High fish richness associated with the southernmost Atlantic coral reef and adjacencies: comments on conservation and management for sustainable use

Ana Clara S. Athayde¹, Fernanda A. Rolim¹, Otto B. F. Gadig², Caio R. Pimentel¹, Rafael R. Munhoz¹, Maisha Gragnolati¹, Fernando Z. Gibran³, Leonardo M. Neves⁴, Domingos Garrone-Neto⁵, Gabriel R. S. Souza⁵, Luiza D. Chelotti¹, Guilherme H. Pereira‑Filho¹ and Fábio S. Motta¹

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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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The southeastern coast of Brazil is located in a subtropical transition zone of the southwestern Atlantic Ocean, characterized by a high richness of fishes (e.g., Floeter et al., 2001; Cord et al., 2022) and benthic organisms (e.g., Aued et al., 2018), as well as an extensive marine biodiversity, including tropical and temperate organisms as also endemic species (Eschmeyer et al., 2010; Pinheiro et al., 2018; Quimbayo et al., 2021). The subtropical Brazilian reefs are composed mainly of rocky reefs (Floeter et al., 2006; Pinheiro et al., 2018), which are the main habitat of reef fish species (Ferreira et al., 2001) and occur extensively between 19ºS and 29ºS (Anderson et al., 2019). Although rocky reefs generally present less topographic complexity than biogenic reefs, these environments support a rich fish fauna (Ferreira et al., 2001, Luiz et al., 2008; Gibran, Moura, 2012; Pinheiro et al., 2018; Souza et al., 2018; Motta et al., 2021; Rolim et al., 2017, 2019; Hoff et al., 2023), in addition to being important marine ecosystems worldwide for providing an array of ecosystem services, including food security, the survival of marine species and the maintenance of biodiversity, as well as the contribution to human well-being by providing leisure and recreation (Bevilaqcua et al., 2021; Riofrío-Lazo et al., 2022). In contrast, more structurally complex and biodiverse biogenic subtropical reefs are restricted to a few areas within the subtropical Brazilian region, including both rhodolith beds and coral reefs (Pereira-Filho et al., 2019).

Recently mapped, a coral reef associated with a rhodolith bed in the subtropical waters of São Paulo state, southeastern Brazil (around the Queimada Grande Island – QGI; 24°29’S 46°41’W), expanded the southern limit of the Atlantic coral reefs from 17ºS to 24°S (Pereira-Filho et al., 2019, 2021). Structurally complex coral reefs can enhance the abundance and richness of reef fishes (Darling et al., 2017), playing a key role in the trophic and non-trophic interactions between fish and benthic communities (Pereira et al., 2014; Longo et al., 2019). In this regard, the QGI has a high diversity of reef fishes, which may be associated with the three distinct reef habitats present in the region (i.e., rocky reefs, high coral cover, and rhodolith bed) (Pereira-Filho et al., 2019; Grillo et al., 2021; Becker et al., 2024). In addition to the QGI, located 16 km away, the coastal island of Queimada Pequena – QPI (24º22’S 46º48’W) is part of the Tupiniquins Ecological Station (ESEC, IUCN Category Ia: Strict Nature Reserve). Despite being a small no-take area (0.12 km²), the proximity of QPI to the QGI makes it particularly relevant for studying reef fish assemblages, as the short distance can allow ecological connectivity between these islands. The QPI has an exclusion radius of 1 km for marine protection, with only scientific research permitted, providing an important comparison to QGI and helping to understand how local protection and habitat diversity influence reef fish communities (e.g., Gragnolati et al., 2024).

The Brazilian subtropical coast has the highest number of threatened marine species in Brazil (Magris et al., 2021). In this context, coastal islands are important for biodiversity conservation, particularly in areas facing cumulative impacts, where high species diversity coincides with intense human activities. Coastal islands are also often associated with tourism and recreation due to their proximity to the coast (Ghermandi, Nunes, 2013; Freire et al., 2016; Marconi et al., 2020; Santos et al., 2023). However, the context of broad interests and multiple uses of these areas present a challenge to effective management, contributing to adverse environmental impacts and influencing the local economy (Tratalos, Austin, 2001; Coleman et al., 2004; Marconi et al., 2020). This reality results in an alarming scenario, given that Brazilian reef fishes play key ecological roles and their fisheries account for up to 15% of the total marine catches in Brazil (Eggertsen et al., 2024).

Considering the socioecological relevance of the southernmost Atlantic coral reef (see Marconi et al., 2020; Motta et al., 2022; Niz et al., 2023) and the limited number of studies on the fish fauna associated with rocky reefs in the subtropical Brazilian region (e.g., Luiz et al., 2008; Gibran, Moura, 2012; Souza et al., 2018; Motta et al., 2021), the aim of this study is to provide a comprehensive checklist of fish species (actinopterygians and chondrichthyans) occurring in the coastal areas around two nearby islands, Queimada Grande and Queimada Pequena, in southeastern Brazil. We also include information on the conservation status, geographic distribution, and trophic category of the recorded species. Altogether, this dataset provides an important baseline for conservation planning, monitoring, and management.

Material and methods

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Study sites. Sampling was carried out in Queimada Grande (QGI) and Queimada Pequena (QPI) islands (Fig. 1). The QGI (24°29’S 46°41’W) is located off the state of São Paulo, southeastern Brazil, at a distance of 34.8 km from the coast. The QGI has an emerging area of 0.78 km² and its marine surroundings are part of the Environmental Protected Area of the central coast of São Paulo State (EPA, category V IUCN), a multiple-use Marine Protected Area (MPA). The terrestrial area of the island belongs to the Area of Relevant Ecological Interest (ARIE, category IV IUCN) of Queimada Grande and Queimada Pequena islands. Tourism and fishing are only allowed within the EPA, with some restrictions. For instance, in 2021, through the EPA management plan a zone of the 19 km² around the QGI was established aiming to making recreational uses (e.g., fishing and scuba diving) compatible with biodiversity conservation (Motta et al., 2021; Niz et al., 2023). Despite this recent zoning, QGI is a historically targeted area for commercial and recreational fisheries, including spearfishing (Niz et al., 2023). QGI also is an important destination for recreational scuba diving (Marconi et al., 2020) and despite these multiple uses, the conservation of local marine biodiversity is compromised due to the lack of effective enforcement and fishing management initiatives (Marconi et al., 2020).

FIGURE 1| Study area in the Southwestern Atlantic. Brazil in light gray and São Paulo State in dark gray on the maps above. Queimada Grande Island (QGI) and Queimada Pequena Island (QPI) represented by black points. Below are the Queimada Grande and Queimada Pequena islands, and the coral and rocky reefs that occur in the area, respectively.

The QPI marine surroundings are protected by the Tupiniquins Ecological Station (Tupiniquins ESEC), a no-take MPA (IUCN category Ia) that covers a radius of 1 km from the coast of the island. QPI is located about 22 km from the coast (24°22’S 46°48’W) with an area of approximately 0.12 km². Visitation and recreational diving in this area are prohibited, with only scientific research allowed. The proximity to the coast exposes the island to various disturbances, mainly due to the easy access for artisanal fishers and recreational anglers (Giglio et al., 2022).

