Reef fishes biodiversity and conservation at the largest Brazilian coastal Marine Protected Area (MPA Costa dos Corais)

Abstract

Coral reefs harbor one of the largest fish biodiversity on earth; yet information on reef fishes is still absent for many regions. We analyzed reef fish richness, distribution, and conservation on the largest Brazilian multiple use coastal MPA; which cover a large extent of coral reefs at the SWA. A total of 325 fish species have been listed for MPA Costa dos Corais, including Chondrichthyes (28 species) and Actinopterygii (297). Fish species were represented by 81 families and the most representative families were Carangidae (23 species), Labridae (21) and Gobiidae (15). The MPA fish richness represented 44% of all recorded fish species of the Southwestern Atlantic Ocean (SWA) highlighting the large-scale importance of this MPA. A total of 40 species (12%) are registered at Near Threatened (NT), Vulnerable (VU), Endangered (EN) or Critically Endangered (CR). This study reinforces the importance of MPA Costa dos Corais on reef fish biodiversity and conservation and emphasize the urgent need of conservation strategies.

Keywords:
Conservation Unit; Coral Reefs; Fish community; Ichthyofauna; Management Plan

Resumo

Os recifes de coral abrigam uma das maiores biodiversidades de peixes do planeta; no entanto, as informações sobre peixes de recife ainda estão ausentes em muitas regiões. Analisamos a riqueza, distribuição e a conservação de peixes recifais na maior Área de Proteção Ambiental (APA) costeira de uso múltiplo do Brasil; área que possui grande extensão de recifes de corais no SWA. Um total de 325 espécies de peixes foram listadas para APA Costa dos Corais, incluindo Chondrichthyes (28 espécies) e Actinopterygii (297). As espécies de peixes foram representadas por 81 famílias e as famílias mais representativas foram Carangidae (23 espécies), Labridae (21) e Gobiidae (15). A riqueza de peixes da APA representou 44% de todas as espécies de peixes registradas no Oceano Atlântico Sudoeste (SWA), destacando a importância em grande escala desta APA. Um total de 40 espécies (12%) estão registradas como Quase Ameaçada (NT), Vulnerável (VU), Em Perigo (EN) ou Criticamente Em Perigo (CR). Este estudo reforça a importância da APA Costa dos Corais na biodiversidade e conservação dos peixes recifais e enfatiza a necessidade urgente de estratégias de conservação.

Palavras-chave:
Comunidade de peixes; Ictiofauna; Plano de manejo; Recifes de coral; Unidade de conservação

INTRODUCTION

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.

MATERIAL AND METHODS

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).

FIGURE 1 |
Long-term sampling sites at Marine Protected Area (MPA) “Costa dos Corais” – South America.

FIGURE 2 |
Multiple habitats at MPA Costa dos Corais highlighting ecologically important areas for reef fishes biodiversity within the MPA. A. Deeper reefs (> 30 m depth) with predominance of the hard coral Montastraea cavernosa B. Shallow reefs (< 30 m depth) with dominance of hydrocoral Millepora alcicornis and Brazilian endemic brain coral Mussumila hartii. C. Sponge reefs with predominance of tube sponges from the genus Aplysina.

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.

RESULTS

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).

TABLE 1 |
Reef fishes biodiversity at the largest Brazilian coastal marine protection area (MPA Costa dos Corais). Depth category – VSHALL, 0–10 m; SHALL, 10–25 m; MID, 25–50 m; DEEP, 50–100 m; VDEEP, > 100 m. Dispersal potential - LIV: Live birth, pelagic larvae/Young; DNP: Demersal egg, no pelagic phase; DEG: Demersal egg; BAL, Balistid-type, demersal egg; BRO, Brooded egg; PAL: Semipelagic to Pelagic adults; PEL: Pelagic eggs; UK: Unknown. IUCN category - DD: Data deficient; LC: Least Concern; NT: Near Threatened; VU: Vulnerable; EN: Endangered; CR: Critically Endangered; NE: Not evaluated. Trophic category - MCAR: Macro-carnivores; MINV: Mobile benthic invertivores/cleaners; SINV: Sessile invertivores; OMNI: Omnivores; HERB: Herbivore/detritivores; PLANK: Planktivores. Functional group - MCAR: Macro-carnivores; PIS: Strict Piscivores; MINV: Mobile benthic invertivores/cleaners; SINV: Sessile invertivores; SAND: Sand invertivores; SPON: Spongivore; DPLA: Diurnal planktivores; NPLA: Nocturnal planktivores; THER: Territorial Algae/detritos; TRUF: Turf grazing; SCRP: Scrapers; EXCV: Excavator/eroders; MALG: Macroalgae browser; OMNI: General omnivores.

