The marine and estuarine bony fishes (Teleostei) of the Golfão Maranhense on the eastern Amazon coast, northern Brazil

Os peixes ósseos (Teleostei) marinhos e estuarinos do Golfão Maranhense, costa leste da Amazônia, norte do Brasil

Abstract

The Golfão Maranhense is an extensive estuarine complex that provides nursery grounds and other habitats for several fish species, as well as a wealth of resources for the coastal and estuarine artisanal fisheries of the Maranhão state, in northern Brazil. The present study provides a comprehensive inventory of marine and estuarine teleost fishes of the Golfão Maranhense based on field samplings, analyses of voucher specimens from local collections and literature data. This inventory presents a total of 163 species of teleosts belonging to 107 genera, 57 families and 27 orders. The order Perciformes prevailed in terms of richness (27 species), followed by Acanthuriformes, Carangiformes, Clupeiformes and Siluriformes. The most diverse family was Sciaenidae (19 species), followed by Carangidae, Ariidae, Engraulidae and Gerreidae. Among the 163 species recorded, 18 are new occurrences for the Golfão Maranhense. According to the IUCN Red List, seven species are classified as Vulnerable. Considering the official Brazilian list of threatened species, five species appear as Vulnerable and one – Epinephelus itajara – as Critically Endangered. Four of the recorded species (Butis koilomatodon, Trypauchen vagina, Omobranchus sewalli and Oreochromis niloticus) are exotic, and were probably introduced into the study area by shipping, fish farms or by the extension of their range along the Brazilian coast. The present study organizes, updates and expands the known and relatively limited data on the fish fauna of the Golfão Maranhense, highlighting the need for further research towards gathering data for conservation and management of the ichthyofauna of this important estuarine complex in the Northern Brazilian Coast.

Keywords:
conservation status; estuarine fish; estuary; fish fauna; marine fish; São Marcos Bay

Resumo

O Golfão Maranhense é um extenso complexo estuarino caracterizado como uma fundamental área de berçário e habitat para diversas espécies de peixes, fornecendo grande riqueza de recursos naturais para a pesca artesanal estuarina e costeira do estado do Maranhão. O presente estudo constitui um inventário abrangente dos peixes teleósteos marinhos e estuarinos do Golfão Maranhense baseado em coletas de campo, análise de vouchers de espécimes de coleções locais e dados da literatura. Este inventário apresenta um total de 163 espécies de teleósteos pertencentes a 107 gêneros, 57 famílias e 27 ordens. A ordem Perciformes dominou em termos de riqueza (27 espécies), seguida por Acanthuriformes, Carangiformes, Clupeiformes e Siluriformes. A família mais diversificada foi Sciaenidae (19 espécies), seguida por Carangidae, Ariidae, Engraulidae e Gerreidae. Entre as 163 espécies registradas, 18 são novas ocorrências para o Golfão Maranhense. Sete espécies estão classificadas como Vulneráveis de acordo com a lista vermelha da IUCN. Considerando a lista brasileira oficial de espécies ameaçadas, cinco espécies têm status Vulnerável e uma – Epinephelus itajara – está Criticamente em Perigo. Do total de registros, quatro espécies (Butis koilomatodon, Trypauchen vagina, Omobranchus sewalli e Oreochromis niloticus) são exóticas e provavelmente foram introduzidas na área de estudo por água de lastro de navios cargueiros, pela piscicultura ou pela extensão de sua área de ocorrência ao longo da costa brasileira. O presente estudo organiza, atualiza e amplia os dados conhecidos e relativamente limitados sobre a ictiofauna do Golfão Maranhense, destacando a necessidade de mais pesquisas para conservação e gestão da ictiofauna deste importante complexo estuarino da costa norte brasileira.

Palavras-chave:
estado de conservação; peixes estuarinos; estuário; fauna de peixes; peixes marinhos; Baía de São Marcos

1. Introduction

Tropical estuaries typically have an extremely diverse fauna and are socioeconomically important for local coastal communities (Barletta et al., 2023; Leung et al., 2024). As many fish use these environments at some stage of their life cycle, for feed or reproduce, or even as a permanent home, estuaries are fundamental for the survival of many marine species (Pessanha et al., 2021; Harrison and Whitfield, 2024). The fish fauna of an estuary can influence the composition, abundance, and distribution of other groups of organisms that inhabit these environments, which means that shifts in the structure of the fish assemblage can provide systematic insights into the quality of the environment (Borges et al., 2010; Viana and Lucena-Frédou, 2014). Given this, inventories of the fauna that inhabits an estuarine ecosystem are fundamental to the establishment of sound public policies for the conservation and management of these environments, in particular those that are most threatened by human impacts (Silveira et al., 2010; Stephenson and Stengel, 2020; Gouvêa et al., 2023), as are typical of many tropical estuaries (Azevedo et al., 2016).

The northern coast of Brazil extends over more than 2,500 km from the mouth of the Oiapoque river, at the border with French Guiana to the Parnaíba River, which forms the border between the states of Maranhão and Piauí (Ekau and Knoppers, 1999; Marceniuk et al., 2021a). This coastline is highly indented in many places by the formation of bays, inlets, estuaries, and continental islands, and encompasses the largest continuous tract of mangrove forest under legal protection found anywhere in the world (Souza-Filho, 2005; Brasil, 2006; Souza-Filho et al., 2009; ICMBio, 2018). This is also one of the most productive sectors of the Brazilian coast, with major global fishing grounds off the coasts of the states of Amapá, Pará, and Maranhão (Brasil, 2006; Corrêa and Martinelli, 2009; Marceniuk et al., 2021a). This region has unique oceanographic characteristics, which are shaped by the mixing of the freshwater and sediments discharged by the Amazon and Orinoco rivers, that form the Amazon-Orinoco plume (Ekau and Knoppers, 1999; Rocha, 2003; Brasil, 2007; Marceniuk et al., 2013). The coastline of the state of Maranhão is approximately 640 km long, extending from the mouth of the Gurupi River, in the west, to the Parnaíba Delta in the east (Rios, 2001; Feitosa, 2006). The Golfão Maranhense is located in the center of this coast, encompassing a major estuarine complex, which is of considerable importance for the Brazilian economy, due to the presence of industrial installations and three major shipping terminals – Itaqui, Ponta da Madeira, and Alumar – for the transportation of cargo and minerals (Amaral and Alfredin, 2010; Queiroz et al., 2022).

Previous studies of the fish fauna of the northern Brazilian coast have highlighted the diversity of this region, including numerous rare and endemic species, many of which are adapted to life in estuaries or occur in areas dominated by the phenomenal discharge of freshwater from the Amazon River (Camargo and Isaac, 2001; Caires et al., 2019; Rotundo et al., 2019; Marceniuk et al., 2021a).

Up to now, however, the composition of the fish fauna of the Golfão Maranhense has been poorly documented, given that most studies have focused on either commercial species (Santana et al., 2016; Santana et al., 2019a) or only a few locations (Martins-Juras et al., 1987; Pinheiro-Junior et al., 2005; Carvalho Neta and Castro, 2008; Silva et al., 2018). Some of these studies have also employed outdated nomenclature (e.g. Martins-Juras et al., 1987; Pinheiro-Junior et al., 2005) or have provided no information on the voucher material (e.g. Carvalho Neta and Castro, 2008; Silva Júnior et al., 2013; Silva et al., 2018). Together, these shortcomings impose severe limitations on the potential of these studies for the understanding of the composition of the local fish fauna (Winker, 2004; Marceniuk et al., 2013, 2017; Gurgel-Lourenço et al., 2023).