Data compilation. The checklist was compiled from data collected between 1999 to 2025, composed by: Censos Visuais Subaquáticos Estacionários (UVC), protocol adapted from Minte-Vera et al. (2008); landings of gillnet fisheries from the Cação Project; Baited Remote Underwater Video (BRUV), method widely used to evaluate fish assemblages (Cappo et al., 2006; Mallet, Pelletier, 2014; Goetze et al., 2015); personal observation; photographic; and scientific collection vouchers (Queimada Grande Island, NUPEC Collection – Research and Study Center in Chondrichthyes – 984); citizen science monitoring program from the “Viu Raia? Project”, and literature (LIT; Moura, 2002; Moura et al., 2003; Pivetta et al., 2012).

The method by which species were recorded and documented: UVC = Underwater Visual Census; BRUV = Baited Remote Underwater Video; CP = landings of gillnet fishing (Cação Project); PHO = Photographs; OBS = Personal Observation; VOU = scientific collection vouchers of the Center for Research and Study in Chondrichthyes (NUPEC); Moura et al., 2002, 2003; Pivetta et al., 2012; and Anderson et al., 2020. The record types were categorized in two types of font: LIT = Literature and Present Study (PS) (Tabs. 1, S1;Fig. S2).

TABLE 1 | Record type, method and font. Literature (LIT) and Present Study (PS). Underwater Visual Census (UVC), Baited Remote Underwater Video (BRUV), landings of gillnet fishing (Cação Project), Photographs (PHO), Personal observation (DGN; OBFG; ACSA) and scientific collection vouchers (NUPEC).

Record type	Method	Sampling effort	Font
Underwater Visual Census (2015)	UVC	196 visual census	PS
Baited Remote Underwater Video (2015)	BRUV	6 deployments	PS
Baited Remote Underwater Video (2017)	BRUV	26 deployments	PS
Baited Remote Underwater Video (2020–2021)	BRUV	33 deployments	PS
Landings of gillnet fishing (1996–2005 and 2012–2017)	Cação Project	326 landing samples	PS
Photographs (2019–2021)	PHO	83 photographs	PS
Domingos Garrone-Neto – DGN (2017–2024)	Personal observation	25 years of field experience	PS
Otto B. F. Gadig – OBFG (1996–2017)	Personal observation	40 years of field experience	PS
Ana Clara S. Athayde – ACSA (2024)	PHO	10 dives	PS
Center for Research and Study in Chondrichthyes (NUPEC)	Scientific Collection Vouchers	1 species	PS
Recreational Scuba Diver sightings (2023–2024)	“Viu Raia? Project”	58 diving operations	PS
Moura (2002)	UVC	29 visual census	LIT
Moura et al. (2003)	UVC	62 visual census	LIT
Pivetta et al. (2012)	UVC	30 visual census	LIT

Fishes were grouped and listed in the phylogenetic order of families, following Eschmeyer’s Catalog of Fishes (Fricke et al., 2026). Species were identified according to Menezes, Figueiredo (1980, 1985), Figueiredo, Menezes (1980), Carvalho-Filho (1999, 2024), Rolim et al. (2017) and Floeter et al. (2023).

The general diet of species, the geographical distributions and endemism of Southwestern Atlantic were adapted from the literature (Pinheiro et al., 2018; Floeter et al., 2023). Trophic group was categorized as follows: MINV: mobile invertebrate feeders; MCAR: macro carnivores; HERB: herbivores; PLAN: planktivores; OMNI: omnivores; and SINV: sessile invertebrate feeders. About geographical distributions: WA, Western Atlantic; OIB, Oceanic Islands of Brazil; CT, Circumtropical; EA, Eastern Atlantic; BR, Brazilian Coast Endemics; MAR, Mid Atlantic Ridge and SA, Southwestern Atlantic.

The conservation category was based on the International Red List of Threatened Species (International Union for Conservation of Nature – IUCN, 2021), and the National Red List of Threatened Species (ICMBio, 2018; MMA, 2022, 2023).

Results​

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We compiled a total of 212 taxa belonging to 71 families and 209 species (202 species belonging to 71 families and 89 species belonging to 44 families were recorded in Queimada Grande and Queimada Pequena, respectively). Five species were only recorded in QPI but not recorded in QGI, which were Scomberomorus brasiliensis, Alectis ciliaris, Bothus robinsi, Caranx lugubris, and Selene vomer (Tab. 2). A total of 184 genus (85.38%) included in 54 families was associated to actinopterygians, and 31 genus (14.62%) of 17 families to chondrichthyans (Figs. 2–4). We also recorded a Holocephali (chimaera) for the first time in these areas. The families of actinopterygians with the highest species richness were Carangidae (n = 21), followed by Labridae (n = 19), Epinephelidae (n = 9) and Pomacentridae (n = 9). Regarding the chondrichthyans, the richest families were Carcharhinidae, Dasyatidae and Mobulidae (n = 7, n = 4 and n = 3, respectively).

TABLE 2 | Checklist of the fish species recorded at the Queimada Grande and Queimada Pequena Islands. IUCN: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; and DD = Data Deficient. National Category – ICMBio: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; and DD = Data Deficient. Geographic Range: WA = Western Atlantic; OIB = Oceanic Islands of Brazil; CT = Circumtropical; EA = Eastern Atlantic; BR = Brazilian Province Endemics; MAR = Mid Atlantic Ridge; and SA = Southwestern Atlantic. Trophic group: MINV = mobile invertebrate feeders; MCAR = macro-carnivores; HERB = herbivores; PLAN = planktivores; OMNI = omnivores; and SINV = sessile invertebrate feeders. Font record type: Present Study (PS); Literature (LIT). Vouchers used as reference: Diplectrum radiale ACF74110003, Haemulon atlanticus ACF77110001, Halichoeres bathyphilus ACF76060026, Caranx latus ACF78040101, Acanthurus sp. ACF78040101, Gymnothorax ocellatus HEM57022001, Serranus baldwini RLO91092003, Synodus foetens RLO91112303, Bothus sp. RLO91112303, Scorpaena isthmensis RLO93050203, Sphoeroides camila RLO93050203, Canthigaster figueiredoi RLO93050203, Coryphopterus glaucofrenum RLO91061503, Emblemariopsis signifer RLO91061503, Apogon pseudomaculatus RLO91061503, Stegastes pictus RLO91061503, Chromis jubauna RLO91121503, Azurina multilineata RLO92040501, Trachinotus falcatus RPL66120001, Squatina guggenheim IQG001.

