Deep-sea anglerfishes (Lophiiformes: Ceratioidei) from off northeastern Brazil, with remarks on the ceratioids reported from the Brazilian Exclusive Economic Zone

The deep-sea anglerfishes of the suborder Ceratioidei (Lophiiformes) are represented by about 170 valid species with some of the most extraordinary morphological and reproductive adaptations among vertebrates, including extreme sexual dimorphism and male parasitism. Here we report on the diversity and distribution of rare ceratioids collected during the ABRACOS (Acoustics along the BRAzilian COaSt) expeditions off northeastern Brazil and the Fernando de Noronha Ridge (Rocas Atoll, Fernando de Noronha Archipelago, and associated seamounts). Chaenophryne ramifera , Oneirodes anisacanthus , O. carlsbergi , Gigantactis watermani , and unidentified specimens of Caulophryne , Dolopichthys , and Rhynchactis are recorded for the first time in the Brazilian Exclusive Economic Zone. Ceratias uranoscopus , Melanocetus johnsonii , and Chaenophryne draco have their distributions extended in Brazilian waters. Caulophryne , O. anisacanthus , and G. watermani are also recorded for the first time in the western South Atlantic. The specimen of G. watermani reported here represents the third known specimen of the species, and variations of its escal anatomy in relation to the holotype are described. Based on specimens examined and a review of records in the literature, 20 species of the Ceratioidei, in addition to unidentified species of Caulophryne , Dolopichthys , and Rhynchactis , are confirmed in the Brazilian Exclusive Economic Zone.


INTRODUCTION
The Ceratioidei (Lophiiformes) includes 11 families, 35 genera and about 170 valid species of fishes commonly known as deep-sea anglerfishes (Pietsch, Orr, 2007;Pietsch, 2009;Ho, Shao, 2019;Fricke et al., 2021). Adults and larvae of the group are remarkable in a number of features, including their anatomical diversity and extreme sexual dimorphism in which males are often obligatorily attached to females, with fusion of tissues and sharing of circulatory systems (Pietsch, 2009;Swann et al., 2020). Members of the Ceratioidei are also externally recognized by the absence of pelvic fins, scales usually absent (but prickles, spines or plates variably present), gill openings narrowly constricted and forming a tube-like structure that opens posteriorly, denticular bones present in dwarf males, usually 12-28 pectoral-fin rays, 8-9 caudal-fin rays, and females with an illicial apparatus usually tipped by a modified esca containing a globular, bacteriafilled photophore (Pietsch, 2009). Female ceratioids are also typically short and deep, with a nearly globular body in the Caulophrynidae, Melanocetidae, Himantolophidae, Diceratiidae, and Linophrynidae, or with a more elongate and somewhat laterally compressed body in the Centrophrynidae, Ceratiidae, Gigantactinidae, Neoceratiidae, Thaumatichthyidae, and some members of the Oneroididae (Pietsch, 2009).
Despite representing one of the most ubiquitous deep-sea groups of vertebrates in the meso-and bathypelagic zones, new species of deep-sea anglerfishes continue to 3/28 ni.bio.br | scielo.br/ni be described, mostly from still poorly explored regions of the world (Ho, Shao, 2004;Stewart, Pietsch, 2010;Pietsch, Kenaley, 2011;Prokofiev, 2014a,b;Ho et al., 2016;Rajeeshkumar et al., 2017;Ho, Shao, 2019). With about 7,500 km of coastline in addition to some biogeographically relevant oceanic island complexes (Reis et al., 2016), the Brazilian Exclusive Economic Zone (EEZ) stands out in the western South Atlantic as one of such poorly known regions in terms of its deep-sea biota. Indeed, this area includes several Ecologically or Biologically Significant Marine Areas that encompass hotspots of biodiversity and endemism (CBD, 2014).
Part of the northeastern Brazilian coast and adjacent oceanic islands and seamounts were recently explored by the RV Antea, resulting in the collection of more than 9,000 specimens of mesopelagic fishes (Bertrand, 2015(Bertrand, , 2017. Studies based on these collections have contributed significantly to the understanding of the deep-sea fauna of different groups in the region (Eduardo et al., 2018(Eduardo et al., , 2019(Eduardo et al., , 2020aMincarone et al., 2019Mincarone et al., , 2020Afonso et al., in press). This study, part of an ongoing effort to report on the still puzzling deep-sea fauna of the western South Atlantic, focus on the diversity of deep-sea anglerfishes collected off northeastern Brazil, including oceanic islands and seamounts. Remarks on the taxonomy and distribution of previous records of the Ceratioidei in the Brazilian EEZ are also presented.