However, some studies carried out in the last decade have contributed to changing this scenario and increasing knowledge of the species from the Golfão Maranhense. These studies began to include data from specimens deposited in scientific collections that allow the accuracy of the information to be verified by other researchers (e.g. Santana et al., 2019b; Aguiar et al., 2021; Brito et al., 2019; Vidigal et al., 2023). Another study that contributed to increasing knowledge of the ichthyofauna from the Golfão Maranhense was the study by Marques et al. (2024), who carried out a systematic review for several groups of local fauna and flora, establishing a standardized protocol for obtaining data from information in published articles, monographs, dissertations, theses, and digital repositories of collections.

Despite this effort, more extensive and comprehensive studies are still needed to provide a more complete inventory of the fish fauna of the Golfão Maranhense. In addition to the basic scientific knowledge, these data are fundamental to the understanding of the impacts of fisheries and shipping on the local fish assemblage. In this context, the present study aimed to provide the most comprehensive possible list of marine and estuarine bony fish species found in the Golfão Maranhense, based on the collection of new specimens, the analysis of vouchers deposited in local ichthyological collections, and published data. It is hoped that these findings will contribute to the best understanding of the changes that have occurred in the characteristics of the local fish community over the past few decades.

2. Material and Methods

2.1. Study area

The Golfão Maranhense (Figure 1) is an extense estuarine complex influenced by the waters of the western South Atlantic and the discharge of a number of major rivers, mainly the Itapecuru, Mearim, Munim and Pindaré, which flow into São Marcos, São José and Arraial bays (Stride, 1992; Azevedo, 1999). This gulf is 90 km wide, with 490 km of open coast and 144 beaches (Bandeira, 2013; Pereira et al., 2016), stretching from Ponta do Guajuru, in the municipality of Cedral, in the west, to Santaninha Island, in the municipality of Primeira Cruz, in the east, which is also the western limit of the adjacent Lençóis Maranhenses National Park (Feitosa, 2006; Teixeira and Souza Filho, 2009). The gulf also encompasses a number of major islands, including Upaon-Açu (São Luís Island), Medo, Pequena, Livramento, Caranguejos, Duas Irmãs, Tauá-Redonda, Tauá-Mirim and Ponta Grossa islands, as well as five principal bays, including the three mentioned above, and the Cumã and Tubarão bays (Feitosa, 2006; Chaves et al., 2016; Pereira et al., 2016; Teixeira and Souza Filho, 2009).

Figure 1
Location of the fish sampling sites (blue dots) surveyed during the present study in the Golfão Maranhense, in Maranhão, northern Brazil. Green dots = represent the collecting localities of the specimens examined from the COFAUMA, black dots = represent the collecting localities of the specimens examined from the CPUFMA.

The local tides are semi-diurnal and have the greatest tidal amplitude of any location on the Brazilian coast, reaching 7.2 m, with a mean amplitude of approximately 6.6 meters (Caroli et al., 2017). The region has a humid tropical climate, with maximum temperatures varying between 28°C and 34°C along the year, and two well-defined seasons: a dry season from July through December, and a rainy season between January and June. Annual precipitation ranges between 1,600 mm and 2,000 mm (Teixeira and Souza Filho, 2009; UEMA, 2016; IMESC, 2021).

2.2. Sampling sites and collection methods

Fish specimens were collected between May 2022 and January 2024 at 18 sites in the Golfão Maranhense, including the mouth of the Jaguarema River (P1), Raposa (P2, P3), Curupu Bay (P4, P5), the estuary of the Santo Antônio River (P6, P7, P8, P9, P10), the estuary of the Paciência River (P11, P12, P13, P14), Porto do Barbosa (P15), and the estuaries of the Tibiri (P16), Munim (P17) and Periá (P18) rivers (Figure 1). The specimens were captured using trawls, gillnets and hand lines, and were euthanized immediately by immersion in ice water. Specimen collection was authorized by the Chico Mendes Institute for Biodiversity Conservation (ICMBio) through the Biodiversity Authorization and Information System (SISBio), licenses 26334-4 and 83715-1. The study was also approved by the Ethical Committee for Animal Research (CEEA) of Maranhão State University, under protocol 45/2022, and the Ethics Committee for Animal Research of the Federal University of Maranhão (CIAEP: 02.0341.2019), under protocol number 23115.016180/2020-85.

2.3. Specimen identification, taxonomic classification and storage

The specimens were identified using taxonomic keys and the descriptions provided by Chao (2002), Munroe (2002), Smith-Vaniz (2002), Mol (2012), Menezes et al. (2015), Chao et al. (2021), Marceniuk et al. (2021b) and Carvalho-Filho (2023). The species nomenclature followed Fricke et al. (2024) and the taxonomic classification was based on Nelson et al. (2016). Tissue samples of the muscle, gills or fins were extracted from each specimen and stored in microtubes containing 99.5% ethanol for eventual molecular analysis. Whole specimens were designated as vouchers in 10% formalin or 70% ethanol and deposited in the tissue and DNA collection of the fauna of Maranhão (COFAUMA) of Maranhão State University (UEMA) and in the UEMA Ichthyological collection (CIUEMA).

2.4. Habitat classification and conservation status

The habitats occupied by each species were classified as marine, estuarine, marine-estuarine, or freshwater based on Carpenter (2002), Marceniuk et al. (2017) and Froese and Pauly (2023). The current conservation status of each species is also presented here, based on both the global criteria of the IUCN Red List (IUCN, 2023) and the national perspective available on the online platform Biodiversity Extinction Risk Assessment System − SALVE (ICMBio, 2024), which are based on IUCN criteria, but refer only to the Brazilian populations of each species.

2.5. Additional specimen examination and literature review

In addition to the specimens collected during the present study, specimens from the COFAUMA and the ichthyology collection of the Federal University of Maranhão (CPUFMA) were also examined to supplement the checklist. This material was collected from São Marcos Bay (sampling points P19–P34) and Curupu Bay (P35; Figure 1). Specific details on this material, including the 421 voucher numbers to 120 species already deposited in these collections, can be found in the Supplementary Material (Table S1).

