Deep-sea dragonfishes (Teleostei: Stomiiformes) collected from off northeastern Brazil, with a review of the species reported from the Brazilian Exclusive Economic Zone

Bárbara T. Villarins1, Fabio Di Dario2, Leandro N. Eduardo3,4, Flávia Lucena-Frédou3, Arnaud Bertrand3,4, Artem M. Prokofiev5,6 and Michael M. Mincarone2

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Abstract​


EN

The Stomiiformes includes about 455 species of mostly mesopelagic fishes in the families Diplophidae, Gonostomatidae, Phosichthyidae, Sternoptychidae, and Stomiidae. Here we report 55 species of the order collected during the ABRACOS (Acoustics along the BRAzilian COaSt) expeditions off northeastern Brazil, including islands and seamounts of Fernando de Noronha Ridge. Triplophos hemingi (Diplophidae), Pachystomias microdon and Melanostomias biseriatus (Stomiidae) are recorded for the first time in the western South Atlantic. Eustomias bibulbosus, Grammatostomias ovatus and Photonectes achirus (Stomiidae) are recorded for the first time in the South Atlantic. Eustomias minimus (Stomiidae), previously known from four poorly preserved specimens, is reported for the first time in the Atlantic. Occurrences of 18 species are confirmed or recorded for the first time in the Brazilian Exclusive Economic Zone: Triplophos hemingi (Diplophidae), Gonostoma denudatum (Gonostomatidae), Aristostomias grimaldii, Astronesthes gudrunae, Bathophilus nigerrimus, Borostomias elucens, Eustomias bibulbosus, E. braueri, E. minimus, E. schmidti, Grammatostomias ovatus, G. dentatus, Leptostomias gladiator, Melanostomias biseriatus, M. bartonbeani, Pachystomias microdon, Photonectes achirus, and Photostomias goodyeari (Stomiidae). Diagnostic and distributional data for all species recorded are provided, with new anatomical information presented for Melanostomias bartonbeani and Grammatostomias ovatus. A checklist of the 108 species of Stomiiformes confirmed in Brazilian waters is also presented.

Keywords: Fernando de Noronha Archipelago, Mesopelagic fishes, Rocas Atoll, Seamounts, Southwestern Tropical Atlantic.

PT

Stomiiformes inclui cerca de 455 espécies válidas de peixes principalmente mesopelágicos, distribuídos nas famílias Diplophidae, Gonostomatidae, Phosichthyidae, Sternoptychidae e Stomiidae. Aqui, relatamos a ocorrência de 55 espécies de Stomiiformes coletadas durante as expedições ABRACOS (Acoustics along the BRAzilian COaSt) no nordeste do Brasil, incluindo as ilhas e montes submarinos da Cadeia de Fernando de Noronha. Triplophos hemingi (Diplophidae), Pachystomias microdon e Melanostomias biseriatus (Stomiidae) são registradas pela primeira vez no Atlântico Sul ocidental. Eustomias bibulbosus, Grammatostomias ovatus e Photonectes achirus (Stomiidae) são registradas pela primeira vez no Atlântico Sul. Eustomias minimus (Stomiidae), anteriormente conhecida a partir de quatro espécimes mal preservados, é reportada pela primeira vez no Atlântico. A ocorrência de 18 espécies é confirmada ou registrada pela primeira vez na Zona Econômica Exclusiva brasileira: Triplophos hemingi (Diplophidae), Gonostoma denudatum (Gonostomatidae), Aristostomias grimaldii, Astronesthes gudrunae, Bathophilus nigerrimus, Borostomias elucens, Eustomias bibulbosus, E. braueri, E. minimus, E. schmidti, Grammatostomias ovatus, G. dentatus, Leptostomias gladiator, Melanostomias biseriatus, M. bartonbeani, Pachystomias microdon, Photonectes achirus, e Photostomias goodyeari (Stomiidae). Dados diagnósticos e de distribuição de todas as espécies registradas são fornecidos, com novas informações anatômicas para Melanostomias bartonbeani e Grammatostomias ovatus. Uma lista preliminar das 108 espécies de Stomiiformes confirmadas em águas brasileiras também é apresentada.

Palavras-chave: Arquipélago Fernando de Noronha, Atol das Rocas, Atlântico Sudoeste Tropical, Montes submarinos, Peixes mesopelágicos.

Introduction​


The Stomiiformes is one of the most diverse and abundant groups of fishes in the meso- and bathypelagic zones (Gjøsaeter, Kawaguchi, 1980; Marks et al., 2020). Although they vary in body shape, fishes of the order are mostly characterized by having serial luminescent organs of unique derived morphology, an elongate body (except in most members of the Sternoptychidae), the pectoral fins situated low on body, the mouth extending beyond eye in most groups, with the maxilla partly included into mouth gape, teeth present on the maxilla and premaxilla, and chin barbel present in some groups (Morrow, 1964a; Harold, Weitzman, 1996; Prokofiev, 2005; Nelson et al., 2016). Currently, the Stomiiformes comprises about 455 valid species commonly known as bristlemouths, lightfishes, hatchetfishes, and barbeled dragonfishes, in the families Diplophidae (10 species), Gonostomatidae (24), Phosichthyidae (24), Sternoptychidae (78), and Stomiidae (319), respectively (Fink, 1985; Kenaley, Stewart, 2015a; Nelson et al., 2016; Fricke et al., 2022).

Species of the order are distributed throughout the world’s oceans (Morrow, Gibbs, 1964; Harold, Weitzman, 1996; Kenaley, Stewart, 2015a), but data on the species composition of the group and other aspects of its diversity are still scarce in the western South Atlantic. In the Brazilian Exclusive Economic Zone, which includes a substantial portion of the western South Atlantic, about 100 species of the Stomiiformes were previously reported (Gibbs, 1969; Mukhacheva, 1972; Gibbs et al., 1983; Gomon, Gibbs, 1985; Serét, Andreata, 1992; Haimovici et al., 1994; Clarke, 2000; Parin, Borodulina, 1997, 2000; Parin et al., 1999; Figueiredo et al., 2002; Mincarone et al., 2004; Bernardes et al., 2005; Bonecker, Castro, 2006; Braga et al., 2007; Haimovici et al., 2008; Lima et al., 2011; Costa, Mincarone, 2010; Goçalo et al., 2011; Bonecker et al., 2012; Perez et al., 2013; Lins Oliveira et al., 2015; Judkins, Haedrich, 2018; Eduardo et al., 2018a; Melo et al., 2020), but there are still doubts about the taxonomy, occurrence, and distribution of several of those species in the region.

In recent oceanographic cruises conducted during the ABRACOS (Acoustic along the BRAzilian COast) expeditions off northeastern Brazil and adjacent oceanic islands and seamounts, more than 9,000 specimens of mesopelagic fishes were collected, resulting in a significant increase of knowledge on the diversity and distribution of several deep-sea groups (e.g., Eduardo et al., 2018a,b, 2019, 2020a,b,c, 2021; Mincarone et al., 2019, 2020, 2021; Afonso et al., 2021). Here we report on new records and further distributional and anatomical information on the stomiiforms collected during the ABRACOS expeditions. In addition, we provide a checklist of the species of the group confirmed in the Brazilian Exclusive Econonic Zone (EEZ).

Material and methods


Specimens examined are part of a large collection of mesopelagic invertebrates and fishes made during the ABRACOS expeditions, carried out between 29 September and 21 October 2015 (ABRACOS 1 – AB1; Bertrand, 2015), and between 9 April and 10 May 2017 (ABRACOS 2 – AB2; Bertrand, 2017). Both expeditions were conducted using the French RV Antea off northeastern Brazil (Rio Grande do Norte, Paraíba, Pernambuco, and Alagoas states) and along the Fernando de Noronha Ridge, which includes the Rocas Atoll, the Fernando de Noronha Archipelago, and associated seamounts. Sampling was conducted using micronekton (body mesh 40 mm, cod-end mesh 10 mm) and mesopelagic (body mesh 30 mm, cod-end mesh 4 mm) nets. Trawl depth was continuously recorded using a Scanmar depth sensor fitted to the upper part of the trawl mouth. Open-mouth nets were employed, but pre-established target depths 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 at 2–3 kt. Therefore, collection of specimens most likely occurred at target depths, which are indicated as capture depths in the species accounts.

The survey comprised 82 fishing stations between the surface and 1,113 m depth (Fig. 1; Tab. 1), resulting in the collection of 3,113 specimens of the Stomiiformes out of a total of about 9,000 specimens of fishes. Among the stomiiforms collected, 2,220 were identified to species level and 893 were identified only to genus due to their small size and/or poor condition of preservation (S1). After capture, specimens were fixed in a 10% formalin solution and then preserved in 70% ethanol. Families and subfamilies were considered according to Kenaley, Stewart (2015a,e). Specimens were identified according to Regan, Trewavas (1929, 1930), Grey (1964), Schultz (1964), Gibbs (1964, 1971), Morrow (1964b,c,d), Morrow, Gibbs (1964, 1969), Baird (1971), Gibbs, Craddock (1973), Gibbs et al. (1983, 1984), Gomon, Gibbs (1985), Parin, Borodulina (1995, 1996, 2000, 2002, 2003), Clarke (1998, 1999, 2000, 2001), McEachran, Fechhelm (1998), Harold (2003), Kenaley, Hartel (2005), Kenaley (2007, 2009), Liao et al. (2006), Lima et al. (2011), Kenaley, Stewart (2015a,b,c,d,e,f,g,h), Harold et al. (2015), Stewart, Kenaley (2015a,b) and Sutton et al. (2020). All specimens examined are deposited in the Fish Collection of the Instituto de Biodiversidade e Sustentabilidade, Universidade Federal do Rio de Janeiro (NPM, Macaé, Brazil), except some specimens of the Sternoptychidae (Eduardo et al., 2020a) and Chauliodontinae (Eduardo et al., 2020b), which were selected for biological analyses. Fin rays and vertebrae counts follow Hubbs, Lagler (1958). Photophore terminology, barbel morphology, measurements, teeth and vertebrae counts for Eustomias Vaillant, 1884 follow Morrow, Gibbs (1964) and Gibbs et al. (1983). Vertebrae counts were accessed through radiographs taken on a Faxitron LX-60 Digital x-ray cabinet. Radiographs were also used to confirm fin ray counts. Unless stated otherwise, gill raker count refers to the total number of rakers in the first gill arch. Institutional abbreviations follow Sabaj (2020).

FIGURE 1 | Sampling stations (n = 80) of the ABRACOS 1 (squares; 29 September to 21 October 2015) and ABRACOS 2 (circles; 9 April to 10 May 2017) expeditions. Black symbols indicate stations with occurrence of Stomiiformes. Oceanic islands: RA, Rocas Atoll; FN, Fernando de Noronha Archipelago.

TABLE 1 | Sampling stations conducted by French RV Antea off northeastern Brazil during the ABRACOS 1 (AB1) and ABRACOS 2 (AB2) expeditions. Only stations with occurrence of Stomiiformes are listed.