Family/Species	QGI	QPI	IUCN	National	Geographic range	Trophic category	Font
CHONDRICHTHYES
Rhincodontidae
Rhincodon typus Smith, 1828	x		EN	VU	CT, OIB	PLAN	PS
Carchariidae
Carcharias taurus Rafinesque, 1810	x	x	CR	CR	–	MCAR	PS
Lamnidae
Isurus oxyrinchus Rafinesque, 1810	x		EN	CR	–	MCAR	PS
Carcharhinidae
Carcharhinus brevipinna (Valenciennes, 1839)	x	x	VU	VU	–	MCAR	PS
Carcharhinus falciformis (Bibron, 1839)	x	x	VU	CR	–	MCAR	PS
Carcharhinus limbatus (Valenciennes, 1839)	x	x	VU	NT	–	MCAR	PS
Carcharhinus obscurus (Lesueur, 1818)	x	x	EN	EN	OIB	MCAR	PS
Carcharhinus plumbeus (Nardo, 1827)	x	x	EN	CR	OIB	MCAR	PS
Rhizoprionodon lalandii (Valenciennes, 1839)	x	x	VU	NT	WA	MCAR	PS
Rhizoprionodon porosus (Poey, 1861)	x	x	VU	DD	WA	MCAR	PS
Galeocerdidae
Galeocerdo cuvier (Péron & Lesueur, 1822)	x	x	NT	NT	CT	MCAR	PS
Sphyrnidae
Sphyrna lewini (Griffith & Smith, 1834)	x	x	CR	CR	OIB	MCAR	PS
Sphyrna zygaena (Linnaeus, 1758)	x	x	VU	CR	OIB	MCAR	PS
Squatinidae
Squatina guggenheim Marini, 1936	x	x	EN	CR	SA	–	PS
Narcinidae
Narcine brasiliensis (Olfers, 1831)	x	x	NT	VU	SA	MINV	PS
Trygonorrhinidae
Zapteryx brevirostris (Müller & Henle, 1841)	x	x	EN	VU	SA	MINV	PS
Rhinobatidae
Pseudobatos horkelii (Müller & Henle, 1841)	x	x	CR	CR	SA	MCAR	PS
Pseudobatos percellens (Walbaum, 1792)	x	x	EN	VU	WA	MCAR	PS
Dasyatidae
Bathytoshia centroura (Mitchill, 1815)	x	x	VU	CR	WA, OIB, EA	MCAR	PS
Dasyatis hypostigma Santos & Carvalho, 2004	x	x	EN	DD	SA	MCAR	PS
Hypanus berthalutzae Petean, Naylor & Lima, 2020	x	x	VU	–	WA, OIB	MCAR	PS
Hypanus guttatus (Bloch & Scheider, 1801)	x	x	NT	LC	WA	MCAR	PS
Gymnuridae
Gymnura altavela (Linnaeus, 1758)	x	x	EN	CR	WA, EA	MCAR	PS
Aetobatidae
Aetobatus narinari (Euphrasen, 1790)	x	x	EN	DD	WA, EA	MCAR	PS, LIT
Myliobatidae
Myliobatis freminvillei Lesueur, 1824	x	x	VU	EN	WA	–	PS
Rhinopteridae
Rhinoptera bonasus (Mitchill, 1815)	x	x	VU	DD	WA	–	PS
Rhinoptera brasiliensis Müller, 1836	x	x	VU	CR	WA	–	PS
Mobulidae
Mobula birostris (Walbaum, 1792)	x		EN	VU	CT, OIB	PLAN	LIT
Mobula hypostoma (Bancroft, 1831)	x	x	EN	–	WA, OIB	–	LIT
Mobula thurstoni (Lloyd, 1908)	x	x	EN	VU	CT, OIB	–	PS
Callorhinchidae
Callorhinchus callorynchus (Linnaeus, 1758)	x	x	VU	DD	SA	–	PS
ACTINOPTERYGII
Elopidae
Elops smithi McBride, Rocha, Ruiz-Carus, & Bowen, 2010	x		DD	–	–	MCAR	PS
Megalopidae
Megalops atlanticus Valenciennes, 1847	x		VU	VU	WA, EA, EP	MCAR	PS
Muraenidae
Gymnothorax funebris Ranzani, 1839	x		LC	DD	WA, OIB	MCAR	PS, LIT
Gymnothorax moringa (Cuvier, 1829)	x	x	LC	DD	WA, OIB, MAR	MCAR	PS, LIT
Gymnothorax ocellatus Agassiz, 1831	x		LC	DD	WA	MCAR	PS, LIT
Gymnothorax vicinus (Castelnau, 1855)	x		LC	DD	WA, OIB, MAR	MCAR	PS, LIT
Ophichthidae
Myrichthys breviceps (Richardson, 1848)	x		LC	LC	WA, OIB	MINV	LIT
Myrichthys ocellatus (Lesueur, 1825)	x		LC	LC	WA, OIB	MINV	PS, LIT
Ophichthus ophis (Linnaeus, 1758)	x		LC	LC	WA, OIB, EA	MCAR	LIT
Clupeidae
Sardinella aurita Valenciennes, 1847	x		DD	DD	WA, EA	PLAN	PS
Synodontidae
Synodus foetens (Linnaeus, 1766)	x		LC	LC	WA	MCAR	PS, LIT
Synodus intermedius (Spix & Agassiz, 1829)	x		LC	LC	WA	MCAR	PS, LIT
Synodus synodus (Linnaeus, 1758)	x		LC	LC	WA, OIB, MAR	MCAR	PS
Holocentridae
Holocentrus adscensionis (Osbeck, 1765)	x	x	LC	LC	WA, OIB, MAR	MINV	PS, LIT
Myripristis jacobus Cuvier, 1829	x		LC	LC	WA, OIB, MAR	MINV	PS, LIT
Plectrypops retrospinis (Guichenot, 1853)	x		LC	LC	WA, OIB	MINV	PS
Apogonidae
Apogon americanus Castelnau, 1855	x		–	LC	BR, OIB	PLAN	PS, LIT
Apogon quadrisquamatus Longley, 1934	x		LC	DD	WA	PLAN	LIT
Apogon pseudomaculatus Longley, 1932	x		LC	LC	WA, EA	PLAN	LIT
Gobiidae
Bathygobius soporator (Valenciennes, 1837)	x		LC	LC	BR	MINV	LIT
Coryphopterus dicrus Böhlke & Robins, 1960	x	x	LC	LC	WA	MINV	LIT
Coryphopterus glaucofraenum Gill, 1863	x		LC	LC	WA	MINV	PS, LIT
Ctenogobius saepepallens (Gilbert & Randall, 1968)	x		LC	LC	WA	MINV	LIT
Elacatinus figaro Sazima, Moura & Rosa, 1997	x	x	–	VU	BR	MINV	PS, LIT
Gnatholepis thompsoni Jordan, 1904	x		LC	LC	WA	MINV	LIT
Lythrypnus brasiliensis Greenfield, 1988	x		–	LC	BR, OIB	MINV	LIT
Ptereleotris randalli Gasparini, Rocha & Floeter, 2001	x		LC	LC	BR	PLAN	PS, LIT
Dactylopteridae
Dactylopterus volitans (Linnaeus, 1758)	x		LC	LC	WA, OIB, EA	MINV	LIT
Mullidae
Pseudupeneus maculatus (Bloch, 1793)	x	x	LC	LC	WA, OIB	MINV	PS, LIT
Callionymidae
Callionymus bairdi Jordan, 1888	x		LC	LC	WA, OIB, MAR	MINV	PS
Syngnathidae
Hippocampus erectus Perry, 1810	x		VU	VU	WA, MAR	MINV	LIT
Hippocampus reidi Ginsburg, 1933	x		NT	VU	WA, OIB	MINV	LIT
Pomatomidae
Pomatomus saltatrix (Linnaeus, 1766)	x		VU	NT	CT	MCAR	PS
Scombridae
Katsuwonus pelamis (Linnaeus, 1758)	x		LC	LC	CT	MCAR	PS
Scomberomorus brasiliensis Collette, Russo & Zavala-Camin, 1978		x	LC	LC	WA	MCAR	PS
Trichiuridae
Trichiurus lepturus Linnaeus, 1758	x	x	LC	LC	CT	MCAR	PS
Centropomidae
Centropomus undecimalis (Bloch, 1792)	x		LC	LC	WA	MCAR	PS
Centropomus spp.	x		LC	LC	WA	MCAR	LIT
Sphyraenidae
Sphyraena barracuda (Edwards, 1771)	x		LC	LC	CT	MCAR	LIT
Sphyraena guachancho Cuvier, 1829	x		LC	LC	WA, OIB, EA	MCAR	PS, LIT
Bothidae
Bothus lunatus (Linnaeus, 1758)	x		LC	LC	WA, OIB, MAR	MCAR	LIT
Bothus ocellatus (Agassiz, 1831)	x		LC	LC	WA	MINV	PS, LIT
Bothus robinsi Topp & Hoff, 1972		x	LC	LC	WA	MINV	PS
Xiphiidae
Xiphias gladius Linnaeus, 1758	x		LC	NT	CT	MCAR	PS
Carangidae
Alectis ciliaris (Bloch, 1787)		x	LC	LC	CT	MCAR	PS
Caranx bartholomaei Cuvier, 1833	x	x	LC	LC	WA, OIB, EA	MCAR	PS, LIT
Caranx crysos (Mitchill, 1815)	x	x	LC	LC	WA, OIB, MAR	MCAR	PS, LIT
Caranx hippos (Linnaeus, 1766)	x	x	LC	LC	WA, EA	MCAR	PS
Caranx latus Agassiz, 1831	x	x	LC	LC	WA, OIB, MAR	MCAR	PS, LIT
Caranx lugubris Poey, 1860		x	LC	LC	CT	MCAR	PS
Caranx ruber (Bloch, 1793)	x		LC	LC	WA, OIB, MAR	MCAR	LIT
Chloroscombrus chrysurus (Linnaeus, 1766)	x	x	LC	LC	WA, EA	PLAN	PS, LIT
Decapterus macarellus (Cuvier, 1833)	x		LC	LC	CT	PLAN	PS
Decapterus punctatus (Cuvier,1829)	x		LC	LC	WA, MAR, EA	PLAN	PS
Elagatis bipinnulata (Quoy & Gaimard, 1825)	x		LC	LC	CT	MCAR	PS
Oligoplites spp.	x		LC	LC	WA	MCAR	PS
Pseudocaranx dentex (Bloch & Schneider, 1801)	x	x	LC	LC	CT	MINV	PS, LIT
Selene vomer (Linnaeus, 1758)		x	LC	LC	WA	PLAN	PS
Seriola dumerili (Risso, 1810)	x	x	LC	LC	CT	MCAR	PS, LIT
Seriola fasciata (Bloch, 1793)	x		LC	DD	WA	MCAR	LIT
Seriola lalandi Valenciennes, 1833	x		LC	LC	CT	MCAR	PS, LIT
Seriola rivoliana Valenciennes, 1833	x		LC	LC	CT	MCAR	PS, LIT
Trachinotus carolinus (Linnaeus, 1766)	x		LC	LC	WA	MINV	PS
Trachinotus falcatus (Linnaeus, 1758)	x		LC	LC	WA, OIB	MINV	PS
Trachinotus goodei Jordan & Evermann, 1896	x	x	LC	LC	WA	MINV	PS, LIT
Rachycentridae
Rachycentron canadum (Linnaeus, 1766)	x	x	LC	LC	CT	MCAR	PS, LIT
Coryphaenidae
Coryphaena equiselis Linnaeus, 1758	x		LC	LC	CT	MCAR	PS
Coryphaena hippurus Linnaeus, 1758	x		LC	LC	CT	MCAR	PS
Belonidae
Tylosurus acus (Lacepède, 1803)	x		LC	LC	WA, EA	MCAR	PS
Mugilidae
Mugil spp.	x		–	–	–	HERB	LIT
Pomacentridae
Abudefduf saxatilis (Linnaeus, 1758)	x	x	LC	LC	WA, OIB, MAR	OMNI	PS, LIT
Azurina multilineata (Guichenot, 1853)	x	x	LC	LC	WA, OIB, MAR	PLAN	PS, LIT
Chromis flavicauda (Günther, 1880)	x		DD	LC	BR, OIB	PLAN	LIT
Chromis jubauna Moura, 1995	x		–	LC	BR, OIB	PLAN	PS, LIT
Chromis limbata (Valenciennes, 1833)	x		LC	–	SA, EA	PLAN	PS, LIT
Chromis vanbebberae McFarland, Baldwin, Robertson, Rocha & Tornabene, 2020	x		LC	LC	BR, OIB	PLAN	LIT
Stegastes fuscus (Cuvier, 1830)	x	x	LC	LC	BR, OIB	HERB	PS, LIT