Additionally, we consulted species records documented in previously published literature for the Golfão Maranhense, between 1987 and mid-November 2024, using the Google Scholar platform. The search terms included the following combinations: “ictiofauna Golfão Maranhense”, “peixes Ilha de São Luís” “ictiofauna Ilha de São Luís”, “exotic fish São Marcos Bay”, “exotic fish São Luís”, “estuarine fishes Golfão Maranhense”, “fish Golfão Maranhense”. For compiling these records, we used 24 studies that reported one or more species present in the area [see: Martins-Juras et al. (1987), Batista and Rego (1996), Castro (1997, 2001), Gomes et al. (2003), Pinheiro-Junior et al. (2005), Carvalho-Neta and Castro (2008), Piorski et al. (2009), Castro et al. (2010), Coelho et al. (2013), Silva Júnior et al. (2013), Guimarães et al. (2017a, b), Bonfim et al. (2017), Silva et al. (2018), Brito et al. (2019), Santana et al. (2019b), Lima et al. (2020), Fernandes et al. (2020), Aguiar et al. (2021), Trevisan et al. (2022), Vidigal et al. (2023), Franco et al. (2024) and Marques et al. (2024)]. The species Bairdiella ronchus (Cuvier 1830), Elops saurus Linnaeus 1766, Haemulon steindachneri (Jordan & Gilbert 1882), Menticirrhus americanus (Linnaeus 1758), Menticirrhus littoralis (Holbrook 1847) and Peprilus paru (Linnaeus 1758) published in Marques et al. (2024), were not considered because the records are incorrect for the study area, according to recent studies carried out in the western Atlantic, which include description of new species, revalidation of synonyms and updating of areas of occurrence (See discussion in section 4.2). Several species were not available in the consulted literature or collected in the present study, but the existence of vouchers from Golfão Maranhense was verified in the international network Global Biodiversity Information Facility (GBIF, 2024), specifically from datasets provided by Harvard University (2024a, b) and California Academy of Sciences (Catania and Fong, 2024a, b, c).

3. Results

The checklist of the bony fish of the Golfão Maranhense presented here includes a total of 163 species representing 107 genera, 57 families and 27 orders (Table 1, Figures 2, 3 and 4). The most representative order was Perciformes (n = 27 species), followed by the Acanthuriformes (n = 19), Carangiformes (n = 16), Clupeiformes (n = 15), and Siluriformes, with 15 species (Figure 5A). The most diverse families were the Sciaenidae (n = 19 species), Carangidae (n = 14), Ariidae (n = 11), Engraulidae and Gerreidae, with seven species each (Figure 5B).

Table 1
Fish species recorded from the Golfão Maranhense in the present study.