Station

Date

Hour

Net

Coordinates

Target

Depth (m)

Initial

Final

Initial

Final

AB1#04

2 Oct 2015

14:00

14:30

Micro

03°54’26”S 32°20’21”W

03°53’17”S 32°19’25”W

90

AB1#09

4 Oct 2015

21:17

21:47

Meso

03°28’14”S 32°45’37”W

03°27’35”S 32°46’42”W

105

AB1#12

5 Oct 2015

21:24

21:54

Meso

03°56’19”S 33°30’41”W

03°56’36”S 33°32’00”W

130

AB1#14

6 Oct 2015

21:40

22:26

Meso

03°59’58”S 34°03’22”W

03°57’42”S 34°04’52”W

510

AB1#15

7 Oct 2015

11:34

12:14

Meso

03°44’02”S 33°59’58”W

03°43’07”S 34°01’17”W

537

AB1#20

7 Oct 2015

21:13

21:37

Meso

03°45’19”S 33°53’11”W

03°44’32”S 33°54’05”W

60

AB1#22

8 Oct 2015

21:32

22:12

Meso

04°07’43”S 33°47’27”W

04°07’00”S 33°48’59”W

525

AB1#25

11 Oct 2015

21:30

21:54

Meso

05°48’08”S 34°57’07”W

05°46’38”S 34°57’30”W

75

AB1#26

12 Oct 2015

12:48

13:08

Meso

06°09’13”S 34°34’35”W

06°09’49”S 34°33’45”W

560

AB1#31

13 Oct 2015

21:24

21:54

Meso

06°44’04”S 34°26’23”W

06°44’47”S 34°25’59”W

50

AB1#34

14 Oct 2015

21:30

22:00

Meso

07°11’26”S 34°15’56”W

07°12’06”S 34°15’19”W

90–100

AB1#36

15 Oct 2015

21:47

22:17

Meso

07°36’08”S 34°20’15”W

07°36’44”S 34°19’23”W

60

AB1#51

19 Oct 2015

22:09

23:35

Meso

08°56’29”S 34°29’03”W

08°59’07”S 34°28’37”W

45–200

AB1#52

20 Oct 2015

11:44

11:24

Meso

09°03’55”S 34°48’05”W

09°01’54”S 34°48’11”W

570

AB2#16

14 Apr 2017

21:53

22:39

Micro

07°36’15”S 33°59’30”W

07°36’49”S 33°57’19”W

680

AB2#21

16 Apr 2017

11:25

12:12

Micro

06°50’21”S 34°18’24”W

06°48’24”S 34°20’30”W

>800

AB2#28

18 Apr 2017

22:33

23:10

Micro

05°37’02”S 34°47’07”W

05°38’02”S 34°49’55”W

70/130

AB2#35

20 Apr 2017

22:35

23:15

Micro

04°19’37”S 35°29’52”W

04°18’32”S 35°32’20”W

630

AB2#39

24 Apr 2017

21:49

22:37

Micro

04°52’27”S 34°35’23”W

04°50’53”S 34°51’05”W

650–800

AB2#40A

26 Apr 2017

10:43

11:06

Micro

03°31’21”S 32°31’40”W

03°31’31”S 32°30’41”W

440

AB2#41A

26 Apr 2017

21:44

22:06

Micro

03°19’59”S 32°24’42”W

03°19’32”S 32°25’05”W

430

AB2#41B

26 Apr 2017

23:16

23:31

Micro

03°19’16”S 32°25’42”W

03°19’14”S 32°25’11”W

25

AB2#42A

27 Apr 2017

12:23

12:26

Micro

03°15’28”S 31°48’29”W

03°15’28”S 31°50’41”W

780

AB2#42B

27 Apr 2017

14:52

15:26

Micro

03°15’44”S 31°48’59”W

03°15’26”S 31°48’23”W

50

AB2#44A

28 Apr 2017

12:44

13:17

Micro

03°52’53”S 32°17’33”W

03°52’13”S 32°16’28”W

<850

AB2#44B

28 Apr 2017

14:47

15:02

Micro

03°52’18”S 32°17’58”W

03°52’46”S 32°18’20”W

130

AB2#45B

28 Apr 2017

22:53

23:15

Meso

04°14’19”S 32°12’52”W

04°14’23”S 32°07’00”W

50

AB2#46A

29 Apr 2017

11:20

11:45

Meso

04°08’31”S 32°18’15”W

04°08’57”S 32°17’29”W

360

AB2#48A

30 Apr 2017

10:30

10:58

Micro

04°25’05”S 32°57’52”W

04°25’25”S 32°56’56”W

505

AB2#49A

30 Apr 2017

21:17

21:52

Micro

04°10’38”S 33°16’07”W

04°10’58”S 33°15’04”W

770–1020

AB2#49B

30 Apr 2017

23:27

23:56

Micro

04°10’34”S 33°15’34”W

04°10’10”S 33°16’36”W

90

AB2#50A

1 May 2017

10:48

11:29

Micro

03°49’01”S 32°35’56”W

03°47’33”S 32°36’51”W

615

AB2#52A

2 May 2017

11:47

12:18

Micro

03°43’16”S 33°25’10”W

03°42’14”S 33°24’36”W

822–984

AB2#52B

2 May 2017

14:00

14:30

Micro

03°41’56”S 33°23’29”W

03°42’34”S 33°22’36”W

385

AB2#53A

2 May 2017

22:08

22:40

Micro

03°48’59”S 33°59’17”W

03°50’06”S 33°58’47”W

610

AB2#53B

2 May 2017

23:57

00:27

Micro

03°49’47”S 33°57’42”W

03°48’28”S 33°57’44”W

65

AB2#54B

3 May 2017

03:11

13:47

Micro

03°45’17”S 34°41’04”W

03°44’39”S 34°40’05”W

830–1030

AB2#56B

4 May 2017

12:19

 02:52

Micro

03°57’44”S 35°24’22”W

03°56’23”S 35°24’41”W

260

AB2#56C

4 May 2017

14:03

14:34

Micro

03°58’47”S 35°23’04”W

03°58’33”S 35°22’25”W

260

AB2#58A

5 May 2017

11:58

 02:29

Micro

03°56’54”S 36°06’14”W

03°57’57”S 36°09’28”W

520

AB2#59A

5 May 2017

21:57

22:37

Micro

03°38’02”S 36°31’46”W

03°38’36”S 36°17’50”W

700–1113

AB2#59B

5 May 2017

00:00

00:26

Micro

03°38’36”S 36°22’38”W

03°38’08”S 36°23’46”W

110

AB2#60B

6 May 2017

12:49

13:19

Micro

03°31’43”S 36°21’20”W

03°31’47”S 36°22’26”W

700–670

 

Results​


DIPLOPHIDAE

The Diplophidaeas considered here includes ten valid species in the genera Diplophos Günther, 1873, Manducus Goode & Bean, 1896, and Triplophos Brauer, 1902 (Nelson, 2006; Kenaley, Stewart, 2015b). Fishes of the family are characterized by having an elongate body, two rows of photophores along the body (ventral and lateral series), dorsal adipose fin absent, chin barbel absent, dorsal-fin origin in advance of anal-fin origin, 10–15 dorsal-fin rays, 36–68 anal-fin rays, more than 61 vertebrae, teeth in premaxilla uniseral and biserial in lower jaw, presence of photophores on isthmus, one orbital photophore and IC 70–115 (Grey, 1964; Johnson, 1970; Ozawa et al., 1990; Kenaley, Stewart, 2015b; Koeda, Ho, 2019).

Three species of the Diplophidae were previously reported in the Brazilian EEZ in the genera Diplophos and Manducus (Mukhacheva, 1978; Smith et al., 1991; Figueiredo et al.,2002; Menezes et al., 2003; Braga et al., 2007; Costa, Mincarone, 2010; Pinheiro et al., 2015; Judkins, Haedrich, 2018; Eduardo et al., 2020c), all of which were identified among specimens examined in this study. In addition, the occurrence of a fourth species of the family, Triplophos hemingi (McArdle, 1901), is reported for the first time in Brazilian waters.

Diplophos Günther, 18732C447C

Diagnosis. Diplophos can be distinguished in the Diplophidae by the presence of 10–15 dorsal-fin rays; 43–68 anal-fin rays; more than 74 vertebrae; IC 83–115; OA 78–97 (Grey, 1964; Ozawa et al., 1990; Kenaley, Stewart, 2015b; Koeda, Ho, 2019). Seven valid species (Fricke et al., 2022).

Diplophos australis Ozawa, Oda & Ida, 1990

(Fig. 2A)

FIGURE 2 | A. Diplophos australis, NPM 4564, 99 mm SL; B. Diplophos taenia, NPM 4566, 129 mm SL; C. Manducus maderensis, NPM 4530, 63 mm SL; D. Triplophos hemingi, NPM 4561, 196 mm SL. Scale bars = 10 mm.

Diagnosis. Diplophos australis is distinguished from congeners by the presence of 84–91 vertebrae; 12–13 gill rakers; 61–67 anal-fin rays; IV 41–44; OA 80–87; IC 99–105 (Ozawa et al., 1990; Koeda, Ho, 2019).

Geographical distribution. South Atlantic and South Pacific oceans, between 20ºS and 40ºS (Ozawa et al., 1990). In the South Atlantic, D. australis was reported from off Brazil, South Africa, Saint Helena, Ascension, and Tristan da Cunha islands (Ozawa et al., 1990; Judkins, Haedrich, 2018). In Brazilian waters, the species was previously reported from off São Paulo to Rio Grande do Sul States (Figueiredo et al., 2002; Menezes et al., 2003;Judkins, Haedrich, 2018). The specimens reported here, collected from off Fernando de Noronha Archipelago at depths between 780 and 850 m, represent the northernmost record of the species in the Atlantic Ocean (about 3º S; Fig. 3A).

Specimens examined. 3 (71–99 mm SL): NPM 4564, 2 (75–99 mm), AB2#42A; NPM 4920, 1 (71 mm), AB2#44A.

Diplophos taenia Günther, 1873

(Fig. 2B)

Diagnosis. Diplophos taenia is distinguished from congeners by the presence of 90–100 vertebrae; 11–13 gill rakers; 56–72 anal-fin rays; IV 42–48; OA 85–97; IC 104–115 (Ozawa et al., 1990; Kenaley, Stewart, 2015b; Koeda, Ho, 2019).

Geographical distribution. Circumglobal, between 40ºN and 55ºS (Mukhacheva, 1978; Pequeño, 1989; Ozawa et al., 1990; Quéro et al., 1990a; Kenaley, Stewart, 2015b; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, D. taenia is reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (McAllister, 1990; Ozawa et al., 1990; McEachran, Fechhelm, 1998; Harold, 2003; Judkins, Haedrich, 2018). Previous records of the species in Brazilian waters were made off the northeastern coast (including oceanic islands), the Vitória-Trindade Chain (Montague and Davis seamounts), and off Rio de Janeiro, Santa Catarina and Rio Grande do Sul States (Figueiredo et al.,2002; Menezes et al., 2003; Braga et al., 2007; Pinheiro et al., 2015; Eduardo et al., 2020c). Additional records are also known off northeastern Brazil, outside the EEZ (Parin et al., 1974; Olivar et al., 2017). Larvae were reported along the Vitória-Trindade Chain and off Rio de Janeiro State (Bonecker, Castro, 2006, 2014; Stocco, Joyeux, 2015). The specimens reported here were collected off the Rocas Atoll, Fernando de Noronha Archipelago, and Paraíba and Rio Grande do Norte States (including the seamounts), between depths of 25 and 1,113 m (Fig. 3A).