MATERIAL AND METHODS
Specimens examined in this study are part of a large collection of mesopelagic invertebrates and fishes collected during the ABRACOS expeditions (Acoustics along the BRAzilian COaSt), carried out between 30 September and 20 October 2015 (ABRACOS 1 -AB1; Bertrand, 2015), and between 9 April and 6 May 2017 (ABRACOS 2 -AB2; Bertrand, 2017). Both expeditions were conducted onboard the French RV Antea off Rio Grande do Norte to Pernambuco States and along the Fernando de Noronha Ridge, formed by the Fernando de Noronha Archipelago, the Rocas Atoll, and the seamounts off Rio Grande do Norte and Ceará States (Jinno, Souza, 1999). The survey comprised 82 fishing stations, between the surface and 1,113 m depth. Sampling was conducted using micronekton (body mesh 40-80 mm, cod-end mesh 10 mm, height 24 m, width 24 m) and mesopelagic (body mesh 30 mm, cod-end mesh 4 mm, height 8.4 m, width 12.6 m) nets. Trawl depth was continuously recorded using a Scanmar depth sensor fitted on the upper part of the trawl mouth. An open-mouth net was employed, but collection of specimens most likely occurred at pre-established target depths, which were defined for each trawl according to the presence of an acoustic scattered layer or patches detected with a Simrad EK60 split-beam scientific echo sounder. At the target depths, trawling activity lasted for about 30 minutes. Target depth is therefore indicated as capture depth in the species accounts presented herein. Specimens were identified based on Pietsch (2009). Only female specimens were examined and, unless stated otherwise, distributional data refers to female specimens. All specimens collected were deposited in the Fish Collection of the Instituto de Biodiversidade e Sustentabilidade, Universidade Federal do Rio de Janeiro (NPM, Macaé, Brazil). Other institutional abbreviations follow Sabaj (2020).
In addition to Ceratias uranoscopus, C. holboelli and C. tentaculatus were previously reported in the western South Atlantic (Sutton et al., 2008;Porteiro et al., 2017). Ceratias holboelli is widely distributed in the Atlantic and Indo-Pacific, with records in the Atlantic ranging between 68ºN and 14ºS. The species was recorded in Brazilian waters based on a single specimen collected off Ilhéus, Bahia State (MNRJ 30701, 14º36'36"S 38º49'21"W;Pietsch, 2009;Fig. 2). This specimen was previously identified as C. uranoscopus by Costa et al. (2007). Melo et al. (2020) also listed C. uranoscopus in Brazilian waters based on this misidentification. Ceratias tentaculatus is restricted to the Southern Hemisphere with two records in the western South Atlantic, one off northern Argentina (ISH 435/71, 38º20'S 54º33'W), and another off Rio Grande do Sul State, close to the Brazilian EEZ (ISH 1657/68, 35°16'S 49°26'W) (Pietsch, 1986). Ceratias tentaculatus has also been briefly mentioned as occurring off Uruguay (Nión et al., 2016).

HIMANTOLOPHIDAE
Females of the Himantolophidae are distinguished by having a short, deep, globular body; lower jaw unusually blunt, extending anteriorly beyond the upper jaw; illicium thick and stout, esca unusually large and anatomically complex, the pterygiophore of the illicium fully embedded in the dermis of the head; low and rounded wart-like papilla covering the snout and chin; sphenotic spines well developed, spines absent on quadrate, articular, angular and preopercular bones; jaw teeth numerous and short, arranged in several close-set longitudinal series, vomer broad and toothless; skin of specimens larger than 30-40 mm SL, with large, widely spaced bony plates, each bearing a single median spine; 5-6 dorsal-fin rays, 4 anal-fin rays, 14-18 pectoral fin-rays, 9 caudal-fin rays; males free-living, apparently never parasitic on females (Bertelsen, Krefft, 1988;Pietsch, 2009).

