	Cryptotomus roseus Cope, 1871	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Doratonotus megalepis Günther, 1862	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Halichoeres bivittatus (Bloch, 1791)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Halichoeres brasiliensis (Bloch, 1791)	Marine	DD	DD		CIUFPB, Li
Labriformes	Labridae	Halichoeres dimidiatus (Agassiz, 1831)	Marine	LC	LC	Co(Or)	CIUFPB, Li
Labriformes	Labridae	Halichoeres penrosei Starks, 1913	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Halichoeres poeyi (Steindachner, 1867)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Halichoeres radiatus (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Nicholsina usta (Valenciennes, 1840)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Scarus trispinosus Valenciennes, 1840	Marine	EN	CR		CIUFPB, Li
Labriformes	Labridae	Scarus zelindae Moura, Figueiredo & Sazima, 2001	Marine	DD	EN		CIUFPB, Li
Labriformes	Labridae	Sparisoma amplum (Ranzani, 1841)	Marine	LC	VU		Li
Labriformes	Labridae	Sparisoma axillare (Steindachner, 1878)	Marine	DD	VU		CIUFPB, Li
Labriformes	Labridae	Sparisoma frondosum (Agassiz, 1831)	Marine	DD	VU		Li
Labriformes	Labridae	Sparisoma radians (Valenciennes, 1840)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Thalassoma noronhanum (Boulenger, 1890)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Xyrichtys martinicensis Valenciennes, 1840	Marine	LC	Ne		Li
Labriformes	Labridae	Xyrichtys novacula (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Labriformes	Labridae	Xyrichtys splendens Castelnau, 1855	Marine	LC	LC		CIUFPB, Li
Labriformes	Uranoscopidae	Astroscopus y-graecum (Cuvier, 1829)	Marine	LC	Ne		HVASF
Centrarchiformes	Kyphosidae	Kyphosus sectatrix (Linnaeus, 1758)	Marine	LC	LC	Co	CIUFPB, Li
Centrarchiformes	Cirrhitidae	Amblycirrhitus pinos (Mowbray, 1927)	Marine	LC	DD		Li
Acropomatiformes	Pempheridae	Pempheris schomburgki Müller & Troschel, 1848	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Gerreidae	Diapterus auratus Ranzani, 1842	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Diapterus rhombeus (Cuvier, 1829)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus argenteus Baird & Girard, 1855	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus gula (Quoy & Gaimard, 1824)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus havana (Nichols, 1912)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus lefroyi (Goode, 1874)	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus melanopterus (Bleeker, 1863)	Marine-Estuary-Freshwater	LC	LC	Co	CIUFPB, MNRJ, Li
Acanthuriformes	Gerreidae	Eugerres brasilianus (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Genyatremus luteus (Bloch, 1790)	Marine-Estuary	DD	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Gerres cinereus (Walbaum, 1792)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Gerreidae	Eucinostomus jonesii (Günther, 1879)	Marine	LC	LC		Li
Acanthuriformes	Ephippidae	Chaetodipterus faber (Broussonet, 1782)	Marine-Estuary	LC	LC	Co, Lc	CIUFPB, Li
Acanthuriformes	Sciaenidae	Bairdiella goeldi Marceniuk, Molina, Caires, Rotundo, Wosiacki & Oliveira, 2019	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Ctenosciaena gracilicirrhus (Metzelaar, 1919)	Marine	LC	LC		CIUFPB
Acanthuriformes	Sciaenidae	Cynoscion acoupa (Lacepède, 1801)	Marine-Estuary	VU	DD	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Cynoscion jamaicensis (Vaillant & Bocourt, 1883)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Cynoscion leiarchus (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Cynoscion microlepidotus (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Cynoscion virescens (Cuvier, 1830)	Marine-Estuary	LC	DD	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Eques lanceolatus (Linnaeus, 1758)	Marine	Ne	LC	Co	Li
Acanthuriformes	Sciaenidae	Isopisthus parvipinnis (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Larimus breviceps Cuvier, 1830	Marine	LC	LC	Co	CIUFPB, MNRJ, Li
Acanthuriformes	Sciaenidae	Macrodon ancylodon (Bloch & Schneider, 1801)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Menticirrhus cuiaranensis Marceniuk, Caires, Rotundo, Cerqueira, Siccha-Ramirez, Wosiacki & Oliveira, 2020	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Menticirrhus martinicensis (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB
Acanthuriformes	Sciaenidae	Micropogonias furnieri (Desmarest, 1823)	Marine-Estuary	LC	VU	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Nebris microps Cuvier, 1830	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Odontoscion aff. dentex (Cuvier, 1830)	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Paralonchurus brasiliensis (Steindachner, 1875)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Pareques lineatus (Cuvier, 1830)	Marine	LC	Ne	Co	CIUFPB, Li
Acanthuriformes	Sciaenidae	Plagioscion squamosissimus (Heckel, 1840)*	Freshwater	LC	LC	Co, Lc	CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer brasiliensis (Schultz, 1945)	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer collettei Chao, Carvalho-Filho & Andrade Santos, 2021	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer gomezi (Cervigón, 2011)	Marine	Ne	LC		CIUFPB
Acanthuriformes	Sciaenidae	Stellifer menezesi Chao, Carvalho-Filho & Andrade Santos, 2021	Marine-Estuary	LC	LC		CIUFPB
Acanthuriformes	Sciaenidae	Stellifer musicki Chao, Carvalho-Filho & Andrade Santos, 2021	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer microps (Steindachner, 1864)	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer naso (Jordan, 1889)	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer punctatissimus (Meek & Hildebrand, 1925)	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer rastrifer (Jordan, 1889)	Marine-Estuary	LC	LC		CIUFPB, Li
Acanthuriformes	Sciaenidae	Stellifer stellifer (Bloch, 1790)	Marine-Estuary	DD	LC		CIUFPB, MNRJ, Li
Acanthuriformes	Sciaenidae	Umbrina coroides Cuvier, 1830	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Anisotremus surinamensis (Bloch, 1791)	Marine	DD	DD	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Anisotremus virginicus (Linnaeus, 1758)	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Conodon nobilis (Linnaeus, 1758)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Haemulon atlanticus Carvalho, Marceniuk, Oliveira & Wosiacki, 2021	Marine	LC	LC	Co	CIUFPB, MNRJ, Li
Acanthuriformes	Haemulidae	Haemulon aurolineatum Cuvier, 1830	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Haemulon bonariense Cuvier, 1830	Marine	LC	Ne		Li
Acanthuriformes	Haemulidae	Haemulon melanurum (Linnaeus, 1758)	Marine	LC	LC	Co	Li
Acanthuriformes	Haemulidae	Haemulon parra (Desmarest, 1823)	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Haemulon plumierii (Lacepède, 1801)	Marine	LC	DD	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Haemulon squamipinna Rocha & Rosa, 1999	Marine	LC	LC	Co	CIUFPB, MZUSP, Li
Acanthuriformes	Haemulidae	Haemulon striatum (Linnaeus, 1758)	Marine	LC	LC		Li
Acanthuriformes	Haemulidae	Haemulopsis corvinaeformis (Steindachner, 1868)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Orthopristis rubra? (Cuvier, 1830)	Marine-Estuary	LC	LC	Co	CIUFPB, MNRJ