Specimens examined. 25 (42–129 mm SL): NPM 3291, 1 (76 mm), AB1#4; NPM 3292, 1 (51 mm), AB1#15; NPM 3293, 1 (46 mm), AB1#31; NPM 4562, 1 (100 mm), AB2#39; NPM 4563, 7 (49–106 mm), AB2#40A; NPM 4566, 1 (129 mm), AB2#59A; NPM 4567, 6 (42–73 mm), AB2#41B; NPM 4569, 1 (92 mm), AB2#49B; NPM 4614, 3 (49–77 mm), AB2#41A; NPM 6066, 3 (53–61 mm), AB2#45B.

Manducus Goode & Bean, 1896

Diagnosis. Manducus can be distinguished in the Diplophidae by the presence of 12–13 dorsal-fin rays; 36–59 anal-fin rays; IC 70–86; photophore series along lateral line (LLP) 65–78; total vertebrae 61–76 (Grey, 1964; Johnson, 1970). Two valid species (Fricke et al., 2022).

Manducus maderensis (Johnson, 1890)

(Fig. 2C)

Diagnosis. Manducus maderensis differs from Manducus greyae (Johnson, 1970) by the number of anal-fin rays (36–41 vs. 58–59); number of OA photophores (45–48 vs. 50–54); number of IC photophores (70–75 vs. 84–86); number of photophore series along lateral line (65–68 vs. 76–78); total vertebrae (61–63 vs. 76) (Grey, 1964; Johnson, 1970).

Geographical distribution. Atlantic Ocean, from 40ºN to 30ºS (Smith et al., 1991; Figueiredo et al.,2002; Sutton et al., 2020). In the western Atlantic, the species is reported from off United States to southeastern Brazil, including the Gulf of Mexico and the Caribbean Sea (Smith et al., 1991; McEachran, Fechhelm, 1998, Harold, 2003). The species was previously recorded off Rio Grande do Norte and Paraíba States, and from Bahia to Rio de Janeiro States, Fernando de Noronha Archipelago, Rocas Atoll, and the Vitória-Trindade Chain (Davis Seamount) in Brazilian waters (Mukhacheva, 1978; Smith et al., 1991; Figueiredo et al., 2002; Menezes et al., 2003; Braga et al., 2007; Costa, Mincarone, 2010; Pinheiro et al.,2015; Judkins, Haedrich, 2018). Additional records outside the Brazilian EEZ are known along the western South Atlantic (Smith et al., 1991). Manducus maderensis is reported here based on specimens collected off the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 90 and 615 m (Fig. 3A).

Specimens examined. 3 (42–65 mm SL): NPM 3259, 1 (42 mm), AB1#22; NPM 4122, 1 (65 mm), AB2#50A; NPM 4530, 1 (63 mm), AB2#49B.

Triplophos Brauer, 1902

Diagnosis. Triplophos can be distinguished in the Diplophidae by the trunk much shorter than tail; dorsal-fin origin well in advance of anal-fin origin; 10–11 dorsal-fin rays; 57–63 anal-fin rays; 9–10 pectoral-fin rays; 23–25 gill rakers; two rows of conspicuous photophores along the body; IC 68–70; OA 50–55; first 9 OA sightly lower than the remaining photophores of the series; additional head photophores above premaxilla and maxilla; presence of photophores on isthmus; maxilla with a short toothed portion, with 5–6 uniserial teeth; premaxilla with two rows of small teeth (Grey, 1964; Schaefer et al., 1986a). A single valid species (Fricke et al., 2022).

Triplophos hemingi (McArdle, 1901)

(Fig. 2D)

Diagnosis. Same as for the genus.

Geographical distribution. Atlantic, Indian, and western Pacific oceans, between 23ºN and 20ºS (Grey, 1964; Lloris, 1982; Schaefer et al., 1986a; Quéro et al., 1990a; Randall, Lim, 2000; Harold, 2003; Sutton et al., 2020). In the western Atlantic, reported from the Caribbean Sea and Suriname (Grey, 1964; Harold, 2003; Saavedra-Díaz et al., 2004). The specimen of T. hemingi identified here was collected near the seamounts off Rio Grande do Norte State, between depths of 700 and 1,113 m (Fig. 3A), and represents the first record in the western South Atlantic.

Specimen examined. 1: NPM 4561, 1 (196 mm), AB2#59A.

FIGURE 3 | Records of species of Diplophidae (A), Gonostomatidae (B, C) and Sternoptychidae (D, E, F) collected during the ABRACOS expeditions off northeastern Brazil. Oceanic islands: RA, Rocas Atoll; FN, Fernando de Noronha Archipelago.

GONOSTOMATIDAE

The Gonostomatidae includes 24 valid species in the genera Cyclothone Goode & Bean, 1883, Gonostoma Rafinesque, 1810, Margrethia Jespersen & Tåning, 1919, Sigmops Gill, 1883, and Zaphotias Goode & Bean, 1898, with a wide distribution in the Atlantic, Indian, Pacific, and Southern oceans (Gon, 1990a; Duhamel et al., 2014; Nelson et al., 2016; Koeda, Ho, 2019; Fricke et al., 2022). Fishes of the family are characterized by having an elongate and laterally compressed body, at least one row of serial photophores, chin barbel and postorbital photophore absent, adipose fins present or absent, 16–67 anal-fin rays, and 12–16 branchiostegal rays (Harold, 1998, 2003; Kenaley, Stewart, 2015c; Nelson et al., 2016). Gonostomatids also have the dorsal-fin origin near the middle of body or slightly posterior to it, one row of premaxillary teeth, and two rows of dentary teeth near the symphysis (Harold, 1998, 2003).

Fourteen species in all the five currently recognized genera of the Gonostomatidae were previously reported in Brazilian waters (Mukhacheva, 1972, 1974, 1978; Séret, Andreata, 1992; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Braga et al., 2007; Haimovici et al., 2008; Costa, Mincarone, 2010; Bonecker et al., 2014; Pinheiro et al., 2015; Stocco, Joyeux, 2015; Mincarone et al., 2017; Olivar et al., 2017; Eduardo et al., 2018a, 2020c; Judkins, Haedrich, 2018; Melo et al., 2020; Pinheiro et al., 2020; Tab. 2). Cyclothone signata Garman, 1899 was included in a list of Brazilian marine deep-sea fishes by Melo et al. (2020), likely based on NHMUK 1969.6.26.254-257 (four specimens collected near the Fernando de Noronha Archipelago). However, records of the species in the Atlantic require verification, since they actually may refer to C. alba Brauer, 1906 (Carneiro et al., 2019). Here we also report for the first time on the occurrence of Gonostoma denudatum Rafinesque, 1810in the Brazilian EEZ.

Cyclothone Goode & Bean, 1883

Diagnosis. Cyclothone can be distinguished in the Gonostomatidae by the absence of adipose fin;dorsal- and anal-fin origins about opposite; 12–15 dorsal-fin rays; 16–21 anal-fin rays; 29–35 vertebrae; maxillary teeth close-set, subequal, increasing in size posteriorly; vomerine teeth present or absent, palatine teeth only in an anterior patch; photophores present or absent, SO and isthmus series always absent; when present, photophores along the body in ventral and lateral series, OP 2, OA 6–10; caudal glands present or absent; anus closer to pelvic-fin than to anal-fin origin, or midway between origin of those fins (Grey, 1964; Mukhacheva, 1964, 1974; Badcock, 1982; Nazarkin, 2015). A total of 13 valid species (Fricke et al., 2022).

Remarks. Nine species of Cyclothone were previously reported in the Brazilian EEZ (Tab. 2). Unfortunately, all collected specimens were severely damaged and could not be identified at the species level (S1).

Gonostoma Rafinesque, 1810

Diagnosis. Gonostoma can be distinguished in the Gonostomatidae by the anal opening close to the anal-fin origin; anal-fin origin in advance of, or opposite to, dorsal-fin origin; absence of elongate rays in dorsal and anal fins; head and trunk longer than tail; 15–18 gill-rakers; no photophores on isthmus; no minute photophores on body or head; two rows of conspicuous photophores along the body (lateral and ventral series); first 5–6 IV forming an arch; upper jaw teeth in a single row, with a series of long, slender, subequal teeth interspaced with shorter teeth (Grey, 1964; Mukhacheva, 1972; Schaefer et al., 1986a). Two valid species (Schaefer et al., 1986a; Miya, Nishida, 2000; Fricke et al., 2022).

Gonostoma atlanticum Norman, 1930

(Fig. 4A)

FIGURE 4 | A. Gonostoma atlanticum, NPM 4615, 61 mm SL; B. Gonostoma denudatum, NPM 6519, 122 mm SL; C. Margrethia obtusirostra, NPM 3384, 27 mm SL; D. Sigmops bathyphilus, NPM 4764, 155 mm SL; E. Sigmops elongatus, NPM 3182, 173 mm SL; F. Zaphotias pedaliotus, NPM 4407, 62 mm SL. Scale bars = 10 mm.

Diagnosis. Gonostoma atlanticum differs from Gonostoma denudatum by the absence of adipose fin (vs. adipose fin present); 5–6+11=16–17 gill rakers (vs. 5+10=15); teeth present on vomer (vs. teeth absent on vomer); first 1 or 2 AC photophores elevated and the others at the same level (vs. first 2 AC elevated and the following 2 AC below the others) (Grey, 1964; Mukhacheva, 1972; Schaefer et al., 1986a).

Geographical distribution. Circumglobal, between 48ºN and 36ºS (Grey, 1964; Mukhacheva, 1972; Clarke, 1974; Nakabo, 2002; Hutchins, 2001; Judkins, Haedrich 2018; Burton, Lea, 2019; Sutton et al., 2020; Love et al., 2021). In the Atlantic Ocean, reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al.,2003; Judkins, Haedrich, 2018). In Brazil, the species was previously recorded off Pará, Maranhão, and Rio Grande do Norte States, and off the Saint Peter and Saint Paul Archipelago (Mukhacheva, 1972; Judkins, Haedrich, 2018). Additional records are also known off northeastern and southeastern Brazil, outside the country’s EEZ (Parin et al., 1974; Menezes et al., 2003; Stocco, Joyeux, 2015; Judkins, Haedrich, 2018). Gonostoma atlanticum is reported here based on specimens collected off Rio Grande do Norte (including seamounts), Pernambuco, and Paraíba States, the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 130 and 1,113 m (Fig. 3B).

Specimens examined. 67 (19–68 mm SL): NPM 3498, 2 (damaged), AB1#14; NPM 4532, 3 (51–57 mm), AB2#44A; NPM 4612, 5 (30–59 mm), AB2#42A; NPM 4615, 13 (45–68 mm), AB2#41A; NPM 4617, 9 (46–67 mm), AB2#40A; NPM 4619, 15 (39–57 mm), AB2#52B; NPM 4779, 5 (49–59 mm), AB2#35; NPM 4780, 1 (50 mm), AB2#52A; NPM 4781, 7 (41–56 mm), AB2#39; NPM 4897, 3 (53–60 mm), AB2#59A; NPM 4927, 2 (46–58 mm), AB2#21; NPM 5064, 1 (19 mm), AB2#28; NPM 5940, 1 (49 mm), AB2#16.

