Diversity and biogeography of fishes in the Arinaga-Gando area, east coast of Gran Canaria (Canary Island)

9 DIVERSITY AND BIOGEOGRAPHY OF FISHES IN THE ARINAGA-GANDO AREA, EAST COAST OF GRAN CANARIA (CANARY ISLANDS) 1*Espino, F., 2González, J. A., 3Boyra, A., 3Fernández, C., 1Tuya, F. & 4Brito, A. 1Centro de Investigación en Biodiversidad y Gestión Ambiental (BIOGES) Departamento de Biología, Facultad de Ciencias del Mar, Universidad de Las Palmas de Gran Canaria Campus de Tafira. 35017 Las Palmas. Islas Canarias, España. 2Grupo de Investigación en Ecología Marina Aplicada y Pesquerías Departamento de Biología, Facultad de Ciencias del Mar, Universidad de Las Palmas de Gran Canaria Campus de Tafira. 35017 Las Palmas. Islas Canarias, España. 3Oceanográfica: Divulgación, Educación y Ciencia. Peregrinos 30. Polígono Industrial El Goro 35219 Telde, Las Palmas. Islas Canarias, España. 4Grupo de Investigación en Biología, Ecología Marina y Conservación (BIOECOMAC) Departamento de Biología Animal (Ciencias Marinas), Facultad de Biología, Universidad de La Laguna 38206 La Laguna, Tenerife. Islas Canarias, España. * Corresponding author: Fernando Espino email: [email protected] ABSTRACT A check-list of fishes regularly present in waters of the Gando-Arinaga area, in the East coast of Gran Canaria, is presented herein. This inventory includes 175 species, 22 of which were of chondrichthyans (belonging to 19 genera and 13 families), and 153 were of actinopterygiians (121 genera and 60 families); 77.15% of the species have stable populations in the area and 61.15% have commercial interest. The zoogeographic analysis showed a dominance of the Atlantic-Mediterranean distribution species (26%), followed by the Warmtemperate in the Eastern Atlantic distribution species (16%), while Eastern Central Atlantic Oceanic Islands (9%), Guinean (8%) and Macaronesian (4%) species showed smaller values, but they were representative of the general biogeographic pattern of the canarian ichthyofauna. This work pointed the presence of warm affinity fish species in the area, like it occurs in other areas of the Canary Islands. This coastal area hosts a high diversity of fish species, which could be explained by a variety of complex habitats, its geographic location and oceanographic conditions. This area meets the conservation criteria for the establishment of a marine protected area. Key words: ichthyofauna, biodiversity, marine protected area, zoogeography, Canary Islands. Rev. Acad. Canar. Cienc., Vol. XXVI, 9-25 (diciembre de 2014) 10 RESUMEN El inventario de la ictiofauna del sector costero Gando-Arinaga, en el litoral Este de la isla de Gran Canaria, incluye 175 especies, 22 especies de condrictios (agrupados en 19 géneros y 13 familias) y 153 de actinopterigios (121 géneros y 60 familias); el 77,15% de las especies presentan poblaciones estables en la zona y un 61,15% de las especies tienen interés comercial. El análisis zoogeográfico mostró un predomino de las especies de distribución Atlántico-Mediterránea (26%), seguidas por las especies de distribución en áreas Cálido-templadas del Atlántico Oriental (16%), mientras que las especies de las Islas Oceánicas del Atlántico Centroriental (9%), Guineanas (8%) y Macaronésicas (4%) mostraron valores menores, pero representativos del patrón biogeográfico general de la ictiofauna canaria. Se destaca la presencia de especies de afinidad tropical en la zona, como sucede en otras áreas del Archipiélago Canario. Esta zona del litoral alberga una alta diversidad de especies de peces, que puede ser explicada por la variedad de hábitats complejos que presenta, su localización geográfica y sus condiciones oceanográficas. En conclusión, esta zona cumple con los criterios de conservación para poder establecer un área marina protegida. Palabras clave: ictiofauna, biodiversidad, área marina protegida, zoogeografía, Islas Canarias. 