Himantolophus Reinhardt, 1837
Diagnosis. Himantolophus is the only genus in the family. In addition to the diagnostic features of the Himantolophidae, females and males are distinguished by the absence of the parietal bone throughout life (vs. parietal present or lost during metamorphosis in females of the gigantactinid genus Rhynchactis), and by the presence of a triradiate pelvic bone (sometimes also present in the oneirodid genus Chaenophryne) (Pietsch, 2009 Remarks. Due to the immature nature of the specimens collected in this study, identification was possible only to genus. It is also possible that those specimens represent more than one species. Himantolophus currently includes 20 species distributed among five species groups, with representatives of all groups occurring in the Atlantic Ocean (Bertelsen, Krefft, 1988;Pietsch, 2009;Stewart, Pietsch, 2010;Pietsch, Kenaley, 2011;Fricke et al., 2021). Two species of Himantolophus were previously reported in Brazilian waters: Himantolophus macroceras Bertelsen & Krefft, 1988, known from five specimens reported from the central Atlantic, including one collected off Saint Peter and Saint Paul Archipelago (MCZ 58177, 0º10'N 27º30'W;Bertelsen, Krefft, 1988); and Himantolophus groenlandicus Reinhardt, 1837, widely distributed in the Atlantic, with one specimen reported off Saint Peter and Saint Paul Archipelago (MCZ 49841, 1º02'N 29º04'W;Bertelsen, Krefft, 1988;Fig. 2

THAUMATICHTHYIDAE
Females of the Thaumatichthyidae are distinguished by having an elongate body; esca bearing 1-3 large toothlike denticles (bony hooks); upper jaw extending forward far beyond the lower jaw; premaxillae bearing numerous hooked teeth; upper arm of opercle divided into two or more branches; males and larvae of Lasiognathus are unknown, metamorphosed males of Thaumatichthys are unusually slender and elongate, apparently never parasitic on females (Pietsch, 2009).

Remarks. Thaumatichthys has three valid species, with only Thaumatichthys binghami
Parr, 1927 reported from the western Atlantic, in the Bahamas, Gulf of Mexico, Caribbean Sea, and off Espírito Santo State, Brazil (MNRJ 30710, 19º45'S 39º30'W;Pietsch, 2009;Fig. 3). The single juvenile specimen recorded here could not be identified to species, but might be T. binghami.

ONEIRODIDAE
Females of the Oneirodidae are distinguished by having a short, deep to moderately elongate and laterally compressed body; mouth oblique to nearly horizontal, jaws equal anteriorly; illicium with a bulbous distal light organ; pterygiophore of the illicium usually emerging anteriorly on the snout, extending posteriorly on the back behind the head only in Oneirodes; top of head usually bearing sharp sphenotic spines, absent only in Chaenophryne and short in Ctenochirichthys; quadrate and articular spines usually well developed; skin smooth, dermal spines or spinules absent except in Spiniphryne; 4-8 dorsal-fin rays, 4-7 anal fin-rays, 13-30 pectoral-fin rays; a narrow, spatulate, anterodorsally directed process that overlaps the posterolateral surface of the respective sphenotic present in metamorphosed females; males usually free-living, non-parasitic, but two species apparently with facultative sexual parasitism (Pietsch, 2009).

Geographical distribution.
Chaenophryne ramifera occurs in the Atlantic, Indian and Pacific Oceans. In the Atlantic, the species has been reported between 35°N off North Carolina and 12°S off Angola, with records near the Brazilian EEZ off Saint Peter and Saint Paul Archipelago (Pietsch, 1975(Pietsch, , 2009. Chaenophryne ramifera is recorded for the first time in the Brazilian EEZ based on specimens collected off Fernando de Noronha Archipelago, Rocas Atoll, and Rio Grande do Norte and Pernambuco States, between depths of 505 and 850 m (Fig. 4). 14/28 ni.bio.br | scielo.br/ni Remarks. Two other specimens of Chaenophryne (NPM 4963, 28 mm SL; NPM 5219, 17 mm SL) could not be identified to species due to their extremely small sizes. They were collected from off Fernando de Noronha Archipelago and Rocas Atoll, between depths of 510 and 850 m (Fig. 4).