Gonostoma denudatum Rafinesque, 1810

(Fig. 4B)

Diagnosis. See Gonostoma atlanticum.

Geographical distribution. Atlantic Ocean from 40ºN to 40ºS (apparently more common in the eastern Atlantic), including the Mediterranean Sea (De Groot, Nijssen, 1971; Mukhacheva, 1972; Parin et al., 1974; Schaefer et al., 1986a; Quéro et al., 1990a; Golani, 1996; Santos et al.,1997; Bordes et al., 1999; Harold, 2003; Lipej, Dulčić, 2010; Béarez et al., 2017; Bayhan, Ergüden, 2019; Elbaraasi et al., 2019; Bariche, Fricke, 2020; Akbora et al., 2020; Sutton et al., 2020). Gonostoma denudatum is reported for the first time in Brazilian waters based on one specimen collected near the Fernando de Noronha Archipelago, at 440 m depth (Fig. 3B).

Specimen examined. 1: NPM 6519, 1 (122 mm), AB2#40A.

Margrethia Jespersen & Tåning, 1919

Diagnosis. Margrethia can be distinguished in the Gonostomatidae by the presence of adipose fin; dorsal-fin origin in advance of anal-fin origin; anterior rays of the dorsal- and anal-fin elongated; 14–16 pectoral-fin rays; 15–16 dorsal-fin rays; 21–26 anal-fin rays; no photophores on isthmus; a single row of large and irregular ventral photophores; SO absent; BR 10–13; OP 3; IV 13–16; AC 13–17 (Grey, 1964; Mukhacheva, 1976; Schaefer et al., 1986a). Two valid species (Fricke et al., 2022).

Margrethia obtusirostra Jespersen & Tåning, 1919

(Fig. 4C)

Diagnosis. Margrethia obtusirostra differs from M. valentinae Parin, 1982 by having the last photophores of the anterior AC group above the anal-fin base (vs. last photophores of the anterior AC group behind the anal-fin base); last 2 IV displaced ventrally (vs. last 2 IV in a straight line with anterior IV) (Grey, 1964; Schaefer et al., 1986a).

Geographical distribution. Circumglobal, from 43ºN to 45ºS, except the eastern Pacific Ocean (Grey, 1964; Bordes et al., 1999; Quéro et al., 1990a; Hutchins, 2001; Moore et al., 2003; Kenaley, Stewart, 2015c; Fricke et al., 2018; Sutton et al., 2020). In the western Atlantic, known from off Canada to southern Brazil, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; Parin et al., 1974; Vinnichenko, 1997; McEachran, Fechhelm, 1998; Figueiredo et al., 2002; Harold, 2003; Judkins, Haedrich, 2018). In Brazilian waters, the species was previously reported from off Bahia to Rio de Janeiro States and off Rio Grande do Sul State (Figueiredo et al., 2002; Menezes et al., 2003; Braga et al., 2007), with additional records off the Trindade and Martin Vaz Archipelago, outside the Brazilian EEZ (Parin et al., 1974; Judkins, Haedrich, 2018). Larvae were also recorded off Rio de Janeiro State (Bonecker et al., 2012, 2014). The specimen of M. obtusirostra examined here was collected off the Rocas Atoll, at 525 m depth (Fig. 3B).

Specimen examined. 1: NPM 3384, 1 (27 mm), AB1#22.

Sigmops Gill, 1883

Diagnosis. The genus was reappraised by Miya, Nishida (2000), who included in Sigmops five species previously allocated in Gonostoma: Sigmops bathyphilus (Vaillant, 1884), S. ebelingi (Grey, 1960), S. elongatus (Günther, 1878), S. gracilis (Günther, 1878),and S. longipinnis (Mukhacheva, 1972). Sigmops can be distinguished in the Gonostomatidae by the absence of elongate rays in dorsal and anal fins; head and trunk nearly as long as tail; no photophores on isthmus; two rows (lateral and ventral) of photophores; SO present; OA 11–21; IC 32–43; first 4–6 IV forming an arch or an ascending line; palatine teeth uniserial; upper jaw teeth in a single row, with a series of long, slender, subequal teeth interspaced with shorter teeth (Grey, 1964; Mukhacheva, 1972; Schaefer et al., 1986a).

Sigmops bathyphilus (Vaillant, 1884)

(Fig. 4D)

Diagnosis. Sigmops bathyphilus can be distinguished from congeners by the presence of adipose fin; anal-fin origin in line with dorsal-fin origin; 10–14 pectoral-fin rays; 11–15 dorsal-fin rays; 7–9 pelvic-fin rays; 21–26 anal-fin rays; 28–32 gill rakers; no photophores on isthmus; two rows (lateral and ventral) of minute and obscure photophores; IC 32–38; OA 14–16, in an irregular row (Grey, 1964; Mukhacheva, 1972; Kenaley, Stewart, 2015c; Coad, 2018a).

Geographical distribution. Circumglobal between 70ºN and 45ºS, except the western Indian Ocean (Grey, 1964; Schaefer et al., 1986a; Pequeño, 1989; Hutchins, 2001; Møller et al., 2010; Kenaley, Stewart, 2015c; Béarez et al., 2017; Sutton et al., 2020). In the western Atlantic, the species is reported from off Greenland, Canada, United States, the Bermudas and the Caribbean Sea, and off Rio de Janeiro State, Brazil (Mukhacheva, 1972; Grey, 1964; Harold, 2003; Moore et al., 2003; Costa, Mincarone, 2010; Møller et al., 2010; Judkins, Haedrich, 2018). Here we report on one specimen of S. bathyphilus collected off Rio Grande do Norte State, near the seamounts of the region, between depths of 700 and 1,113 m (Fig. 3C).

Specimen examined. 1: NPM 4764, 1 (155 mm), AB2#59A.

Sigmops elongatus (Günther, 1878)

(Fig. 4E)

Diagnosis. Sigmops elongatus can be distinguished from congeners by the presence of adipose fin; anal-fin origin in line with dorsal-fin origin; paired fins short; 10–13 pectoral-fin rays; 12–15 dorsal-fin rays; 28–32 anal-fin rays; 6 pelvic-fin rays; 19–21 gill rakers; two rows (lateral and ventral) of conspicuous photophores; OP 3; OA 13–15; AC 20–24; IC 41–43; first 1–2 OA elevated; first 5–6 IV forming an arch; presence of a large luminous gland behind ORB; presence of 2 infracaudal and 1–2 supracaudal luminous glands; adult size larger than 200 mm SL (Grey, 1964; Mukhacheva, 1972; Kenaley, Stewart, 2015c).

Geographical distribution. Circumglobal between 65ºN and 45ºS, including the Mediterranean Sea (Mukhacheva, 1972; Grey, 1964; Schaefer et al., 1986a; Pequeño, 1989; Quéro et al., 1990a; Nakabo, 2002; Román et al., 2000; Hutchins, 2001; Møller et al.,2010; Kenaley, Stewart, 2015c; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; Parin et al., 1974; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Judkins, Haedrich, 2018; Ramírez et al., 2019). In Brazilian waters, the species was previously reported from off Rio Grande do Norte to Rio Grande do Sul States, including the Rocas Atoll, and the Fernando de Noronha and Trindade and Martin Vaz archipelagos (Mukhacheva, 1972; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Braga et al., 2007; Haimovici et al., 2008; Mincarone et al., 2017; Eduardo et al., 2018a; Judkins, Haedrich, 2018). Larvae were also recorded off Rio de Janeiro State (Bonecker et al., 2012, 2014). Specimens of S. elongatus examined here were collected off Rio Grande do Norte (including seamounts), Paraíba, and Pernambuco States, the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 70 and 984 m (Fig. 3C).

Specimens examined. 41 (45–207 mm SL): NPM 3182, 3 (49–173 mm), AB1#51; NPM 3284, 1 (45 mm), AB1#14; NPM 4539, 4 (93–198 mm), AB2#39; NPM 4540, 4 (90–205 mm), AB2#52A; NPM 4542, 5 (134–190 mm), AB2#50A; NPM 4543, 3 (186–192 mm), AB2#44A; NPM 4544, 9 (51–184 mm), AB2#21; NPM 4546, 5 (164–207 mm), AB2#16; NPM 4547, 2 (128–190 mm), AB2#53A; NPM 4548, 2 (186–196 mm), AB2#42A; NPM 4783, 2 (47–78 mm), AB2#56C; NPM 5056, 1 (49 mm), AB2#28.

Zaphotias Goode & Bean, 1898

Diagnosis. Zaphotias can be distinguished in the Gonostomatidae by the absence of adipose fin; anal-fin origin in advance of dorsal-fin origin; anteriormost anal-fin rays greatly elongated; 14–16 pectoral-fin rays; 17–20 dorsal-fin rays; no photophores on isthmus; a single row of conspicuous ventral photophores; SO present; BR 11–13; upper jaw teeth in a single row, with 10–12 long teeth and 2–4 small teeth in each interspace (Grey, 1964; Mukhacheva, 1976). A single valid species (Fricke et al., 2022).

Zaphotias pedaliotus (Goode & Bean, 1896)

(Fig. 4F)

Diagnosis. As for the genus.

Geographical distribution. Atlantic and Indian oceans, from 43ºN to 40ºS (Grey, 1964; Mukhacheva, 1976; Schaefer et al., 1986a; Quéro et al., 1990a; Porteiro et al., 1999; Hutchins, 2001; Olivar et al., 2017; Judkins, Haedrich, 2018; Bernal et al., 2020; Sutton et al., 2020). In the western Atlantic, the species was recorded from off Canada to northeastern Brazil, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Judkins, Haedrich, 2018). In Brazilian waters, the species was previously recorded off Maranhão State, the Rocas Atoll, and the Saint Peter and Saint Paul and Fernando de Noronha archipelagos (Mukhacheva, 1976; Eduardo et al., 2018a; Judkins, Haedrich, 2018). Zaphotias pedaliotus is herein recorded from off Rio Grande do Norte State (near the seamounts), Paraíba State, Rocas Atoll, and Fernando de Noronha Archipelago, based on specimens collected between depths of 130 and 984 m (Fig. 3C).

Remarks. Until recently, this species was usually recognized as Bonapartia pedaliota Goode & Bean, 1896, including in Brazilian waters (e.g., Eduardo et al., 2018a; Judkins, Haedrich, 2018; Melo et al., 2020). However, Bonapartia Goode & Bean, 1896 is a junior homonym of Bonapartia Büttikofer, 1896 (Aves: Pycnonotidae). The species should therefore be recognized in Zaphotias Goode & Bean, 1898 (Sangster, 2021).

Specimens examined. 141 (37–81 mm SL): NPM 4229, 47 (37–72 mm), AB2#52B; NPM 4392, 28 (42–71 mm), AB2#41A; NPM 4407, 3 (53–62 mm), AB2#56B; NPM 4408, 23 (40–81 mm), AB2#42A; NPM 4409, 6 (58–61 mm), AB2#52A; NPM 4423, 29 (49–71 mm), AB2#40A; NPM 4529, 1 (56 mm), AB2#21; NPM 4611, 1 (damaged), AB2#59A; NPM 5059, 2 (57–60 mm), AB2#44A; NPM 6106, 1 (damaged), AB2#44B.