Number	Group	Family	Species	Authorship	Depth category	Dispersal potential	IUCN category	Trophic category	Functional group
1	Chondrichthyes	Ginglymostomatidae	Ginglymostoma cirratum	(Bonnaterre, 1788)	MID	LIV	DD	MCAR	MCAR
2	Chondrichthyes	Rhincodontidae	Rhincodon typus	Smith, 1828	VDEEP	LIV	EN	PLANK	DPLA
3	Chondrichthyes	Triakidae	Mustelus higmani	Springer & Lowe, 1963	SHALL	LIV	LC	MINV	MINV
4	Chondrichthyes	Carcharhinidae	Carcharhinus acronotus	(Poey, 1860)	DEEP	LIV	NT	MCAR	MCAR
5	Chondrichthyes	Carcharhinidae	Carcharhinus falciformis	(Bibron, 1839)	VDEEP	LIV	VU	MCAR	MCAR
6	Chondrichthyes	Carcharhinidae	Carcharhinus leucas	(Valenciennes, 1839)	MID	LIV	NT	MCAR	MCAR
7	Chondrichthyes	Carcharhinidae	Carcharhinus limbatus	(Valenciennes, 1839)	VDEEP	LIV	NT	MCAR	MCAR
8	Chondrichthyes	Carcharhinidae	Carcharhinus obscurus	(LeSueur, 1818)	VDEEP	LIV	EN	MCAR	MCAR
9	Chondrichthyes	Carcharhinidae	Carcharhinus perezi	(Poey, 1876)	MID	LIV	NT	MCAR	MCAR
10	Chondrichthyes	Carcharhinidae	Carcharhinus plumbeus	(Nardo, 1827)	DEEP	LIV	VU	MCAR	MCAR
11	Chondrichthyes	Galeocerdonidae	Galeocerdo cuvier	(Péron & LeSueur, 1822)	DEEP	LIV	NT	MCAR	MCAR
12	Chondrichthyes	Carcharhinidae	Negaprion brevirostris	(Poey, 1868)	MID	LIV	NT	MCAR	MCAR
13	Chondrichthyes	Carcharhinidae	Rhizoprionodon lalandii	(Valenciennes, 1839)	MID	LIV	DD	MCAR	MCAR
14	Chondrichthyes	Carcharhinidae	Rhizoprionodon porosus	(Poey, 1861)	MID	LIV	LC	MCAR	MCAR
15	Chondrichthyes	Sphyrnidae	Sphyrna lewini	(Griffith & Smith, 1834)	VDEEP	LIV	CR	MCAR	MCAR
16	Chondrichthyes	Sphyrnidae	Sphyrna mokarran	(Rüppel, 1837)	VDEEP	LIV	CR	MCAR	MCAR
17	Chondrichthyes	Sphyrnidae	Sphyrna tiburo	(Linnaeus, 1758)	SHALL	LIV	LC	MCAR	MCAR
18	Chondrichthyes	Pristidae	Pristis pristis	(Linnaeus, 1758)	SHALL	LIV	CR	MCAR	MCAR
19	Chondrichthyes	Dasyatidae	Hypanus berthalutzae	Petean, Naylor & Lima, 2020	SHALL	LIV	DD	MINV	SAND
20	Chondrichthyes	Dasyatidae	Hypanus guttatus	(Bloch & Schneider, 1801)	SHALL	LIV	DD	MINV	SAND
21	Chondrichthyes	Dasyatidae	Hypanus marianae	(Gomes, Rosa & Gadig, 2000)	SHALL	LIV	DD	MINV	SAND
22	Chondrichthyes	Gymnuridae	Gymnura micrura	(Bloch & Schneider, 1801)	MID	LIV	DD	MINV	MINV
23	Chondrichthyes	Urotrygonidae	Urotrygon microphthalmum	Delsman, 1941	MID	LIV	LC	MINV	SAND
24	Chondrichthyes	Mobulidae	Mobula birostris	(Walbaum, 1792)	DEEP	PEL	VU	PLANK	DPLA
25	Chondrichthyes	Mobulidae	Mobula mobular	(Bonnaterre, 1788)	DEEP	PEL	EN	PLANK	DPLA
26	Chondrichthyes	Mobulidae	Mobula tarapacana	(Philippi, 1892)	DEEP	PEL	EN	PLANK	DPLA
27	Chondrichthyes	Aetobatidae	Aetobatus narinari	(Euprasen, 1790)	MID	LIV	NT	MCAR	SAND
28	Chondrichthyes	Rhinopteridae	Rhinoptera bonasus	(Mitchill, 1815)	VDEEP	LIV	NT	MINV	SAND
29	Osteichthyes	Elopidae	Elops saurus	Linnaeus, 1766	MID	PEL	LC	MCAR	MCAR
30	Osteichthyes	Megalopidae	Megalops atlanticus	Valenciennes, 1847	MID	PEL	VU	MCAR	MCAR
31	Osteichthyes	Albulidae	Albula nemoptera	(Fowler, 1911)	MID	PEL	DD	MINV	SAND
32	Osteichthyes	Albulidae	Albula vulpes	(Linnaeus, 1758)	SHALL	PEL	NT	MINV	SAND
33	Osteichthyes	Muraenidae	Enchelycore carychroa	Böhlke & Böhlke, 1976	MID	PEL	LC	MCAR	MCAR
34	Osteichthyes	Muraenidae	Enchelycore nigricans	(Bonnaterre, 1788)	SHALL	PEL	LC	MCAR	MCAR
35	Osteichthyes	Muraenidae	Gymnothorax funebris	Ranzani, 1839	SHALL	PEL	LC	MCAR	MCAR
36	Osteichthyes	Muraenidae	Gymnothorax miliaris	(Kaup, 1856)	SHALL	PEL	LC	MCAR	MCAR
37	Osteichthyes	Muraenidae	Gymnothorax moringa	(Cuvier, 1829)	SHALL	PEL	LC	MCAR	MCAR
38	Osteichthyes	Muraenidae	Gymnothorax ocellatus	Agassiz, 1831	SHALL	PEL	LC	MCAR	MCAR
39	Osteichthyes	Muraenidae	Gymnothorax vicinus	(Castelnau, 1855)	SHALL	PEL	LC	MCAR	MCAR
40	Osteichthyes	Muraenidae	Muraena pavonina	Richardson, 1845	SHALL	PEL	LC	MCAR	MCAR
41	Osteichthyes	Ophichthidae	Ahlia egmontis	(Jordan, 1884)	VSHALL	PEL	LC	MINV	MINV
42	Osteichthyes	Ophichthidae	Myrichthys breviceps	(Richardson, 1848)	VSHALL	PEL	LC	MINV	MINV
43	Osteichthyes	Ophichthidae	Myrichthys ocellatus	(Lesueur, 1825)	VSHALL	PEL	LC	MINV	MINV
44	Osteichthyes	Ophichthidae	Ophichthus cylindroideus	(Ranzani,1839)	SHALL	PEL	LC	MINV	MINV
45	Osteichthyes	Ophichthidae	Ophichthus ophis	(Linnaeus, 1758)	VSHALL	LIV	LC	MINV	MINV
46	Osteichthyes	Congridae	Heteroconger camelopardalis	(Lubbock, 1980)	MID	PEL	LC	PLANK	DPLA
47	Osteichthyes	Engraulidae	Anchoa filifera	(Fowler, 1915)	SHALL	PEL	LC	PLANK	DPLA
48	Osteichthyes	Engraulidae	Anchoa januaria	(Steindachner, 1879)	SHALL	PEL	LC	PLANK	DPLA
49	Osteichthyes	Engraulidae	Anchoa spinifer	(Valenciennes, 1848)	SHALL	PEL	LC	PLANK	DPLA
50	Osteichthyes	Engraulidae	Anchoa tricolor	(Spix & Agassiz, 1829)	SHALL	PEL	LC	PLANK	DPLA
51	Osteichthyes	Engraulidae	Anchovia clupeoides	(Swainson, 1839)	SHALL	PEL	LC	PLANK	DPLA
52	Osteichthyes	Engraulidae	Anchoviella lepidentostole	(Fowler, 1911)	SHALL	PEL	LC	PLANK	DPLA
53	Osteichthyes	Engraulidae	Lycengraulis grossidens	(Spix & Agassiz, 1829)	SHALL	PEL	LC	PLANK	DPLA
