Coral reefs are amongst the most relevant ecosystems on Earth with the greatest biodiversity of all marine habitats (Moberg, Folke, 1999; Adey, 2000). In addition to harbor a rich biodiversity, coral reefs are crucial both ecologically and economically, providing several services, such as fish stocks and coastal protection (Brander et al., 2007; Paula et al., 2018). The use of coral reefs, whether for tourism or fishing, is a notable attraction for the industry (Davenport, Davenport, 2006; Cowburn et al., 2018), raising billions of dollars annually and benefiting around 450 million people in 109 countries (Moberg, Folke, 1999; Pandolfi et al., 2011; Spalding et al., 2017).

Brazil has the unique coral reefs with many representative endemics to the southwest Atlantic Ocean (Ferreira et al., 2013; Leão et al., 2016). These ecosystems differ from reefs around the world due to the absence of natural disasters, low species richness; yet they present high endemism and high levels of sedimentation (Leão et al., 2003; Leão et al., 2016; Soares et al., 2021). Despite this ecological relevance, Brazilian reefs are still poorly investigated and data on biodiversity assessment of corals and reef fishes are still absent for several regions, including many Marine Protected Areas (MPAs) (Sampaio et al., 2016).

Reef ecosystems harbor a high diversity of fish species (Reaka-Kudla, 1997) and the close relationship of fish and reef environments has been extensively described in the last decades (see Mora, 2015 for a review). Reef fish communities are driven by multiple factors such as depth, wave exposure, latitude, and benthic structure (Gibran, Moura, 2012; Pereira et al., 2014; Leal et al., 2015; Pinheiro et al., 2018). However, human influence has also been currently considered one important factor affecting reef fish community and how management strategies are implemented seem to directly drive reef fish diversity on coral reefs (Graham et al., 2017; Ruppert et al., 2018). In this scenario, the creation of MPAs along with adequate zoning process such as multiple use areas, or restricting any type of exploration, such as no-take zones, rises as a vital reef fish conservation strategy worldwide (Mora et al., 2006; Francini-Filho, Moura, 2008; Emslie et al., 2015; Hall et al., 2021).

During the last decades, the number of studies regarding reef fishes in Brazilian waters extensively increased. Many recent studies described several new fishes, from cryptic species (Smith-Vaniz et al., 2018; Carvalho-Filho et al., 2020) to large elasmobranchs (Gomes et al., 2000; Petean et al., 2020) and aimed to understand the relationship of community dynamics and environmental factors (Gibran, Moura, 2012; Pereira et al., 2014; Andrades et al., 2018; Matheus et al., 2019), fish­ing effects (Floeter et al., 2006; Rolim et al., 2019) and human induced behavior (Benevides et al., 2019; Pereira et al., 2020). In this context, Pinheiro et al., (2018) recently revealed a very rich reef fish fauna at Southwestern Atlantic Ocean (SWA) with a total of 733 fish species, 405 are SWA resident reef fishes, of which 111 (27%) are endemics and 78 (19%) are globally threatened.

Nevertheless, despite substantial increase in knowledge regarding reef fish systematic, ecology, biogeography and evolution on the Southwestern Atlantic (Floeter et al., 2008; Morais et al., 2017; Smith-Vaniz et al., 2018; Lellys et al., 2019; Cordeiro et al., 2021; Moura et al., 2021), several large reef areas in the SWA remained unknown and poorly studied until recently (Freitas, Lotufo, 2014; Pinheiro et al., 2014; Anderson et al., 2014 Pinheiro et al., 2015; Pereira et al., 2018; Guabiroba et al., 2020; Araújo et al., 2020). Hence, this study analyzed reef fishes biodiversity on the largest Brazilian multiple use coastal MPA and provided insights on fishes depth distribution, dispersal potential, IUCN red list categories and trophic/functional groups. Additionally, aspects of conservation and distribution of reef fishes inside MPA Costa dos Corais are also discussed.

Study area. Marine Protect Area (MPA) “Costa dos Corais” (APACC) is the largest Brazilian coastal MPA created to protect coral reef systems on Brazilian waters. This MPA stretches from 120 km in the Northeast Brazil encompassing two states and 12 municipalities (Maida, Ferreira, 1997; Miranda et al., 2020) (Fig. 1). MPA Costa dos Corais covers a large range of different ecosystems such as shallow reefs, mangroves, seagrass beds, rhodolith/sponge beds and mesophotic reefs (Fig. 2) from the coast to the break of the continental shelf (Maida, Ferreira, 1997; Pereira et al., 2018) (Figs. 1–2).

The multiple use MPA corresponds to an IUCN category VI protected area, where sustainable use is admitted according to its management plan, published in 2013 (ICMBio, 2013). A new version of the management plan elaborated by managers in partnership with local NGOs, researchers, fishers, and touristic trade has just been published (ICMBio, 2021). APACC zoning plan is probably the best strategy to promote coral reefs conservation locally with some areas selected as “no take zones” (all human activities, except research, are prohibited), some touristic areas (only low impact visiting activities allowed) and large multiple use locations (fishing and tourists allowed).