STERNOPTYCHIDAE

The Sternoptychidae comprises meso- and bathypelagic fishes distributed in the Atlantic, Indian, Pacific, and Southern oceans (Schultz, 1964; Gon, 1990b; Lima et al., 2011; Duhamel et al., 2014; Harold et al., 2015). The family includes 78 valid species in 10 genera and two subfamilies: Maurolicinae, with the genera Araiophos Grey, 1961, Argyripnus Gilbert & Cramer, 1897, Danaphos Bruun, 1931, Maurolicus Cocco, 1838, Sonoda Grey, 1959, Thorophos Bruun 1931, and Valenciennellus Jordan & Evermann, 1896; and Sternoptychinae, with the genera Argyropelecus Cocco, 1829, Polyipnus Günther, 1887, and Sternoptyx Hermann, 1781 (Lima et al., 2011; Nelson et al., 2016; Fricke et al., 2022). Species of the Sternoptychidae are typically small and characterized in the Stomiiformes by the pseudobranch present, ventral photophores series united in groups of two or more photophores, and chin barbel absent (Harold, 2003; Harold et al., 2015).

Fifteen species of the Sternoptychidae were reported in Brazilian waters, in the genera Argyripnus, Argyropelecus, Maurolicus, Polyipnus, Sternoptyx, and Valenciennellus (Baird, 1971; Borodulina, 1978; Séret, Andreata, 1992; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Braga et al., 2007; Haimovici et al., 2008; Costa, Mincarone, 2010; Lima et al., 2011; Bonecker et al., 2014; Lins Oliveira et al., 2015; Mincarone et al., 2017; Eduardo et al., 2018a; Judkins, Haedrich, 2018; Melo et al., 2020; Pinheiro et al., 2020; Tab. 2). An undescribed species of Polyipnus was also reported off the central coast of Brazil (Lima et al., 2011). Maurolicus javanicus Parin & Kobyliansky, 1993and M. muelleri (Gmelin, 1789) were also reported from off the north and northeastern coasts of Brazil (Judkins, Haedrich, 2018; Melo et al.,2020), but according to Rees et al. (2020) the distribution of these species is likely restricted to the eastern Indian Ocean and the Arctic Circle, respectively. Therefore, the occurrence of those species in Brazilian waters needs confirmation. All sternoptychids collected during the ABRACOS expeditions, listed below, were previously reported in Brazilian waters.

Maurolicinae

Members of the Maurolicinae can be recognized in the Sternoptychidae by an elongate body, not extremely compressed laterally, dorsal blade absent, and 19–27 anal-fin rays (Harold, 2003; Nelson et al., 2016).

Valenciennellus Jordan & Evermann, 1896

Diagnosis. Valenciennellus can be distinguished in the Maurolicinae by the eyes moderately tubular and directed upwards; 7–8 dorsal-fin rays; 24–25 anal-fin rays; AC in 3 to 6 groups; BR 6 (Bruun, 1931; Grey, 1964; McEachran, Fechhelm, 1998). Two valid species (Fricke et al., 2022).

Valenciennellus tripunctulatus (Esmark, 1871)

(Fig. 5A)

FIGURE 5 | A. Valenciennellus tripunctulatus, NPM 3301, 29 mm SL; B. Argyropelecus aculeatus, NPM 4133, 82 mm SL; C. Argyropelecus affinis, NPM 4089, 53 mm SL; D. Argyropelecus gigas, NPM 4090, 88 mm SL; E. Argyropelecus hemigymnus, NPM 3203, 26 mm SL; F. Argyropelecus sladeni, NPM 4102, 60 mm SL; G. Sternoptyx diaphana, NPM 4160, 28 mm SL; H. Sternoptyx pseudobscura, NPM 4103, 51 mm SL; I. Sternoptyx pseudodiaphana, NPM 4160, 59 mm SL. Scale bars = 10 mm.

Diagnosis. Valenciennellus tripunctulatus differs from V. carlsbergi Bruun 1931 by the AC in 4 to 6 (typically 5) groups (vs. AC in 3 groups), and 5 OA photophores (vs. 2) (Bruun, 1931; Grey, 1964).

Geographical distribution. Circumglobal between 55ºN and 44ºS (Grey, 1964; Parin et al., 1974; Quéro et al., 1990b; Lipej, Dulčić, 2010; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species was reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Judkins, Haedrich, 2018). In Brazilian waters, V. tripunctulatus was previously reported off Pará, Maranhão, Santa Catarina, and Rio Grande do Sul States, also in the vicinities of the Fernando de Noronha, Saint Peter and Saint Paul, and Trindade and Martin Vaz archipelagos (Figueiredo et al., 2002; Olivar et al., 2017; Judkins, Haedrich, 2018). Additional records in the western Atlantic outside the Brazilian EEZ are also known (Menezes et al., 2003; Olivar et al., 2017; Judkins, Haedrich, 2018). Larvae were recorded off Rio de Janeiro State (Bonecker et al., 2014). Specimens of V. tripunctulatus examined here were collected off Pernambuco State, Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 45 and 537 m (Fig. 3D).

Specimens examined. 19 (23–32 mm SL): NPM 3285, 7 (23–28 mm), AB1#51; NPM 3301, 3 (29–30 mm), AB1#15; NPM 3302, 3 (28 mm), AB1#14; NPM 3303, 2 (23–29 mm), AB1#22; NPM 4243, 1 (32 mm), AB2#52B; NPM 4438, 1 (31 mm), AB2#41A; NPM 4439, 2 (damaged), AB2#46A.

Sternoptychinae

Members of the Sternoptychinae can be recognized in the Sternoptychidae by the body extremely compressed laterally, presence of dorsal blade and 11–19 anal-fin rays (Harold, 2003; Nelson et al., 2016).

Argyropelecus Cocco, 1829

Diagnosis. Argyropelecus can be distinguished in the Sternoptychinae by the eyes tubular, directed upwards; dorsal blade with several spines anterior to dorsal-fin rays; abdominal keel developed; post-abdominal spines present; AB 12; SAB 6; SP 2; PAN 4 (Baird, 1971, 1986). A total of 10 valid species (Fricke et al., 2022).

Argyropelecus aculeatus Valenciennes, 1850

(Fig. 5B)

Diagnosis. Argyropelecus aculeatus differs from congeners by the presence of denticles in the SC photophores; PAN, AN and SC in separated groups; two post-abdominal spines with different sizes, the posterior one markedly larger than the anterior; posteriormost abdominal keel scale slanting forward; dorsal blade height and width approximately equal; 9–10 dorsal-fin rays; lower jaw with a pair of canine teeth (Baird, 1971; Borodulina, 1978; Lima et al., 2011; Harold et al., 2015).

Geographical distribution. Circumglobal in tropical and temperate waters, between 43ºN and 40ºS (Baird, 1971; Parin et al., 1974; Borodulina, 1978; Pequeño, 1989; Quéro et al., 1990b; McAllister, 1990; Harold, 2003; Harold et al., 2015; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species was reported from off the United States to Uruguay, including the Gulf of Mexico and the Caribbean Sea (McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Judkins, Haedrich, 2018; Ramírez et al., 2019). In the Brazilian EEZ, A. aculeatus was previously reported from off the northeast to southeast coast, including the Rocas Atoll and the Fernando de Noronha and Saint Peter and Saint Paul archipelagos (Borodulina, 1978; Figueiredo et al., 2002; Menezes et al.,2003; Bernardes et al., 2005; Braga et al.,2007; Haimovici et al., 2008; Costa, Mincarone, 2010; Lima et al., 2011; Judkins, Haedrich, 2018; Eduardo et al., 2018a, 2020a). Larvae were also recorded off Rio de Janeiro State (Bonecker et al.,2014). Specimens of A. aculeatus examined here were collected off Rio Grande do Norte (including seamounts), Paraíba, and Pernambuco States, in addition to the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 430 and 1,113 m (Fig. 3D).

Specimens examined. 43 (30–82 mm SL): NPM 4112, 6 (32–46 mm), AB2#16; NPM 4113, 1 (46 mm), AB2#39; NPM 4114, 1 (45 mm), AB2#60B; NPM 4120, 1 (42 mm), AB2#59A; NPM 4125, 1 (48 mm), AB2#52B; NPM 4131, 22 (34–82 mm), AB2#58A; NPM 4133, 1 (82 mm), AB2#41A; NPM 4142, 1 (36 mm), AB2#21; NPM 4187, 9 (30–67 mm), AB2#56C.

Argyropelecus affinis Garman, 1899

(Fig. 5C)

Diagnosis. Argyropelecus affinis differs from congeners by the PAN, AN and SC not distinctly separated in groups, organized in a nearly straight line; dorsal blade reduced, its height less than one-third of its length; ventral keel scales not extending below abdominal photophores; laterally directed postorbital spine absent (Baird, 1971, 1986; Borodulina, 1978).

Geographical distribution. Circumglobal in tropical and temperate waters, between 43ºN and 35ºS (Norman, 1939; Baird, 1971; Pequeño, 1989; Quéro et al., 1990b; Harold, 2003; Vélez, 2009; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species was reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (Uyeno, Aizawa, 1983; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al.,2003; Judkins, Haedrich, 2018). In Brazilian waters, A. affinis was previously reported off the northeastern coast and adjacent oceanic islands (Borodulina, 1978; Eduardo et al., 2018a, 2020a), with additional records in the western Atlantic outside the Brazilian EEZ (Parin et al., 1974; Menezes et al., 2003; Olivar et al., 2017; Judkins, Haedrich, 2018). The specimens examined here were collected from off the Paraíba and Rio Grande do Norte States, including seamounts, and off the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 50 and 850 m (Fig. 3D).

Specimens examined. 378 (28–78 mm SL): NPM 4089, 31 (28–68 mm), AB2#52B; NPM 4094, 7 (45–61 mm), AB2#58A; NPM 4110, 24 (45–59 mm), AB2#48A; NPM 4111, 4 (damaged), AB2#42A; NPM 4134, 2 (53–65 mm), AB2#60B; NPM 4135, 27 (33–68 mm), AB2#45B; NPM 4180, 46 (33–78 mm), AB2#41A; NPM 4210, 14 (44–65 mm), AB2#39; NPM 4211, 210 (35–73 mm), AB2#35; NPM 4426, 11 (48–57 mm), AB2#44A; NPM 4428, 1 (48 mm), AB2#40A; NPM 4436, 1 (39 mm), AB2#21.

Argyropelecus gigas Norman, 1930

(Fig. 5D)

Diagnosis. Argyropelecus gigas differs from congeners by the PAN, AN, and SC not distinctly separated in groups; dorsal blade developed, its height more than one-third its length; a prominent, laterally directed, postorbital (sphenotic) spine near the postero-dorsal margin of the eye (Baird, 1971; Borodulina, 1978; Harold et al., 2015).