1. INTRODUCTION The knowledge of the distribution patterns of the species and the areas that host high diversity rates is crucial for a better management and conservation of the marine biological resources. In a global context, these resources are suffering a severe depletion and the marine space is poorly represented in the global network of marine protected areas (hereafter MPA’s), only 0.01% of the world’s ocean is effectively protected (PAULY et al., 2002). This implies that the identification and protection of the most representative areas for the marine biodiversity are priorities (BRITO, 2010). On the Canarian Archipelago, especially in the fisheries marine reserves, several studies and species inventories have been carried out. Most of these studies have focused on fishery species and pointed the importance of the MPA’s in the conservation of the marine biodiversity (e.g., FALCÓN et al., 2002). The Gando-Arinaga marine area, located in the East coast of Gran Canaria, has been recognized in some works (BACALLADO et al., 1989; HERRERA et al., 1993; ESPINO, 1997; PORTILLO & PÉREZ, 1998) for its special natural resources. In fact, the protection of this area has been proposed by some authors (BACALLADO et al., 1989; AGUILAR et al., 2010), and many previous works have been carried out to establish a fishery marine reserve (see CASTRO et al., 2001; PÉREZ et al., 2001; LUQUE et al., 2001; MEDINA et al., 2001; TUYA et al., 2004). Today, there are two legal protected areas in Gando-Arinaga: The Special Area of Conservation (hereafter SAC) SAC 26-GC, namely ‘Bahía de Gando’ located at north, and the SAC 34-GC, namely ‘Playa del Cabrón’ located at south, both established under the EU Habitat Directive. The Gando-Arinaga area is located in the East coast of Gran Canaria (27º 50’ N - 27º 57’ N y 15º 19’ W - 15º 24’ W) (Fig. 1), with a surface of ca. 7,998.5 ha and a total perimeter of ca. 48.5 km. The geomorphology of the coast is abrupt in the north and south parts and smoother in the central part. The littoral shelf is relatively wide, reaching 50 and 100 m bathymetric lines at 2.7 and 7.2 km from the line coast in the central part of the area, respectively. 11 The basaltic hard bottoms can be seen to 20-30 m depth in ‘Tufia’, ‘Península de Gando’, ‘Baja de Gando’, ‘Punta de La Sal’, ‘Veril del Cabrón’ and ‘Roque de Arinaga’; from here the volcanic materials are covered by sediments. The factors influencing the coast relief had given up a high variety of biotopes that support a diversity of biological communities (BACALLADO et al., 1989; HERRERA et al., 1993; ESPINO, 1997; PORTILLO & PÉREZ, 1998; PÉREZ et al., 2001). For example: 1). In the intertidal zone: rocky flats, rock pools, cliffs, sandy and rocky beaches; 2). In the infralittoral zone: rocky reefs with photophilic macroalgae communities dominated by Cystoseira abies-marina (S. G. Gmelin) C. Agardh, 1820 and Sargassum spp., rocky bottoms without macroalgae or barren grounds dominated by Diadema africanum Rodríguez, Hernández & Clemente, 2010, isolated rocky outcrops, vertical cliffs or ‘veriles’, hollows and caves, mixed sandy-rocky bottoms, sandy bottoms with seagrass meadows constituted by Cymodocea nodosa (Ucria) Ascherson, 1870 or ‘sebadales’, communities of the green rhizophytic algae Caulerpa prolifera (Forsskål) J. V. Lamouroux, 1809, mixed meadows of Figure 1.- Map of the Gando-Arinaga area (East coast of Gran Canaria) with the sector studied (broken line). 12 C. nodosa-C. prolifera, and sandy sediments without vegetation; 3). In the infralittoralcircalittoral ecotone there are communities of free-living coralline algae (maerl) characterized by Lithothamnion corallioides P. L. Crouan & H. M. Crouan, 1867, and deep corals communities (gorgonians, anthipatarians). In this sector of the Gran Canaria coast, there are some wrecks, artificial reefs and sea-cage fish farms that exert attraction-production effects on some fish species. In addition, the littoral water mass can be considered as the habitat for many pelagic fish species. The Gando-Arinaga area is directly exposed to the Trade Winds from NNE; these reach their maximum intensity in the June-August period, with values of 9.7-10.7 m s-1. The topo - graphy of the coasts and bottoms exerts a strong influence on the local marine currents, which are originated by winds and tides (PORTILLO & PÉREZ, 1998). The dominant current shows the direction of the Canary Current (SW), followed in frequency by the direction of the opposite current (NE), their speeds in surface varied between 5.57 to 24 cm s-1. The water mass of the area is temperate, with sea surface temperature varying between 18 to 18.6 ºC in Fe bruary-March and 22.5 to 24.1 ºC in September-October. The luminic intensity shows a seasonal variation between 