Taxonomy	Local common names	Life habitat	IUCN Status	ICMBio Status	Ref
ELOPIFORMES
Elopidae
Elops smithi McBride et al. (2010)	Ubarana	M, E	DD	LC	24
Megalopidae
Megalops atlanticus Valenciennes 1847	Camurupim	M, E, R	VU	VU	1; 2; 3; 4; 6; 8; 9; 11; 24
ALBULIFORMES
Albulidae
Albula vulpes (Linnaeus 1758)	Urubarana	M, E, R	NT	DD	1; 24
ANGUILLIFORMES
Muraenidae
Gymnothorax funebris Ranzani 1839	Moreia-verde	M, E, R	LC	DD	1; 3; 8; 9; 11; 20; 24
Gymnothorax ocellatus Agassiz 1831	Moreia	M, E, R	LC	DD	22
Gymnothorax vicinus (Castelnau 1855) (NR)	Moreia	M, R	LC	DD	-
Ophichthidae
Ophichthus cylindroideus (Ranzani 1839)	Jucutuca	M	LC	LC	1; 24
Myrophis punctatus Lütken 1852	Cobrinha da lama	M, E, R	LC	LC	22
CLUPEIFORMES
Pristigasteridae
Odontognathus mucronatus Lacepède 1800	Sardinha	M, E	LC	LC	8; 24
Pellona castelnaeana Valenciennes 1847	Sardinhão	FW, E	LC	LC	8; 15; 24
Pellona flavipinnis (Valenciennes 1837)	Sardinha-dourada	FW, E	LC	LC	1; 3; 24
Pellona harroweri (Fowler 1917)	Sardinha	M, E	LC	LC	8
Engraulidae
Anchoa spinifer (Valenciennes 1848)	Sardinha-vermelha	FW, M, E	LC	LC	1; 2; 3; 6; 7; 9; 11; 15; 24
Anchovia clupeoides (Swainson 1839)	Sardinha-gulelê	FW, M, E	LC	LC	1; 3; 4; 11; 24
Anchoviella elongata (Meek & Hildebrand 1923)	Sardinha	FW, M, E	LC	NE	18; 24
Anchoviella lepidentostole (Fowler 1911)	Sardinha	FW, M, E	LC	LC	8; 24
Cetengraulis edentulus (Cuvier 1829)	Sardinha boca-torta	M, E	LC	LC	1; 2; 3; 4; 6; 7; 8; 11; 15; 24
Lycengraulis grossidens (Spix & Agassiz 1829)	Sardinha-manjuba	FW, E	LC	LC	1; 3; 4; 24
Pterengraulis atherinoides (Linnaeus 1766)	Sardinha-de-gato	FW, E	LC	LC	1; 3; 4; 6; 9; 11; 24
Clupeidae
Lile piquitinga (Schreiner & Miranda Ribeiro 1903)	Pititinga	FW, E	LC	LC	1; 15; 24
Opisthonema oglinum (Lesueur 1818)	Cupeú	M, R	LC	LC	1; 2; 3; 4; 24
Rhinosardinia amazonica (Steindachner 1879)	Sardinha-de-serra	FW, E	LC	LC	1; 3; 4; 24
Sardinella brasiliensis (Steindachner 1879)	Sardinha	M, E	DD	LC	6; 7; 9; 11
SILURIFORMES
Ariidae
Bagre bagre (Linnaeus 1766)	Bandeirado	M, E	LC	NT	1; 2; 3; 4; 7; 8; 9; 11; 15; 19; 24
Cathorops agassizii (Eigenmann & Eigenmann 1888)	Uriacica-branco	FW, E	NE	LC	7; 17; 24
Cathorops arenatus (Valenciennes 1840)	Uriacica-amarelo	M, E	LC	LC	17; 24
Cathorops spixii (Agassiz 1829)	Uriacica-amarelo	M, E	NE	LC	1; 2; 3; 4; 7; 11; 15; 17; 24
Notarius grandicassis (Valenciennes 1840)	Cambéu	FW, M, E	LC	LC	1; 3; 8; 24
Notarius rugispinis (Valenciennes 1840)	Jurupiranga	M, E	LC	LC	1; 2; 3; 7; 9; 11; 15; 24
Notarius quadriscutis (Valenciennes 1840)	Cangatã	M, E	LC	LC	1; 3; 4; 8; 11; 15; 24
Sciades couma (Valenciennes 1840)	Bagre-catinga	E	LC	DD	1; 3; 24
Sciades herzbergii (Bloch 1794)	Guribu	E	LC	LC	1; 2; 3; 4; 5; 6; 7; 9; 11; 24
Sciades parkeri (Traill 1832)	Gurijuba	M	VU	VU	1; 3
Sciades proops (Valenciennes 1840)	Uritinga	M	NE	DD	1; 3; 7; 9; 24
Aspredinidae
Aspredo aspredo (Linnaeus 1758)	Viola	M, E	LC	LC	1; 3; 18; 24
Aspredinichthys filamentosus (Valenciennes 1840)	Viola	M, E	LC	LC	18; 24
Aspredinichthys tibicen (Valenciennes 1840)	Viola	M, E	LC	LC	1; 2; 3; 4; 7; 9; 11; 15; 24
Auchenipteridae
Pseudauchenipterus nodosus (Bloch 1794)	Papista	FW, E	NE	LC	1; 2; 3; 4; 5; 6; 7; 9; 11; 24
AULOPIFORMES
Synodontidae
Synodus foetens (Linnaeus 1766)	Peixe-lagarto	M, E, R	LC	LC	1; 3; 24
BATRACHOIDIFORMES
Batrachoididae
Amphichthys cryptocentrus (Valenciennes 1837)	Pacamão	M, E, R	LC	LC	10
Batrachoides surinamensis (Bloch & Schneider 1801)	Pacamão	M, E	LC	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 24
Thalassophryne nattereri Steindachner 1876	Niquim	M, E	LC	LC	1; 3; 11; 24
GOBIIFORMES
Eleotridae
Butis koilomatodon (Bleeker 1849) +	Muré	FW, M, E	LC	NE	12; 14; 20
Guavina guavina (Valenciennes 1837)	Muré	FW, M, E	LC	LC	1; 3; 4; 24
Oxudercidae
Ctenogobius boleosoma (Jordan & Gilbert 1882)	Muré	FW, M, E, R	LC	LC	13
Ctenogobius smaragdus (Valenciennes 1837)	Muré	FW, M, E	LC	LC	16; 24
Gobioides broussonnetii Lacepède 1800	Muré	FW, M, E	LC	DD	18; 24
Gobionellus oceanicus (Pallas 1770)	Muré	FW, M, E	LC	LC	1; 24
Trypauchen vagina (Bloch & Schneider 1801) +	-	M, E	LC	NE	21
Gobiidae
Bathygobius soporator (Valenciennes 1837)	Muré	FW, M, E	LC	LC	1; 20; 24
Pomacentridae
Abudefduf saxatilis (Linnaeus 1758)	Sargento	M, R	LC	LC	20
MUGILIFORMES
Mugilidae
Mugil brevirostris Miranda Ribeiro 1915	Tainha-pitiu	M, E	NE	DD	23
Mugil curema Valenciennes 1836	Tainha-sajuba	M, E	LC	DD	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 24
Mugil incilis Hancock 1830	Tainha-urixoca	M, E	LC	DD	1; 2; 3; 4; 6; 7; 9; 11; 15; 20; 24
Mugil liza Valenciennes 1836	Tainha-curimã	M, E	DD	NT	1; 3; 11; 24
Mugil rubrioculus Harrison et al. (2007) (NR)	Tainha-sajuba	M, E	LC	DD	-
BLENNIIFORMES
Blenniidae
Omobranchus sewalli (Fowler 1931) +	-	M, E	LC	NE	20
Scartella cristata (Linnaeus 1758)	-	M, R	LC	LC	20
GOBIESOCIFORMES
Gobiesocidae
Gobiesox cf. barbatulus Starks 1913	-	M	LC	LC	20
CICHLIFORMES
Cichlidae
Oreochromis niloticus (Linnaeus 1758) +	Tilápia	FW, E	LC	NE	23; 24
ATHERINIFORMES
Atherinopsidae
Atherinella brasiliensis (Quoy & Gaimard 1825)	João-duro	M, E	LC	LC	1; 24
BELONIFORMES
Exocoetidae
Cheilopogon melanurus (Valenciennes 1847)	Voador	M	LC	LC	1
Hemiramphidae
Hyporhamphus roberti (Valenciennes 1847)	Agulhinha	M, E	LC	LC	25
Belonidae
Strongylura marina (Walbaum 1792)	Peixe-agulha	FW, M, E, R	LC	LC	1; 3; 4; 6; 7; 8; 11; 24
Strongylura timucu (Walbaum 1792)	Peixe-agulha	FW, M, E, R	LC	LC	1; 3; 9; 24
CYPRINODONTIFORMES
Anablepidae
Anableps anableps (Linnaeus 1758)	Tralhoto	FW, E	LC	LC	1; 2; 3; 4; 5; 6; 7; 9; 11; 24
Anableps microlepis Müller & Troschel 1844	Tralhoto	FW, E	LC	LC	1; 3
CARANGIFORMES
Rachycentridae
Rachycentron canadum (Linnaeus 1766) (NR)	Beijupirá	M, R	LC	LC	-
Echeneidae
Echeneis naucrates Linnaeus 1758	Rêmora	M, E, R	LC	LC	1; 3; 4; 8; 24
Carangidae
Caranx crysos (Mitchill 1815)	Garajuba	M, E, R	LC	LC	1; 3; 4; 8; 24
Caranx hippos (Linnaeus 1766)	Xaréu	M, E	LC	LC	1; 3; 4; 6; 24
Caranx latus Agassiz 1831	Xaréu	M, E, R	LC	LC	1; 3; 4; 9; 11; 15; 24
Chloroscombrus chrysurus (Linnaeus 1766)	Favinha	M, E	LC	LC	1; 3; 4; 8; 11; 24
Hemicaranx amblyrhynchus (Cuvier 1833)	Xixarro	M	LC	LC	1; 3; 4; 24
Oligoplites palometa (Cuvier 1832)	Tibiro	M, E	LC	LC	1; 3; 4; 11; 15; 24
Oligoplites saliens (Bloch 1793)	Tibiro	M, E	LC	LC	19
Oligoplites saurus (Bloch & Schneider 1801)	Tibiro branco	M, E, R	LC	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 24
Selene brownii (Cuvier 1816)	Peixe-galo	M, E	LC	LC	24; 26
Selene setapinnis (Mitchill 1815)	Peixe-galo	M, E	LC	LC	24
Selene vomer (Linnaeus 1758)	Peixe-galo	M, E	LC	LC	1; 2; 3; 4; 8; 11; 24
Trachinotus carolinus (Linnaeus 1766)	Pampo / Canguira	M, E	LC	LC	1
Trachinotus cayennensis Cuvier 1832	Pampo / Canguira	M, E	LC	LC	1; 24
Trachinotus falcatus (Linnaeus 1758)	Pampo / Canguira	M, E, R	LC	LC	1; 3; 4; 11; 24
ISTIOPHORIFORMES
Sphyraenidae
Sphyraena guachancho Cuvier 1829 (NR)	Barracuda	M, E	LC	LC	-
PLEURONECTIFORMES
Paralichthyidae
Paralichthys brasiliensis (Ranzani 1842)	Linguado / Solha	M, E	LC	LC	7; 24
Citharichthys spilopterus Günther 1862	Solha-urumaçara	M, E	LC	LC	1; 3; 4; 24
Etropus crossotus Jordan & Gilbert 1882	Solha-urumaçara	M, E	LC	LC	1; 3; 4; 24
Achiridae
Achirus achirus (Linnaeus 1758)	Linguado / Solha	FW, E	LC	LC	1; 2; 3; 4; 6; 7; 9; 24
Achirus declivis Chabanaud 1940 (NR)	Linguado / Solha	FW, E	LC	LC	-
Achirus lineatus (Linnaeus 1758)	Linguado / Solha	FW, E	LC	LC	8; 11; 15; 24
Apionichthys dumerili Kaup 1858	Solha	FW, M, E	LC	LC	24; 27
Trinectes paulistanus (Miranda Ribeiro 1915)	Linguado / Solha	FW, E	LC	LC	1; 24
Cynoglossidae
Symphurus plagusia (Bloch & Schneider 1801)	Linguado	M, E	LC	LC	1; 3; 9; 11; 24
Symphurus tessellatus (Quoy & Gaimard 1824) (NR)	Linguado	M, E	LC	LC	-
SYNGNATHIFORMES
Syngnathidae
Hippocampus reidi Ginsburg 1933	Cavalo-marinho	M, E, R	NT	VU	24
Microphis lineatus (Kaup 1856) (NR)	Cachimbo	FW, M, E	NE	NE	-
Dactylopteridae
Dactylopterus volitans (Linnaeus 1758) (NR)	Coió	M, E, R	LC	LC	-
SCOMBRIFORMES
Trichiuridae
Trichiurus lepturus Linnaeus 1758	Guaravira	M, E	LC	LC	1; 3; 4; 6; 8; 9; 11; 24
Scombridae
Euthynnus alletteratus (Rafinesque 1810) (NR)	Bonito-cachorro	M, E	LC	LC	-
Scomberomorus brasiliensis Collette et al. (1978)	Peixe-serra	M, R	LC	NT	8
Scomberomorus cavalla (Cuvier 1829)	Cavala	M	LC	NT	24; 28
Scomberomorus regalis (Bloch 1793)	Peixe-serra	M, R	LC	LC	1; 3
Stromateidae
Peprilus crenulatus Cuvier 1829	Arriba-saia	M, E	NE	DD	19
PERCIFORMES
Centropomidae
Centropomus parallelus Poey 1860	Camurim-branco	FW, M, E	LC	DD	1; 2; 3; 4; 6; 7; 8; 9; 11; 15; 24
Centropomus pectinatus Poey 1860	Camurim	FW, M, E	LC	DD	11; 15
Centropomus undecimalis (Bloch 1792)	Camurim-preto	FW, M, E, R	LC	DD	1; 3; 4; 5; 24
Gerreidae
Diapterus auratus Ranzani 1842	Carapeba	FW, M, E	LC	LC	1; 3; 4; 8; 9; 24
Diapterus rhombeus (Cuvier 1829)	Carapeba	FW, M, E	LC	LC	1; 3; 4; 5; 6; 7; 11; 15; 24
Eucinostomus argenteus Baird & Girard 1855	Escrivão	M, E, R	LC	LC	1; 3; 4; 6; 24
Eucinostomus gula (Quoy & Gaimard 1824)	Escrivão	M, E, R	LC	LC	1; 3; 4; 24
Eucinostomus harengulus Goode & Bean 1879 (NR)	Escrivão	M, E	LC	LC	-
Eucinostomus melanopterus (Bleeker 1863)	Escrivão	M, E	LC	LC	1; 3; 4; 24
Eugerres brasilianus (Cuvier 1830)	Escrivão	M, E	LC	LC	1; 3
Serranidae
Diplectrum radiale (Quoy & Gaimard 1824)	Papa-terra	M, E	LC	LC	1; 3; 4; 24
Epinephelus itajara (Lichtenstein 1822)	Mero	M, E, R	VU	CR	1; 2; 3; 4; 6; 9; 11; 20; 24
Mycteroperca bonaci (Poey 1860)	Sirigado	M, R	NT	VU	1; 24
Rypticus randalli Courtenay 1967	Peixe-sabão	M	LC	LC	1; 2; 3; 4; 6; 24
Pomatomidae
Pomatomus saltatrix (Linnaeus 1766) (NR)	Enchova	M, E	VU	LC	-
Pomacanthidae
Pomacanthus paru (Bloch 1787)	Frade	M, R	LC	DD	1; 24
Haemulidae
Anisotremus virginicus (Linnaeus 1758) (NR)	Frade	M, R	LC	LC	-
Conodon nobilis (Linnaeus 1758)	Jiquiri-listrado	M	LC	LC	1; 8
Genyatremus luteus (Bloch 1790)	Peixe-pedra	M, E	DD	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 20; 24
Haemulon atlanticus Carvalho et al. (2020) (NR)	Cororoca	M, E	LC	LC	-
Haemulopsis corvinaeformis (Steindachner 1868)	Jiquiri-branco	M, E	LC	LC	1; 3; 4; 11; 24
Lutjanidae
Lutjanus campechanus (Poey 1860) (NR)	Pargo	M, R	VU	NE	-
Lutjanus cyanopterus (Cuvier 1828)	Caranha	M, E, R	VU	VU	24; 29
Lutjanus jocu (Bloch & Schneider 1801)	Carapitanga	M, E, R	DD	NT	1; 3; 4; 11; 15; 20; 24
Lutjanus synagris (Linnaeus 1758)	Ariacó	M, R	NT	DD	1; 3; 4; 6; 24
Polynemidae
Polydactylus oligodon (Günther 1860)	Barbudo	M, E	LC	LC	1; 3; 4; 24
Polydactylus virginicus (Linnaeus 1758)	Barbudo	M, E	LC	LC	1; 3; 6; 9; 11; 24
SCORPAENIFORMES
Scorpaenidae
Scorpaena plumieri Bloch 1789	Mangangá	M, R	LC	LC	1; 3; 4; 24
Triglidae
Prionotus beanii Goode 1896	Cabrinha	M, E	LC	NE	8
Prionotus punctatus (Bloch 1793)	Cabrinha	M, E	LC	LC	24
MORONIFORMES
Ephippidae
Chaetodipterus faber (Broussonet 1782)	Paru	M, E, R	LC	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 24
ACANTHURIFORMES
Sciaenidae
Bairdiella goeldi Marceniuk et al. (2019) (NR)	Cororoca	M, E	LC	LC	-
Cynoscion acoupa (Lacepède 1801)	Pescada-amarela	FW, M, E	VU	DD	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 24
Cynoscion jamaicensis (Vaillant & Bocourt 1883)	Pescadinha / Goete	M, E	LC	LC	15; 24
Cynoscion leiarchus (Cuvier 1830)	Pescada-branca	M, E	LC	LC	1; 2; 3; 4; 6; 7; 8; 9; 11; 15; 24
Cynoscion microlepidotus (Cuvier 1830)	Corvina-uçu	M, E	LC	LC	1; 2; 3; 4; 6; 9; 11; 15; 24
Cynoscion steindachneri (Jordan 1889)	Juruapara	FW, M, E	LC	LC	1; 3; 4; 6; 7; 9; 24
Isopisthus parvipinnis (Cuvier 1830)	Curvitinga	M, E	LC	LC	1; 3; 4; 6; 9; 24
Larimus breviceps Cuvier 1830	Pirucaia	M, E	LC	LC	8; 24
Macrodon ancylodon (Bloch & Schneider 1801)	Pescada-gó	M, E	LC	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 24
Menticirrhus cuiaranensis Marceniuk et al. (2020) (NR)	Boca-de-rato	M, E	NE	LC	-
Menticirrhus martinicensis (Cuvier 1830) (NR)	Boca-de-rato	M, E	NE	LC	-
Micropogonias furnieri (Desmarest 1823)	Cururuca	M, E	LC	LC	1; 2; 3; 4; 6; 9; 11; 15; 24
Nebris microps Cuvier 1830	Amor-sem-olho	M, E	LC	LC	1; 3; 9; 11; 15; 19; 24
Stellifer brasiliensis (Schultz 1945)	Cabeçudo	M, E	LC	LC	1; 3; 6; 9; 24
Stellifer microps (Steindachner 1864)	Cabeçudo	M, E	LC	LC	8; 24
Stellifer naso (Jordan 1889)	Cabeçudo-preto	M, E	LC	LC	1; 2; 3; 4; 6; 7; 9; 11; 24
Stellifer punctatissimus (Meek & Hildebrand 1925)	Cabeçudo-preto	M, E	LC	LC	6; 24
Stellifer rastrifer (Jordan 1889)	Cabeçudo-vermelho	M, E	LC	LC	1; 2; 3; 6; 7; 8; 11; 24
Stellifer stellifer (Bloch 1790)	Cabeçudo-vermelho	M, E	DD	LC	1; 3; 24
SPARIFORMES
Lobotidae
Lobotes surinamensis (Bloch 1790)	Crauaçu	M, E	LC	LC	1; 3; 6; 8; 15; 24
Sparidae
Archosargus probatocephalus (Walbaum 1792)	Sargo	M, E, R	LC	LC	1; 3; 4; 24
LOPHIIFORMES
Ogcocephalidae
Ogcocephalus vespertilio (Linnaeus 1758)	Bacacuá	M, E, R	LC	LC	1; 3; 4; 8; 15; 24
TETRAODONTIFORMES
Ostraciidae
Lactophrys trigonus (Linnaeus 1758) (NR)	Baiacu-cofre	M, R	LC	LC	-
Monacanthidae
Aluterus monoceros (Linnaeus 1758)	Cangulo	M, E, R	LC	DD	1; 3; 15; 24
Tetraodontidae
Lagocephalus laevigatus (Linnaeus 1766)	Baiacu-guará	M, E	LC	LC	1; 3; 8; 24
Sphoeroides greeleyi Gilbert 1900	Baiacu-areia	M, E, R	LC	LC	1; 3; 18; 24
Sphoeroides psittacus (Bloch & Schneider 1801)	Baiacu-açu	FW, M, E	LC	LC	1; 2; 3; 4; 5; 6; 7; 8; 9; 11; 15; 20; 24
Sphoeroides testudineus (Linnaeus 1758)	Baiacu-pininga	M, E, R	LC	LC	1; 3; 4; 6; 8; 11; 20; 24
Diodontidae
Chilomycterus antillarum Jordan & Rutter 1897	Baiacu-de-espinho	M, E, R	LC	LC	1; 3; 24