Acanthuriformes	Haemulidae	Orthopristis scapularis Fowler, 1915	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Haemulidae	Paranisotremus moricandi (Ranzani, 1842)	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Haemulidae	Pomadasys ramosus (Poey, 1860)	Marine-Estuary	Ne	LC		CIUFPB
Acanthuriformes	Haemulidae	Rhonciscus approximans (Bean & Dresel, 1884)	Marine-Estuary-Freshwater	Ne	Ne		Li
Acanthuriformes	Haemulidae	Rhonciscus crocro (Cuvier, 1830)	Marine-Estuary	DD	LC	Co	CIUFPB
Acanthuriformes	Lobotidae	Lobotes surinamensis (Bloch, 1790)	Marine-Estuary	LC	LC	Co(Rf)	CIUFPB, Li
Acanthuriformes	Lutjanidae	Lutjanus alexandrei Moura & Lindeman, 2007	Marine	Ne	DD	Co	CIUFPB, Li
Acanthuriformes	Lutjanidae	Lutjanus analis (Cuvier, 1828)	Marine-Estuary	NT	NT	Co	CIUFPB, MNRJ, Li
Acanthuriformes	Lutjanidae	Lutjanus buccanella (Cuvier, 1828)	Marine	DD	DD	Co	CIUFPB, Li
Acanthuriformes	Lutjanidae	Lutjanus cyanopterus (Cuvier, 1828)	Marine-Estuary	VU	VU	Co	CIUFPB, MNRJ, Li
Acanthuriformes	Lutjanidae	Lutjanus jocu (Bloch & Schneider, 1801)	Marine-Estuary	DD	NT	Co	OBIS_BR
Acanthuriformes	Lutjanidae	Lutjanus campechanus (Poey, 1860)	Marine	VU	EN	Co	Li
Acanthuriformes	Lutjanidae	Lutjanus synagris (Linnaeus, 1758)	Marine-Estuary	NT	DD	Co	CIUFPB, Li
Acanthuriformes	Lutjanidae	Lutjanus vivanus (Cuvier, 1828)	Marine	LC	DD	Co	CIUFPB, Li
Acanthuriformes	Lutjanidae	Ocyurus chrysurus (Bloch, 1791)	Marine	DD	DD	Co	CIUFPB, Li
Acanthuriformes	Lutjanidae	Rhomboplites aurorubens (Cuvier, 1829)	Marine	VU	DD	Co	CIUFPB, Li
Acanthuriformes	Malacanthidae	Malacanthus plumieri (Bloch, 1786)	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Pomacanthidae	Centropyge aurantonotus Burgess, 1974	Marine	LC	DD	Co(Or)	CIUFPB, Li
Acanthuriformes	Pomacanthidae	Holacanthus ciliaris (Linnaeus, 1758)	Marine	LC	DD	Co(Or)	CIUFPB, Li
Acanthuriformes	Pomacanthidae	Holacanthus tricolor (Bloch, 1795)	Marine	LC	DD	Co(Or)	CIUFPB, Li
Acanthuriformes	Pomacanthidae	Pomacanthus arcuatus (Linnaeus, 1758)	Marine	LC	DD	Co(Or)	Li
Acanthuriformes	Pomacanthidae	Pomacanthus paru (Bloch, 1787)	Marine	LC	DD	Co(Or)	CIUFPB, Li
Acanthuriformes	Chaetodontidae	Chaetodon ocellatus Bloch, 1787	Marine	LC	DD	Co(Or)	CIUFPB, Li
Acanthuriformes	Chaetodontidae	Chaetodon sedentarius Poey, 1860	Marine	LC	LC	Co(Or)	Li
Acanthuriformes	Chaetodontidae	Chaetodon striatus Linnaeus, 1758	Marine	LC	LC	Co(Or)	CIUFPB, Li
Acanthuriformes	Acanthuridae	Acanthurus bahianus Castelnau, 1855	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Acanthuridae	Acanthurus chirurgus (Bloch, 1787)	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Acanthuridae	Acanthurus coeruleus Bloch & Schneider, 1801	Marine	LC	LC	Co(Or)	CIUFPB, Li
Acanthuriformes	Sparidae	Archosargus probatocephalus (Walbaum, 1792)	Marine-Estuary	LC	LC	Co	OBIS_BR, Li
Acanthuriformes	Sparidae	Archosargus rhomboidalis (Linnaeus, 1758)	Marine-Estuary	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Sparidae	Calamus calamus (Valenciennes, 1830)	Marine	LC	DD	Co	Li
Acanthuriformes	Sparidae	Calamus penna (Valenciennes, 1830)	Marine	LC	LC		CIUFPB, Li
Acanthuriformes	Sparidae	Calamus pennatula Guichenot, 1868	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Priacanthidae	Heteropriacanthus cruentatus (Lacepède, 1801)	Marine	LC	LC	Co	Li
Acanthuriformes	Priacanthidae	Priacanthus arenatus Cuvier, 1829	Marine	LC	LC	Co	CIUFPB, Li
Acanthuriformes	Antigoniidae	Antigonia capros? Lowe, 1843	Marine	LC	LC		CIUFPB
Lophiiformes	Ogcocephalidae	Ogcocephalus parvus Longley & Hildebrand, 1940	Marine	LC	DD		Li
Lophiiformes	Ogcocephalidae	Ogcocephalus vespertilio (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, MZUSP, Li
Lophiiformes	Antennariidae	Antennarius multiocellatus (Valenciennes, 1837)	Marine	LC	LC		CIUFPB, Li
Lophiiformes	Antennariidae	Antennarius scaber (Cuvier, 1817)	Marine	LC	Ne		Li
Lophiiformes	Antennariidae	Antennarius striatus (Shaw, 1794)	Marine	LC	LC		CIUFPB, Li
Lophiiformes	Antennariidae	Histrio histrio (Linnaeus, 1758)	Marine	LC	LC		Li
Tetraodontiformes	Molidae	Mola mola (Linnaeus, 1758)	Marine	VU	LC		CIUFPB, Li
Tetraodontiformes	Diodontidae	Chilomycterus antennatus (Cuvier, 1816)	Marine	LC	Ne		Li
Tetraodontiformes	Diodontidae	Chilomycterus antillarum (Jordan & Rutter, 1897)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Diodontidae	Chilomycterus spinosus (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, Li
Tetraodontiformes	Diodontidae	Diodon holocanthus (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, Li
Tetraodontiformes	Diodontidae	Diodon hystrix (Linnaeus, 1758)	Marine-Estuary	LC	LC		CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Canthigaster figueiredoi (Moura & Castro, 2002)	Marine	LC	LC		Li
Tetraodontiformes	Tetraodontidae	Sphoeroides psittacus (Bloch & Schneider, 1801)	Marine-Estuary	LC	LC	Co, Lc	Li
Tetraodontiformes	Tetraodontidae	Lagocephalus laevigatus (Linnaeus, 1766)	Marine-Estuary	LC	LC	Co, Lc	CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Sphoeroides camila Carvalho-Filho, Rotundo, Pitassy & Sazima, 2023	Marine	Ne	Ne		Li
Tetraodontiformes	Tetraodontidae	Sphoeroides greeleyi Gilbert, 1900	Marine-Estuary	LC	LC		CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Sphoeroides pachygaster (Muller & Troschel, 1848)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Sphoeroides spengleri (Bloch, 1785)	Marine-Estuary	LC	LC		CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Sphoeroides testudineus (Linnaeus, 1758)	Marine-Estuary	LC	LC	Lc	CIUFPB, Li
Tetraodontiformes	Tetraodontidae	Sphoeroides tyleri (Shipp, 1972)	Marine	LC	LC		Li
Tetraodontiformes	Ostraciidae	Acanthostracion polygonius (Poey, 1876)	Marine	LC	LC		Li
Tetraodontiformes	Ostraciidae	Acanthostracion quadricornis (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Ostraciidae	Lactophrys trigonus (Linnaeus, 1758)	Marine	LC	LC		Li
Tetraodontiformes	Monacanthidae	Aluterus heudelotii (Hollard, 1855)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Aluterus monoceros (Linnaeus, 1758)	Marine	LC	DD		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Aluterus schoepfi (Walbaum, 1792)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Aluterus scriptus (Osbeck, 1765)	Marine	LC	LC		Li
Tetraodontiformes	Monacanthidae	Cantherhines macrocerus (Hollard, 1853)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Cantherhines pullus (Ranzani, 1842)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Monacanthus ciliatus (Mitchill, 1818)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Stephanolepis hispida (Linnaeus, 1766)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Monacanthidae	Stephanolepis setifer (Bennett, 1831)	Marine	LC	LC		Li
Tetraodontiformes	Balistidae	Balistes capriscus (Gmelin, 1789)	Marine	VU	EN	Co, Me	CIUFPB, Li
Tetraodontiformes	Balistidae	Balistes vetula (Linnaeus, 1758)	Marine	NT	DD	Co	CIUFPB, MZUSP, Li
Tetraodontiformes	Balistidae	Canthidermis sufflamen (Mitchill, 1815)	Marine	LC	DD		Li
Tetraodontiformes	Balistidae	Melichthys niger (Bloch, 1786)	Marine	LC	LC		CIUFPB, Li
Tetraodontiformes	Balistidae	Xanthichthys ringens (Linnaeus, 1758)	Marine	LC	LC		CIUFPB, Li