379 to 2,819 µE m-2 s-1, reaching 1% of this intensity at 38 and 61 m depth, respectively. Like in the Canary Islands region, these waters are oligotrophic. The primary production shows maximum values between 2.0 mg C m-3 h-1 during the DecemberFebruary period and 4.9 mg C m-3 h-1 in March, and points a peak of production in late winter and early spring (MEDINA, 1995; MEDINA et al., 2001). There are many references to the ichthyofauna of the Gando-Arinaga area. For example: FALCÓN et al. (1996) recorded 20 coastal fish species and found high mean richness and diversity. ESPINO (1997) pointed that more than 100 species inhabit this area, and classified 76 species as common and 30 as occasional. PORTILLO & PÉREZ (1998) recorded 53 fish species in their underwater guide and indicated high fish richness. PÉREZ et al. (2001) recorded 93 species. CASTRO et al. (2001) found that 29 fish species are exploited by the artisanal fisheries in the area. TUYA et al. (2004) registered 36 species during an ecologic evaluation. Finally, AGUILAR et al. (2010) recorded 59 species; some of them were observed more than 100 m depth. The aims of this work were: 1) to make a check-list of fish species that can be found in the Arinaga-Gando area; 2) to realize a comparative zoogeographic analysis of the ichthyofauna; and 3) to propose the establishment of an MPA to preserve the marine fish biodiversity. 2. MATERIAL AND METHODS To make the inventory of the ichthyofauna, first we reviewed the scientific papers published and the grey literature works. Second, we interviewed professional fishermen of the area (Melenara, El Burrero and Arinaga), sport fishermen and divers, with special attention to the dive centres that regularly visit the area. Observational data of some cryptic and rare fish species corresponding to marine biologists, scientific divers and specialized underwater photographers. The species recorded were grouped in genera and families, and ordered following the criteria of Nelson (2006). Most of the scientific names of the species are those recorded in WHITEHEAD et al. (1984-1986) and QUERÓ et al. (1990), and updated according to the recent scientific literature (FROESE & PAULY, 2014). Within each family, genera and species are in alphabetical order. We classified each fish species in a spatial distribution category, following those established in the ‘Catalogue of the Canary Islands Fishes’ by BRITO et al. (2002). To real- 13 ize the comparative zoogeographic analysis of the ichthyofauna, the fish species were classified in 11 groups according to their geographic distribution area: 1. Amphi-Atlantic species of wide distribution; 2. Amphi-Atlantic species of warm affinity; 3. Species of Wide distribution in the Eastern Atlantic; 4. Warm-temperate species in the Eastern Atlantic; 5. Coldtemperate species in the Eastern Atlantic; 6. Atlantic-Mediterranean species; 7. Cosmopolitan species; 8. Guinean species; 9. Eastern Central Atlantic Oceanic Islands species (i.e. species distributed from the Azores to Cape Verde Islands); 10. Macaronesian species (i.e. species of Azores, Madeira and Canary Islands); and 11. Pantropical species. Here, the classification model proposed by BRITO et al. (1996) and applied by FALCÓN et al. (2002) was followed, but taking into account more recent biogeographic analyses that separate the marine fauna of Cape Verde Archipelago from the marine Azores, Madeira and Canaries ecoregión (BRITO et al., 2007; SPALDING et al., 2007; FLOETER et al., 2008; BRITO, 2010; WIRTZ et al., 2013). 3. RESULTS The inventory of the Gando-Arinaga ichthyofauna includes 175 species, 22 of which were of chondrichthyans (belonging to 19 genera and 13 families), and 153 were of actinopterygiians (121 genera and 60 families). Within the chondrichthyans, the families with more species richness were Myliobatidae (5 spp.), Carcharhinidae (3) and Dasyatidae (3); while in the actinopterygiians were Sparidae (17), Carangidae (9), Scombridae (8), Blenniidae (7), Gobiidae (7) and Labridae (7) (Table 1); 77.15% of the species have stable populations in the area and 61.15% have commercial interest. The zoogeographic analysis of the ichthyofauna showed that Atlantic-Mediterranean distribution species were dominant (22%), followed by Warm-temperate in the Eastern Atlantic distribution species (13%), Wide