Stegastes pictus (Castelnau, 1855)	x	x	–	LC	BR, WA, OIB	OMNI	PS, LIT
Stegastes variabilis (Castelnau, 1855)	x	x	–	LC	BR	HERB	PS, LIT
Tripterygiidae
Enneanectes altivelis Rosenblatt, 1960	x		LC	LC	WA, OIB	MINV	LIT
Blenniidae
Hypleurochilus fissicornis (Quoy & Gaimard, 1824)	x		LC	LC	SA, EA	SINV	LIT
Hypsoblennius invemar Smith-Vaniz & Acero, 1980	x		LC	LC	WA	MINV	LIT
Parablennius marmoreus (Poey, 1876)	x	x	LC	LC	WA	OMNI	PS, LIT
Parablennius pilicornis (Cuvier, 1829)	x		LC	LC	SA, EA	OMNI	PS LIT
Scartella cristata (Linnaeus, 1758)	x		LC	LC	WA, EA	HERB	LIT
Labrisomidae
Gobioclinus kalisherae (Jordan, 1904)	x	x	LC	LC	WA, OIB	MINV	LIT
Labrisomus nuchipinnis (Quoy & Gaimard, 1824)	x	x	LC	LC	WA, OIB, EA	MCAR	PS, LIT
Malacoctenus delalandii (Valenciennes, 1836)	x	x	LC	LC	WA	MINV	LIT
Malacoctenus zaluari Carvalho-Filho, Gasparini & Sazima, 2020	x		LC	LC	–	–	LIT
Starksia brasiliensis (Gilbert, 1900)	x		LC	LC	BR	MINV	LIT
Chaenopsidae
Emblemariopsis signifer (Ginsburg, 1942)	x	x	LC	LC	BR	MINV	LIT
Serranidae
Diplectrum formosum (Linnaeus, 1766)	x		LC	LC	WA, OIB	MCAR	PS
Diplectrum radiale (Quoy & Gaimard, 1824)	x		LC	LC	WA	MCAR	PS, LIT
Serranus atrobranchus (Cuvier, 1829)	x		LC	LC	WA	MCAR	PS
Serranus baldwini (Evermann & Marsh, 1899)	x	x	LC	LC	WA, OIB	MCAR	PS, LIT
Serranus flaviventris (Cuvier, 1829)	x	x	LC	LC	WA	MCAR	PS, LIT
Epinephelidae
Cephalopholis furcifer (Valenciennes, 1828)	x		LC	LC	WA, OIB, MAR	PLAN	PS, LIT
Epinephelus itajara (Lichtenstein, 1822)	x		VU	CR	WA, OIB, EA	MCAR	LIT
Epinephelus marginatus (Lowe, 1834)	x	x	VU	VU	SA, EA, WA	MCAR	PS, LIT
Epinephelus morio (Valenciennes, 1828)	x		VU	VU	WA	MCAR	LIT
Hyporthodus niveatus (Valenciennes, 1828)	x		VU	VU	WA	MCAR	PS, LIT
Mycteroperca acutirostris (Valenciennes, 1828)	x	x	LC	DD	WA	MCAR	PS, LIT
Mycteroperca bonaci (Poey, 1860)	x		NT	VU	WA, OIB	MCAR	PS, LIT
Mycteroperca interstitialis (Poey, 1860)	x		VU	VU	WA, OIB	MCAR	LIT
Mycteroperca microlepis (Goode & Bean, 1879)	x		VU	DD	WA	MCAR	LIT
Scorpaenidae
Scorpaena isthmensis Meek & Hildebrand, 1928	x		LC	LC	WA, OIB	MCAR	LIT
Scorpaena plumieri Bloch, 1789	x		LC	LC	WA, OIB, MAR	MCAR	PS
Labridae
Bodianus pulchellus (Poey, 1860)	x		LC	LC	WA, OIB, EA	MINV	PS, LIT
Bodianus rufus (Linnaeus, 1758)	x	x	LC	LC	WA, OIB	MINV	PS, LIT
Clepticus brasiliensis Heiser, Moura & Robertson, 2000	x		LC	LC	BR, OIB	PLAN	PS, LIT
Cryptotomus roseus Cope, 1871	x		LC	LC	WA, OIB, MAR	HERB	PS, LIT
Doratonotus megalepis Günther, 1862	x		LC	LC	WA, OIB	MINV	LIT
Halichoeres brasiliensis (Bloch, 1791)	x	x	DD	LC	BR, OIB	MINV	PS, LIT
Halichoeres dimidiatus (Agassiz, 1831)	x	x	LC	LC	BR, OIB	MINV	PS, LIT
Halichoeres poeyi (Steindachner, 1867)	x	x	LC	LC	WA, OIB	MINV	PS, LIT
Halichoeres sazimai Luiz, Ferreira & Rocha, 2009	x	x	–	LC	BR	MINV	PS, LIT
Nicholsina usta (Valenciennes, 1840)	x		LC	LC	WA	HERB	LIT
Scarus trispinosus Valenciennes, 1840	x		EN	EN	BR	HERB	LIT
Scarus zelindae Moura, Figueiredo & Sazima, 2001	x		DD	VU	BR, OIB	HERB	PS, LIT
Sparisoma amplum (Ranzani, 1841)	x		LC	NT	BR, OIB	HERB	PS, LIT
Sparisoma axillare (Steindachner, 1878)	x	x	DD	VU	BR, OIB	HERB	PS, LIT
Sparisoma frondosum (Agassiz, 1831)	x	x	DD	VU	BR, OIB	HERB	PS, LIT
Sparisoma radians (Valenciennes, 1840)	x		LC	LC	WA	HERB	LIT
Sparisoma tuiupiranga Gasparini, Joyeux & Floeter, 2003	x	x	LC	LC	BR, OIB	HERB	PS
Thalassoma noronhanum (Boulenger, 1890)	x		LC	LC	BR, OIB	MINV	LIT
Xyrichthys novacula (Linnaeus, 1758)	x		LC	LC	WA, OIB, EA	MINV	LIT
Kyphosidae
Kyphosus sectatrix (Linnaeus, 1758)	x	x	LC	–	WA, OIB, MAR	HERB	PS, LIT
Kyphosus vaigiensis (Quoy & Gaimard, 1825)	x		–	–	WA, EA	HERB	LIT
Pempheridae
Pempheris schomburgkii Müller & Troschel, 1848	x	x	LC	LC	WA, OIB	PLAN	PS, LIT
Gerreidae
Eucinostomus melanopterus (Bleeker, 1863)	x		LC	LC	WA, EA	MINV	LIT
Ephippidae
Chaetodipterus faber (Broussonet, 1782)	x	x	LC	LC	WA	OMNI	PS, LIT
Sciaenidae
Cynoscion acoupa (Lacepède, 1801)	x		VU	NT	–	–	PS
Cynoscion leiarchus (Cuvier, 1830)	x		LC	LC	–	–	PS
Odontoscion dentex (Cuvier, 1830)	x	x	LC	LC	WA	MCAR	PS, LIT
Pareques lineatus (Cuvier, 1830)	x	x	LC	DD	WA	MINV	PS, LIT
Pogonias cromis (Linnaeus, 1766)	x		LC	EN	–	–	PS
Haemulidae
Anisotremus surinamensis (Bloch, 1791)	x	x	DD	DD	WA, OIB	MINV	PS, LIT
Anisotremus virginicus (Linnaeus, 1758)	x	x	LC	LC	WA	MINV	PS, LIT
Haemulon atlanticus Carvalho, Marceniuk, Oliveira & Wosiacki, 2020	x	x	LC	LC	WA	MINV	PS, LIT
Haemulon aurolineatum Cuvier, 1830	x	x	LC	LC	WA	MINV	PS, LIT
Haemulon parra (Desmarest, 1823)	x	x	LC	LC	WA, OIB	MINV	PS
Haemulon plumierii (Lacepède, 1801)	x		LC	DD	WA	MINV	PS, LIT
Orthopristis rubra (Cuvier, 1830)	x		LC	LC	WA	MINV	PS
Lobotidae
Lobotes surinamensis (Bloch, 1790)	x		LC	LC	CT	MCAR	PS
Lutjanidae
Lutjanus analis (Cuvier, 1828)	x		NT	NT	WA	MCAR	PS, LIT
Lutjanus cyanopterus (Cuvier, 1828)	x	x	VU	VU	WA	MCAR	PS, LIT
Lutjanus jocu (Bloch & Schneider, 1801)	x		DD	NT	WA, OIB, MAR	MCAR	PS, LIT
Lutjanus synagris (Linnaeus, 1758)	x		NT	NT	WA	MCAR	PS
Ocyurus chrysurus (Bloch, 1791)	x		DD	NT	CT	MCAR	PS
Pristipomoides freemani Anderson, 1966	x		LC	LC	WA	MCAR	PS
Rhomboplites aurorubens (Cuvier, 1829)	x	x	VU	NT	WA, OIB	MCAR	PS, LIT
Malacanthidae
Malacanthus plumieri (Bloch, 1786)	x		LC	LC	WA, OIB, MAR	MCAR	PS, LIT
Pomacanthidae
Centropyge aurantonotus Burgess, 1974	x		LC	DD	BR, WA, OIB	HERB	LIT
Holacanthus ciliaris (Linnaeus, 1758)	x		LC	DD	WA, OIB	SINV	LIT
Holacanthus tricolor (Bloch, 1795)	x		LC	DD	WA, OIB	SINV	PS, LIT
Pomacanthus arcuatus (Linnaeus, 1758)	x		LC	DD	WA, OIB	OMNI	LIT
Pomacanthus paru (Bloch, 1787)	x	x	LC	DD	WA, OIB, MAR	OMNI	PS, LIT
Chaetodontidae
Chaetodon sedentarius Poey, 1860	x		LC	LC	WA, EA	SINV	LIT
Chaetodon striatus Linnaeus, 1758	x	x	LC	LC	WA, OIB	SINV	PS, LIT
Acanthuridae
Acanthurus bahianus Castelnau, 1855	x	x	LC	LC	BR, WA, OIB	HERB	PS, LIT
Acanthurus chirurgus (Bloch, 1787)	x	x	LC	LC	WA, OIB, MAR	HERB	PS, LIT
Acanthurus coeruleus Bloch & Schneider, 1801	x	x	LC	LC	WA, OIB, MAR	HERB	LIT
Sparidae
Calamus penna (Valenciennes, 1830)	x		LC	LC	WA	MINV	PS, LIT
Calamus pennatula Guichenot, 1868	x		LC	LC	WA	MINV	PS, LIT
Diplodus argenteus (Valenciennes, 1830)	x	x	LC	LC	BR, OIB	OMNI	PS, LIT
Pagrus pagrus (Linnaeus, 1758)	x		LC	DD	WA, EA, OIB	MINV	PS, LIT
Priacanthidae
Heteropriacanthus cruentatus (Lacepède, 1801)	x		LC	LC	CT	MCAR	LIT
Priacanthus arenatus Cuvier, 1829	x		LC	LC	WA, OIB, EA	MCAR	PS, LIT
Ogcocephalidae
Ogcocephalus vespertilio (Linnaeus, 1758)	x	x	LC	LC	WA	MINV	PS, LIT
Diodontidae
Diodon holocanthus Linnaeus, 1758	x		LC	LC	CT	MINV	LIT
Diodon hystrix Linnaeus, 1758	x		LC	LC	CT	MINV	LIT
Tetraodontidae
Canthigaster figueiredoi Moura & Castro, 2002	x	x	LC	LC	BR, WA, OIB	OMNI	PS, LIT
Sphoeroides camila Carvalho-Filho, Rotundo, Pitassy & Sazima, 2023	x	x	LC	LC	WA, OIB, EA	MINV	PS, LIT
Sphoeroides testudineus (Linnaeus, 1758)	x		LC	DD	WA	MINV	PS
Ostraciidae
Acanthostracion quadricornis (Linnaeus, 1758)	x		LC	LC	WA, OIB, EA	SINV	LIT
Acanthostracion polygonius Poey, 1876	x		LC	LC	WA, OIB	SINV	LIT
Lactophrys trigonus (Linnaeus, 1758)	x		LC	LC	WA, OIB	OMNI	LIT
Monacanthidae
Aluterus scriptus (Osbeck, 1765)	x		LC	LC	CT	OMNI	LIT
Cantherhines macrocerus (Hollard, 1843)	x		LC	LC	WA, OIB, EA	OMNI	PS
Cantherhines pullus (Ranzani, 1842)	x		LC	LC	WA, OIB, EA	OMNI	PS, LIT
Stephanolepis hispida (Linnaeus, 1766)	x		LC	LC	WA, OIB, EA	OMNI	LIT
Balistidae
Balistes vetula Linnaeus, 1758	x		NT	NT	WA, OIB, MAR	MINV	PS, LIT