FIGURE 4| Schematic overview of the ichthyofauna of Paraíba State and their associated habitats: Freshwater (1–2), estuarine (3) and marine (4). A. Parotocinclus jacksoni; B. Pimelodella enochi; C. Hippocampus reidi; D. Hypanus marianae; E. Lutjanus cyanopterus; F. Haemulon squamipinna; G. Ginglymostoma cirratum; H. Emblemaria australis; I. Scarus trispinosus; J. Coryphaena hippurus. The number indicate the biome and realms: 1. Caatinga, 2. Atlantic Forest, 3. Mangrove Forest, and 4. Marine region. Illustration by Lukas M. Silva.

We identified 467 marine and estuarine teleosts, distributed across 26 orders and 96 families. The most representative orders are Acanthuriformes (87 spp., 18.67%), Carangiformes (61, 13.09%), and Gobiiformes (42, 9.01%). The most speciose families are Sciaenidae (29, 6.12%), Carangidae (27, 5.70%), and Labridae (23, 4.85%) (Tab. 2; Fig. 5A). Only one species is non-native, the lionfish Pterois volitans, an invasive species from the family Scorpaenidae (Tab. 2; Fig. 6F).

FIGURE 5| Absolute frequency of A. Marine-estuarine teleost species; B. Freshwater teleost species; and C. Elasmobranch species in the Paraíba State, Brazil. Marine teleost fishes are further categorized by their most representative orders, families and species.

FIGURE 6| Fish species of Paraíba State. A. Apareiodon davisi; B. Parotocinclus jacksoni, endemic to the Mamanguape River; C. Parotocinclus spilosoma, endemic to the Paraíba River(Photos by Telton Ramos); D. Haemulon squamipinna, endemic to the Tropical Southwestern Atlantic province(Photo by Luis A. Rocha), and E. Hypanus marianae,endemic to Tropical Southwestern Atlanticprovince(Photo by Tiego Costa). F. The invasive marine species Pterois volitans (Photo by Vannesa Mieko).

For elasmobranch fishes, we identified 35 species across 7 orders and 17 families, all inhabiting marine environments, with some occasionally visiting estuaries. The most representative elasmobranchs fish orders are Carcharhiniformes (19 species, 54.30%) and Myliobatiformes (9 species, 25.71%), together accounting for 80% of all species. The families with the highest diversity are Carcharhinidae (11 species, 31.43%) and both Sphyrnidae and Dasyatidae (4 species each, 11.43%, see Tab. 2; Fig. 5C).

In freshwater environments, we identified a total of 63 teleost fish species, including exclusively freshwater species and some that may occasionally enter marine waters. These species belong to nine orders and 22 families (Tab. 2). Additionally, one species, the rivulid fish Hypsolebias sp., is currently being described (in prep.). The most speciose orders are Characiformes (30 spp., 47.69%), Siluriformes (15 spp., 23.08%), and Cichliformes (10 spp., 15.38%), collectively accounting for 80% of all freshwater species. The most speciose families are Cichlidae (10, 15.40%), Acestrorhamphidae (9, 13.84%), and Loricariidae (7, 12.31%) (Tab. 2; Fig. 5B). Of the 63 species, 13 are non-native, belonging to six families: Cichlidae (6 species), Serrasalmidae (1), Poeciliidae (2), and Acestrorhamphidae, Engraulidae, Cyprinidae, and Sciaenidae (1 each). Regarding species richness per basin, the Paraíba River is the most diverse, holding 47 species, followed by the Piranhas River (45 species), Mamanguape River (31), Abiaí River (29), Gramame River (29), Curimataú River (21), Camaratuba River (18), and Miriri River (12) (see Tab. 2; Fig. 3A).

Endemism and threat status. A total of nine freshwater fish species is endemic to the NCCD ecoregion, encompassing two orders and three families (Tab. 2). To date, no species have been reported exclusively as endemic to the Atlantic Forest biome in Paraíba. The family Loricariidae exhibits the highest richness of endemic species. Parotocinclus cearensis, P. seridoensis, and P. jacksoni; are endemic to the NCCD. Additionally, P. jacksoni is endemic to Mamanguape River basin and P. spilosoma is endemic to the Paraíba River basin. In marine environments, one marine species is endemic to the Northeastern Brazil Marine ecoregion (NBM): the teleost fish Emblemaria australis. The species Thalassophryne punctata; Haemulon squamipinna, and the stingray Hypanus marianae are endemic to the Tropical Southwestern Atlantic province and are restricted to the coastal zone (Tab. 2; Fig. 6).

In this study, we report that 44 species are threatened according to the IUCN, comprising 5 Critically Endangered (CR) species, 13 Endangered (EN) species, and 26 Vulnerable (VU) species, all of which inhabit marine and estuarine environments. According to the Sistema de Avaliação do Risco de Extinção da Biodiversidade (SALVE), Paraíba State is home to 43 threatened species, including 11 CR species, such as the Atlantic goliath grouper, Epinephelus itajara. Nine EN species occur in the state, including the red grouper, Epinephelus morio, and only two species from freshwater environments, the loricariids Parotocinclus seridoensis and P. spilosoma. Additionally, Paraíba has 23 VU species, including the large-eye marine stingray, Hypanus marianae, endemic to the Tropical Southwestern Atlantic province (Tab. 2; Fig. 7). Of the total number of threatened fish species in Paraíba, 51.20% are elasmobranch species, including 14 sharks (e.g., the great hammerhead Sphyrna mokarran) and eight rays (e.g., the Brazilian guitarfish Pseudobatos horkelii), both classified as CR species (Tab. 2). Among the data-deficient species (DD), 38 (61%) are marine and 24 (39%) are estuarine. Additionally, 28 (82%) marine and 3 (9%) estuarine and freshwater species remain non-evaluated (NE) (Tab. 2; Fig. 7).

FIGURE 7| A. Extinction risk categories of fish species of the Paraíba State, according to national assessment lists; and B. The threatened species. CR: Critically Endangered; EN: Endangered; VU: Vulnerable; NT: Near Threatened; LC: Least Concern; DD: Data Deficient; NE: Not Evaluated.

Human use. Around 200 species are used by humans for trade (including ornamental purpose), local consumption, and traditional medicine (Tab. 2). In marine environments, commercially valuable species include the “camurin” Centropomus undecimalis, the “pescada” Cynoscion acoupa,the “vermelhos” Lutjanus spp., the “serras” Scomberomorus cavalla and S. brasiliensis, and the “dourado” Coryphaena hippurus. Some elasmobranch species are also caught and consumed locally, such as “cação rabo seco” Rhizoprionodon porosus and other species of “cação”(e.g., Carcharhinus spp.). Among the species captured, some are classified as threatened, including Lutjanus cyanopterus and L. campechanus, as well as some shark and ray species such as Carcharhinus porosus and Urotrygon microphthalmum, which are incidentally captured and consumed by local communities. In freshwater environments, around 14 species are used by local communities, including the native “curimatã” Prochilodus brevis and the “piau” Leporinus piau, which are important food and income sources (Tab. 2). Additionally, several exotic cichlid species, such as “tilápia do Nilo” Oreochromis niloticus, “tucunaré” Cichla monoculus and C. ocellaris are also relevant food sources.