Geographical distribution. Circumglobal in tropical and temperate waters, between 65ºN and 40ºS (Baird, 1971; Borodulina, 1978; Pequeño, 1989; Quéro et al., 1990b; Randall, Lim, 2000; Harold, 2003; Møller et al., 2010; Harold et al., 2015; Sutton et al., 2020). In the western Atlantic, reported from off Greenland to Uruguay, including the Gulf of Mexico and the Caribbean Sea (McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Møller et al., 2010; Nión et al., 2016; Afonso et al., 2018; Judkins, Haedrich, 2018). In Brazilian waters the species was previously reported off Saint Peter and Saint Paul Archipelago, and Rio Grande do Norte and Rio Grande do Sul States (Borodulina, 1978; Bernardes et al.,2005; Haimovici et al., 2008). Additional records in the western Atlantic outside the Brazilian EEZ are also known (Parin et al., 1974; Borodulina, 1978; Judkins, Haedrich, 2018). Specimens of A. gigas examined here were collected off Rio Grande do Norte State, near the seamounts, and the Rocas Atoll, between depths of 610 and 700 m (Fig. 3E).

Specimens examined. 9 (78–91 mm SL): NPM 4090, 2 (78–88 mm), AB2#35; NPM 4096, 5 (84–90 mm), AB2#53A; NPM 4132, 2 (83–91 mm), AB2#60B.

Argyropelecus hemigymnus Cocco, 1829

(Fig. 5E)

Diagnosis. Argyropelecus hemigymnus differs from congeners by the PAN, AN and SC in separated groups; AN series length shorter than the gap between AC and SC; a single post-abdominal spine with serrated margins; an elongated, slender posterior body trunk; 8 dorsal-fin rays (Baird, 1971; Borodulina, 1978; Lima et al., 2011; Harold et al., 2015).

Geographical distribution. Circumglobal in tropical and temperate waters, between 66ºN and 60ºS, including the Mediterranean Sea (Baird, 1971; Borodulina, 1978; Pequeño, 1989; Quéro et al., 1990b; Harold, 2003; Møller et al., 2010; Harold et al., 2015; Elbaraasi et al., 2019; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species was reported from off Greenland to Argentina, including the Gulf of Mexico and the Caribbean Sea (Parin et al., 1974; Borodulina, 1978; Baird, 1971; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Møller et al., 2010; Judkins, Haedrich, 2018). Argyropelecus hemigymnus was also reported along almost the entire Brazilian coast, and off Saint Peter and Saint Paul Archipelago (Borodulina, 1978; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Haimovici et al., 2008; Costa, Mincarone, 2010; Lima et al., 2011; Olivar et al., 2017; Judkins, Haedrich, 2018; Eduardo et al., 2020a). Larvae were recorded off Rio de Janeiro State (Bonecker et al., 2014). Specimens of A. hemigymnus examined here were collected off Pernambuco, Alagoas, and Rio Grande do Norte States (including the seamounts), and off the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 260 and 1,113 m (Fig. 3E).

Specimens examined. 53 (8–31 mm SL): NPM 3201, 7 (8–20 mm), AB1#15; NPM 3202, 3 (27–30 mm), AB1#14; NPM 3203, 5 (21–26 mm), AB1#22; NPM 3204, 11 (16–22 mm), AB1#26; NPM 3999, 2 (11–13 mm), AB1#52; NPM 4136, 4 (14–28 mm), AB2#59A; NPM 4137, 3 (23–26 mm), AB2#16; NPM 4138, 2 (26–31 mm), AB2#42A; NPM 4139, 1 (25 mm), AB2#52A; NPM 4140, 5 (22–28 mm), AB2#41A; NPM 4188, 2 (17–21 mm), AB2#56C; NPM 4429, 1 (22 mm), AB2#44A; NPM 4432, 5 (19–22 mm), AB2#46A; NPM 4456, 1 (21 mm), AB2#46B; NPM 5734, 1 (15 mm), AB2#52B.

Argyropelecus sladeni Regan, 1908

(Fig. 5F)

Diagnosis. Argyropelecus sladeni differs from congeners by the PAN, AN and SC in separated groups; denticles in SC photophores absent; two post-abdominal spines with similar sizes; a long, outwardly hooked upper preopercular spine; lower jaw teeth uniform in size; dorsal blade low, its height about three or more times its length; 9–10 dorsal-fin rays (Baird, 1971; Borodulina, 1978; Lima et al., 2011; Harold et al., 2015).

Geographical distribution. Circumglobal in tropical and temperate waters, between 60ºN and 41ºS (Baird, 1971; Borodulina, 1978; Pequeño, 1989; Quéro et al., 1990b; Aizawa, 2002; Harold, 2003; Harold et al., 2015; Orlov, Tokranov, 2019; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Baird, 1971; Borodulina, 1978; Uyeno, Aizawa, 1983; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Judkins, Haedrich, 2018). In Brazilian waters, the species was previously reported off Pará, Rio Grande do Norte, and Bahia States, in addition to the Fernando de Noronha and Saint Peter and Saint Paul archipelagos (Baird, 1971; Borodulina, 1978; Menezes et al., 2003; Lima et al., 2011; Olivar et al., 2017; Eduardo et al., 2018a; Judkins, Haedrich, 2018). Larvae were recorded off Rio de Janeiro State (Bonecker et al., 2014). The specimens of A. sladeni examined here were collected off Rio Grande do Norte State, including seamounts, and the Fernando de Noronha Archipelago, between depths of 50 and 850 m (Fig. 3E).

Specimens examined. 14 (32–65 mm SL): NPM 4091, 1 (41 mm), AB2#48A; NPM 4102, 1 (60 mm), AB2#42A; NPM 4106, 2 (51–64 mm), AB2#39; NPM 4116, 1 (54 mm), AB2#58A; NPM 4141, 2 (39–40 mm), AB2#41A; NPM 4182, 6 (32–61 mm), AB2#45B; NPM 4427, 1 (65 mm), AB2#44A.

Sternoptyx Hermann, 1781

Diagnosis. Sternoptyx can be distinguished in the Sternoptychidae by the body extremely compressed; eyes not tubular; dorsal blade with one spine; anterior margin of dorsal blade serrated; abdominal keel developed; post-abdominal spines present; AB 10; BR 3; I 5; PO externally indistinct and ventral to orbit (Baird, 1971; Borodulina, 1978; Lima et al., 2011). Four valid species (Fricke et al., 2022).

Sternoptyx diaphana Hermann, 1781

(Fig. 5G)

Diagnosis. Sternoptyx diaphana differs from congeners by the SAN slightly elevated, its height less than half the distance between the ventral body margin and its midline; posterior margin of AN and ventral margin of the anal-fin base forming a narrow V shaped line; body depth larger than standard length; 9–11 dorsal-fin rays, usually less than 11; 28 vertebrae, occasionally 27 or 29 (Baird, 1971; Borodulina, 1978; Aizawa, 2002; Lima et al., 2011).

Geographical distribution. Circumglobal in tropical and temperate waters, between 55ºN and 55ºS (Baird, 1971; Borodulina, 1978; Pequeño, 1989; Quéro et al., 1990b; Lima et al.,2011; Harold, 2003; Harold et al., 2015; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species is reported from off Canada to Argentina, including the Gulf of Mexico and the Caribbean Sea (Baird, 1971; Parin et al., 1974; Uyeno, Aizawa, 1983; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Nión et al., 2016; Judkins, Haedrich, 2018; Ramírez et al., 2019). The species was previously recorded off Maranhão, Rio Grande do Norte, Bahia, Espírito Santo, Rio de Janeiro, and Rio Grande do Sul States in Brazilian waters, in addition to the Fernando de Noronha and Saint Peter and Saint Paul archipelagos (Baird, 1971; Borodulina, 1978; Séret, Andreata, 1992; Figueiredo et al., 2002; Menezes et al., 2003; Braga et al., 2007; Costa, Mincarone, 2010; Lima et al., 2011; Lins Oliveira et al., 2015; Olivar et al., 2017; Judkins, Haedrich, 2018; Eduardo et al., 2018a, 2020a; Pinheiro et al., 2020). Larvae were recorded off Rio de Janeiro State (Bonecker et al., 2012, 2014). Sternoptyx diaphana is reported here based on specimens collected off Paraíba, Pernambuco, and Rio Grande do Norte States, including the seamounts, in addition to the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 130 and 1,113 m (Fig. 3F).

Specimens examined. 1,046 (11–43 mm SL): NPM 4152, 95 (14–43 mm), AB2#54B; NPM 4154, 2 (27–28 mm), AB2#56B; NPM 4155, 3 (26–31 mm), AB2#35; NPM 4156, 12 (19–35 mm), AB2#21; NPM 4157, 80 (14–35 mm), AB2#44A; NPM 4158, 10 (23–29 mm), AB2#58A; NPM 4160, 91 (13–38 mm), AB2#59A; NPM 4161, 17 (16–42 mm), AB2#16; NPM 4167, 23 (18–33 mm), AB2#41A; NPM 4185, 1 (21 mm), AB2#50A; NPM 4186, 466 (15–33 mm), AB2#39; NPM 4189, 1 (22 mm), AB2#56C; NPM 4215, 65 (11–36 mm), AB2#42A; NPM 4231, 173 (14–34 mm), AB2#53A; NPM 4327, 2 (12–17 mm), AB2#52A; NPM 4453, 1 (26 mm), AB2#44B; NPM 4506, 3 (17–32 mm), AB2#60B; NPM 5011, 1 (15 mm), AB2#53B.

Sternoptyx pseudobscura Baird, 1971

(Fig. 5H)

Diagnosis. Sternoptyx pseudobscura differs from congeners by the SAN distinctly elevated, positioned higher or near mid-trunk line; posterior anal-fin pterygiophores nearly as long as the anterior ones; 9–11 dorsal-fin rays; usually 29 vertebrae, occasionally 27, 28, 30 or 31 (Baird, 1971, 1986; Borodulina, 1978; Badcock, Baird, 1980; Lima et al., 2011).

Geographical distribution. Circumglobal in tropical and temperate waters, between 40ºN and 40ºS (Baird, 1971; Borodulina, 1977, 1978; Quéro et al., 1990b; Harold, 2003; Lima et al.,2011; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species is reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (Baird, 1971; Uyeno, Aizawa, 1983; McAllister, 1990; Harold, 2003; Milkova et al., 2016; Judkins, Haedrich, 2018; Ramírez et al., 2019). In Brazilian waters, S. pseudobscura was previously reported off the northeastern coast and adjacent oceanic islands, and from Bahia to Rio de Janeiro States, with additional records outside the Brazilian EEZ off the northeast and south regions (Baird, 1971; Borodulina, 1978; Menezes et al., 2003; Braga et al.,2007; Costa, Mincarone, 2010; Lima et al.,2011; Eduardo et al., 2018a, 2020a; Judkins, Haedrich, 2018). The speciesis reported here based on specimens collected between depths of 520 and 1,113 m off Paraíba, Pernambuco, and Rio Grande do Norte States, including the seamounts, in addition to the Fernando de Noronha Archipelago (Fig. 3F).