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Sampling design. Present study data represents a decadal field work effort (initiated in 2010) collecting reef fish information inside the MPA with multiple sampling techniques such as, remotely operated underwater vehicle (ROV), baited remote underwater video (BRUV) and scuba diving/transects. A total of 125 sampling sites distributed on MPA from 3 to 70 m depth have been investigated in more than 500 dives and 1000 belt transects (Fig. 1). Species were listed from our database (long term monitoring throughout the MPA since 2010), artisanal fisheries landing and based on literature (Pinheiro et al., 2018), and authors personal information. Vouchers specimens are listed in Tab. S1.

Species categorization. All recorded species were categorized according to depth distribution, dispersal potential, IUCN (2020) red list category, trophic and functional group as follow: Depth distribution: According to depth distribution species were categorize as: Very shallow (0–10 m), Shallow (10–25 m), Middle (25–50 m), Deep (50–100 m), and Very deep (> 100 m). Dispersal potential: Live birth, pelagic larvae/young, Demersal egg, no pelagic phase, Demersal egg, Balistid-type, demersal egg, Brooded egg, Semipelagic to Pelagic adults, Pelagic eggs and Unknown. IUCN Red list Categories: According to 2020 IUCN red list of threatened species, taxa are classified as: Not Evaluated (NE), Data Deficient (DD), Least Concern (LC), Near Threatened (NT), Vulnerable (VU), Endangered (EN), and Critically Endangered (CR). Trophic Categories: Species were classified according to their trophic categories based on literature (Ferreira et al., 2004; Pinheiro et al., 2018) and complementary data: Macro-carnivores, Mobile benthic invertivores/cleaners, Sessile invertivores, Omnivores, Herbivore/detritivores, Piscivores and Planktivores. Functional groups: Species were classified according to their functional groups based on the literature (Pinheiro et al., 2018) and complementary data: Macro-carnivores, Strict Piscivores, Mobile benthic invertivores/cleaners, Sessile invertivores, Sand invertivores, Spongivores, Diurnal planktivores, Nocturnal planktivores, Territorial Algae/detritus, Turf grazing, Scrapers, Excavator/eroders, Macroalgae browser and General omnivores.

A total of 325 fish species have been listed for MPA Costa dos Corais, including Chondrichthyes (28 species) and Actinopterygii (297). This total was represented by 81 families, being the most representative: Carangidae (23 species), Labridae (21), Gobiidae (15), Haemulidae (14), Carcharhinidae (11) and Gerreidae/Ephinephelidae (10) (Tab. 1; Tab. S1). According to depth categories, most species inhabited Middle (120 species) and Shallow (108) areas, while the remaining categories account for 97 species altogether (Fig. 3A).

Taxa were grouped according to the dispersion potential, the most abundant categories were: Pelagic egg (205), followed by Demersal egg (40), Live birth, pelagic larvae/young (31) and Semipelagic to Pelagic adults (27) (Fig. 3B).

Considering IUCN red list categories, most of the species were classified as Least Concern (254 species), followed by Data Deficient (23) and Near Threatened (17). Species classified within threatened categories were mainly Vulnerable (15). Only eight species were Endangered (5) and Critically Endangered (3) (Fig. 3C).

For trophic groups, the most abundant category was Mobile benthic invertivores/cleaners – MINV (133 species), followed by Macro-carnivores – MCAR (107) and Planktivores (29) (Fig. 3D).

Regarding functional groups, the most abundant category was Mobile benthic invertivores/cleaners (115 species), followed by Macro-carnivores (87), Diurnal planktivores (26), Strict Piscivores/Sand invertivores (19 each) and General omnivores (18) (Fig. 3E).

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Our checklist encompassed a total of 325 reef fish species at MPA Costa dos Corais. Pinheiro et al., (2018) recently compiled reef fish fauna at Southwestern Atlantic Ocean (SWA) with a total of 733 fish species. Our data represents 44% of all the SWA fish biodiversity inside the MPA territory. We highlight the vital large-scale significance of the largest Brazilian coastal MPA as one of the richest reef fish community on the SWA and reinforce the importance of this MPA on reef fish biodiversity and conservation. Additionally, we emphasize the importance of large scale and long-term surveys analyzing reef fish community compositions inside Brazilian MPAs. Initiatives as the present study, which often demonstrate unprecedented rich communities in otherwise poorly assessed and underestimated areas, are vital to enhance the effectiveness of MPAs acting as milestones for species monitoring and conservation (Figs. 4, 5, 6, 7, 8, 9, 10, 11, 12 and 13).