54	Osteichthyes	Ariidae	Bagre marinus	(Mitchill, 1815)	MID	BRO	LC	MCAR	MCAR
55	Osteichthyes	Synodontidae	Synodus foetens	(Linnaeus, 1766)	VSHALL	PEL	LC	MCAR	MCAR
56	Osteichthyes	Synodontidae	Synodus intermedius	(Spix & Agassiz, 1829)	VSHALL	PEL	LC	MCAR	MCAR
57	Osteichthyes	Synodontidae	Synodus synodus	(Linnaeus, 1758)	VSHALL	PEL	LC	MCAR	MCAR
58	Osteichthyes	Synodontidae	Trachinocephalus myops	(Forster, 1801)	MID	PEL	LC	MCAR	MCAR
59	Osteichthyes	Holocentridae	Holocentrus adscensionis	(Osbeck, 1765)	SHALL	PEL	LC	MINV	MINV
60	Osteichthyes	Holocentridae	Myripristis jacobus	Cuvier, 1829	SHALL	PEL	LC	MINV	MINV
61	Osteichthyes	Holocentridae	Plectrypops retrospinis	(Guichenot, 1853)	MID	PEL	LC	MINV	MINV
62	Osteichthyes	Bythitidae	Petrotyx sanguineus	(Meek & Hildebrand, 1928)	MID	PEL	LC	MINV	MINV
63	Osteichthyes	Batrachoididae	Amphichthys cryptocentrus	(Valenciennes, 1837)	SHALL	DNP	LC	MCAR	MCAR
64	Osteichthyes	Batrachoididae	Thalassophryne nattereri	Steindachner, 1876	SHALL	DNP	LC	MINV	MINV
65	Osteichthyes	Apogonidae	Apogon americanus	Castelnau, 1855	SHALL	BRO	NE	PLANK	NPLA
66	Osteichthyes	Apogonidae	Astrapogon puncticulatus	(Poey, 1867)	SHALL	BRO	LC	PLANK	NPLA
67	Osteichthyes	Apogonidae	Phaeoptyx pigmentaria	(Poey, 1860)	SHALL	BRO	LC	PLANK	NPLA
68	Osteichthyes	Eleotridae	Eleotris pisonis	(Gmelin, 1789)	SHALL	PEL	LC	MINV	MINV
69	Osteichthyes	Eleotridae	Erotelis smaragdus	(Valenciennes, 1837)	SHALL	PEL	LC	MINV	MINV
70	Osteichthyes	Gobiidae	Barbulifer ceuthoecus	(Jordan & Gilbert, 1884)	VSHALL	DEG	LC	MINV	MINV
71	Osteichthyes	Gobiidae	Bathygobius geminatus	Tornabene, Baldwin & Pezold, 2010	VSHALL	DEG	DD	MINV	MINV
72	Osteichthyes	Gobiidae	Bathygobius soporator	(Valenciennes, 1837)	VSHALL	DEG	LC	MINV	MINV
73	Osteichthyes	Gobiidae	Coryphopterus dicrus	Böhlke & Robins, 1960	VSHALL	DEG	LC	MINV	MINV
74	Osteichthyes	Gobiidae	Coryphopterus glaucofraenum	Gill, 1863	SHALL	DEG	LC	MINV	MINV
75	Osteichthyes	Gobiidae	Ctenogobius boleosoma	(Jordan & Gilbert, 1882)	SHALL	DEG	LC	MINV	MINV
76	Osteichthyes	Gobiidae	Ctenogobius saepepallens	(Gilbert & Randall, 1968)	VSHALL	DEG	LC	MINV	MINV
77	Osteichthyes	Gobiidae	Elacatinus figaro	Sazima, Moura & Rosa, 1997	MID	DEG	VU	MINV	MINV
78	Osteichthyes	Gobiidae	Gnatholepis thompsoni	Jordan, 1904	SHALL	DEG	LC	MINV	MINV
79	Osteichthyes	Gobiesocidae	Gobiesox barbatulus	Starks, 1913	VSHALL	DEG	LC	MINV	MINV
80	Osteichthyes	Gobiesocidae	Gobiesox strumosus	Cope, 1870	VSHALL	DEG	LC	MINV	MINV
81	Osteichthyes	Gobiidae	Gobionellus stomatus	Starks, 1913	SHALL	DEG	LC	MINV	MINV
82	Osteichthyes	Gobiidae	Lythrypnus brasiliensis	Greenfield, 1988	SHALL	DEG	NE	MINV	MINV
83	Osteichthyes	Gobiidae	Microgobius carri	Fowler, 1945	SHALL	DEG	LC	MINV	MINV
84	Osteichthyes	Microdesmidae	Ptereleotris randalli	Gasparini, Rocha & Floeter, 2001	MID	DEG	LC	PLANK	DPLA
85	Osteichthyes	Grammatidae	Gramma brasiliensis	Sazima, Gasparini & Moura, 1998	MID	BRO	NT	MINV	MINV
86	Osteichthyes	Pomacanthidae	Centropyge aurantonotus	Burgess, 1974	VDEEP	PEL	LC	HERB	THER
87	Osteichthyes	Pomacanthidae	Holacanthus ciliaris	(Linnaeus, 1758)	MID	PEL	LC	SINV	SPON
88	Osteichthyes	Pomacanthidae	Holacanthus tricolor	(Bloch, 1795)	MID	PEL	LC	SINV	SPON
89	Osteichthyes	Pomacanthidae	Pomacanthus arcuatus	(Linnaeus, 1758)	MID	PEL	LC	SINV	SPON
90	Osteichthyes	Pomacanthidae	Pomacanthus paru	(Bloch, 1787)	SHALL	PEL	LC	SINV	SPON
91	Osteichthyes	Pomacentridae	Abudefduf saxatilis	(Linnaeus, 1758)	SHALL	DEG	LC	OMNI	OMNI
92	Osteichthyes	Pomacentridae	Chromis flavicauda	(Günther, 1880)	VDEEP	PEL	DD	PLANK	DPLA
93	Osteichthyes	Pomacentridae	Chromis jubauna	Moura, 1995	VDEEP	PEL	NE	PLANK	DPLA
94	Osteichthyes	Pomacentridae	Azurina multilineata	(Guichenot, 1853)	SHALL	PEL	LC	PLANK	DPLA
95	Osteichthyes	Pomacentridae	Microspathodon chrysurus	(Cuvier, 1830)	SHALL	DEG	LC	HERB	THER
96	Osteichthyes	Pomacentridae	Stegastes fuscus	(Cuvier, 1830)	SHALL	DEG	LC	HERB	THER
97	Osteichthyes	Pomacentridae	Stegastes pictus	(Castelnau, 1855)	MID	DEG	NE	HERB	THER
98	Osteichthyes	Pomacentridae	Stegastes variabilis	(Castelnau, 1855)	SHALL	DEG	LC	HERB	THER
99	Osteichthyes	Mugilidae	Mugil curema	Valenciennes, 1836	SHALL	PEL	LC	HERB	TURF
100	Osteichthyes	Mugilidae	Mugil curvidens	Valenciennes, 1836	SHALL	PEL	LC	HERB	TURF
101	Osteichthyes	Mugilidae	Mugil liza	Valenciennes, 1836	SHALL	PEL	DD	HERB	TURF
102	Osteichthyes	Mugilidae	Mugil rubrioculus	Harrison, Nirchio, Oliveira, Ron & Gavíria, 2007	SHALL	PEL	LC	HERB	TURF
103	Osteichthyes	Mugilidae	Mugil trichodon	Poey, 1875	SHALL	PEL	LC	HERB	TURF
104	Osteichthyes	Polynemidae	Polydactylus virginicus	(Linnaeus, 1758)	SHALL	PEL	LC	OMNI	OMNI
105	Osteichthyes	Tripterygiidae	Enneanectes altivelis	Rosenblatt, 1960	SHALL	DEG	LC	MINV	MINV
106	Osteichthyes	Dactylopteridae	Dactylopterus volitans	(Linnaeus, 1758)	MID	BRO	LC	MINV	SAND
107	Osteichthyes	Dactyloscopidae	Dactyloscopus foraminosus	Dawson, 1982	SHALL	BRO	LC	MINV	SAND