distribution in the Eastern Atlantic species group (12%), Amphi-Atlantic species of warm affinity (11%) and Cosmopolitan species (11%), if all species were included in the analysis (Fig. 2a). When pelagic and benthopelagic species were excluded of the analysis, the percentage of the Atlantic-Mediterranean species (26%) and Warm-temperate in the Eastern Atlantic species (16%) increased. The Eastern Central Atlantic Oceanic Islands (9%), Guinean (8%) and Macaronesian (4%) species slightly increased too. In opposite, Pantropical (3%) and Cosmopolitan (1%) species decreased significantly (Fig. 2b). The endemism of the Eastern Atlantic Islands (i.e. from the Azores to the Cape Verde islands) up today, were represented in the Gando-Arinaga area: Raja maderensis (Rajidae), Muraena augusti (Muraenidae), Gaidropsarus guttatus (Phycidae), Scorpaena canariensis (Scorpaenidae), Mycteroperca fusca (Serranidae), Abudefduf luridus (Pomacentridae), Bodianus scrofa (Labridae), Centrolabrus trutta (Labridae), Ophioblennius atlanticus (Blenniidae), Canthigaster capistrata (Tetraodontidae), Diplecogaster pectoralis (Gobiesocidae), Didogobius kochi (Gobiidae), Bothus podas maderensis (Bothidae), Symphurus insularis (Cynoglossidae) and Mauligobius maderensis (Gobiidae) (BRITO et al., 1999; BRITO et al., 2002; BRITO et al., 2007; WIRTZ et al., 2013); of these species, R. maderensis, G. guttatus, S. canariensis, C. trutta and M. maderensis were exclusive of the Macaronesian ecoregion (i.e. Azores, Madeira and Canaries) (BRITO et al., 2002; BRITO et al., 2007; SPALDING et al., 2007; WIRTZ et al., 2013). 14 Figure 2.- Biogeographic composition of the ichthyofauna of the Gando-Arinaga area including all species. b. Excluding pelagic and benthopelagic species. For biogeographic categories see material and methods. 15 4. DISCUSSION The Gando-Arinaga littoral hosts a high diversity of fish species, especially taking into account that this area only represents 9.5% of the line coast and 18.5% of the insular shelf surface, to 100 m depth, off Gran Canaria. The high diversity could be explained by many factors. First, by a diversity of natural and artificial habitats and its structural complexity. These factors have been pointed as determinant in the richness, abundance and composition of fish assemblages (GRATWICKE & SPEIGHT, 2005, and references therein). Second, other factors influencing fish species richness are the location and orientation of this area. Gando-Arinaga is located in a channel between islands and orientated parallel to the direction of the dominant current, this allows that some migratory fish species can be seen here (e.g., Rhincodon typus, Cetorhinus maximus, Mobula spp., Manta birostris, Mola mola and Thunnus spp.). The situation of the area facilitates that fish larvae of septentrional sectors of the islands could be carried by the Canary Current, and later settle here. In addition, some human activities, like marine aquaculture could explain the presence of some species, like Sparus aurata, Dicentrarchus labrax and Argyrosomus regius. The biogeographic composition of the Gando-Arinaga ichthyofauna is according with the model proposed by BRITO et al. (1996) for the ichthyofauna of the Canary Islands that shares the littoral and upper bathyal bottoms. This can be explained by the high percentage of benthic and demersal species that could extend their bathymetric distribution range to the upper bathyal bottoms versus the percentage of benthic and demersal species that inhabit exclusively littoral bottoms (49.15% vs. 17.15%). The Macaronesian species recorded a small percentage according to previous analysis (BRITO et al., 1996; FALCÓN et al., 2002). Nevertheless, ca. 50% of the endemic Macaronesian fish species were recorded in Gando-Arinaga, and ca. 75% of the endemic fish species of the Eastern Central Atlantic Oceanic Islands, excluding endemism of Cape Verde Islands. In this group, the wide-eyed flounder Bothus podas maderensis (Bothidae) must be reviewed, according with BRITO et al. (2007), and the small tongue fish Symphurus insularis (Cynoglossidae), was recently recorded on the NGor Island (Senegal) (WIRTZ, 2012), outside the eastern Atlantic Islands. The gradual displacement of warm affinity fishes toward septentrional latitudes had been pointed in many works, for example in