• New records are designated “^NR”, and exotic species as “+”, while the references (Ref.) indicate previous studies conducted in the Golfão Maranhense. Habitats: E = Estuarine, FW = Freshwater, M = Marine, R = Reef-dwelling. Conservation status (IUCN and ICMBio): NE = Not Evaluated; DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; CR = Critically Endangered. 1 - Martins-Juras et al. (1987); 2 - Batista and Rego (1996); 3 - Castro (1997); 4 - Castro (2001); 5 - Gomes et al. (2003); 6 - Pinheiro-Junior et al. (2005); 7 - Carvalho-Neta and Castro (2008); 8 - Piorski et al. (2009); 9 - Castro et al. (2010); 10 - Coelho et al. (2013); 11 - Silva Júnior et al. (2013); 12 - Guimarães et al. (2017a); 13- Guimarães et al. (2017b); 14 - Bonfim et al. (2017); 15 - Silva et al. (2018); 16 - Brito et al. (2019); 17 - Santana et al. (2019b); 18 - Lima et al. (2020); 19 - Fernandes et al. (2020); 20 - Aguiar et al. (2021); 21 - Trevisan et al. (2022); 22 - Vidigal et al. (2023); 23 - Franco et al. (2024); 24 - Marques et al. (2024); 25 - Catania and Fong (2024a); 26 - Harvard University (2024a); 27 - Catania and Fong (2024b); 28 - Harvard University (2024b); 29 - Catania and Fong (2024c).