Dolopichthys Garman, 1899
Diagnosis. Females of Dolopichthys differ from those of Chaenophryne by the presence of sphenotic spines (vs. absence of sphenotic spines), opercle deeply notched posteriorly (vs. opercle not deeply notched posteriorly), pelvic bones rod shaped, with or without slight distal expansion (vs. pelvic bones triradiate or greatly expanded distally); from Oneirodes, Tyrannophryne, Phyllorhinichthys, Microlophichthys, and Danaphryne by having the dorsal margin of frontal bones nearly straight (vs. dorsal margin of frontal bones strongly convex) and subopercle long and narrow, ventral end strongly oval (vs. subopercle short and broad, ventral end nearly circular); from Ctenochirichthys, Leptacanthichthys, Chirophryne and Puck by the pectoral-fin lobe broad, shorter than the longest pectoral-fin rays (vs. pectoral-fin lobe narrow, longer than longest pectoral-fin rays); from Bertella by having the hyomandibula with a double head (vs. hyomandibula with a single head); from Dermatias by the depth of caudal peduncle less than 20% SL (vs. greater than 20% SL); from Lophodolos by the illicial apparatus emerging near the tip of snout, between the frontal bones (vs. illicial apparatus emerging from the dorsal surface of head, between or behind sphenotic spines); from Pentherichthys by having the lower jaw with a symphysial spine (vs. lower jaw without a symphysial spine, ventral margin of dentaries at symphysis concave), and caudal-fin rays without internal pigment (vs. caudal-fin rays internally pigmented); and from Spiniphryne by the skin naked or the presence of only minute, widely spaced dermal spinules, visible only with the aid of a microscope in cleared and stained specimens (vs. skin covered with close-set dermal spinules) (Pietsch, 2009 Regan, 1926, and D. pullatus Regan & Trewavas, 1932(Pietsch, 1972, 2009). The small specimen of Dolopichthys reported here and identified only to genus, however, represents the first record of the genus in Brazilian waters. The specimen was collected near the seamounts off Rio Grande do Norte State, between depths of 830 and 1,030 m (Fig. 4).

Oneirodes Lütken, 1871
Diagnosis. Oneirodes is the largest genus of the Ceratioidei, with 35 currently recognized species. Metamorphosed females of Oneirodes differ from those of all other genera of the Oneirodidae by having the posterior end of the pterygiophore of the illicium protruding from the dorsal midline of the trunk behind the head (vs. posterior end of the pterygiophore of the illicium not protruding from the dorsal midline of the trunk behind the head) (Pietsch, 2009 posterior escal appendage about onethird the length of escal bulb (vs. posterior escal appendage minute), anterior appendage anterodorsally directed, bearing numerous short filaments, and 2 unpigmented tapering filaments on anterior margin near the distal tip (vs. anterior appendage narrow, elongate, and anteroventrally directed, bearing a single short distal filament); from O. kreffti and O. posti by the esca without elongate medial appendages (vs. esca with 2 or 3 medial filaments more than twice the length of escal bulb); from O. rosenblatti and O. dicromischus by the lower jaw with fewer than 90 teeth in specimens greater than 45 mm, fewer than 60 teeth in specimens greater than 25 mm (vs. lower jaw with more than 90 teeth in specimens greater than 45 mm, more than 60 teeth in specimens greater than 25 mm), 3-9 (usually fewer than 8) teeth on vomer in specimens greater than 25 mm (vs. 8-14, usually more than 9 teeth); from O. luetkeni and O. carlsbergi by the presence of teeth on the epibranchial of the first gill arch (vs. epibranchial teeth absent); and from species of the O. schmidti group by the anterior escal appendage internally pigmented, anterolateral appendages absent (vs. anterior appendage without internal pigment, usually two pairs of filamentous anterolateral appendages) (Pietsch, 1974(Pietsch, , 2009Orr, 1991;Prokofiev, 2014a,b;Ho et al., 2016;Rajeeshkumar et al., 2017;Ho, Shao, 2019).

Geographical distribution. Oneirodes anisacanthus is widespread in the Atlantic
Ocean, with records from off eastern Greenland, the Caribbean Sea, Madeira, Gulf of Guinea, and off Cape Town, South Africa (Pietsch, 1974(Pietsch, , 2009). The two specimens collected around the Fernando de Noronha Archipelago and the seamounts off Rio Grande do Norte State, between depths of 505 and 1,030 m (Fig. 4), represent the first record of the species in Brazilian waters and in the western South Atlantic.  (Pietsch, 1974(Pietsch, , 2009Orr, 1991;Prokofiev, 2014a,b;Ho et al., 2016;Rajeeshkumar et al., 2017;Ho, Shao, 2019).