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Deeper reefs (> 30 m depth) have been considered less impacted from anthropogenic effects compared to shallow reefs (< 30 m depth) (Jankowski et al., 2015; Pereira et al., 2018); although it is not universal statement (Rocha et al., 2018). On the studied multiple use MPA, several species once frequently recorded on shallow areas such as barracudas (Sphyraena barracuda), snappers (Lutjanus spp.) and large parrotfishes (Scarus spp. and Sparisoma spp.) are currently recorded nearly unique at deeper reefs. For instance, Pereira et al., (2021) demonstrated by local ecological knowledge (LEK) that fishers used to catch the endemic and endangered parrotfish Scarus trispinosus on shallow reefs three decades ago. Yet, the species is currently rarely recorded at shallow sites; with the remaining populations inhabiting deeper reefs (Pereira et al., 2021). Additionally, fish behavior has been altered inside the MPA following human presence (tourism and/or fishing activity). Benevides et al., (2019) indicated that the zoning applied by APACC has a positive effect on the caution of target species, wherein in the tourist zone, where fishing is prohibited, fish allows a closer approach to the diver than in fishing area. Pereira et al., (2020) also suggested that fish species on shallow reefs tend to be less tolerant to human presence compared to individuals inhabiting deeper reefs. This could be supporting the idea that deeper reefs could be currently used by fish as “refuge” areas that are likely to be safeguarding both fish communities and species behavior.

Rezoning process on MPA Costa dos Corais has been recently conducted and published (ICMBio, 2021). During rezoning process, a series of new no-take zones have been selected and thoughtfully discussed with local communities to increase coral reefs conservation and effective local community engagement. However, it should be noted that there are controversies regarding the shared use of multiple use locations by fishers. These differences are mainly due to the dispute over territories between fishers and the tourism industry. Catamarans, speedboats, and jet skis are perceived by artisanal fishers as negative impacts that keep fish away and cause damage to fishing gear (Araújo et al., 2014; Zatan, 2019). In this context, the importance of the management plan aims to regulate nautical activity is appreciated, supporting biodiversity conservation, as well as strengthening artisanal fishing activity and community-based tourism (Zatan, 2019).

This no-take zones effort represents an increase of 900% in no-take zones inside the MPA Costa dos Corais that, together with local engagement, will represent a major conservation outcome on large scale reef fish conservation for the SWA Ocean. Most of these new no-take zones encompass both shallow and deeper reefs allowing a cross-shelf protection of reef biodiversity and ensuring connectivity of fish population between multiple habitats such as shallow and deeper reefs, seagrass and algae beds, mud/sand bottoms and rhodolith beds.

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A total of 12% (40 species) from the present study are registered on the 2020 IUCN red list as Near Threatened (NT), Vulnerable (VU), Endangered (EN) or Critically Endangered (CR). Additionally, 9% (29 species) are on Brazilian red list species (ICMBio, 2018). Overexploitation of many Brazilian reef fish species have been reported by several authors (Araújo, Martins, 2009; Bender et al., 2014; Di Dario et al., 2015; Pereira et al., 2021). Studies suggested that a total of 60% of the red‐listed marine species are primarily jeopardized by overfishing and the remaining 40% are threatened by habitat degradation and other non‐fisheries related impacts (Pinheiro et al., 2015). However, fisheries monitoring along Brazilian coast is nearly inexistent and only 0.8% of the coastal zone is included within no‐take areas (Di Dario et al., 2015; Vila‐Nova et al., 2014); regardless, consistent evidence of no‐take MPAs and strict no‐entry marine reserves demonstrating a fish recovery/spillover potential (Francini, Moura, 2008; Anderson et al., 2014; Motta et al., 2021; PHCP and collaborators, work in progress).

Coral cover and structural complexity have been described as an important variable influencing reef fish abundance and richness worldwide (Pereira, Munday, 2016; Darling et al., 2017). An increase on habitat complexity, food provision and shelter suggest that higher coral cover is likely to increase fish diversity (Leal et al., 2015; Mora, 2015). Previous studies have demonstrated up to 50% of coral cover on some areas of the MPA (PHCP and collaborators, work in progress), a much higher coral cover value compared to Brazilian coast with an average of 4.38% ± 8.17 (Aued et al., 2018). Hence, together with Abrolhos bank (up to 21% of coral cover) (Teixeira et al., 2021), MPA Costa dos Corais stands as one of highest coral cover areas on Brazilian coast. This trend likely to influence and explain such a high reef fish richness, featuring an unique and relevant site for fish biodiversity and conservation on the Southwestern Atlantic Ocean.

Brazilian marine biodiversity has been recently jeopardized due to a series of inconsistent governmental policies (Pinheiro et al., 2015; Miranda et al., 2020). Many SWA biodiversity hotspots are under risk such Fernando de Noronha Archipelago – UNESCO Heritage, Vitória‐Trindade Chain (VTC), Abrolhos Bank and MPA Costa dos Corais (Mazzei et al., 2017; Magris et al., 2020; Pimentel et al., 2020) that would compromise reef fish biodiversity on SWA. Hence, local strategies such as co-management, surveillance and local community compliance/engagement seems to be vital approaches for marine habitat conservation and maintenance of reef fish populations.