Azores, Madeira, Canaries, Galicia, west coast of Portugal and the Mediterranean Sea (see BRITO et al., 2005 and references therein; HORTAe COSTA et al., 2014). BRITO et al. (2005) found out that 80% of the 30 new records of actinopterygiians littoral fishes on the Canary Islands, between 1991 and 2005, corresponded to species of tropical origin. This phenomenon of ‘tropicalization’ has been found in some areas of the Canarian Archipelago (e.g., FALCÓN et al., 2002). Recently, two species of tropical fishes were found in the Arinaga-Gando area: Echiophis punctifer (Ophichthidae) (A. BOYRA pers. obs.), a species distributed in tropical areas of the Eastern and Western Atlantic; and Hypleurochilus sp. (Blenniidae) (A. UBIERNA pers. com.). In addition, other warm affinity species observed here were: Rhincodon typus (Rhincodontidae), Grammonus longhursti (Bythitidae), Holocentrus ascensionis (Holocentridae), Caranx crysos (Carangidae), Lutjanus goreensis (Lutjanidae), Gnatholepis thompsoni (Gobiidae), Acanthurus monroviae (Acanthuridae), Canthidermis sufflamen (Balistidae) and Chilomycterus atringa (Diodontidae). Of these, only G. thompsoni and C. atringa seem to have stable population in the area. The presence of some of these warm species could be related to some human activities, like ships traffic and aquarium collection. Some authors pointed that other species could reach the Canarian waters by their own dispersal abilities, in relation to the sea surface temperature increasing on the Canaries (BRITO et al., 2005). 16 The number of taxa recorded could be increased when more studies would be carried out; especially focused on pelagic species (e.g., Cacharhinidae, Sphyrnidae, Exocoetidae) and deep benthic or demersal species that could be associated with free-living coralline algae habitat, black corals or scleractinian corals. For example, AGUILAR et al. (2010) recorded the presence, between 200 and 500 m depth, of: Squalus megalops (Squalidae), Raja maderensis (Rajidae), Dasyatis pastinaca (Dasyatidae), Conger conger (Congridae), Chlorophthalmus agassizi (Chlorophthalmidae), Merluccius merluccius (Merlucciidae), Cyttopsis rosea (Zeidae), Grammicolepis brachiusculus (Grammicolepididae), Capros aper (Caproidae), Macroramphosus scolopax (Macroramphosidae), Helicolenus dactylopterus (Scorpaenidae) and Arnoglossus cf. imperialis (Bothidae). Of these species, C. agassizi is relevant for its abundance, forming big schools near the bottom and swimming against the current. These species could be present in the area studied, since the 500 m bathymetric line is reached in the NE sector (see Fig. 1). Many species of small epipelagic and mesopelagic fishes (Myctophidae, Gonostomatidae, Photichthyidae and Stomiidae) were not included in the inventory. Species of these families were recorded by MOYANO & HERNÁNDEZ-LEÓN (2009) when ichthyoplankton surveys were carried out on the 100 m bathymetric line of the area. The Gando-Arinaga area is one of the richest in fish species of the Canary Islands. Nevertheless, like in other areas of the archipelago, the fishery overexploitation (AGUILERA et al., 1994; BAS et al., 1995; FALCÓN et al., 1996; GONZÁLEZ, 2008), especially in the most populated central islands like Gran Canaria (BORTONE et al., 1991; TUYA et al., 2006a, b), and the marine contamination (AGUILERA et al., 1994; AGUILAR et al., 2010), are the most important threat factors for the ichthyofauna. The SAC’s established in this area have the aim of conservation determined habitat, and they have not an efficient management up today; due to this, they are not sufficient to protect the marine biodiversity, especially the species and populations of fishes. Despite the many proposals of protection of this area, the public administration involved has not been able to protect this marine biodiversity. If we consider the variety of littoral ecosystems, the richness of fish species, and other taxonomic groups, like algae and marine invertebrates (BACALLADO et al., 1989; HERRERA et al., 1993; ESPINO, 1997; PORTILLO & PÉREZ, 1998) present in the area, it is clear the necessity to establish an MPA to protect the natural resources. This proposal is according with the criteria pointed by BRITO (2010), who recommended the selection of marine spaces with high heterogeneity and complexity habitats. Hereby, the