108	Osteichthyes	Dactyloscopidae	Dactyloscopus tridigitatus	Gill, 1859	SHALL	BRO	LC	MINV	SAND
109	Osteichthyes	Dactyloscopidae	Platygillellus brasiliensis	Feitoza, 2002	SHALL	BRO	LC	MINV	SAND
110	Osteichthyes	Labrisomidae	Gobioclinus kalisherae	(Jordan, 1904)	VSHALL	DEG	LC	MINV	MINV
111	Osteichthyes	Labrisomidae	Labrisomus cricota	Sazima, Gasparini & Moura, 2002	VSHALL	PEL	LC	MINV	MINV
112	Osteichthyes	Labrisomidae	Labrisomus nuchipinnis	(Quoy & Gaimard, 1824)	VSHALL	DEG	LC	MINV	MINV
113	Osteichthyes	Labrisomidae	Malacoctenus delalandii	(Valenciennes, 1836)	VSHALL	DEG	LC	MINV	MINV
114	Osteichthyes	Labrisomidae	Malacoctenus zaluari	Carvalho-Filho, Gasparini & Sazima, 2020	VSHALL	DEG	LC	MINV	MINV
115	Osteichthyes	Labrisomidae	Paraclinus spectator	Guimarães & Bacelar, 2002	VSHALL	PEL	LC	MINV	MINV
116	Osteichthyes	Labrisomidae	Starksia brasiliensis	(Gilbert, 1900)	VSHALL	PEL	LC	MINV	MINV
117	Osteichthyes	Chaenopsidae	Emblemariopsis signifer	(Ginsburg, 1942)	SHALL	DEG	LC	MCAR	MCAR
118	Osteichthyes	Atherinopsidae	Atherinella brasiliensis	(Quoy & Gaimard, 1825)	SHALL	PEL	LC	PLANK	DPLA
119	Osteichthyes	Hemiramphidae	Hemiramphus brasiliensis	(Linnaeus, 1758)	SHALL	PEL	LC	OMNI	OMNI
120	Osteichthyes	Hemiramphidae	Hyporhamphus roberti	(Valenciennes, 1847)	SHALL	PEL	LC	MCAR	MCAR
121	Osteichthyes	Hemiramphidae	Hyporhamphus unifasciatus	(Ranzani, 1841)	SHALL	PEL	LC	MCAR	MCAR
122	Osteichthyes	Belonidae	Strongylura timucu	(Walbaum, 1792)	SHALL	PEL	LC	MCAR	MCAR
123	Osteichthyes	Belonidae	Tylosurus acus	(Lacepède, 1803)	MID	PEL	LC	MCAR	PISC
124	Osteichthyes	Belonidae	Tylosurus crocodilus	(Péron & Lesueur, 1821)	MID	PEL	LC	MCAR	PISC
125	Osteichthyes	Blenniidae	Entomacrodus vomerinus	(Valenciennes, 1836)	SHALL	DEG	LC	HERB	THER
126	Osteichthyes	Blenniidae	Hypleurochilus fissicornis	(Quoy & Gaimard, 1824)	SHALL	DEG	LC	MINV	MINV
127	Osteichthyes	Blenniidae	Hypleurochilus aequipinnis	(Günther, 1861)	SHALL	DEG	LC	MINV	MINV
128	Osteichthyes	Blenniidae	Ophioblennius trinitatis	Miranda Ribeiro, 1919	SHALL	DEG	LC	HERB	THER
129	Osteichthyes	Blenniidae	Parablennius marmoreus	(Poey, 1876)	SHALL	DEG	LC	OMNI	OMNI
130	Osteichthyes	Blenniidae	Parablennius pilicornis	(Cuvier, 1829)	SHALL	DEG	LC	OMNI	OMNI
131	Osteichthyes	Blenniidae	Scartella cristata	(Linnaeus, 1758)	SHALL	DEG	LC	HERB	THER
132	Osteichthyes	Rachycentridae	Rachycentron canadum	(Linnaeus, 1766)	MID	PEL	LC	MCAR	MCAR
133	Osteichthyes	Echeneidae	Echeneis naucrates	Linnaeus, 1758	MID	PEL	DD	MCAR	MCAR
134	Osteichthyes	Carangidae	Alectis ciliaris	(Bloch, 1787)	MID	PAD	LC	MCAR	MCAR
135	Osteichthyes	Carangidae	Caranx bartholomaei	Cuvier, 1833	MID	PAD	LC	MCAR	PISC
136	Osteichthyes	Carangidae	Caranx ruber	(Bloch, 1793)	DEEP	PAD	LC	MCAR	MCAR
137	Osteichthyes	Carangidae	Caranx crysos	(Mitchill, 1815)	MID	PAD	LC	MCAR	MCAR
138	Osteichthyes	Carangidae	Caranx hippos	(Linnaeus, 1766)	MID	PAD	LC	MCAR	MCAR
139	Osteichthyes	Carangidae	Caranx latus	Agassiz, 1831	MID	PAD	LC	MCAR	MCAR
140	Osteichthyes	Carangidae	Caranx lugubris	Poey, 1860	VDEEP	PAD	LC	MCAR	PISC
141	Osteichthyes	Carangidae	Chloroscombrus chrysurus	(Linnaeus, 1766)	MID	PAD	LC	PLANK	DPLA
142	Osteichthyes	Carangidae	Decapterus macarellus	(Cuvier, 1833)	MID	PAD	LC	MCAR	MCAR
143	Osteichthyes	Carangidae	Decapterus punctatus	(Cuvier, 1829)	MID	PAD	LC	MCAR	MCAR
144	Osteichthyes	Carangidae	Decapterus tabl	Berry, 1968	DEEP	PAD	LC	MCAR	MCAR
145	Osteichthyes	Carangidae	Elagatis bipinnulata	(Quoy & Gaimard, 1825)	DEEP	PAD	LC	MCAR	PISC
146	Osteichthyes	Carangidae	Oligoplites saliens	(Bloch, 1793)	MID	PAD	LC	MCAR	MCAR
147	Osteichthyes	Carangidae	Oligoplites saurus	(Bloch & Schneider, 1801)	MID	PAD	LC	MCAR	MCAR
148	Osteichthyes	Carangidae	Selar crumenophthalmus	(Bloch, 1793)	MID	PAD	LC	PLANK	DPLA
149	Osteichthyes	Carangidae	Selene brownii	(Cuvier, 1816)	MID	PAD	LC	MCAR	MCAR
150	Osteichthyes	Carangidae	Selene setapinnis	(Mitchil, 1815)	MID	PAD	LC	MCAR	MCAR
151	Osteichthyes	Carangidae	Selene vomer	(Linnaeus, 1758)	MID	PAD	LC	MCAR	MCAR
152	Osteichthyes	Carangidae	Seriola dumerili	(Risso, 1810)	DEEP	PAD	LC	MCAR	PISC
153	Osteichthyes	Carangidae	Seriola rivoliana	Valenciennes, 1833	VDEEP	PAD	LC	MCAR	PISC
154	Osteichthyes	Carangidae	Trachinotus carolinus	(Linnaeus, 1766)	MID	PAD	LC	MINV	MINV
155	Osteichthyes	Carangidae	Trachinotus falcatus	(Linnaeus, 1758)	MID	PAD	LC	MINV	MINV
156	Osteichthyes	Carangidae	Trachinotus goodei	Jordan & Evermann, 1896	VSHALL	PAD	LC	MINV	MINV
157	Osteichthyes	Sphyraenidae	Sphyraena barracuda	(Edwards, 1771)	MID	PEL	LC	MCAR	MCAR
158	Osteichthyes	Sphyraenidae	Sphyraena guachancho	Cuvier, 1829	MID	PEL	LC	MCAR	MCAR
159	Osteichthyes	Paralichthyidae	Citharichthys arenaceus	Evermann & Marsh, 1900	SHALL	PEL	LC	MINV	MINV
160	Osteichthyes	Paralichthyidae	Citharichthys spilopterus	Günther, 1862	SHALL	PEL	LC	MINV	MINV
161	Osteichthyes	Paralichthyidae	Paralichthys brasiliensis	(Ranzani, 1842)	SHALL	PEL	NE	MINV	MINV