FIGURE 2| Some reef fishes from Queimada Grande and Queimada Pequena islands. A. Gymnothorax moringa; B. Sparisoma tuiupiranga; C. Ptereleotris randalli; D. Myrichthys ocellatus; E. Epinephelus marginatus; F. Chromis limbata; G. Ogcocephalus vespertilio; H. Elacatinus figaro; I. Pomacanthus paru (juvenile); J. Parablennius marmoreus (yellow phase); K. Dasyatis hypostigma; L. Mobula birostris. Photos by Carlo L. Francini.

FIGURE 3| Some shark and chimaera species from Queimada Grande and Queimada Pequena islands. A. Sphyrna lewini; B. Sphyrna zygaena – 86 cm; C. Rhizoprionodon lalandii; D. Rhizoprionodon porosus; E. Carcharhinus brevipinna – 70 cm; F. Carcharhinus falciformis; G. Carcharhinus limbatus; H. Carcharhinus obscurus; I. Carcharhinus plumbeus – 87 cm; J. Carcharias taurus – 93.7 cm; K. Galeocerdo cuvier; L. Isurus oxyrinchus; M. Rhincodon typus; N. Squatina guggenheim; O. Callorhinchus callorynchus. Photos by Otto Bismarck F. Gadig, except photo M by C. L. Sampaio.