Discussion​

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Multiple anthropogenic pressures threaten tropical fish biodiversity, ecosystem resilience, and the sustainability of resource use (Floeter et al., 2006; Costa, Barletta, 2016; Torremorell et al., 2021). Despite significant progress in studying Paraíba’s ichthyofauna, research efforts remain spatially biased, thematically concentrated, and insufficient to face conservation and management demands. Addressing these gaps is crucial for protecting fish diversity and supporting food security and local economies (Floeter et al., 2006; Ribeiro et al., 2009, 2015; Leão et al., 2016; Dala-Corte et al., 2020; Pelicice et al., 2021).

Although research on Paraíba’s fish dates back to the 17th century, contemporary efforts have notably expanded since the 1990s, reflecting broader national trends in ichthyology (e.g., Ramos et al., 2019; Vendel et al., 2022; Villarins et al., 2022; Marceniuk et al., 2023) including reef fishes in the Southwestern Atlantic (Rocha, Sampaio, 2022), stream-dwelling fishes (Lima et al., 2021) and elasmobranchs (Coelho et al., 2021; Viana et al., 2024). This growth results from the expansion of graduate programs, the consolidation of research groups, and the establishment of biological collections such as CIUFPB, now the largest ichthyological collection in Brazil’s Northeast (Azevedo et al., 2010; Viana et al., 2019; Rosa et al., 2023; Márquez Velásquez et al., 2025). However, institutional consolidation contrasts with knowledge asymmetries, such as certain basins and taxonomic groups that remain poorly studied due to inconsistent funding and the lack of a long-term research policy (Crouzeilles et al., 2017; Oliveira et al., 2020).

The increasing number of publications has not led to proportional advances in understanding the life histories and ecological roles of the studied species, some of which are commercially exploited. Although the application of molecular tools and integrative taxonomy (e.g., Berbel-Filho et al., 2016, 2018; Lustosa-Costa et al., 2024) is starting to uncover cryptic diversity and enhancing taxonomical work (e.g., Medeiros, Maltchik, 2001); and comparative anatomical studies have recently begun to be applied to phylogenetic analyses (Medeiros et al., 2025),such studies are still emerging and should be stimulated by funding agencies. Ethnobiological studies increasingly focus on the importance of traditional ecological knowledge in understanding fish ecology (e.g., Rosa et al., 2005; Mourão, Nordi, 2006; Almeida et al., 2014) and supporting participatory governance and management practices at the local level (Müller, Guimbo, 2011; Brasil, 2025). In the future, collaborative governance models that integrate academic, governmental, and community actors will be essential. Long-term funding for underrepresented regions and species groups, and integrating local knowledge into research agendas can help align biodiversity science with conservation and social equity.

Geographical distribution of research

Continental environments. Ichthyological surveys in Paraíba over recent decades produced valuable data for major basins, including the Paraíba (e.g., Rosa, Groth, 2004; Ramos et al., 2018a,b), Piranhas-Açu (Silva et al., 2014), Mamanguape (e.g., Rosa, Groth, 2004; Xavier et al., 2012, Oliveira-Silva et al., 2018), Gramame (Gomes-Filho, Rosa, 2001; Suruagy, Crispim, 2023), Curimataú rivers (Ramos et al., 2005) and smaller coastal basins, such as Jaguaribe river (Ramos et al., 2017). The small coastal rivers within the Tambaba Environmentally Protected Area, including Graú, Mucatú, Bucatú, and Gurugi rivers (Ramos et al., 2019), have also been well sampled. However, research efforts are spatially biased, with more than half of all studies focused on the Paraíba and Mamanguape River basins.

This asymmetry results from biogeographical, logistical, and institutional factors. The Borborema Plateau shapes Paraíba’s hydrography, which encompasses both the Caatinga (northwest-flowing) and Atlantic Forest (eastward-draining) domains (Rosa et al., 2003b). However, the proximity of the Paraíba and Mamanguape River basins to the capital João Pessoa, where research infrastructure, graduate programs, and funding are concentrated, has led to their overrepresentation in the literature. Furthermore, the Mamanguape River flows through the municipality of Rio Tinto, where one of the campuses of the Universidade Federal da Paraíba (UFPB) is located, and its estuary lies within the federal Environmental Protected Area Barra de Mamanguape, thereby enhancing the visibility and research appeal in the river basin (CERHPB, 2004; Azevedo et al., 2010; Junqueira et al., 2020). Systematic inventories covering all of the state’s eleven drainage systems are still lacking.

Smaller basins, such as Gramame and Abiaí Rivers, are relatively well-represented in collections and publications, while others remain poorly known. The Curimataú basin, covering 5% of the state’s area, and the Miriri basin, both under strong degradation pressure, received insufficient attention (CERHPB, 2004; AESA, 2022). Similarly, the interstate river basins of Jacu, Guaju, and Trairi, despite covering only small portions of Paraíba, face intense anthropogenic pressures, including sugarcane expansion, sewage input, and overgrazing (Hunka et al., 2007; Araújo et al., 2019; Marinho et al., 2023). The lack of surveys in these basins, even when they overlap adjacent inventoried areas (e.g., Trairi River in RN; Medeiros et al., 2019), reveals not only logistical and institutional asymmetries but also a misalignment between conservation priorities and research programs.

In addition, ichthyofaunal surveys within Conservation Units (CUs) in Paraíba State have been published. However, these data remain scattered and often scarce for several CUs, hindering a comprehensive assessment of the species distribution and management effectiveness. Of the 34 CUs established, 12 classified as strictly protected and 22 as sustainable use, 14 is entirely covered by Atlantic Forest, 12 by Caatinga, and only the Guaribas Biological Reserve encompasses both biomes (da Costa & da Silva, 2025). Five UCs included terrestrial and coastal marine environments. The freshwater ichthyofaunal data are restricted to some units, including the Environmental Protected Area (APA) of Tambaba (Ramos et al., 2019); the Biological Reserves (REBIO) Guaribas in Camaratuba River (Gouveia et al., 2017); the Private Natural Heritage Reserve (RPPN) Fazenda Pacatuba (Furtado etal., 2018); the State Park Mata do Xem-Xém (Ramos etal., 2018a,b), both in Paraíba River; and the Mata do Buraquinho Wildlife Refuge, crossed by the smaller coastal basin of the Jaguaribe River (Ramos et al., 2017). Overall, these fragmented records highlight the need for broader and more systematic surveys within these units to better inform management and conservation planning.

Marine environments. Marine ichthyological research in Paraíba State has predominantly centered on shallow coastal zones, particularly natural reefs (e.g., Souza et al., 2007; Honório et al., 2010) and artificial structures, such as shipwrecks (e.g., Cardoso et al., 2020; Medeiros et al., 2022), largely due to the logistical ease of sampling nearshore habitats. Initial efforts began in the 1970s with estuarine surveys (Oliveira, 1972), followed by studies on shark fisheries (Lima, Ferreira, 1975) and broader inventories in the 1980s (Rosa 1980a, b). From the late 1990s, taxonomic efforts (e.g., Rosa et al., 1997; Rocha et al., 1998, 2000) expanded knowledge of the coastal ichthyofauna and compilations validated new records of species (Marceniuk et al., 2023; Silva Nascimento, Ramos, 2024).