162	Osteichthyes	Paralichthyidae	Syacium micrurum	Ranzani, 1842	SHALL	PEL	LC	MINV	MINV
163	Osteichthyes	Paralichthyidae	Syacium papillosum	(Linnaeus, 1758)	SHALL	PEL	LC	MINV	MINV
164	Osteichthyes	Bothidae	Bothus lunatus	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
165	Osteichthyes	Bothidae	Bothus ocellatus	(Agassiz, 1831)	DEEP	BRO	LC	MINV	MINV
166	Osteichthyes	Achiridae	Achirus lineatus	(Linnaeus, 1758)	SHALL	PEL	LC	MINV	SAND
167	Osteichthyes	Achiridae	Gymnachirus nudus	Kaup, 1858	MID	PEL	LC	MINV	SAND
168	Osteichthyes	Cynoglossidae	Symphurus diomedeanus	(Goode & Bean, 1885)	MID	PEL	LC	MINV	MINV
169	Osteichthyes	Cynoglossidae	Symphurus tessellatus	(Quoy & Gaimard, 1824)	MID	PEL	LC	MINV	MINV
170	Osteichthyes	Syngnathidae	Hippocampus erectus	Perry, 1810	VSHALL	LIV	VU	MINV	MINV
171	Osteichthyes	Syngnathidae	Hippocampus patagonicus	Piacentino & Luzzatto, 2004	VSHALL	LIV	VU	MINV	MINV
172	Osteichthyes	Syngnathidae	Hippocampus reidi	Ginsburg, 1933	VSHALL	LIV	NT	MINV	MINV
173	Osteichthyes	Syngnathidae	Halicampus crinitus	(Jenyns, 1842)	VSHALL	PEL	LC	MINV	MINV
174	Osteichthyes	Syngnathidae	Microphis lineatus	(Kaup, 1856)	VSHALL	PEL	DD	MINV	MINV
175	Osteichthyes	Syngnathidae	Syngnathus pelagicus	Linnaeus, 1758	VSHALL	PEL	LC	MINV	MINV
176	Osteichthyes	Aulostomidae	Aulostomus strigosus	Wheeler, 1955	MID	BAL	LC	MCAR	PISC
177	Osteichthyes	Fistulariidae	Fistularia tabacaria	Linnaeus, 1758	SHALL	DEG	LC	MCAR	PISC
178	Osteichthyes	Trichiuridae	Trichiurus lepturus	Linnaeus, 1758	MID	PEL	LC	MCAR	MCAR
179	Osteichthyes	Scombridae	Acanthocybium solandri	(Cuvier, 1832)	VDEEP	PEL	LC	MCAR	PISC
180	Osteichthyes	Scombridae	Auxis thazard	(Lacepède, 1800)	MID	PAD	LC	MCAR	MCAR
181	Osteichthyes	Scombridae	Euthynnus alletteratus	(Rafinesque, 1810)	MID	PAD	LC	MCAR	MCAR
182	Osteichthyes	Scombridae	Scomberomorus brasiliensis	Collette, Russo & Zavala-Camin, 1978	MID	PEL	LC	MCAR	PISC
183	Osteichthyes	Scombridae	Scomberomorus cavalla	(Cuvier, 1829)	MID	PEL	LC	MCAR	PISC
184	Osteichthyes	Scombridae	Scomberomorus regalis	(Bloch, 1793)	MID	PEL	LC	MCAR	PISC
185	Osteichthyes	Scombridae	Thunnus albacares	(Bonnaterre, 1788)	VDEEP	PEL	NT	MCAR	PISC
186	Osteichthyes	Scombridae	Thunnus atlanticus	(Lesson, 1831)	VDEEP	PEL	LC	MCAR	PISC
187	Osteichthyes	Scombridae	Thunnus obesus	(Lowe, 1839)	VDEEP	PEL	VU	MCAR	PISC
188	Osteichthyes	Labridae	Bodianus pulchellus	(Poey, 1860)	VDEEP	PEL	LC	MINV	MINV
189	Osteichthyes	Labridae	Bodianus rufus	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
190	Osteichthyes	Labridae	Clepticus brasiliensis	Heiser, Moura & Robertson, 2000	DEEP	PEL	LC	PLANK	DPLA
191	Osteichthyes	Labridae	Doratonotus megalepis	Günther, 1862	SHALL	PEL	LC	MINV	MINV
192	Osteichthyes	Labridae	Halichoeres bivittatus	(Bloch, 1791)	MID	PEL	LC	MINV	MINV
193	Osteichthyes	Labridae	Halichoeres brasiliensis	(Bloch, 1791)	SHALL	PEL	DD	MINV	MINV
194	Osteichthyes	Labridae	Halichoeres dimidiatus	(Agassiz, 1831)	MID	PEL	LC	MINV	MINV
195	Osteichthyes	Labridae	Halichoeres penrosei	Starks, 1913	SHALL	PEL	LC	MINV	MINV
196	Osteichthyes	Labridae	Halichoeres poeyi	(Steindachner, 1867)	SHALL	DEG	LC	MINV	MINV
197	Osteichthyes	Labridae	Lachnolaimus maximus	(Walbaum, 1792)	SHALL	PEL	VU	MINV	MINV
198	Osteichthyes	Labridae	Thalassoma noronhanum	(Boulenger, 1890)	SHALL	PEL	LC	PLANK	DPLA
199	Osteichthyes	Labridae	Xyrichtys novacula	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
200	Osteichthyes	Labridae	Xyrichtys splendens	Castelnau, 1855	MID	PEL	LC	MINV	MINV
201	Osteichthyes	Scaridae	Cryptotomus roseus	Cope, 1871	SHALL	DEG	LC	HERB	SCRP
202	Osteichthyes	Scaridae	Nicholsina usta	(Valenciennes, 1840)	SHALL	DEG	LC	HERB	EXCV
203	Osteichthyes	Scaridae	Scarus trispinosus	Valenciennes, 1840	MID	DEG	EN	HERB	EXCV
204	Osteichthyes	Scaridae	Scarus zelindae	Moura, Figueiredo & Sazima, 2001	MID	PEL	DD	HERB	SCRP
205	Osteichthyes	Scaridae	Sparisoma amplum	(Ranzani, 1841)	MID	PEL	LC	HERB	SCRP
206	Osteichthyes	Scaridae	Sparisoma axillare	(Steindachner, 1878)	MID	PEL	DD	HERB	SCRP
207	Osteichthyes	Scaridae	Sparisoma frondosum	(Agassiz, 1831)	MID	PEL	DD	HERB	SCRP
208	Osteichthyes	Scaridae	Sparisoma radians	(Valenciennes, 1840)	SHALL	PEL	LC	HERB	MALG
209	Osteichthyes	Centropomidae	Centropomus parallelus	Poey, 1860	SHALL	PEL	LC	MCAR	MCAR
210	Osteichthyes	Centropomidae	Centropomus undecimalis	(Bloch, 1792)	SHALL	PEL	LC	MCAR	MCAR
211	Osteichthyes	Centropomidae	Centropomus pectinatus	Poey, 1860	SHALL	PEL	LC	MCAR	MCAR
212	Osteichthyes	Gerreidae	Diapterus auratus	Ranzani, 1842	VSHALL	PEL	LC	MINV	MINV
213	Osteichthyes	Gerreidae	Diapterus rhombeus	(Cuvier, 1829)	VSHALL	PEL	LC	MINV	MINV
214	Osteichthyes	Gerreidae	Eucinostomus argenteus	Baird & Girard, 1855	VSHALL	PEL	LC	MINV	MINV
215	Osteichthyes	Gerreidae	Eucinostomus gula	(Quoy & Gaimard, 1824)	VSHALL	PEL	LC	MINV	MINV
216	Osteichthyes	Gerreidae	Eucinostomus havana	(Nichols, 1912)	VSHALL	PEL	LC	MINV	MINV