FIGURE 4| Some ray species from Queimada Grande and Queimada Pequena islands. A. Mobula thurstoni; B. Modula birostris; C. Mobula hypostoma; D. Narcine brasiliensis; E. Aetobatus narinari; F. Myliobatis freminvillei; G. Rhinoptera brasiliensis; H. Rhinoptera bonasus; I. Pseudobatos percellens; J. Pseudobatos horkelii; K. Zapteryx brevirostris; L. Gymnura altavela; M. Dasyatis hypostigma; N. Bathytoshia centroura; O. Hypanus guttatus; P. Hypanus berthalutzae. Photos by Otto Bismarck F. Gadig, except photo B by William White, F by C. E. Rangel, N by H. Santos, and P by F. Petean.

Regarding global conservation status, 42 species (~20% of the total) were classified in threatened categories (CR, EN, or VU; Tab. 2; Fig. 5). Three chondrichthyan species (~10% of this group of fish) were listed as Critically Endangered (CR) by the IUCN: Carcharias taurus, Sphyrna lewini, and Pseudobatos horkelii; all were recorded in both islands. Among the 14 Endangered (EN) species identified, 13 were chondrichthyans (42%) and one is an actinopterygian. Four species were recorded exclusively in QGI: Rhincodon typus, Isurus oxyrinchus, Mobula birostris, and Scarus trispinosus. The remaining species were observed on both islands. Twenty-four species were classified as Vulnerable (VU), including 12 chondrichthyans (38.7%) and 12 actinopterygians (~6.5%). Of these, 15 species were recorded in both islands, while nine were observed exclusively in QGI: Megalops atlanticus, Pomatomus saltatrix, Hippocampus erectus, Cynoscion acoupa, Epinephelus morio, E. itajara, Hyporthodus niveatus, Mycteroperca interstitialis, and M. microlepis. Regarding the national conservation status, ~6% of species were classified as Critically Endangered (CR), ~2% as Endangered (EN) and ~9% as Vulnerable (VU).