Despite their ecological and socioeconomic significance, providing food, income, and tourism services (Costa et al., 2007; Ilarri et al., 2008), coastal reef systems are increasingly threatened by pollution, mass tourism, and unregulated coastal development (Ilarri et al., 2008; Queiroz Pereira, 2020). In contrast, deeper marine zones remain markedly understudied as a consequence of cost and access constraints. Once considered buffered from anthropogenic impacts (Pulido Mantas et al., 2024), these environments are subject to overfishing, sedimentation, debris, and invasive species (Rocha et al., 2018; Soares et al., 2019). Existing data for deeper areas result from oceanographic surveys (Ramos, 1994; Rincon et al., 2017; Mincarone et al., 2022; Villarins et al., 2022) and technical SCUBA dives (Rocha et al., 2000; Feitoza et al., 2005). Only recently, advanced methods, such as trimix diving, remotely operated vehicles (ROVs), baited remote underwater video systems (BRUVS), and sidescan sonar, begun to reveal mesophotic reef biodiversity (e.g., Medeiros et al., 2021; Silva et al., 2021). Scaling up the use of these tools is essential to minimize depth-related sampling gaps and inform conservation strategies across the full marine gradient.

Some of these marine environments in Paraíba State are currently under the protection of CUs, and some ichthyofaunal surveys have been carried out within and around these protected areas. Most of the five federal protected areas designated for sustainable use have ichthyofaunal studiesdocumented in the literature: the Restinga de Cabedelo National Forest (FLONA), the only federally protected area in the Paraíba River estuary (e.g., Vendel et al., 2022); the Acaú-Goiana Extractive Reserve (RESEX), located along the coastal boundary between Paraíba and Pernambuco States (e.g., Barleta, Costa 2009); the Area of Relevant Ecological Interest (AREI) of the Mamanguape River Mangroves, the Environmental Protection Area (APA) of the Mamanguape River (e.g., Xavier et al., 2012; Oliveira-Silva et al., 2018), and the most recently established Queimado Shipwreck Marine Protected Area (Medeiros et al., 2022). Additionally, the state hosts the strictly protected Areia Vermelha Marine State Park, a CU for which an ichthyofaunal survey exists but remains unpublished (Querino, 2011). Given that these CUs play a key role in conserving marine biodiversity and mitigating anthropogenic pressures, systematic surveys are crucial to support effective monitoring and adaptive management.

Species richness and taxonomy trends

Freshwater ecosystems. Paraíba’s freshwater fish fauna comprises about 8% of the species recorded in Northeastern Brazil, 52% of which occur in the Northeastern Caatinga and Coastal Drainagesecoregion (Carvalho-Rocha 2023), 73% in the Mid-Northeastern Caatinga Ecoregion (MNCE) (Lima et al., 2017; Silva et al., 2020), and only 5% of species endemic to the Pernambuco Endemism Centre in the Northern Atlantic Forest, which hosts approximately 58 species (Marinho et al., 2023). These patterns suggest that Paraíba’s ichthyofauna has a strong biogeographic alignment with the Caatinga, despite comprising only a small portion of the MNCE (56,467.24 km²; IBGE, 2022). The MNCE remains one of Brazil’s least studied regions for freshwater fishes, primarily due to its intermittent flows (Lévêque et al., 2008; Lima et al., 2017; Berbel‐Filho et al., 2018), and Northeast Brazil remains underrepresented in global ichthyofaunal databases (Tedesco et al., 2017).

Fish diversity in Paraíba is influenced by historical marine transgressions, aridification, and drainage reorganization (Ab’Sáber, 1957; Lundberg et al., 1998), as well as by human-induced introductions and extinctions (Rosa et al., 2003b; Ribeiro 2006). As is typical in most Neotropical basins (Buckup et al., 2007; Lima et al., 2017; Menezes et al., 2007), Paraíba’s fish fauna is dominated by Characiformes and Siluriformes with Acestrorhamphidae, Cichlidae, and Loricariidae being the most speciose families. Among the 62 recorded species, 13 are non-native (21%), mostly cichlids. Such cichlid dominance represents 73% of all exotic species reported for the Caatinga (Lima et al., 2017), raising concerns about biotic homogenization, competitive displacement, and long-term impacts on the structure of ecological communities.

The absence of Rivulidae records in Paraíba State is notable, given their endemism in the MNCE and commonness in Caatinga habitats (Myers, 1952; Lima et al., 2017; Berbel‐Filho et al., 2018; Abrantes et al., 2020). As annual fish species adapted to ephemeral ponds, rivulids survive through diapause and are typically active only during the rainy season (Berois et al., 2016). Their detectability is sensitive to sampling season, a factor that likely explains their omission from most dry-season surveys (Rosa et al., 2023). More than 50 annual fish species have been described from the Caatinga (Lima et al., 2017), but Paraíba has only one known record of a Rivulid (TPAR, 2025, pers. obs.). This points to the importance of seasonal sampling in temporary aquatic habitats, which may contain endemic lineages relevant for systematics, biogeography, and conservation planning. Eight of the 49 Caatinga endemics occur in Paraíba, including the “cascudinho” Parotocinclus jacksoni, whose distribution is restricted to the Caatinga (upper) section of the Mamanguape River basin (Oliveira-Silva et al., 2018; Ramos et al., 2021a). Another “cascudinho”, Parotocinclus spilosoma, is endemic to the Paraíba River basin. These narrow-range endemics contribute to the state’s ecological and evolutionary distinctiveness in freshwater fish assemblages and suggest the presence of microendemism shaped by isolation.

The full extent of freshwater fish diversity in Paraíba State remains unknown. Underestimation of species richness is the most likely scenario for most taxonomic groups, as supported by: (i) undescribed species in collections (e.g., Hypsolebias sp.; (ii) unresolved taxonomy of widespread complexes (e.g., Astyanax bimaculatus, Hoplias malabaricus, Synbranchus marmoratus); iii) limited digitization and few recent revisions (Lima et al., 2017; Marinho et al., 2023), and iv) under-sampled sub-basins.

Marine ecosystems. Despite its relative small coastline (138 km, Ab’Saber, 2001), Paraíba’s fish diversity comprises ~83% of species known from tropical northeastern Brazil (Rosa et al., 2023), and 14% of South American marine fishes (Reis et al., 2016). Forty of them are Brazilian endemics and three are restricted to the northeastern coast. Recent estimates suggest the existence of 418–427 coastal teleost species (Marceniuk et al., 2023; Rosa et al., 2023), and surveys in mesophotic zones and other under-sampled habitats continue to reveal additional diversity (e.g., Silva et al., 2021).

Elasmobranch fishes recorded in Paraíba account for about 19% of Brazil’s elasmobranch diversity (Kotas et al., 2023) with the stingray Hypanus marianae representing a coastal Tropical Southwestern Atlantic endemic. In Paraíba, sharks are the most common group in elasmobranch records, particularly Carcharhiniformes, as in other Brazilian regions (e.g., Rosa, Gadig, 2014; Bezerra et al., 2021) and worldwide (Compagno, 2003; Ebert et al., 2021). Myliobatiformes is the most species-rich order among batoids (Fricke et al., 2025). Within Myliobatiformes, records of Mobula birostris reported in the literature for Paraíba likely refer to Mobula yarae Bucair & Marshall, 2025. The regional pattern of more shark than ray species, which diverges from global patterns (Fricke et al., 2025), may result from research focus, greater catchability, and higher intrinsic diversity of sharks within Brazil’s elasmobranch fauna (e.g., Meneses et al., 2005; Rosa, Gadig, 2014; Bezerra et al., 2021). It may also be partly explained by the exclusion of grey literature in this study, which includes theses and undergraduate monographs that hold unpublished information on the elasmobranch fish fauna of Paraíba and the Brazilian Northeast.

Paraíba’s marine fish assemblages reflect a complex interplay of biogeographic processes that shape patterns of endemism and species turnover. Situated within the north-northeastern subprovince of the Brazilian Province (Briggs, 1974; Pinheiro et al., 2018), the state lies at a transitional biogeographic zone. The intersection of ecological processes and historical events that have shaped Paraíba’s marine biota includes dispersal capacity (Floeter et al., 2008; Cowman, Bellwood, 2013), environmental conditions and local adaptation (Robertson and Cramer 2014), oceanographic currents (Spalding et al., 2007), habitat discontinuities (Riginos, Nachman, 2001), isolation-by-distance and gene flow (Rocha et al., 2008), and vicariance events (Joyeux et al., 2001).