Specimens examined. 58 (13–58 mm SL): NPM 4101, 4 (46–54 mm), AB2#21; NPM 4103, 1 (51 mm), AB2#39; NPM 4104, 4 (36–51 mm), AB2#16; NPM 4108, 15 (22–59 mm), AB2#59A; NPM 4117, 2 (30–45 mm), AB2#58A; NPM 4153, 5 (39–45 mm), AB2#54B; NPM 4159, 12 (17–56 mm), AB2#42A; NPM 4214, 1 (55 mm), AB2#60B; NPM 4468, 14 (13–42 mm), AB2#44A.

Sternoptyx pseudodiaphana Borodulina, 1977

(Fig. 5I)

Diagnosis. Sternoptyx pseudodiaphana differs from congeners by the SAN slightly elevated; body depth approximately equal to SL; SAN photophore height 33–50% body depth; posterior ACB photophore adjacent to or adjoining the anal-fin base; 9–13 dorsal-fin rays, usually 11 or 12; 30–32 vertebrae, rarely 29 (Borodulina, 1978; Badcock, Baird, 1980; Lima et al., 2011; Harold et al., 2015).

Geographical distribution. Circumglobal in the Southern Hemisphere to 50º S, with a few records in the North Atlantic, between 15ºN and 20ºN (Badcock, Baird, 1980; Pequeño, 1989; Lima et al., 2011; Harold et al, 2015; Judkins, Haedrich, 2018; Sutton et al., 2020). In the western Atlantic, the species was reported off Brazil, in Rio Grande do Norte and Rio de Janeiro States, and near the Saint Peter and Saint Paul Archipelago (Costa, Mincarone, 2010; Lima et al., 2011; Lins Oliveira et al., 2015; Judkins, Haedrich, 2018). Sternoptyx pseudodiaphana is reported here based on specimens collected between depths of 822 and 984 m off the Rocas Atoll and Fernando de Noronha Archipelago (Fig. 3F).

Specimens examined. 3 (42–59 mm SL): NPM 4166, 1 (59 mm), AB2#52A; NPM 4467, 2 (42–46 mm), AB2#44A.

PHOSICHTHYIDAE

The Phosichthyidae is a family of meso- and bathypelagic fishes with a wide distribution in the Atlantic, Indian, and Pacific oceans (Kenaley, Stewart, 2015d). A total of 24 species are recognized in the genera Ichthyococcus Bonaparte, 1840, Phosichthys Hutton, 1872, Pollichthys Grey, 1959, Polymetme McCulloch, 1926, Vinciguerria Jordan & Evermann, 1896, Woodsia Grey, 1959, and Yarrella Goode & Bean, 1896 (Fricke et al., 2022). Phosichthyids are characterized by an elongate and laterally compressed body, with two rows of serial photophores, photophores present on isthmus, and chin barbel absent (Harold, 2003; Kenaley, Stewart, 2015d). Except for Polymetme and Yarrella, the species of Phosichthyidae have two orbital photophores (suborbital and postorbital) (Harold, 2003).

Nine species of the Phosichthyidae were previously reported in Brazilian waters, in the genera Ichthyococcus, Phosichthys, Pollichthys, Polymetme,and Vinciguerria (Gorbunova, 1972; Krefft, 1983; Haimovici et al., 1994; Vasconcelos-Filho et al., 1996; Figueiredo et al., 2002; Menezes et al., 2003; Mincarone et al., 2004; Bernardes et al., 2005; Bonecker, Castro, 2006; Braga et al., 2007; Haimovici et al., 2008; Costa, Mincarone, 2010; Bonecker et al., 2012, 2014, 2019; Perez et al., 2013; Lins Oliveira et al., 2015; Pinheiro et al., 2015, 2020; Mincarone et al., 2017; Judkins, Haedrich, 2018; Melo et al., 2020; Tab. 2). Melo et al. (2020: 201) included Polymetme corythaeola (Alcock, 1898) in a list of Brazilian deep-sea fishes. However, as far as we were able to check, there is no confirmed records of that species in the Brazilian EEZ. The record of Haimovici et al. (1994) of P. corythaeola in southern Brazil actually refers to P. thaeocoryla Parin & Borodulina, 1990 (Menezes et al., 2003). All species of the Phosichthyidae identified in the ABRACOS expeditions and listed below were previously reported for Brazilian waters.

Ichthyococcus Bonaparte, 1840

Diagnosis. Ichthyococcus is distinguished in the Phosichthyidae by the relatively small mouth, reaching to about mid-eye; premaxilla excluded from mouth gape; jaw teeth uniserial and closely spaced; two ORB (1 anterior to eye and 1 mid-ventral or posterior to it); anal-fin base well behind dorsal-fin base; adipose fin present; 10–15 dorsal-fin rays; 13–17 anal-fin rays; eyes tubular and somewhat upward directed; two rows of serial photophores (ventral and lateral series); BR 11–13; IV 25–28; VAV 9–14; AC 12–14; OA 23–31 (Grey, 1964; Mukhacheva, 1980; Fujii, 1983a). Seven valid species (Fricke et al., 2022).

Ichthyococcus polli Blache, 1964

(Fig. 6A)

FIGURE 6 | A. Ichthyococcus polli, NPM 4119, 51 mm SL; B. Phosichthys argenteus, NPM 4560, 64 mm SL; C. Pollichthys mauli, NPM 3290, 38 mm SL; D. Vinciguerria nimbaria, NPM 3305, 27 mm SL. Scale bars = 10 mm.

Diagnosis. Ichthyococcus polli differs from congeners by the presence of the abdominal adipose fin; interorbital space 7.5–10.0% HL; body depth 26.9–31.9% SL; 20–27 gill-rakers; SO present; IV photophores in a straight line, only 8th IV elevated; AC in a straight line; OA 28–29; VAV 11–13; IC 50–53 (Mukhacheva, 1980; Fujii, 1983a; Schaefer et al., 1986b).

Geographical distribution. Atlantic Ocean, between 44ºN and 20ºS (Krefft, 1983; Quéro et al., 1990c; Sutton et al., 2020). In the western Atlantic, previously recorded off United States, Suriname, French Guiana, and Brazil (off Saint Peter and Saint Paul Archipelago) (Blache, 1963; Mukhacheva, 1980; Fujii, 1983a; Krefft, 1983; Schaefer et al., 1986b). Ichthyococcus polli is reported here based on specimens collected off Paraíba State, Rio Grande do Norte State (near the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 110 and 1,113 m (Fig. 7A).

FIGURE 7 | Records of species of Phosichthyidae (A), Astronesthinae (B, C), Chauliodontinae (D), Stomiinae (E), and Malacosteinae (F) collected during the ABRACOS expeditions off northeastern Brazil. Oceanic islands: RA, Rocas Atoll; FN, Fernando de Noronha Archipelago.

Specimens examined. 14 (41–72 mm SL): NPM 3258, 1 (41 mm), AB1#14; NPM 4148, 1 (69 mm), AB2#59A; NPM 4149, 1 (51 mm), AB2#21; NPM 4150, 7 (44–59 mm), AB2#52B; NPM 4151, 1 (72 mm), AB2#58A; NPM 4339, 1 (50 mm), AB2#42A; NPM 4410, 1 (53 mm), AB2#40A; NPM 4926, 1 (53 mm), AB2#52A.

Phosichthys Hutton, 1872

Diagnosis. Phosichthys is a monotypic genus distinguished in the Phosichthyidae by the presence of two ORB (one anterior to eye and one mid-ventral or posterior to it); eyes not tubular; lower jaw teeth uniserial; teeth on vomer and palatine; dorsal-fin origin posterior to pelvic-fin origin; dorsal adipose fin present; dorsal-fin base and dorsal adipose-fin base much shorter than anal-fin base; 12–13 dorsal-fin rays; 23–26 anal-fin rays; 15–18 gill rakers; IC 57–58; AC 16–18 (Schaefer et al., 1986b; Kenaley, Stewart, 2015d; Sutton et al., 2020).

Phosichthys argenteus Hutton, 1872

(Fig. 6B)

Diagnosis. Same as for the genus.

Geographical distribution. Cincumglobal in the Southern Hemisphere, except the eastern Pacific Ocean (Quéro et al., 1990c; Hutchins, 2001; Kenaley, Stewart, 2015d; Balushkin, 2017; Sutton et al., 2020). In the western Atlantic, the species was recorded off Brazil (from Bahia to Rio Grande do Sul States), Uruguay, and off the Malvinas Islands (Parin et al., 1974; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Braga et al., 2007; Haimovici et al., 2008; Nión et al., 2016). Phosichthys argenteus is reported here based on one specimen collected at 63 m depth off Rio Grande do Norte State, near the seamounts (Fig. 7A).

Specimen examined. 1: NPM 4560, 1 (64 mm), AB2#35.

Pollichthys Grey, 1959

Diagnosis. Pollichthys is a monotypic genus that can be distinguished in the Phosichthyidae by the presence of two ORB (one anterior to eye and one mid-ventral or posterior to it); eyes not tubular; premaxillary teeth uniserial; 15–17 gill rakers; pseudobranchiae absent; pelvic-fin origin in advance of dorsal-fin origin; anal-fin base length more than twice the dorsal-fin base length; 10–12 dorsal-fin rays; 22–30 anal-fin rays; photophores present on isthmus; ORB 2; OP 3; SO present; IC 47–50; OA 19–21 (Grey, 1964; Schaefer et al., 1986b; Sutton et al., 2020).

Pollichthys mauli (Poll, 1953)

(Fig. 6C)

Diagnosis. Same as for the genus.

Geographical distribution. Atlantic, western and central Pacific, and western Indian oceans, between 47ºN and 36ºS (Grey, 1964; Schaefer et al., 1986b; Quéro et al., 1990c; Aizawa, 2002; Mundy, 2005; Fricke et al., 2009; Judkins, Haedrich, 2018; Sutton et al., 2020). In the western Atlantic, the species is reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; Bekker et al., 1975; McAllister, 1990; McEachran, Fechhelm, 1998; Schmitter-Soto et al., 2000; Harold, 2003; Nión et al., 2016; Judkins, Haedrich, 2018). In Brazil, Pollichthys mauliwas previously recorded from off Rio Grande do Norte to Rio Grande do Sul States (Vasconcelos-Filho et al., 1996; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al.,2005; Braga et al., 2007; Haimovici et al., 2008; Costa, Mincarone, 2010), and near the Vitória-Trindade Chain (Montague and Davis seamounts; Pinheiro et al., 2015). Larvae were recorded between Bahia and São Paulo States, including the Vitória-Trindade Chain (Bonecker, Castro, 2006; Goçalo et al., 2011; Bonecker et al.,2012, 2014). Pollichthys mauli is reported here based on one specimen collected between depths of 75 and 525 m off Rio Grande do Norte State (Fig. 7A).

Specimen examined. 1: NPM 3290, 1 (38 mm), AB1#25.

Vinciguerria Jordan & Evermann, 1896

Diagnosis. Vinciguerria can be distinguished in the Phosichthyidae by the presence of two ORB (one anterior to eye and one mid-ventral or posterior to eye); eyes round or slightly tubular; teeth uniserial in upper jaw and biserial on anterior portion of lower jaw; pseudobranchiae absent; pelvic-fin origin in advance of dorsal-fin origin; anal-fin base length similar to or slightly shorther than dorsal-fin base; dorsal adipose fin present, above posterior portion of anal; 12–16 dorsal-fin rays; 12–17 anal-fin rays; photophores present on isthmus; OP 3; IC 39–48; OA 18–25; 14–36 gill rakers (Grey, 1964; Gorbunova, 1972; Johnson, Feltes, 1984; Schaefer et al., 1986b). Five valid species (Fricke et al., 2022).