FIGURE 5| IUCN threatened species for chondrichthyans and actinopterygians registered in the Queimada Grande and Queimada Pequena Islands. Critically Endangered (CR); Endangered (EN); Vulnerable (VU), Near Threat (NT); Least Concern (LC); Data Deficient (DD) and Not Evaluated (NE).

Regarding to the trophic groups, most species recorded were macro carnivores (~40%; n = 85), mobile invertebrate feeders (~26.4%; n = 56), herbivores (~8.9%; n = 19) and planktivores (~8.9%; n = 19) (Tab. 1). Considering only actinopterygians, the proportion of species belonging to each trophic group were very similar between the studied islands (Tab. 1). Among to the chondrichthyes recorded, 64.5% of the species were macro carnivores (mostly Carcharhinidae), while 6.5% were mobile invertebrate feeders (batoids), and 6.5% are planktivores (e.g., Rhincodon typus and Mobula birostris). Most species (~25%; n = 53) are exclusively distributed across the Western Atlantic, followed by circumtropical distribution (~11%; n = 23) (Tab. 1). Furthermore, ~4% of the species (n = 8) are endemic to Brazil, and one non-native species, Chromis limbata, was also recorded.

Discussion​

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Here we demonstrated that the habitats associated with the southernmost Atlantic coral reef off QGI, along with the rocky reefs of QPI, support a remarkably high reef fish species richness (n = 208). Despite the relatively small reef area, we recorded nearly one third of the 733 reef fish species found along the 8,000 km of the Brazilian coast, seamounts and oceanic islands (Pinheiro et al., 2018), highlighting the significant socioecological and conservation values of these insular environments. The reef fish richness of QGI is comparable to many southeastern Atlantic no-take MPAs (Luiz et al., 2008; Rolim et al., 2017, 2019; Hoff et al., 2023). However, concerning biomass, QGI presents lower values than that observed in no-take MPAs located in São Paulo State coast, probably in the face of fishing effects (see Motta et al., 2021; Gragnolati et al., 2024).

The high fish species richness recorded in this relatively small study area can be explained by a mosaic of reef environments, including a unique condition of QGI at this subtropical region, with different types of complexity and sizes of submerged rocky reefs, a coral reef, rhodolith beds and sandy channels (Pereira-Filho et al., 2019, 2021; Grillo et al., 2021; Motta et al., 2022). Furthermore, seasonal (austral summer) upwellings may create favorable oceanographic conditions such as nutrient enrichment for the occurrence, spawning, and persistence of fish species (Katsuragawa et al., 2014; Motta et al., 2022). The disparity in fish species richness between QGI (202 species) and QPI (89 species) may be related to the larger reef area and higher habitat heterogeneity at QGI, in addition to having a 75,000 m² coral reef at 7–12 m depth, built mainly by M. decactis colonies cemented by crustose coralline algae (CCA) (Pereira-Filho et al., 2019). However, further studies are needed to confirm the influence of these habitat differences on fish richness.

Nearly 20% (40 species) of the recorded fish species are classified as threatened with extinction. Considering only the Chondrichthyes (31 species), this proportion increases to approximately 90% and 55%, according to global (IUCN) and national (ICMBio) assessments, respectively. These findings align with previous studies indicating that the coast of São Paulo harbors the highest number of threatened marine fish species along the Brazilian coastline (Ceretta et al., 2020; Magris et al., 2021). This aspect can be explained by two factors: the São Paulo coast is located in a faunal transition zone and is subject to high levels of cumulative impacts, especially overfishing (Magris et al., 2021). Moreover, these results reinforce, at a local scale, the high number of threatened species of Chondrichthyes in Brazil. For instance, approximately 33% of Chondrichthyes along the Brazilian coast are classified as threatened, similar to an estimated global rate of 32.6% (Dulvy et al., 2021).

Although most recorded actinopterygians species (81%) are categorized as Least Concern, some commercially important families include threatened species, such as Epinephelidae (e.g., Epinephelus itajara – CR; and E. marginatus – VU), the subfamily Scarinae (e.g., Scarus trispinosus – EN), and Lutjanidae represented by Lutjanus cyanopterus (Cuvier, 1828), classified as Vulnerable (VU). According to Niz et al. (2023), jacks, such as Seriola spp. and Alectis ciliaris, dusky grouper (Epinephelus marginatus) and cubera snapper (Lutjanus cyanopterus) are among the main targets of recreational fishing (onboard and spearfishing) around QGI. Regarding cubera snapper, the first in situ validation of a spawning aggregation of this species in subtropical waters off the southwestern Atlantic was recorded in close proximity to QGI and QPI (Motta et al., 2022), reinforcing the importance of local management and conservation efforts.

The high richness of macro carnivores (39.4%) and mobile invertebrate feeders (27.3%), as well as the predominance of species exclusively distributed in the Western Atlantic (24.7%), is consistent with previous studies conducted in the subtropical southwestern Atlantic (Ferreira et al., 2004; Luiz et al., 2008; Daros et al., 2012; Gibran, Moura, 2012). Still regarding the trophic groups, it is also worth highlighting the high richness of macro carnivore, which may be directly related to the various sources of information used which, in turn, include sampling through multiple techniques (e.g., UVC, BRUV, gillnet landings, citizen science) across different habitats (e.g., coral and rocky reef, water column, and sandy and rhodolith beds). By combining these methods and habitats, we detected species that are otherwise difficult to observe, such as large predatory fishes and elasmobranchs. Furthermore, this raises concerns about biodiversity conservation, as the group includes species under intense fishing pressure (Floeter et al., 2023), many of which are currently threatened with extinction, including most epinephelids, some lutjanids, and various species of sharks and rays.

Although the rocky reefs of the QPI are located within a no-take MPA, we observed fishing gear alongside different stages of benthic organism colonization during our field sampling trips, indicating long time frequent illegal fishery. Given the current low management effectiveness of the Tupiniquins ESEC, resulting from the lack of enforcement, the non-implementation of the management plan, and the inactivity of the management council (see Giglio et al., 2019), reactivating the effective implementation of this MPA, particularly through strengthened awareness and enforcement measures, is essential to safeguard the local fish fauna. Alternatively, recategorizing the area from an Ecological Station to a Park would permit public visitation such as recreational diving, which has been shown to help reduce the risk of illegal fishing in popular dive destinations (Steenbergen, 2013; Motta et al., 2021).