Figure 2
Photographs of 16 of the fish species recorded in the Golfão Maranhense in the present study. (A) Elops smithi, CFMPE753; (B) Gymnothorax vicinus, CFMPE666; (C) Ophichthus cylindroideus, CFMPE1024; (D) Odontognathus mucronatus, CFMPE1114; (E) Anchoa spinifer, CFMPE1012; (F) Cetengraulis edentulus, CFMPE1040; (G) Pterengraulis atherinoides, CFMPE1010; (H) Notarius rugispinis, CFMPE1028; (I) Cathorops agassizii, CFMPE1037; (J) Cathorops arenatus, CPUFMA492; (K) Aspredinichthys filamentosus, CFMPE653; (L) Amphichthys cryptocentrus, CFMPE888; (M) Mugil brevirostris, CFMPE865; (N) Oreochromis niloticus, CIUEMA0370; (O) Hyporhamphus roberti, CFMPE809; (P) Echeneis naucrates, CFMPE895. Scale bars = 30 mm.

Figure 3
Photographs of 12 of the fish species recorded in the Golfão Maranhense in the present study. (A) Selene brownii, CFMPE916; (B) Selene setapinnis, CFMPE1038; (C) Sphyraena guachancho, CFMPE901; (D) Apionichthys dumerili, CFMPE1019; (E) Achirus declivis, CFMPE939; (F) Trinectes paulistanus, CFMPE832; (G) Centropomus undecimalis, CIUEMA0377; (H) Dactylopterus volitans, CFMPE959; (I) Hippocampus reidi, CFMPE930; (J) Peprilus crenulatus, CFMPE964; (K) Anisotremus virginicus, CFMPE634; (L) Lutjanus synagris, CFMPE805. Scale bars = 30 mm.

Figure 4
Photographs of 13 of the fish species recorded in the Golfão Maranhense in the present study. (A) Polydactylus virginicus, CFMPE904; (B) Prionotus punctatus, CFMPE966; (C) Cynoscion leiarchus, CFMPE199; (D) Macrodon ancylodon, CFMPE955; (E) Menticirrhus cuiaranensis, CFMPE907; (F) Menticirrhus martinicensis, CFMPE670; (G) Stellifer stellifer, CFMPE718; (H) Stellifer rastrifer, CFMPE159; (I) Stellifer naso, CFMPE881; (J) Lactophrys trigonus, CFMPE925; (K) Sphoeroides psittacus, CFMPE798; (L) Sphoeroides testudineus, CFMPE886; (M) Chilomycterus antillarum, CFMPE819. Scale bars = 30 mm.

Figure 5
The number of fish species registered in the Golfão Maranhense in the present study per taxonomic (A) order and (B) family. The families are color-coded by order.

Based on the IUCN classification (IUCN, 2023), the conservation status of seven of the species recorded here is Vulnerable (VU): Megalops atlanticus Valenciennes 1847, Sciades parkeri (Traill 1832), Epinephelus itajara (Lichtenstein 1822), Pomatomus saltatrix (Linnaeus 1766), Lutjanus campechanus (Poey 1860), Lutjanus cyanopterus (Cuvier 1828) and Cynoscion acoupa (Lacepède 1801). The official Brazilian list (ICMBio, 2024) also cites three of these species – Megalops atlanticus, S. parkeri, and L. cyanopterus – as VU, and one (E. itajara) as Critically Endangered (CR), while also listing two other species – Hippocampus reidi Ginsburg 1933 and Mycteroperca bonaci (Poey 1860) – as VU. None of the species listed in the present study are currently classified as either regionally extinct or extinct in the wild (EW).