Conservation concerns

Aquatic ecosystems in Paraíba State hold numerous endangered species whose ecological functions are tightly linked with ecosystem resilience. In marine habitats, threatened taxa include teleosts of the families Epinephelidae, Labridae, Lutjanidae, Carangidae, and Ariidae, and elasmobranchs of the families Sphyrnidae, Carcharhinidae, and Rhinobatidae. These species play diverse ecological roles. For example, parrotfishes (Labridae) promote reef diversity through grazing and bioerosion (Bonaldo et al., 2014) and sharks control prey dynamics (Heithaus et al., 2008). The critically endangered endemic parrotfish Scarus trispinosus (ICMBio, 2026) experienced population declines over the past three decades due to overexploitation by small-scale fisheries (Roos et al., 2016). Additionally, the demographic decline of the critically threatened mesopredator Epinephelus itajara (Epinephelidae) affects epibenthic and demersal food webs of marine rocky shores (Artero et al., 2015; Freitas et al., 2015). This is a widespread effect as the species ranks among the most endangered taxa in the whole Atlantic Ocean (IUCN, 2025).

Chondrichthyans, which play key ecological roles (Heithaus et al., 2012; Dedman et al., 2024), rank among the most threatened vertebrate groups worldwide (Dulvy et al., 2021; Pacoureau et al., 2021). Paraíba State is no exception, as more than 40% of local chondrichthyan species face severe extinction risks (ICMBio, 2026). Widespread ecological impact derived from disruption of size-structured predation and nutrient flow chains arise from population declines of top predators, such as the great hammerhead shark Sphyrna mokarran, the smalltail shark Carcharhinus porosus and the Brazilian guitarfish Pseudobatos horkelii.

Major anthropogenic processes that may be driving marine population declines in Paraíba include habitat degradation, pollution, coastal development, climate change (Hillebrand et al., 2018; Prates et al., 2010), overfishing (Pinheiro et al., 2010; Bender et al., 2014), and biological invasions, particularly by the lionfish Pterois volitans (Luiz et al., 2021; Soares et al., 2022, Maggioni et al., 2023). The spread of lionfish in a variety of habitats, from mangroves to deep reefs and in marine protected areas, raises concerns as this invasive species imposes strong predation on juvenile reef fishes, which is likely to erode the biotic architecture of local food webs (Albins, Hixon, 2008; Soares et al., 2022, 2023). It is important to note that many of the introduced species documented in the Southwestern Atlantic and along the Brazilian coast since the mid-2000s (Schwindt et al., 2020) exhibit cryptobenthic behavior (Ferreira et al., 2009), which may hinder earlier detection and facilitate unnoticed establishment, highlighting the potential occurrence of cryptic invasive species in Paraíba State.

Despite their ecological and economic importance, estuaries are among the most threatened ecosystems (Whitfield et al., 2022). Estuaries are critical habitats for many fish species, playing a key role as nurseries by providing juveniles with favorable conditions, such as high temperatures, prey availability, and refuge from predators, enhancing growth and survival (Potter et al., 2015; Xavier et al., 2022). Anthropogenic pressures, including human occupation and economic activities, resulting in sewage discharges (Cortez et al., 2023), sugarcane plantations, and intensive shrimp aquaculture in the surrounding area (Marcelino et al., 2005; Barletta et al., 2010) are among the main threats, further compounded by impacts related to climate change. More than 160 species have been recorded in the estuaries of Paraíba State (mainly in Mamanguape and Paraiba Rivers) including threatened species such as Lutjanus cyanopterus and the Atlantic tarpon Megalops atlanticus Valenciennes, 1847, which occurs in both estuarine and reef areas (Xavier et al., 2012; Vendel et al., 2022), highlighting ecosystem connectivity. The Critically Endangered Carcharhinus porosus was reported in the Paraiba estuary and at Tambaú and Bessa beaches, but it has not been captured there or elsewhere in Paraíba State since the 1980s, emphasizing the need for continuous monitoring of fish populations. The diversity and structural complexity of these habitats highlight the importance of the Mangrove-Reef System as a priority area for conservation and research.

Regarding freshwater environments, the conservation status of many species remains unclear as substantial knowledge gaps persist. To date, only two species, the loricariids Parotocinclus seridoensis and P. spilosoma,are listed as endangered, with population declines linked to habitat loss and fragmentation (ICMBio, 2026). Three native species have not been evaluated, including Hemigrammus rodwayi, Pseudancistrus papariae and the endemic Parotocinclus jacksoni (Ramos et al., 2021a). Missing data constraints extinction risk assessments and conservation planning, reinforcing the need for substantial investments supporting systematic surveys in freshwater systems.

Freshwater ecosystems and their fishes are also among the most threatened habitats and species worldwide (Dudgeon et al., 2006; Sayer et al., 2025). Paraíba mirrors these global trends, which are driven by the interplay of diverse regional processes, such as: i) hydrologic changes caused by dams, weirs and channels (Brito et al., 2020; Carvalho et al., 2023); ii) habitat destruction from intensive agriculture, especially sugarcane plantations, along with removal of riparian vegetation, livestock and shrimp farming (Sousa et al., 2020; Marinho et al., 2023); iii) pollution and eutrophication of water bodies (Correia et al., 2015); and iv) introductions of invasive species (Leão et al., 2011).

To date, 13 non-native freshwater fish species have been reported in Paraíba (21%) (Latini et al., 2016), indicating a strong trend of biotic homogenization. This results from accidental or deliberate introductions via aquaculture, aquarium trade (Leão et al., 2011), biological control (Leão et al., 2011; Latini et al., 2016), and interbasin water transfer (Ramos et al., 2021b). The “tucunaré” (Cichla ocellaris) and tilapias (Oreochromis niloticus and Coptodon rendalli) were introduced for food safety by government programs or for sport fishing (Rosa, Groth, 2004; Leão et al., 2011). Aquarium species were accidentally released in the wild, such as Xiphophorus maculatus, Astronotus ocellatus, and Parachromis managuensis (Ramos et al., 2020; Marinho et al., 2023). Species used for mosquito control, such as the“barrigudinho” Poecilia reticulata, used against the dengue vector, Aedes aegypti (Levis et al., 2013; Azevedo-Santos et al., 2016). Biotic homogenization is expected to increase due to large-scale interbasin water transfer, such as illustrated by “tetra-fortuna” Moenkhausia costae, the first record of an allochthonous species in the Paraíba river arising from its novel connections with the São Francisco River since 2018 (Ramos et al., 2021b).

Introductions correlate with habitat degradation, which amplifies the impacts of invasive species (Poff et al., 2007), leading to the regional extinction of native species (Rosa, Groth, 2004; Charvet et al., 2021); biotic homogenization (Latini, Petrere, 2004; Rosa, Groth, 2004; Orlandi Neto et al., 2024); food web changes arising from novel predation and competition pressures (Zaret, Paine, 1973; Pyke, 2008); hybridization (Oliveira et al., 2006), and the spillover of parasites and diseases (Dobson, May, 1987; Williams et al., 2013). Emerging host-parasite dynamics under strong anthropogenic changes deserve particular attention (Chalkowski et al., 2018; De Lima et al., 2025; Moraes et al., 2025). For example, it is still unclear whether nematodes parasitizing M. costae in the Paraíba river are also exotic species (Sousa et al., 2025), with potential to reaching the native fauna and leading to community changes (Lacerda et al., 2013).