217	Osteichthyes	Gerreidae	Eucinostomus lefroyi	(Goode, 1874)	VSHALL	PEL	LC	MINV	MINV
218	Osteichthyes	Gerreidae	Eucinostomus melanopterus	(Bleeker, 1863)	VSHALL	PEL	LC	MINV	MINV
219	Osteichthyes	Gerreidae	Eugerres brasilianus	(Cuvier, 1830)	VSHALL	PEL	LC	MINV	MINV
220	Osteichthyes	Gerreidae	Gerres cinereus	(Walbaum, 1792)	VSHALL	PEL	LC	MINV	MINV
221	Osteichthyes	Gerreidae	Ulaema lefroyi	(Goode, 1874)	VSHALL	PEL	LC	MINV	MINV
222	Osteichthyes	Mullidae	Mulloidichthys martinicus	(Cuvier, 1829)	MID	PEL	LC	MINV	SAND
223	Osteichthyes	Mullidae	Pseudupeneus maculatus	(Bloch, 1793)	MID	PEL	LC	MINV	SAND
224	Osteichthyes	Mullidae	Upeneus parvus	Poey, 1852	VDEEP	PEL	LC	MINV	SAND
225	Osteichthyes	Pempheridae	Pempheris schomburgkii	Müller & Troschel, 1848	SHALL	PEL	LC	PLANK	DPLA
226	Osteichthyes	Clupeidae	Harengula clupeola	(Cuvier, 1829)	SHALL	PEL	LC	PLANK	DPLA
227	Osteichthyes	Clupeidae	Lile piquitinga	(Schreiner & Miranda Ribeiro, 1903)	SHALL	PEL	LC	PLANK	DPLA
228	Osteichthyes	Clupeidae	Opisthonema oglinum	(Lesueur, 1818)	SHALL	PEL	LC	PLANK	DPLA
229	Osteichthyes	Kyphosidae	Kyphosus sectatrix	(Linnaeus, 1758)	MID	PEL	LC	HERB	MALG
230	Osteichthyes	Serranidae	Alphestes afer	(Bloch, 1793)	SHALL	PEL	LC	MCAR	MCAR
231	Osteichthyes	Serranidae	Cephalopholis fulva	(Linnaeus, 1758)	MID	PEL	LC	MCAR	MCAR
232	Osteichthyes	Serranidae	Epinephelus adscensionis	(Osbeck, 1765)	MID	PEL	LC	MCAR	MCAR
233	Osteichthyes	Serranidae	Epinephelus itajara	(Lichtenstein, 1822)	DEEP	PEL	VU	MCAR	MCAR
234	Osteichthyes	Serranidae	Epinephelus morio	(Valenciennes, 1828)	VDEEP	PEL	VU	MCAR	MCAR
235	Osteichthyes	Serranidae	Gonioplectrus hispanus	(Cuvier, 1828)	VDEEP	PEL	LC	MCAR	MCAR
236	Osteichthyes	Serranidae	Hyporthodus niveatus	(Valenciennes, 1828)	VDEEP	PEL	VU	MCAR	MCAR
237	Osteichthyes	Serranidae	Mycteroperca bonaci	(Poey, 1860)	DEEP	DEG	NT	MCAR	PISC
238	Osteichthyes	Serranidae	Mycteroperca interstitialis	(Poey, 1860)	VDEEP	PAD	VU	MCAR	PISC
239	Osteichthyes	Serranidae	Mycteroperca venenosa	(Linnaeus, 1758)	VDEEP	PAD	NT	MCAR	PISC
240	Osteichthyes	Serranidae	Paranthias furcifer	(Valenciennes, 1828)	MID	PEL	LC	PLANK	DPLA
241	Osteichthyes	Serranidae	Dermatolepis inermis	(Valenciennes, 1833)	VDEEP	PEL	DD	MCAR	MCAR
242	Osteichthyes	Serranidae	Diplectrum formosum	(Linnaeus, 1766)	MID	PEL	LC	MCAR	MCAR
243	Osteichthyes	Serranidae	Diplectrum radiale	(Quoy & Gaimard, 1824)	MID	PEL	LC	MCAR	MCAR
244	Osteichthyes	Serranidae	Rypticus bistrispinus	(Mitchill, 1818)	VSHALL	PEL	LC	MINV	MINV
245	Osteichthyes	Serranidae	Rypticus saponaceus	(Bloch & Schneider, 1801)	VSHALL	PEL	LC	MINV	MINV
246	Osteichthyes	Serranidae	Rypticus subbifrenatus	Gill, 1861	VSHALL	PEL	LC	MINV	MINV
247	Osteichthyes	Serranidae	Serranus baldwini	(Evermann & Marsch, 1899)	VSHALL	PEL	LC	MINV	MINV
248	Osteichthyes	Serranidae	Serranus flaviventris	(Cuvier, 1829)	VSHALL	PEL	LC	MINV	MINV
249	Osteichthyes	Priacanthidae	Heteropriacanthus cruentatus	(Lacepède, 1801)	MID	PEL	LC	MINV	MINV
250	Osteichthyes	Priacanthidae	Priacanthus arenatus	Cuvier, 1829	MID	PEL	LC	MINV	MINV
251	Osteichthyes	Chaetodontidae	Chaetodon ocellatus	Bloch, 1787	MID	PEL	LC	SINV	SINV
252	Osteichthyes	Chaetodontidae	Chaetodon sedentarius	Poey, 1860	MID	PEL	LC	SINV	SINV
253	Osteichthyes	Chaetodontidae	Chaetodon striatus	Linnaeus, 1758	MID	PEL	LC	SINV	SINV
254	Osteichthyes	Malacanthidae	Malacanthus plumieri	(Bloch, 1786)	MID	LIV	LC	MCAR	MCAR
255	Osteichthyes	Haemulidae	Paranisotremus moricandi	(Ranzani, 1842)	MID	PEL	LC	MINV	MINV
256	Osteichthyes	Haemulidae	Anisotremus surinamensis	(Bloch, 1791)	MID	PEL	DD	MINV	MINV
257	Osteichthyes	Haemulidae	Anisotremus virginicus	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
258	Osteichthyes	Haemulidae	Conodon nobilis	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
259	Osteichthyes	Haemulidae	Genyatremus luteus	(Bloch, 1790)	SHALL	PEL	DD	MINV	MINV
260	Osteichthyes	Haemulidae	Haemulon aurolineatum	Cuvier, 1830	MID	PEL	LC	MINV	MINV
261	Osteichthyes	Haemulidae	Haemulon parra	(Desmarest, 1823)	MID	PEL	LC	MINV	MINV
262	Osteichthyes	Haemulidae	Haemulon melanurum	(Linnaeus, 1758)	DEEP	PEL	LC	MINV	MINV
263	Osteichthyes	Haemulidae	Haemulon plumieri	(Lacepède, 1801)	DEEP	PEL	LC	MINV	MINV
264	Osteichthyes	Haemulidae	Haemulon squamipinna	Rocha & Rosa, 1999	MID	PEL	NE	MINV	MINV
265	Osteichthyes	Haemulidae	Haemulon atlanticus	Carvalho, Marceniuk, Oliveira & Wosiacki, 2020	MID	PEL	LC	MINV	MINV
266	Osteichthyes	Haemulidae	Orthopristis scapularis	Fowler, 1915	MID	PEL	LC	MINV	MINV
267	Osteichthyes	Haemulidae	Pomadasys ramosus	(Poey, 1860)	SHALL	PEL	NE	OMNI	OMNI
268	Osteichthyes	Haemulidae	Haemulopsis corvinaeformis	(Steindachner, 1868)	MID	PEL	LC	OMNI	OMNI
269	Osteichthyes	Lutjanidae	Etelis oculatus	(Valenciennes, 1828)	VDEEP	PEL	DD	MCAR	MCAR
270	Osteichthyes	Lutjanidae	Lutjanus alexandrei	Moura & Lindeman, 2007	MID	PEL	NE	MCAR	MCAR
271	Osteichthyes	Lutjanidae	Lutjanus analis	(Cuvier, 1828)	MID	PEL	NT	MCAR	MCAR