4. Discussion

4.1. Species richness

The 163 species recorded here represent the largest inventory of the teleost fish fauna of the Golfão Maranhense compiled to date, and 39,32% more than the 117 species recorded by Martins-Juras et al. (1987) in a survey of São Luís Island. More recent studies have recorded even smaller numbers of species, such as the 75 species registered by Castro (2001) in the estuary of the Paciência River, in São José Bay, the 32 species found by Carvalho Neta and Castro (2008) at Caranguejos Island, in São Marcos Bay, and the 54 species recorded by Silva et al. (2018) in the port area of São Marcos Bay. In a recent study, Marques et al. (2024) carried out an inventory of the diversity of the Golfão Maranhense, based on literature data. This inventory represented 74.23% of the species recorded in our study, based on field collections, voucher analysis and literature data. These species also reported by Marques et al. (2024) can be found in Table 1, column 6 (Ref 24).

The fish species richness recorded here is also greater than the values reported in previous surveys of other estuaries in northern and northeastern Brazil. These estuaries include that of the Sergipe River, in Sergipe state, where Alcântara (2006) recorded 136 species, and the Curu River, in Ceará, where Basílio et al. (2009) found 61 species. In Pará state, which is adjacent to Maranhão, the study areas include the estuaries of São Caetano de Odivelas and Vigia, where Barros et al. (2011) recorded 58 species, and that of the Caeté River, where Marceniuk et al. (2017) found 120 species.

The predominance of the order Perciformes recorded here is typical of many previous inventories of marine and estuarine fish on the Brazilian coast (e.g. Vilar et al., 2011; Barrella et al., 2014; Silva et al., 2018; Vendel et al., 2022). Perciforms tend to be relatively tolerant of variations in salinity (Camargo and Isaac, 2003; Shaha et al., 2022), which may account for their predominance in the Golfão Maranhense, in which salinity varies considerably – from 21‰ to 30‰ – between the rainy and dry seasons (Azevedo et al., 2008). This variation is due to the input from the discharge of more than 20 rivers, including the two largest rivers of the state of Maranhão, the Itapecuru and the Mearim.

The predominance of the families Sciaenidae, Carangidae and Ariidae is also typical of the estuaries of the northern coast of Brazil, in particular those under the influence of the Amazon River plume, with high annual precipitation, an ample tidal range, and high levels of freshwater and nutrient input (Barletta and Blaber, 2007; Vilar et al., 2013; Blaber and Barletta, 2016). Previous studies have identified a similar predominance of these families all along the northern coast of South America, between Venezuela and Maranhão (Manjarrés, 2004; Lasso and Señaris, 2008; Lasso and Sánchez-Duarte, 2011; Silva et al., 2018), for example, and in other estuaries of northern and northeastern Brazil (Lira and Teixeira, 2008; Sanders et al., 2011; Chao et al., 2015; Dantas et al., 2016; Marceniuk et al., 2017). The predominance of these families may be associated with certain characteristics of estuarine environments, such as the composition of the substrates, which are predominantly sandy and muddy, and would thus favor generalist species that prefer these types of habitat (Souza Filho, 2005; Silva et al., 2018).

4.2. New records and taxonomic status

Overall, 18 of the 163 species recorded here are listed for the first time for the Golfão Maranhense. These species are Gymnothorax vicinus (Figure 2B), Mugil rubrioculus, Rachycentron canadum, Sphyraena guachancho (Figure 3C), Achirus declivis (Figure 3E), Symphurus tessellatus, Microphis lineatus, Dactylopterus volitans (Figure 3H), Euthynnus alletteratus, Eucinostomus harengulus, Pomatomus saltatrix, Anisotremus virginicus (Figure 3K), Haemulon atlanticus, Lutjanus campechanus, Bairdiella goeldi, Menticirrhus cuiaranensis (Figure 4E), Menticirrhus martinicensis (Figure 4F) and Lactophrys trigonus (Figure 4J). Among the 18 new occurrences for the Golfão Maranhense, only three species are known from other sites of Maranhão state. These species are G. vicinus, R. canadum and A. virginicus for the Parcel Manuel Luiz Marine State Park (Rocha and Rosa, 2001).

All of these species are known to occur on the northern coast of Brazil (Marceniuk et al., 2021a, b), which implies that certain specific factors may have contributed to the lack of previous records from the Golfão Maranhense. These factors include taxonomic questions, such as the description of new species and the revalidation of synonyms (Harrison et al., 2007; Marceniuk 2007; McBride et al., 2010), as well as sampling effects, including the methods used to collect specimens, limited geographic scope of previous inventories and the seasonal patterns in the occurrence and abundance of the different species (Neira et al., 1992; Batista and Rêgo, 1996; Barletta-Bergan et al., 2002; Copatti et al., 2009; Raju et al., 2016; Brito et al., 2019). Silva et al. (2018) also reported that shipping traffic and other port activities may have impacted the diversity of the fish found in the region.

Since the early 2000s, a number of studies have focused on the diversity of the fish fauna that inhabits the northern and northeastern coasts of Brazil (e.g. Menezes et al., 2015; Chao et al., 2021; Araujo et al., 2023; Marceniuk et al., 2021a, 2023). Some of these studies have included the description of new species, the revalidation of synonyms, and new classifications, as well as the updating of areas of occurrence, all of which have contributed to substantial advances in the understanding of the true diversity of the fish fauna of these areas.

Marceniuk (2007), for example, revalidated Cathorops arenatus (Valenciennes 1840) and Cathorops agassizii (Eigenmann & Eigenmann 1888), which had been identified previously as synonyms of Cathorops spixii (Agassiz 1829). These species occur along the northern coast of Brazil, where they inhabit estuaries and rivers, with C. agassizii being more common in freshwater environments (Marceniuk and van der Sleen, 2017). When analyzing vouchers of a local collection – CPUFMA – Santana et al. (2019b) corrected the identification of specimens from the Golfão Maranhense designated previously as Notarius bonillai (Miles, 1945), which they assigned to C. arenatus. Acero and Betancur-R (2006) and Fricke et al. (2024) have identified N. bonillai as a freshwater catfish (Ariidae) that is endemic to the rivers that drain into the Caribbean coast of Colombia. McBride et al. (2010) described Elops smithi from the western Atlantic, extending from the Gulf of Mexico to the southern coast of Brazil, while Elops saurus Linnaeus 1766 is known to range from southern Massachusetts to Florida in the United States, including the Gulf of Mexico.