maximum protection of vulnerable biodiversity due to fisheries and resources production increase could be reached. Another benefit to adjacent areas is the larvae dispersion and the biomass exportation. On the Canary Islands, a legal figure to protect the area is The Marine Natural Park or Marine Natural Reserve, because they allow the resources conservation and many uses (e.g., touristic, recreational, scientific, educative, sports, etc.) at once. The first objective of the MPA must be the conservation of the four levels of marine biodiversity: genetic, specific, population and ecosystem. The second objective must be the limitation and reduction of marine resources exploitation activities, increasing at the same time others more sustainable. 5. ACKNOWLEDGEMENTS We gratefully thank A. Telle, E. Faber, M. Farray, O. Santana, J. M. Falcón, B. Ramírez, V. Benítez, A. Ubierna, J. Casas, B. Goldthorpe, A. Goldthorpe and E. Vera for helping us with data collection. Special thanks go to M. T. Espino and professional fishermen of Melenara, El Burrero and Arinaga. 17 6. REFERENCES AGUILAR, R., A. DE LA TORRIENTE, J. PEÑALVER, J. LÓPEZ, R. GREENBERG & C. CALZADILLA. 2010. 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Catálogo de los peces de la Reserva 24 Family-Species Spatial category Coris julis (Linnaeus, 1758) demersal Labrus bergylta Ascanius, 1767 demersal littoral Symphodus mediterraneus (Linnaeus, 1758) demersal littoral Thalassoma pavo (Linnaeus, 1758) demersal Xyrichtys novacula (Linnaeus, 1758) demersal Scaridae Sparisoma cretense (Linnaeus, 1758) demersal Trachinidae Trachinus draco Linnaeus, 1758 demersal Trachinus radiatus Cuvier, 1829 demersal Uranoscopidae Uranoscopus scaber Linnaeus, 1758 demersal Tripterygiidae Tripterygion delaisi Cadenat & Blache, 1970 benthic Blenniidae Hypleurochilus sp. benthic Lipophrys pholis (Linnaeus, 1758) benthic Ophioblennius atlanticus (Valenciennes, 1836) benthic Parablennius incognitus (Bath, 1968) benthic Parablennius parvicornis (Valenciennes, 1836) benthic Parablennius pilicornis (Cuvier, 1829) benthic Scartella cristata (Linnaeus, 1758) benthic Labrisomidae benthic Labrisomus nuchipinnis (Quoy & Gaimard, 1824) benthic Gobiesocidae benthic Apletodon pellegrini (Chabanaud, 1925) benthic Diplecogaster pectoralis Briggs, 1955 benthic Lepadogaster candolii Risso, 1810 benthic Lepadogaster lepadogaster (Bonnaterre, 1788) benthic Opeatogenys cadenati Briggs, 1957 benthic Gobiidae Didogobius kochi Van Tassell, 1988 benthic Gnatholepis thompsoni Jordan, 1904 benthic Gobius niger Linnaeus, 1758 benthic Gobius paganellus Linnaeus, 1758 benthic Mauligobius maderensis (Valenciennes, 1837) benthic Thorogobius ephippiatus (Lowe, 1839) benthic Vanneaugobius canariensis Van Tassell, Miller & Brito, 1988 benthic Acanthuridae Acanthurus monroviae Steindachner, 1876 demersal littoral Sphyraenidae Sphyraena viridensis Cuvier, 1829 pelagic littoral Gempylidae Lepidocybium flavobrunneum (Smith, 1843) benthopelagic 25 Family-Species Spatial category Scombridae Acanthocybium solandri (Cuvier, 1832) epipelagic littoral & oceanic Katsuwonus pelamis (Linnaeus, 1758) epipelagic oceanic Sarda sarda (Bloch, 1793) epipelagic littoral & oceanic Scomber colias Gmelin, 1789 pelagic littoral & oceanic Thunnus alalunga (Bonnaterre, 1788) pelagic oceanic Thunnus albacares (Bonnaterre, 1788) epipelagic oceanic Thunnus obesus (Lowe, 1839) pelagic oceanic Thunnus thynnus (Linnaeus, 1758) pelagic oceanic Xiphiidae Xiphias gladius Linnaeus, 1758 pelagic oceanic & littoral Centrolophidae Schedophilus ovalis (Cuvier, 1833) benthopelagic Bothidae Bothus podas (Delaroche, 1809) benthic Soleidae Microchirus azevia (de Brito Capello, 1867) benthic Pegusa lascaris (Risso, 1810) benthic Synapturichthys kleinii (Risso, 1827) benthic littoral Cynoglossidae Symphurus insularis Munroe, Brito & Hernández, 2000 benthic Balistidae Balistes capriscus Gmelin, 1789 benthopelagic Canthidermis sufflamen (Mitchill, 1815) benthopelagic littoral Monacanthidae Aluterus scriptus (Osbeck, 1765) benthopelagic littoral Stephanolepis hispidus (Linnaeus, 1766) demersal Tetraodontidae Canthigaster capistrata (Lowe, 1839) demersal Sphoeroides marmoratus (Lowe, 1838) demersal Diodontidae Chilomycterus atringa (Linnaeus, 1758) demersal littoral Molidae Mola mola (Linnaeus, 1758) epipelagic littoral & oceanic