In 2003, a formal proposal for the establishment of a marine national park around Queimada Grande Island (QGI) was introduced by the Brazilian government. However, conflicts with the fishing sector, particularly the recreational fishing, impeded its advancement. Recreational activities at QGI have historical significance and support a substantial local economic chain. For example, onboard recreational fishing is estimated to generate at least USD 350,000 annually (Niz et al., 2023), while recreational scuba diving contributes approximately USD 160,000 per year to the regional economy (Agune, 2024). Additionally, the expansion of no-take zones of two coastal marine protected areas in São Paulo State (Laje de Santos Marine State Park and Alcatrazes Archipelago Wildlife Refuge) has increased the pressure for more effective management of the multiple-use MPA encompassing QGI. This culminated in the establishment of a tourism zoning regulation in 2021. In this context, Niz et al. (2023) developed spatial models to assess various risk scenarios to biodiversity, habitat integrity, and cultural ecosystem services (e.g., recreational activities) at QGI. Their findings suggest that the most effective management scenario is one that harmonizes conservation goals with recreational uses (Niz et al., 2023). Key management strategies highlighted include measures already outlined in the management plan of the multiple-use MPA (EPA), such as the prohibition of trawling within a 500-meter buffer zone around the island and anchoring on coral reef areas. Other important measures established under federal regulations, including catch quotas for recreational fishing, minimum and maximum catch sizes and prohibitions on the capture of threatened species (Niz et al., 2023).

Given the close relationship between fish populations and ecosystem services (Holmlund, Hammer, 1999), the removal of fish species can alter community structure and interactions, thereby affecting ecosystem functions (Luza et al., 2023; Eggertsen et al., 2024) and reducing both the productivity of recreational fisheries and the quality of recreational diving experience. Therefore, it is important to carry out systematic efforts to raise awareness among these users, as well as to develop and implement a tourism management plan to ensure sustainable use. In fact, these important subtropical reefs (QGI and QPI) still lack policies to safeguard their socioecological attributes, such as high fish richness. Here, we provide new information about the hidden biodiversity of the Brazilian subtropical reefs and highlight the need for measures to promote effective conservation and responsible use of the marine life in this singular mosaic of reef habitats.

Acknowledgments​

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We thank the management staff of the Tupiniquins Ecological Station (ESEC Tupiniquins) and Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) and Fundação Florestal by research permits; Bruno Oliveira, Lawrence Ikeda and Marcos Hamamura for field support and to share some fish records for the QGI.

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Authors

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Ana Clara S. Athayde¹, Fernanda A. Rolim¹, Otto B. F. Gadig², Caio R. Pimentel¹, Rafael R. Munhoz¹, Maisha Gragnolati¹, Fernando Z. Gibran³, Leonardo M. Neves⁴, Domingos Garrone-Neto⁵, Gabriel R. S. Souza⁵, Luiza D. Chelotti¹, Guilherme H. Pereira‑Filho¹ and Fábio S. Motta¹

^[1]    Laboratório de Ecologia e Conservação Marinha, Instituto do Mar, Universidade Federal de São Paulo (UNIFESP), Av. Dr. Carvalho de Mendonça, 144, Encruzilhada, 11070-100, Santos, SP, Brazil. (ACSA) anaclarathy@gmail.com, (FAR) fernandarolim2@gmail.com, (CRP) caiopimentelr@gmail.com, (RRM) raro.munhoz@gmail.com, (MG) mgragnolatifernandes@gmail.com, (LDC) luizadchelotti@gmail.com, (GHPF) pereira.filho@unifesp.br, (FSM) fmotta@unifesp.br (corresponding author).

^[2]    ELASMOBRASIL – Laboratório de Pesquisa em Elasmobrânquios, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Campus do Litoral Paulista, Praça Infante Dom Henrique, s/n, Parque Bitaru, 11330-900, São Vicente, SP, Brazil. (OBFG) otto.gadig@unesp.br.

^[3]    Centro de Ciências Naturais e Humanas, Universidade Federal do ABC, Alameda da Universidade, s/n, Bairro Anchieta, 09606 045, São Bernardo do Campo, SP, Brazil. (FZG) fernando.gibran@ufabc.edu.br.

^[4]    Laboratório de Ecologia Marinha e Costeira, Universidade Federal Rural do Rio de Janeiro, BR-465, km 7, 23897-000, Seropédica, RJ, Brazil. (LMV) leonardomneves@gmail.com.

^[5]    Laboratório de Ictiologia e Conservação de Peixes Neotropicais, Faculdade de Ciências Agrárias do Vale do Ribeira, Universidade Estadual Paulista (UNESP), Campus Registro – Av. Saburo Kameyama, 375, Estrada Municipal RGT-265, Bairro Carapiranga, 11900-000, Registro, SP, Brazil. (DGN) domingos.garrone-neto@unesp.br, (GRSS) gabrielraposo.souza@gmail.com.

Authors’ Contribution

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Ana Clara S. Athayde: Conceptualization, Formal analysis, Investigation, Writing-original draft.

Fernanda A. Rolim: Investigation, Writing-review and editing.

Otto B. F. Gadig: Investigation, Writing-review and editing.

Caio R. Pimentel: Writing-review and editing.

Rafael R. Munhoz: Investigation, Writing-review and editing.

Maisha Gragnolati: Formal analysis, Investigation, Writing-review and editing.

Fernando Z. Gibran: Investigation, Writing-review and editing.

Leonardo M. Neves: Investigation, Writing-review and editing.

Domingos Garrone-Neto: Formal analysis, Investigation, Writing-review and editing.

Gabriel R. S. Souza: Investigation, Writing-review and editing.

Luiza D. Chelotti: Investigation, Writing-review and editing.

Guilherme H. Pereira‑Filho: Investigation, Writing-review and editing.

Fábio S. Motta: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Supervision, Writing-original draft.

Ethical Statement​

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Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) and Fundação Florestal by research permits (SISBIO: 46206 and COTEC: 004.358/2020; 26108–009.754/2014).

Competing Interests

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The author declares no competing interests.

Data availability statement

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The authors confirm that the data supporting the findings of this study are available within the article.

AI statement

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The authors did not use any AI-assisted technologies in the creation of this manuscript or its figures.

Funding

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The authors (ACSA, DGN, RRM, CRP, FAR) thank Petrobras S/A (Tripartite Agreement FapUnifesp-Unifesp-Petrobras number #23089.102938/2019–54) for the scholarships. Financial support was provided by FAPESP (Process numbers 2019/19423–5 and 2023/11845–3), Instituto Linha D’água and Fundação SOS Mata Atlântica. Finally, FSM, GHPF, OBFG acknowledge individual grants from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).

Supplementary Material

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Supplementary material S1

Supplementary material S2

How to cite this article

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Athayde ACS, Rolim FA, Gadig OBF, Pimentel CR, Munhoz RR, Gragnolati M, Gibran FZ, Neves LM, Garrone-Neto D, Souza GRS, Chelotti LD, Pereira‑Filho GH, Motta FS. High fish richness associated with the southernmost Atlantic coral reef and adjacencies: comments on conservation and management for sustainable use. Neotrop Ichthyol. 2026; 24(1):e250090. https://doi.org/10.1590/1982-0224-2025-0090

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 October 17, 2025

Submitted May 13, 2025

Epub March 23, 2026