272	Osteichthyes	Lutjanidae	Lutjanus buccanella	(Cuvier, 1828)	VDEEP	PEL	DD	MCAR	MCAR
273	Osteichthyes	Lutjanidae	Lutjanus cyanopterus	(Cuvier, 1828)	MID	PEL	VU	MCAR	MCAR
274	Osteichthyes	Lutjanidae	Lutjanus jocu	(Bloch & Schneider, 1801)	MID	PEL	DD	MCAR	MCAR
275	Osteichthyes	Lutjanidae	Lutjanus synagris	(Linnaeus, 1758)	MID	PEL	NT	MCAR	MCAR
276	Osteichthyes	Lutjanidae	Lutjanus vivanus	(Cuvier, 1828)	VDEEP	PEL	LC	MCAR	MCAR
277	Osteichthyes	Lutjanidae	Ocyurus chrysurus	(Bloch, 1791)	MID	PEL	DD	MCAR	MCAR
278	Osteichthyes	Lutjanidae	Rhomboplites aurorubens	(Cuvier, 1829)	MID	LIV	VU	MCAR	MCAR
279	Osteichthyes	Cirrhitidae	Amblycirrhitus pinos	(Mowbray, 1927)	MID	PEL	LC	MINV	MINV
280	Osteichthyes	Scorpaenidae	Scorpaena brasiliensis	Cuvier, 1829	DEEP	PEL	LC	MCAR	MCAR
281	Osteichthyes	Scorpaenidae	Scorpaena inermis	Cuvier, 1829	DEEP	PEL	LC	MCAR	MCAR
282	Osteichthyes	Scorpaenidae	Scorpaena plumieri	Bloch, 1789	SHALL	PEL	LC	MCAR	MCAR
283	Osteichthyes	Scorpaenidae	Scorpaenodes caribbaeus	Meek & Hildebrand, 1928	SHALL	PEL	LC	MCAR	MCAR
284	Osteichthyes	Triglidae	Prionotus punctatus	(Bloch, 1793)	MID	PEL	LC	MINV	MINV
285	Osteichthyes	Ephippidae	Chaetodipterus faber	(Broussonet, 1782)	MID	PEL	LC	MINV	MINV
286	Osteichthyes	Sciaenidae	Bairdiella ronchus	(Cuvier, 1830)	SHALL	PEL	LC	MINV	SAND
287	Osteichthyes	Sciaenidae	Cynoscion leiarchus	(Cuvier, 1830)	SHALL	PEL	LC	MINV	SAND
288	Osteichthyes	Sciaenidae	Eques lanceolatus	(Linnaeus, 1758)	VDEEP	PEL	LC	MINV	MINV
289	Osteichthyes	Sciaenidae	Larimus breviceps	Cuvier, 1830	MID	PEL	LC	MINV	MINV
290	Osteichthyes	Sciaenidae	Menticirrhus americanus	(Linnaeus, 1758)	MID	PEL	LC	MINV	MINV
291	Osteichthyes	Sciaenidae	Menticirrhus martinicensis	(Cuvier, 1830)	MID	PEL	LC	MINV	MINV
292	Osteichthyes	Sciaenidae	Menticirrhus cuiaranensis	Marceniuk, Caires, Rotundo, Cerqueira, Siccha-Ramirez, Wosiacki & Oliveira, 2020	MID	PEL	LC	MINV	MINV
293	Osteichthyes	Sciaenidae	Odontoscion dentex	(Cuvier, 1830)	MID	PEL	LC	MCAR	MCAR
294	Osteichthyes	Sciaenidae	Pareques acuminatus	(Bloch & Schneider, 1801)	SHALL	PEL	LC	MINV	MINV
295	Osteichthyes	Acanthuridae	Acanthurus bahianus	Castelnau, 1855	SHALL	PEL	LC	HERB	TURF
296	Osteichthyes	Acanthuridae	Acanthurus chirurgus	(Bloch, 1787)	SHALL	PEL	LC	HERB	TURF
297	Osteichthyes	Acanthuridae	Acanthurus coeruleus	Bloch & Schneider, 1801	SHALL	PEL	LC	HERB	TURF
298	Osteichthyes	Lobotidae	Lobotes surinamensis	(Bloch, 1790)	MID	PEL	LC	MCAR	MCAR
299	Osteichthyes	Sparidae	Archosargus probatocephalus	(Walbaum, 1792)	VSHALL	PEL	LC	HERB	MALG
300	Osteichthyes	Sparidae	Archosargus rhomboidalis	(Linnaeus, 1758)	VSHALL	PEL	LC	HERB	MALG
301	Osteichthyes	Sparidae	Calamus penna	(Valenciennes, 1830)	MID	PEL	LC	MINV	MINV
302	Osteichthyes	Sparidae	Calamus pennatula	Guichenot, 1868	MID	PEL	LC	MINV	MINV
303	Osteichthyes	Antennariidae	Antennarius striatus	(Shaw, 1794)	SHALL	PEL	LC	MCAR	MCAR
304	Osteichthyes	Ogcocephalidae	Ogcocephalus vespertilio	(Linnaeus, 1758)	SHALL	UNK	LC	MINV	MINV
305	Osteichthyes	Ostraciidae	Acanthostracion quadricornis	(Linnaeus, 1758)	MID	BAL	LC	OMNI	OMNI
306	Osteichthyes	Ostraciidae	Acanthostracion polygonius	Poey, 1876	MID	BAL	LC	OMNI	OMNI
307	Osteichthyes	Ostraciidae	Lactophrys trigonus	(Linnaeus, 1758)	MID	BAL	LC	OMNI	OMNI
308	Osteichthyes	Monacanthidae	Aluterus heudelotii	Hollard, 1855	MID	BAL	LC	OMNI	OMNI
309	Osteichthyes	Monacanthidae	Aluterus scriptus	(Osbeck, 1765)	MID	BAL	LC	OMNI	OMNI
310	Osteichthyes	Balistidae	Balistes vetula	Linnaeus, 1758	MID	BAL	NT	MINV	MINV
311	Osteichthyes	Balistidae	Canthidermis sufflamen	(Mitchill, 1815)	DEEP	BAL	LC	OMNI	OMNI
312	Osteichthyes	Balistidae	Melichthys niger	(Bloch, 1786)	DEEP	PEL	LC	OMNI	OMNI
313	Osteichthyes	Monacanthidae	Cantherhines macrocerus	(Hollard, 1853)	MID	DEG	LC	OMNI	OMNI
314	Osteichthyes	Monacanthidae	Cantherhines pullus	(Ranzani, 1842)	SHALL	PEL	LC	OMNI	OMNI
315	Osteichthyes	Monacanthidae	Stephanolepis hispida	(Linnaeus, 1766)	SHALL	PEL	LC	OMNI	OMNI
316	Osteichthyes	Tetraodontidae	Canthigaster figueiredoi	Moura & Castro, 2002	VSHALL	BAL	LC	OMNI	OMNI
317	Osteichthyes	Tetraodontidae	Colomesus psittacus	(Bloch & Schneider, 1801)	MID	PEL	LC	MINV	MINV
318	Osteichthyes	Tetraodontidae	Lagocephalus laevigatus	(Linnaeus, 1766)	MID	PEL	LC	MINV	MINV
319	Osteichthyes	Tetraodontidae	Sphoeroides greeleyi	Gilbert, 1900	VSHALL	PEL	LC	MINV	MINV
320	Osteichthyes	Tetraodontidae	Sphoeroides spengleri	(Bloch, 1785)	VSHALL	PEL	LC	MINV	MINV
321	Osteichthyes	Tetraodontidae	Sphoeroides testudineus	(Linnaeus, 1758)	VSHALL	PEL	LC	MINV	MINV
322	Osteichthyes	Diodontidae	Chilomycterus antillarum	Jordan & Rutter, 1897	SHALL	PEL	LC	SINV	SINV
323	Osteichthyes	Diodontidae	Chilomycterus spinosus	(Linnaeus, 1758)	SHALL	PEL	LC	SINV	SINV
324	Osteichthyes	Diodontidae	Diodon holocanthus	Linnaeus, 1758	SHALL	PEL	LC	SINV	SINV
325	Osteichthyes	Diodontidae	Diodon hystrix	Linnaeus, 1758	SHALL	PEL	LC	SINV	SINV