The distribution of the different mullets of the genus Mugil Linnaeus 1758 in the western Atlantic has been the subject of constant debate over the past 30 years (Thomson, 1997; Nirchio et al., 2003; Fraga et al., 2007; Castro et al., 2007; Heras et al., 2009; Menezes et al., 2010). Given these taxonomic uncertainties, Menezes et al. (2015) conducted a comprehensive review of the species from the Caribbean and western South Atlantic, based on morphological, cytogenetic and molecular data. This study identified Mugil brevirostris Miranda-Ribeiro 1915 as valid nominal species for the Western Atlantic and specimens previously identified as Mugil gaimardianus Desmarest 1831 were reidentified as M. brevirostris, based on the fact that the tip of the pectoral fin reached or extended slightly beyond the vertical line running through the base of the first spine of the first dorsal fin in the specimens examined. Harrison et al. (2007) reported that Mugil gaimardianus Desmarest 1831 is an invalid species name, due to its ambiguous description and the type specimen is apparently lost.

Marceniuk et al. (2016) reviewed the diversity of the species of the genus Peprilus Cuvier 1829, based on morphological and DNA data, revalidating the species Peprilus crenulatus Cuvier 1829 and Peprilus xanthurus (Quoy & Gaimard 1825) from the western Atlantic, which range from the Caribbean to the southern coast of South America, and Peprilus paru (Linnaeus 1758) does not occur in the southern coast of South America, and its exact distribution in the Caribbean is unclear, requiring further investigation. However, there is no record of P. xanthurus from either the Golfão Maranhense or other parts of the coast of Maranhão (Rosa et al., 2023), which reinforces the need for the reassessment of the vouchers identified as P. paru, as well as more extensive sampling effort in the region.

Marceniuk et al. (2019) determined that Bairdiella ronchus (Cuvier 1830), whose occurrence on the Brazilian coast is reported widely, is in fact restricted to the Caribbean region, and described the new species Bairdiella goeldi, which is distributed from the northern coast of Brazil to the coast of the state of Santa Catarina, in southern Brazil. Marceniuk et al. (2020) also demonstrated that both Menticirrhus americanus (Linnaeus 1758) and Menticirrhus littoralis (Holbrook 1847), which are amply distributed in the western Atlantic, represent species complexes, and described the new species Menticirrhus cuiaranensis. This study concluded that M. americanus occurs on the east coast of the United States and in the Gulf of Mexico, while Menticirrhus martinicensis (Cuvier 1830), a revalidated junior synonym, ranges from the Caribbean to Argentina. Like M. americanus, M. littoralis has a more limited distribution, while M. cuiaranensis is the species found off the northern and eastern coasts of Brazil. Carvalho et al. (2020) also redescribed Haemulon steindachneri (Jordan & Gilbert 1882) and restricted its distribution to the Pacific coast, between the Gulf of California and Peru, and describe a new species, Haemulon atlanticus, that occurs on the Atlantic coast, between southern Costa Rica and Santa Catarina, in southern Brazil.

4.3. Exotic species

The Itaqui Port Complex located on the central portion of the Golfão Maranhense is one of the largest shipping terminals on the Brazilian coast and is a major driver of the economic and social development of Maranhão State (Silva et al., 2018; Almeida et al., 2021). However, the development of the shipping sector in this region has led to increasing concerns with regard to its negative impacts on the native fauna, especially through the introduction of xenobiotics and exotic species (Castro et al., 2019; Noleto et al., 2021; Trevisan et al., 2022).

In addition to the Nile tilapia Oreochromis niloticus (Franco et al., 2024), three fish species native to the Indo-Pacific Ocean have been detected in the Golfão Maranhense. These species are the mudsleeper, Butis koilomatodon (Guimarães et al., 2017a; Bonfim et al., 2017; Aguiar et al., 2021), the burrowing goby Trypauchen vagina (Trevisan et al., 2022), and the muzzled blenny Omobranchus sewalli (in some references as Omobranchus punctatus (Valenciennes 1836), e.g. Lasso-Alcalá et al., 2011; Aguiar et al., 2021). Specimens of three of these species – B. koilomatodon, T. vagina, and O. niloticus – were collected during the present study. These species have likely been introduced from two principal sources – the disposal of ballast water by ships arriving from foreign ports and the escape of specimens from the ornamental fish trade or from the fish farms located along the rivers that discharge into the Golfão Maranhense (Fernandes et al., 2021; Trevisan et al., 2022). The introduction of exotic fish and the loss of natural habitats together pose a major threat to aquatic biodiversity worldwide (Wonham et al., 2000; Schofield et al., 2011; Garcia et al., 2021; Khan et al., 2022). In Brazil, a large number of studies have reported the impacts provoked by the presence of exotic species in aquatic environments, including eutrophication, the loss of biodiversity, shifts in community structure, the homogenization of the fauna, and changes in fishing systems (e.g. Figueredo and Giani, 2005; Attayde et al., 2011; Novaes and Carvalho, 2011; Pelicice et al., 2013; Fragoso-Moura et al., 2016; Bezerra et al., 2018; Catelani et al., 2020; Charvet et al., 2021; Vitule et al., 2021). These problems are accentuated in environments such as the Golfão Maranhense, which is a major nursery area for the juveniles of many migratory fish, as well as providing habitats and feeding grounds for both resident and transient species (Castro, 1997; Blaber and Barletta, 2016).

The Golfão Maranhense is also an area of special socioeconomic importance, in particular for the local artisanal fisheries, given the presence of a number of fish species of commercial interest. This means that the collection of reliable data on the composition of the local fish fauna is not only an academic exercise, but also provides a fundamentally important baseline for the assessment of the possible impacts of changes in the fish community and the development of effective measures for the control of fisheries and the protection of fish stocks.

The checklist presented here was enriched considerably by the analysis of specimens deposited in zoological collections, which highlights the importance of transferring the biological material collected in the field to these collections. This source of information enhances the ability of conservationists to implement effective measures for the preservation of the biological diversity of a specific location (Pimenta et al., 2017), as well as providing essential input for taxonomic analyses. The present study organizes, updates, and expands the limited data available about the marine and estuarine fish fauna of the Golfão Maranhense. Despite this advance, further research is needed, particularly in voucher specimens, to reinforce the database available about the fish species of this region. There is also a clear need for the continuous monitoring of exotic, fishery target, and threatened species to ensure a better understanding of the long-term consequences of the ongoing human impacts on the local fish fauna, and the eventual socioeconomic consequences of these impacts.

Supplementary Material

Supplementary material accompanies this paper.

Table S1.

List of the fish species identified in the present study, based on the analysis of the specimens collected in the Golfão Maranhense and the vouchers deposited in the ichthyological collections of Maranhão State University (UEMA) and the Federal University of Maranhão (UFMA). The catalog numbers of the specimens are also provided. PS = Present Study (species collected during the present study), ME = Material Examined, CFMPE = vouchers deposited in the COFAUMA collection, CIUEMA = UEMA ichthyological collection, CPUFMA = UFMA ichthyological collection.

This material is available as part of the online article from https://doi.org/10.1590/1519-6984.286625

Acknowledgements

This study was supported by the Brazilian Coordination for Higher Education Personnel Training - Brazil (CAPES), through a graduate stipend granted to T. C. Santana (process number 88887.687222/2022-00), and the Maranhão State Foundation for Scientific Research and Development (FAPEMA), through a project submitted by L. F. Carvalho-Costa, finance code UNIVERSAL-00902/19.

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Publication Dates

• Publication in this collection
  10 Mar 2025
• Date of issue
  2025

History

• Received
  21 May 2024
• Accepted
  16 Dec 2024