Despite the importance of protected areas for biodiversity conservation, they also face threats and challenges. Exotic species have already been recorded within their area. For example, in APAs of Tambaba (Ramos etal., 2019) the “tucunaré C. monoculus and the “barrigudinho” P. reticulata were recorded, the latter also documented in REBIO Guaribas (Gouveia etal., 2017); the RPPN Fazenda Pacatuba (Furtado etal., 2018) and State Park Mata do Xem-Xém (Ramos etal., 2018a,b). Additionally, the “tilápia do Nilo” O. niloticus has been recorded in the REBIO Guaribas and the State Park Mata do Xem-Xém, within the Paraíba River basin (Gouveia et al., 2017; Ramos et al., 2018a). As previously mentioned, these species pose a threat to the native ichthyofauna, and their populations should be monitored. On the other hand, the need to effectively implement management plans persists. These plans have been established in only eight of the 34 protected areas in the state, although in some cases their preparation is underway, including the holding of public hearings (da Costa, da Silva, 2025). This panorama is consistent with that observed in other regions of Brazil, where the implementation of management plans remains limited: 84% of protected areas established more than five years ago still lack this critical management tool (Riva et al., 2014). Moreover, the establishment of management councils, currently present in 14 of these areas, is critical to ensure effective governance and promote participatory decision-making. Therefore, ichthyological surveys, proper design and management plans, and the establishment of management councils in UCs in Paraiba State are essential to guide the biodiversity conservation actions and an effective management.

Economic perspectives. The freshwater and marine ichthyofauna of Paraíba support local communities through commercial and sport fishing (e.g., Nascimento et al., 2016; Santos et al., 2023) and ecotourism (Costa et al., 2007; Souza, 2007). Particularly, small-scale fisheries and aquaculture are vital for local economies in terms of livelihoods and food security (Paulo Júnior et al., 2012). Key resources include “tilapia”, “traíra”, “tucunaré”, “curimatã”, and “pescada do Piauí”(IBAMA, 2008). Artisanal marine fishing targets mostly Scombridae (e.g., Euthynnus alletteratus, Scomberomorus brasiliensis, S. cavalla) and Carangidae (e.g., Caranx bartholomaei), Lutjanidade and Centropomidae (Centropomus undecimalis) (IBAMA, 2008), with eventual captures of elasmobranchs in the family Carcharhinidae (Rhizoprionodon porosus, Carcharhinus spp.), Sphyrnidae (Sphyrna spp.) (Mariano, Rosa, 2010; Morais, 2016), and rays, including Urotrygon microphthalmum, Rhinoptera bonasus, Aetobatus narinari and Hypanus guttatus (RSR, 2025, pers. comm.; JPCB, 2023, pers. comm.). In general, these elasmobranchs are caught incidentally in artisanal beach seines and gillnets. Catches have strongly declined, for example, with artisanal fishing dropping to half between 2002–2004 (IBAMA, 2007). As a consequence, regulation is urgently needed for both marine and freshwater environments as fish stocks approach collapse (MPA, 2013).

Other income-generating activities involve ornamental species, such as seahorses (Hippocampus spp.),a rare fish species threatened in Brazil, not only by the trade and habitat degradation (Borges et al., 2025a) but also by its biological characteristics as high site fidelity and low reproduction rate (Foster, Vincent, 2004). In this context, over the past decades, researchers at UFPB developed conservation projects investigating biological, ecological, and bioeconomic aspects of the seahorse Hippocampus reidi (e.g., Rosa et al., 2005). Currently, researchers at the Universidade Estadual da Paraíba (UEPB) are conducting studies to fill knowledge gaps regarding seahorse species and determine their conservation priorities in Brazil, using a multidisciplinary approach that integrates biogeographical, ecological, and ethnobiological knowledge (Borges et al., 2025b).

The conservation and management of species and fishery resources are essential not only to preserve biodiversity but also to ensure the continuity of economic activities, social well-being and food security (Tickner et al., 2020; Oguh et al., 2021). Multidisciplinary efforts and participatory management, to improve decision-making and socio-ecological management (Pomeroy, Rivera-Guieb, 2006), as well as in the identification of potential resources. Additionally, empowering local communities through human resource development can support management and productive activities by strengthening local capacities in fisheries management, monitoring and enforcement, civic education, and communication. Such initiatives, typically implemented through training courses or workshops, also contribute to the development of sustainable economic alternatives, such as ecotourism, as observed in other regions of Brazil (WCS-Brasil, https://amazoniacienciaciudadana.org/) and other South American countries (https://www.wwf.org.co).

Some examples that advance in this direction may involve training fishermen in the cultivation and processing techniques of native freshwater fish species, as a means to diversify local livelihoods, reduce pressure on wild stocks, and offer products with greater added value and acceptance in different markets, for example, boneless fillets (Kubitza et al., 2007). Additionally, strengthening the fisheries value chain via training in fish processing and marketing are complementary strategies. Encouraging responsible fish consumption can improve food security while increasing community income. Finally, transforming discarded fishing nets into handcrafted products can also contribute to the mitigation of marine pollution.

Finally, it is relevant to discuss with local communities how the sustainable management of fish populations contributes to biodiversity conservation and long-term profitability for the fishing and tourism activities. This could include practices such as: i) the identification, establishment and control of key areas for reproduction of freshwater species and of marine protected areas, ii) catch limits, iii) the promotion of conservation of native species, and iii) the control of invasive species. Thus, it is essential to utilize traditional knowledge and empower local people through the development of human capital, science, and technological innovation to create sustainable solutions that meet the needs of communities (Alcalá Moya, 2011; Nishi, Subramanian, 2023).

Integrative overview and perspective. This study presents significant advances in the research on the ichthyofauna of freshwater, estuarine and marine environments in Paraíba State; however, there is still much to be done. Our review found biases related to research topics and the spatial distribution of studies and species. Although much of the river basins have already been inventoried, some areas are under-sampled, such as the small coastal rivers. Thus, future sampling efforts should be directed to these under-sampled localities to address these gaps, in addition to the inventories performed at the regional scale to support public conservation policies (Rosa et al., 2003a; Tedesco et al., 2017). Reducing these gaps will require an intensive and joint effort of a diverse team of professionals and institutions. Interdisciplinary and inter-institutional efforts are also applied to biodiversity research and social development. Additionally, infrastructure, equipment, new technologies, and investments in human resources are essential to optimize research and promote projects and initiatives that involve local communities, biodiversity conservation, and social welfare.

To address most of the knowledge gaps of the fish fauna of Paraíba State, and promoting its conservation and sustainable use, we recommend the following actions: i) fish surveys to fill local gaps, mainly in: under-sampled drainages such as Miriri, Curimatau and Camaratuba Rivers as well as the small coastal and interstate basins; headwater areas of drainage, which are more difficult to access; deep reefs, estuaries and in the UCs; ii) to prioritize the monitoring of basins and their estuaries, mainly those harboring the highest number of endemic and threatened species, such as the Paraíba and Mamanguape River basins; iii) to strengthen the control mechanisms of conservation units (CUs); to implement and update management plans in order to improve oversight and enforcement; and to expand existing conservation units to encompass potential key areas for species conservation; iv) to develop projects to recover degraded aquatic habitats, through restoration of riparian forests programs, particularly in estuarine areas such as the Mamanguape and Paraíba River estuaries, as well as the protection and recovery of river headwaters; and coastal reef restoration programs; v) promotion of environmental education projects for the conservation of freshwater and marine ecosystems and their species in schools and communities; vi) restriction and control of introduction of exotic species in natural aquatic environments; and vii) digitalization, curation and availability of biological collection data. It is essential to acknowledge that a substantial amount of unpublished research, stemming from undergraduate and graduate theses from universities in the Northeast, remains unexplored and has yet to be published. This represents valuable sources for research on the ichthyofauna of Paraíba State. The following table (Tab. 3) presents topics and questions that we consider relevant to future research. Advancing these actions will be crucial for strengthening research and conservation of the fish fauna of Paraíba State.

TABLE 3 | Future research on the fish fauna of Paraíba State.