Geographical distribution.
Oneirodes carlsbergi seems to have a circumtropical distribution between approximately 18ºN and 8ºS (Pietsch, 2009;Ho et al., 2016;Ho, Shao, 2019). One specimen recorded far from this presumably circumtropical range was collected off the Irish Atlantic slope (Pietsch, 2009). Other records in the Atlantic Ocean range from 17º49'N to 5º34'S, and include two records near the Brazilian EEZ (ISH 660/66, 5º34'S 26º58'W; ISH 924/68, 3º00'S 26º16'W) (Pietsch, 1974(Pietsch, , 2009). In the present study O. carlsbergi is reported for the first time in Brazilian waters based on two specimens collected off Pernambuco State and Rocas Atoll, between depths of 650 and 800 m (Fig. 4).

CAULOPHRYNIDAE
Females of the Caulophrynidae are distinguished by having a short, deep body, more or less globular; mouth large, lower jaw usually extending posteriorly beyond the base 17/28 ni.bio.br | scielo.br/ni of the pectoral-fin lobe; jaw teeth unusually large; epibranchial and ceratobranchial teeth absent; illicium without a bulbous bacteria-filled light organ, the pterygiophore of the illicium fully embedded beneath skin of head; skin smooth and naked, spines or dermal denticles absent; lateral-line structures unusually well-developed, sense organs at the tips of cutaneous papillae; dorsal-and anal-fin rays apparently free, not interconnected by membrane, and usually longer than 60% SL; and 8 caudal-fin rays. Larvae of the Caulophrynidae are also distinguished in the Ceratioidei by the presence of pelvic fins, which are absent at all stages in other families of the suborder. Males are probably facultative parasites on females (Pietsch, 2009).

Caulophryne Goode & Bean, 1896
Diagnosis. Metamorphosed females of Caulophryne can be distinguished from those of Robia, the only other genus of the family (known from a single, 41 mm SL female collected in the western Central Pacific), by having a considerably shorter illicium (less than 130 mm vs. about 270 mm) and by the number and size of dorsal-and anal-fin rays (14-22 dorsal-fin rays, the longest ray > 70% SL vs. 6 dorsal-fin rays, the longest ray < 65% SL; 12-19 anal-fin rays, the longest ray > 60% SL vs. 5 anal-fin rays, the longest ray < 40% SL) (Pietsch, 2009).

Diagnosis. Same as for genus.
Geographical distribution. Species of Caulophryne have been reported from the Atlantic, Indian and Pacific Oceans between approximately 65ºN and 50ºS (Pietsch, 2009). Three of the four currently recognized species of the genus are known from the Atlantic Ocean: Caulophryne jordani Goode & Bean, 1896, known from the North Atlantic up to about 5ºN; Caulophryne pelagica (Brauer, 1902), recorded in the Atlantic at a single locality off Cape Verde Islands; and Caulophryne polynema Regan, 1930, recorded in the North and South Atlantic to 28ºS off Africa, with no records in the western South Atlantic (Pietsch, 1979(Pietsch, , 2009. The extremely small specimen of Caulophryne sp. reported here was collected off Rio Grande do Norte State, between depths of 35 and 100 m, and represents the first record of the genus in Brazilian waters and in the western South Atlantic (Fig. 5).