FIGURE 3 |
Marine Protected Area (MPA) “Costa dos Corais” fishes biodiversity grouped in categories (see “Species categorization” in Material and Methods section and Tab. 1). A. Depth category. B. Dispersal potential. C. IUCN category. D. Trophic category. E. Functional group.

DISCUSSION

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).

FIGURE 4 |
Fish biodiversity from MPA Costa dos Corais. A. Hypanus marianae (35 cm of total length, TL. B. Ginglymostoma cirratum (~ 110 cm TL). C. Aetobatus narinari (~ 80 cm TL). D. Acanthurus bahianus (~ 15 cm TL). E. Acanthurus chirurgus (~ 20 cm TL). F. Acanthurus coeruleus (~ 30 cm TL). G. Apogon americanus (~ 5 cm TL). H. Ophioblennius trinitatis (~ 5 cm TL).

FIGURE 5 |
Fish biodiversity from MPA Costa dos Corais. A. Bothus lunatus (~ 20 cm TL). B. Caranx bartholomaei (~ 20 cm TL). C. Caranx latus (~ 40 cm TL). D. Chloroscombrus chrysurus (~ 10 cm TL). E. Selene vomer (~ 25 cm TL). F. Centropomus undecimalis (~ 50 cm TL). G. Chaetodon ocellatus (~ 5 cm TL). H. Chaetodon striatus (~ 5 cm TL).

FIGURE 6 |
Fish biodiversity from MPA Costa dos Corais. A. Amblycirrhitus pinos (~ 10 cm TL). B. Dactylopterus volitans (~ 20 cm TL). C. Chaetodipterus faber (~ 30 cm TL). D. Alphestes afer (~ 25 cm TL). E. Cephalopholis fulva (~ 30 cm TL). F. Epinephelus adscensionis (~ 20 cm TL). G. Epinephelus itajara (~ 110 cm TL). H. Mycteroperca bonaci (~ 15 cm TL).

FIGURE 7 |
Fish biodiversity from MPA Costa dos Corais. A. Bathygobius soporator (~ 8 cm TL). B. Coryphopterus glaucofraenum (~ 5 cm TL). C. Ctenogobius saepepallens (~ 4 cm TL). D. Elacatinus figaro (~ 4 cm TL). E. Gramma brasiliensis (~ 5 cm TL). F. Paranisotremus moricandi (~ 15 cm TL). G. Anisotremus surinamensis (~ 20 cm TL). H. Anisotremus virginicus (~ 20 cm TL).

FIGURE 8 |
Fish biodiversity from MPA Costa dos Corais. A. Haemulon aurolineatum (~ 18 cm TL). B. Haemulon parra (~ 25 cm TL). C. Haemulon plumieri (~ 25 cm TL). D. Haemulon squamipinna (~ 18 cm TL). E. Myripristis jacobus (~ 20 cm TL). F. Bodianus rufus (~ 20 cm TL). G. Clepticus brasiliensis (~ 30 cm TL). H. Halichoeres brasiliensis (~ 20 cm TL).

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.

FIGURE 9 |
Fish biodiversity from MPA Costa dos Corais. A. Halichoeres dimidiatus (~ 10 cm TL). B. Halichoeres poeyi (~ 12 cm TL). C. Scarus trispinosus (~ 25 cm TL). D. Scarus zelindae (~ 30 cm TL). E. Sparisoma amplum (~ 35 cm TL). F. Sparisoma axillare (~ 32 cm TL). G. Sparisoma frondosum (~ 30 cm TL). H. Sparisoma radians (~ 15 cm TL).

FIGURE 10 |
Fish biodiversity from MPA Costa dos Corais. A. Gobioclinus kalisherae (~ 5 cm TL). B. Labrisomus cricota (~ 4 cm TL). C. Labrisomus nuchipinnis (~ 5 cm TL). D. Lutjanus alexandrei (~ 20 cm TL). E. Lutjanus analis (~ 22 cm TL). F. Lutjanus jocu (~ 30 cm TL). G. Lutjanus synagris (~ 20 cm TL). H. Ocyurus chrysurus (~ 18 cm TL).

FIGURE 11 |
Fish biodiversity from MPA Costa dos Corais. A. Mulloidichthys martinicus (~ 20 cm TL). B. Pseudupeneus maculatus (~ 18 cm TL). C. Gymnothorax vicinus (~ 30 cm TL). D. Gymnothorax funebris (~ 80 cm TL). E. Muraena pavonina (~ 50 cm TL). F. Ogcocephalus vespertilio (~ 20 cm TL). G. Myrichthys ocellatus (~ 30 cm TL). H. Pempheris schomburgkii (~ 5 cm TL).

FIGURE 12 |
Fish biodiversity from MPA Costa dos Corais. A. Holacanthus ciliaris (~ 30 cm TL). B. Pomacanthus paru (~ 25 cm TL). C. Abudefduf saxatilis (~ 10 cm TL). D. Azurina multilineata (~ 8 cm TL). E. Stegastes fuscus (~ 5 cm TL). F. Stegastes variabilis (~ 5 cm TL). G. Odontoscion dentex (~ 10 cm TL). H. Pareques acuminatus (~ 4 cm TL).

FIGURE 13 |
Fish biodiversity from MPA Costa dos Corais. A. Scorpaena plumieri (~ 20 cm TL). B. Rypticus saponaceus (~ 22 cm TL). C. Serranus flaviventris (~ 8 cm TL). D. Calamus penna (~ 12 cm TL). E. Sphyraena barracuda (~ 60 cm TL). F. Hippocampus reidi (~ 8 cm TL). G. Synodus intermedius (~ 25 cm TL). H. Prionotus punctatus (~ 18 cm TL).

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.

ACKNOWLEDGEMENTS

We would like to thank Julian Caon for map preparation. We also thank APACC management team and Projeto Conservação Recifal (PCR) staff for all the help during field activity. This work is part of the Long-Term Ecological Research – Brazil site PELD-CCAL (Projeto Ecológico de Longa Duração - Costa dos Corais Alagoas) funded by the Brazilian National Council for Scientific and Technological Development – CNPq (#441657/2016-8), the Brazilian Coordination for the Improvement of Higher Education Personnel PELD/CAPES (23038.000452/2017-16) and the Research Support Foundation of the State of Alagoas – FAPEAL (#60030.1564/2016). This study was also supported by: Rufford Small Grants, Instituto Linha D’Água (Edital Linha D’Água Tubarões e Raias), “Reef fish under diver influence: behavioual changes and subsidies for management of underwater tourism” (FAPEAL 600030001270/2017), Conservation Leadership Program (CLP), Marine Conservation Action Fund, Mohamed Bin Zayed Species Conservation and The Prince Bernhard Nature Fund (PBNF) grants for PHCP and PCR. We thank Fundação SOS Mata Atlântica and Fundação Toyota for support during APACC field trips.

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ADDITIONAL NOTES

• HOW TO CITE THIS ARTICLE

  Pereira PHC, Côrtes LGF, Lima GV, Gomes E, Pontes AVF, Mattos F, Araújo ME, Ferreira-Junior F, Sampaio CLS. Reef fishes biodiversity and conservation at the largest Brazilian coastal Marine Protected Area (MPA Costa dos Corais). Neotrop Ichthyol. 2021; 19(4):e210071. https://doi.org/10.1590/1982-0224-2021-0071

Edited-by

Fernando Gibran

Publication Dates

• Publication in this collection
  13 Dec 2021
• Date of issue
  2021

History

• Received
  26 Mar 2021
• Accepted
  5 July 2021