Research area	Future perspectives
Biology and Population Ecology	Studies on the biological aspects and population status of data-deficient and near-threatened species.
	Explore the age and growth, as well as the life history parameters, of fisheries resources, which are key to effective fisheries management.
Ecology	Effects of climate change on demography, physiology, and biological processes, as well as on the distribution patterns of species.
Ecotoxicology	Assessing the effects of pollution in fish populations and the effect of human consumption of the polluted species; screening of contaminants in tissues.
Biogeography-Phylogeography	How species responded to past changes may help to predict how they might cope with the ongoing climatic changes. Evaluation of processes that generated and maintained the diversity of fish fauna of the Caatinga and the Atlantic Forest.
Biological invasions	To assess the degree of invasibility of this species in the basin, the ecological impacts (on the ecosystems and on the native species) are considered. For marine species, to assess the distribution, biology, reproduction, and trophic ecology aspects, and to propose control and management measures.
Genetic-eDNA	DNA collected from environmental samples, such as water, can be used to analyse genetic data and detect the presence of species in the area.
Parasitology	Parasite diversity, parasite-host relationships, effects of parasitism on the host`s condition and how environmental changes and biological invasions may affect these associations.
Fisheries	Monitoring of landing data, description of catches and fishing effort.
Conservation	Delineation of habitat use and critical areas for endangered, endemic species; evaluation of the effectiveness of existing MPAs for the species.
Bioeconomy	Explore ways to sustainably use and manage fish, in a manner that ensures the long-term health of aquatic ecosystems and maximises the benefits for local communities.
Taxonomy and Systematics	Ichthyofaunal surveys at poorly sampled areas; use of integrative taxonomy to better understand widespread species complexes; digitise scientific data from collections and make them available; improve data quality of scientific collections by providing reliable species identification.

Acknowledgments​

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We gratefully acknowledge the Fundação de Apoio à Pesquisa do Estado da Paraíba (FAPESQ) and Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPq for the postdoctoral fellowship to VMV (2413/2023). We thank the curators and technicians of the institutions who kindly sent us their records (UFPB, UFRN, MZUSP, MNRJ, PUCRS). We thank all collaborators of the Ichthyological collections of UFPB, and to all the ichthyologists who contributed in any way with their comments. We are grateful to the collaboration and support of PRONEX/FAPESQ-PB (027–2023), specially to Alexandre Vasconcellos (UFPB). We thank the Instituto Nacional de Ciência e Tecnologia da Biodiversidade e uso Sustentável de Peixes Neotropicais (INCT-Peixes) (MCTIC/CNPq (#405706/2022–7) for funding technician work at CIUFPB. TPAR thanks the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the the Senior Postdoctoral Fellowship (PDS) (102654/2024–7).

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Authors

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Viviana Márquez-Velásquez^1,2 , Ricardo S. Rosa³, Manoela Maria Ferreira Marinho⁴, João Paulo Capretz Batista da Silva³, Telton Pedro Anselmo Ramos⁵, Ana Carolina Figueiredo Lacerda⁶, Gabriel Beltrão³, André Castro³, Marcio Wanderson³, Danielle Cristina Gamba³, Laure Berti-Equille^1,7 and Rafael L. G. Raimundo^1,2,7

^[1]    IDEAL International Joint Lab, Universidade Federal da Paraíba, Campus IV, 58297-000, Rio Tinto, PB, Brazil. (VMV) vmarquez@squalus.org (corresponding author), (RLGR) rafael.raimundo@academico.ufpb.br, (LBE) laure.berti@ird.fr.

^[2]    Departamento de Engenharia e Meio Ambiente and Programa de Pós-Graduação em Ecologia e Monitoramento Ambiental (PPGEMA), Universidade Federal da Paraíba, Campus IV, 58297-000, Rio Tinto, PB, Brazil.

^[3]    Laboratório de Ictiologia, Centro de Ciências Exatas e da Natureza, Universidade Federal da Paraíba, 58051-900, João Pessoa, PB, Brazil. (RSR) rsrosa@dse.ufpb.br, (JPCB) jpzoologia@dse.ufpb.br, (GB) gabrielbeltraoictio@gmail.com, (AC) andre.castro@academico.ufpb.br, (MW) mwvm22@gmail.com, (DCG) danielle.gamba@academico.ufpb.br.

^[4]    Instituto de Biologia, Universidade Federal de Mato Grosso do Sul, 79070-900, Campo Grande, MS, Brazil. (MMFM) manoela.marinho@gmail.com.

^[5]    Laboratório de Ictiologia Sistemática e Evolutiva, Departamento de Botânica e Zoologia, Programa de Pós-Graduação em Sistemática e Evolução, Centro de Biociências, Universidade Federal do Rio Grande do Norte, Av. Senador Salgado Filho, 3000, Lagoa Nova, 59978-970, Natal, RN, Brazil. (TPAR) telton@gmail.com.

^[6]    Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento de Sistemática e Ecologia, Laboratório de Hidrologia, Microbiologia e Parasitologia, Universidade Federal da Paraíba, Cidade Universitária, s/n, 58051-900, João Pessoa, PB, Brazil. (ACFL) acflacerda@dse.ufpb.br.

^[7]    Institut de Recherche pour le Développement (IRD), UMR ESPACE-DEV, Montpellier, France.

Authors’ Contribution

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Viviana Márquez-Velásquez: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Visualization, Writing-original draft.

Ricardo S. Rosa: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Supervision, Validation, Writing-original draft, Writing-review and editing.
Manoela Maria Ferreira Marinho: Data curation, Validation, Writing-original draft, Writing-review and editing.

João Paulo Capretz Batista da Silva: Data curation, Writing-original draft, Writing-review and editing.

Telton Pedro Anselmo Ramos: Data curation, Writing-original draft, Writing-review and editing.

Ana Carolina Figueiredo Lacerda: Writing-original draft, Writing-review and editing.

Gabriel Beltrão: Data curation, Writing-review and editing.

André Castro: Data curation, Writing-review and editing.

Marcio Wanderson: Data curation, Digitization of collection data

Danielle Cristina Gamba: Data curation, Digitization of collection data

Laure Berti-Equille: Writing-original draft, Writing-review and editing.

Rafael L. G. Raimundo: Conceptualization, Funding acquisition, Investigation, Project administration, Validation, Writing-original

Ethical Statement​

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Not applicable.

Competing Interests

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The author declares no competing interests.

Data availability statement

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The data supporting the findings of this study are included in the supplementary material of this article. The datasets generated during the current study are also available in the DATAPB repository (DOI: https://doi.org/10.48472/DATAPB/QL6VGJ) upon request.

AI statement

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The authors used the AI tool Claude Sonnet 4.6for a final review of the reference list, specifically to check formatting consistency and ensure compliance with the journal’s guidelines. No AI tools were used for data analysis, interpretation of results, or generation of scientific content.

Funding

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This work was supported by FAPESQ-CNPq (postdoctoral fellowship 77/2022 to VMV), DATAPB Universal Project FAPESQ 3087–21 and the French Institute for Research on Sustainable Development – IRD via the International Joint Laboratory on Sustainability IDEAL.

Supplementary Material

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Supplementary material SUP

Peer Review

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Peer Review File

How to cite this article

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Márquez-Velásquez V, Rosa RS, Marinho MMF, da Silva JPCB, Ramos TPA, Lacerda ACF, Beltrão G, Castro A, Wanderson M, Gamba DC, Berti-Equille L, Raimundo RLG. Ichthyofauna of Paraíba State, Brazil: diversity patterns, knowledge gaps, and challenges for fish conservation and sustainable use. Neotrop Ichthyol. 2026; 24(2):e250176. https://doi.org/10.1590/1982-0224-2025-0176

Copyright​

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This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Distributed under

Creative Commons CC-BY 4.0

© 2025 The Authors.

Diversity and Distributions Published by SBI

Accepted March 5, 2026

Submitted October 12, 2025

Epub July 20, 2026