GIGANTACTINIDAE
Females of the Gigantactinidae are distinguished by having an elongate, laterally compressed body; a long slender illicium, with highly variable lengths (less than SL to 18/28 ni.bio.br | scielo.br/ni nearly five times SL) emerging from the anteriormost tip of the snout; length of head less than 35% SL; mouth nearly horizontal, upper jaw extending slightly beyond lower jaw; epibranchial and ceratobranchial teeth absent; caudal peduncle unusually long and slender, more than 20% SL; 3-10 dorsal-fin rays, 3-8 anal-fin rays; caudal fin usually incised posteriorly, 9 caudal-fin rays, usually highly elongate. Males are probably free living, never parasitic (Pietsch, 2009). In addition to the species recorded here, two species of the family have been previously reported in Brazilian waters: Gigantactis longicirra Waterman, 1939 andG. vanhoeffeni Brauer, 1902. Gigantactis longicirra is known from the Atlantic and Pacific Oceans. In the Atlantic, it occurs in the Gulf of Saint Lawrence, Canada, south along the New England slope to the Gulf of Mexico, Caribbean Sea, off Venezuela, and in the Gulf of Guinea. A single specimen was also collected off Espírito Santo State, Brazil (MNRJ 30700, 19º48'29"S 39º02'21"W;Pietsch, 2009). Gigantactis vanhoeffeni is known from the Atlantic, Indian and Pacific Oceans, with records in the Atlantic ranging from off western Greenland to the South Atlantic, including the Gulf of Mexico, Caribbean Sea, Cape Verde Islands, Gulf of Guinea, and off South Africa (Bertelsen et al., 1981;Sutton et al., 2008;Pietsch, 2009;Porteiro et al., 2017). In Brazil, G. vanhoeffeni was recorded based on specimens collected off Saint Peter and Saint Paul Archipelago (MCZ 61049, 0º34'N 30º43'W) and off Espírito Santo State (MNRJ 30708, 21º12'18"S 40º00'53"W; Costa et al., 2007;Pietsch, 2009;Mincarone et al., 2017;Melo et al., 2020: 188, as "verhoeffeni") (Fig. 5).

Gigantactis Brauer, 1902
Diagnosis. Metamorphosed females of Gigantactis are distinguished from those of Rhynchactis, the other genus of the family, by the absence of pelvic bones and by having 5-9 dorsal-fin rays (rarely 4-10) and 4-7 anal-fin rays (rarely 8) (vs. 3-4 dorsal-fin rays, rarely 5, and 3-4 anal-fin rays). They further differ from those of Rhynchactis by the following characters: frontal and parietal bones present (vs. absent), premaxilla well developed, with teeth present throughout their length (vs. premaxilla represented by a remnant bearing 0-2 teeth), maxilla reduced to threadlike remnants (vs. maxillae absent), dentary with several rows of strong recurved teeth (vs. dentary toothless or with only minute teeth), a single hypohyal (vs. two hypohyals), all caudal-fin rays unbranched (vs. 9 caudal-fins rays, 2 simple + 4 branched + 3 simple), skin spinulose (vs. skin covered with minute spinules in larger specimens, but juveniles naked), snout produced in front of mouth, illicium originating at its tip (vs. snout truncated, illicium origin slightly behind its tip), and esca consisting of an expanded luminous bulb (vs. absence of bulbous, terminal, escal light organs) (Bertelsen et al., 1981;Pietsch, 2009). Bertelsen, Pietsch & Lavenberg, 1981 (Figs. 5, 6A, 7) Diagnosis. Twenty species of Gigantactis are recognized (two of doubtful validity: G. ovifer Regan &Trewavas, 1932 andG. filibulbosus Fraser-Brunner, 1935), of which 14 are reported for the Atlantic. Gigantactis watermani differs from G. elsmani, G. kreffti, and G. perlatus by the length of the illicium (130-490% SL, rarely less than 200%, vs. 60-120% SL); from G. golovani, G. macronema, and G. gargantua (North Pacific and 20/28 ni.bio.br | scielo.br/ni eastern South Indian Ocean) by the escal filaments (distal escal filaments simple, without posterior filaments on or below its base vs. esca with distal filaments branched, several filaments emerging from and below its base); it further differs from G. gargantua by the pigmentation of distal escal filaments (heavily pigmented for more than one-half their length vs. lightly pigmented for less than one-fifth their length) and position of proximal escal filaments (restricted to the anterior margin of the escal bulb vs. not restricted to the anterior margin of escal bulb); from G. ios, G. longicauda, G. macronema, G. microdontis (eastern Pacific), and G. savagei (eastern North Pacific) by the presence of a group of anterior filaments arising from the base of esca (vs. absence), escal bulb structure (distal part of escal bulb bearing four or five pairs of stout filaments along posterior margin vs. filaments of distal part of escal bulb different from above), and length of caudalfin rays (second and seventh greater than 50% SL vs. longest caudal-fin rays less than 40% SL); from G. herwigi by the number of filaments at esca base (10 vs. less than 10), number of pair of filaments on the distal part of escal bulb (four or five, each with a pigmented swollen base vs. four, each gradually tapering and only faintly pigmented at base); from G. longicirra by the number and length of the dorsal-fin rays (4-7, all about equal in length vs. 8-10, the first and last distinctly longer than intermediate rays) and length of the first and eighth caudal-fin rays (less than 40% vs. 60-100% SL); and from G. gibbsi, G. gracilicauda, G. meadi, G. vanhoeffeni, and G. paxtoni (western South Indian Ocean and western South Pacific) by the absence of a darkly pigmented, spinulose distal prolongation in the esca (vs. presence of dark pigment) (Pietsch, 2009). Geographical distribution. Only two metamorphosed females of Gigantactis watermani were previously known, one from the eastern Tropical Atlantic (ISH 2330/71, 1º04'N 18º22'W) and another from the western Tropical Pacific, off New Caledonia (Pietsch, 2009). The specimen collected off seamounts of Rio Grande do Norte State, between depths of 700 and 1,113 m, represents the third known female specimen of the species and the first record in the South Atlantic (Fig. 5).

Gigantactis watermani
Remarks. Morphological and meristic data of the specimen agree with the description provided by Bertelsen et al. (1981) for the holotype, but some slightly differences were noted in its escal anatomy. The esca is bilaterally asymmetric, with four stout, tapering filaments present on the left side and five filaments present on the right side. The base of the most proximal filament of the right side is, however, reduced, with the structure mostly represented by the swollen, dark pigmented proximal part and a tiny unpigmented narrow tip (Fig. 7). In addition, the left filament of the most distal pair of filaments is secondarily branched, resulting in three filaments for this pair. In the holotype, the filaments of the most distal pair have a single branch. Bertelsen et al. (1981) also indicated the presence of 12 narrow unpigmented filaments on the anterior margin base of the escal bulb, but 14 filaments are present in the specimen examined (Fig. 7), a number that is within the range noted by Pietsch (2009: 467) for the species.
One additional small-sized specimen (NPM 3836, 6 mm SL) of Gigantactis collected off Rio Grande do Norte State, between depths of 35 and 100 m, was identified only to genus (Fig. 5).

Rhynchactis sp.
(Figs. 5, 6B) Geographical distribution. Two specimens were collected off Rio Grande do Norte State and Fernando de Noronha Archipelago between depths of 650 and 800 m (Fig. 5). As discussed below, they could not be identified to species, but represent the first record of the genus in Brazilian waters.

Remarks.
Of the three valid species of Rhynchactis, two occur in the Atlantic: Rhynchactis leptonema Regan, 1925and Rhynchactis macrothrix Bertelsen & Pietsch, 1998(Pietsch, 2009. Both species are poorly represented in collections and their geographic distributions are poorly known (Pietsch, 2009). Rhynchactis leptonema has been collected in a few localities of the Atlantic and Pacific (off Hawaii and Taiwan). In the Atlantic, it is known from the holotype collected in the western Tropical Atlantic (ZMUC P92133, 22/28 ni.bio.br | scielo.br/ni 8º19'N 44º35'W). Rhynchactis macrothrix is also known from widely spread localities in the Atlantic and the western Indian and western Pacific Oceans. In the Atlantic, it is known from three specimens: the holotype collected in central equatorial waters (ISH 605/74,7º55'N 32º41'W), and two specimens collected off Bermuda and in the Gulf of Mexico (Bertelsen, Pietsch, 1998;Pietsch, 2009).
The larger specimen reported here (NPM 4425, 113 mm SL; Fig. 6B) is in overall good condition but while it retains the full length of the illicium, the skin of the structure has been lost. The illicium length (208% SL) clearly indicates that it is not R. leptonema (maximum 177% SL;Bertelsen et al., 1981), being more similar in that respect to R. microthrix (210% SL; Bertelsen, Pietsch, 1998). The smaller specimen (NPM 5014) is an unidentified juvenile.
Summing up, a total of 23 species of the Ceratioidei, across 15 genera and nine families, occur in the Brazilian Exclusive Economic Zone (Tab. 1). Most of those species were reported along the northeastern coast and off oceanic islands. Given that most of the Brazilian coast has not been sufficiently explored in terms of its deep-sea fauna, these numbers are certainly an underestimate, reinforcing the need for more deepwater surveys in the Brazilian EEZ and in the western South Atlantic overall. Exploring deeper waters and trawling for longer distances will certainly result in an increase in the number of deep-water fishes known from the region.