Vinciguerria nimbaria (Jordan & Williams, 1895)

(Fig. 6D)

Diagnosis. Vinciguerria nimbaria differs from congeners by the eyes not tubular, lateral; interorbital width broad; 6–10 small teeth on premaxilla; SO present; IV 21–24; IC 45–46; 17–26 gill rakers; 13–15 dorsal-fin rays; 13–16 anal-fin rays; 9–11 pectoral-fin rays; 7 pelvic-fin rays; dorsal-adipose fin origin near vertical through end of anal-fin base; 39–44 vertebrae (Grey, 1964; Gorbunova, 1972; Schaefer et al., 1986b; Kenaley, Stewart, 2015d).

Geographical distribution. Circumglobal, between 48ºN and 45ºS (Grey, 1964; Pequeño, 1989; Quéro et al., 1990c; Kenaley, Stewart, 2015d; Judkins, Haedrich, 2018; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species is reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Grey, 1964; Parin et al., 1974; McAllister, 1990; McEachran, Fechhelm, 1998; Harold, 2003; Judkins, Haedrich, 2018). In Brazilian waters, V. nimbaria was recorded off Maranhão, Rio Grande do Norte, Paraíba and from off Bahia to Rio Grande do Sul States, in addition to the Trindade and Martin Vaz and Saint Peter and Saint Paul archipelagos, and the Vitória-Trindade Chain, near the Montague Seamount (Gorbunova, 1972; Figueiredo et al., 2002; Menezes et al., 2003; Braga et al., 2007, Judkins, Haedrich, 2018; Pinheiro et al., 2015, 2020). Larvae were recorded from Bahia to São Paulo States, including the Vitória-Trindade Chain (Bonecker, Castro, 2006; Goçalo et al., 2011; Bonecker et al., 2012, 2014, 2019; Stocco, Joyeux, 2015). Vinciguerria nimbaria is reported here based on specimens collected between depths of 50 and 780 m off Rio Grande do Norte, Paraíba, and Pernambuco States, and from the vicinities of the Rocas Atoll (Fig. 7A).

Specimens examined. 22 (17–49 mm SL): NPM 3305, 1 (27 mm), AB1#14; NPM 3385, 8 (21–32 mm), AB1#12; NPM 3386, 1 (21 mm), AB1#20; NPM 3387, 3 (18–25 mm), AB1#31; NPM 3388, 3 (17–21 mm), AB1#26; NPM 4568, 2 (31–38 mm), AB2#49B; NPM 4618, 1 (21 mm), AB1#36; NPM 4623, 1 (22 mm), AB2#42A; NPM 4765, 2 (49 mm), AB2#16.

STOMIIDAE

The Stomiidae is a family of meso- and bathypelagic fishes with a wide distribution in the Atlantic, Indian, Pacific, and Southern oceans (Fink, 1985; Gon, 1990c; Duhamel et al., 2014; Kenaley, Stewart, 2015e). It is the most diverse family of the Stomiiformes, with a total of 28 genera and about 319 valid species (Fink, 1985; Nelson et al., 2016; Fricke et al., 2022). The family is generally regarded as monophyletic (e.g., Fink, 1985; Nelson et al., 2016; Kenaley, Stewart, 2015e), but subfamilies within the assemblage are likely not monophyletic and are sometimes treated as distinct families (e.g., Harold, 2003; Parin, Borodulina, 2003; Prokofiev, 2014, 2015, 2020). Here we follow Kenaley, Stewart (2015e), who provisionally recognized six subfamilies of stomiid fishes: Astronesthinae, Chauliodontinae, Idiacanthinae, Malacosteinae, Melanostomiinae, and Stomiinae. In general, members of the Stomiidae have an elongated body, mouth large to extremely large, teeth often fang-like, sometimes small, no true gill rakers in adults, two rows of photophores along the body, and a chin barbel often ornate and associated with the hyoid apparatus present in most species (Kenaley, Stewart, 2015e; Nelson et al., 2016).

At least 50 species of the Stomiidae were previously reported in Brazillian waters, in the genera Aristostomias Zugmayer, 1913, Astronesthes Richardson, 1845, Bathophilus Giglioli, 1882, Chauliodus Bloch & Schneider, 1801, Echiostoma Lowe, 1843, Eustomias Vaillant 1884, Flagellostomias Parr, 1927, Grammatostomias Goode & Bean, 1896, Heterophotus Regan & Trewavas, 1929, Idiacanthus Peters, 1877, Leptostomias Gilbert, 1905, Malacosteus Ayres, 1848, Melanostomias Brauer, 1902, Photonectes Günther, 1887, Photostomias Collett, 1889, Stomias Cuvier, 1816, and Thysanactis Regan & Trewavas, 1930 (Gibbs, 1969; Parin, Novikova, 1974; Shcherbachev, Novikova, 1976; Gibbs et al., 1983; Gomon, Gibbs, 1985; Séret, Andreata, 1992; Haimovici et al.,1994; Parin, Borodulina, 1996, 1997, 2000; Parin et al., 1999; Clarke, 2000; Figueiredo et al., 2002; Menezes et al., 2003; Mincarone et al., 2004; Bernardes et al., 2005; Kenaley, Hartel, 2005; Bonecker, Castro, 2006; Braga et al., 2007; Haimovici et al., 2008; Costa, Mincarone, 2010; Lins Oliveira et al., 2015; Pinheiro et al., 2015; Mincarone et al., 2017; Eduardo et al., 2018a, 2020c; Judkins, Haedrich, 2018; Klautau et al., 2020; Melo et al., 2020; Tab. 2). Here we report on 10 new records of stomiid fishes in the Brazilian EEZ: Aristostomias grimaldii Zugmayer, 1913, Astronesthes gudrunae Parin & Borodulina, 2002, Bathophilus nigerrimus Giglioli, 1882, Borostomias elucens (Brauer, 1906), Eustomias bibulbosus Parr, 1927, E. minimus Clarke, 1999, Grammatostomias ovatus Prokofiev, 2014, Melanostomias biseriatus Regan & Trewavas, 1930, Pachystomias microdon (Günther, 1878), and Photonectes achirus Regan & Trewavas, 1930. Occurrences of Eustomias braueri Zugmayer, 1911, E. schmidti Regan & Trewavas, 1930, Grammatostomias dentatus Goode & Bean, 1896, Leptostomias gladiator (Zugmayer, 1911), Melanostomias bartonbeani Parr, 1927 and P. goodyeari Kenaley & Hartel, 2005, are also confirmed for Brazilian waters. Records of Idiacanthus fasciola Peters, 1877, Aristostomias polydactylus Regan & Tewavas, 1930, Bathophilus altipinnis Beebe, 1933, and Eustomias metamelas Gomon & Gibbs, 1985 in the Brazilian EEZ (UF 190899, MNRJ 30734, MCZ 132776, and MNRJ 30807, respectively) require verification, but those species likely occur in the region given their wide distribution in the Atlantic (Gomon, Gibbs, 1985; McEachran, Fechhelm, 1998;Stewart, Kenaley, 2015a; Kenaley, Stewart, 2015g).

Astronesthinae

Members of the Astronesthinae can be recognized in the Stomiidae by an elongated body; mouth large; barbel present on chin; absence of scales; dorsal-fin origin well in advance of anal-fin origin; dorsal adipose fin present, except in the genus Rhadinesthes Regan & Trewavas, 1929; ventral adipose fin often present in front of anal fin; 9–21 dorsal-fin rays; 12–28 anal-fin rays (Stewart, Kenaley, 2015b; Nelson et al., 2016).

Astronesthes Richardson, 1845

Diagnosis. Astronesthes can be distinguished in the Astronesthinae by the dorsal-fin origin above or behind the pelvic-fin origin; anal-fin origin after the posterior end of the dorsal-fin base; 11–22 anal-fin rays; both dorsal and ventral adipose fins present; premaxillary teeth fang-like; maxillary teeth closely set and comb-like; ceratohyal teeth absent (Gibbs, 1964; Nakayama et al., 2021). A total of 48 valid species (Fricke et al., 2022).

Astronesthes atlantica Parin & Borodulina, 1996

(Fig. 8A)

FIGURE 8 | A. Astronesthes atlantica, NPM 4019, 32 mm SL; B. Astronesthes gemmifer, NPM 4799, 146 mm SL; C. Astronesthes gudrunae, NPM 4802, 111 mm SL; D. Astronesthes richardsoni, NPM 3194, 71 mm SL; E. Astronesthes similus, NPM 4811, 75 mm SL. Scale bars = 10 mm.

Diagnosis. Astronesthes atlantica differs from congeners by the low counts of widely spaced photophores in the IC (26–28) and OA (13–14; usually 13) series; AC 7–8; IP and PV in a straight line; last photophore of VAL behind anal-fin origin and last VAV; 14–17 dorsal-fin rays; 13–15 anal-fin rays; 7–8 pectoral-fin rays (Parin, Borodulina, 1996; Sutton et al., 2020).

Geographical distribution. Tropical and subtropical portions of the Atlantic Ocean, between 35ºN and 40ºS (Parin, Borodulina, 1996; Harold, 2003; Sutton et al., 2020). In the western Atlantic, the species is reported from the Gulf of Mexico, Caribbean Sea, and off French Guiana and Brazil (Parin, Borodulina, 1996). In Brazilian waters, the species was reported off Saint Peter and Saint Paul and Trindade and Martin Vaz archipelagos, with additional records outside the Brazilian EEZ in the western Atlantic (Parin, Borodulina, 1996; Judkins, Haedrich, 2018; Sutton et al., 2020). Astronesthes atlantica is reported here based on specimens collected off the Rocas Atoll, between depths of 90 and 525 m (Fig. 7B).

Specimens examined. 2 (31–32 mm SL): NPM 4019, 1 (32 mm), AB1#22; NPM 4824, 1 (31 mm), AB2#49B.

Astronesthes gemmifer Goode & Bean, 1896

(Fig. 8B)

Diagnosis. Astronesthes gemmifer differs from congeners by the presence of four fangs and two additional teeth on the premaxilla; serial photophores closely spaced; last 5–6 VAL and VAV above anal-fin base; IV forming an arch below the pectoral-fin base; IC 62–70; OA 42–48; posterior part of elongated terminal bulb of chin barbel black ventrally (Parin, Borodulina, 2000, 2003; Sutton et al., 2020).

Geographical distribution. Atlantic, Indian, and central Pacific oceans, between 62ºN and 40ºS (Gibbs, 1990a; Parin, Borodulina, 2003; Sutton et al., 2020). In the western Atlantic, the species was reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (McAllister, 1990; McEachran, Fechhelm, 1998; Parin, Borodulina, 2000; Harold, 2003; Menezes et al., 2003). In Brazil, A. gemmifer was reported off Saint Peter and Saint Paul Archipelago, with additional records outside the Brazilian EEZ off Rio Grande do Sul State (Parin, Borodulina, 2000; Judkins, Haedrich, 2018; Sutton et al., 2020). Astronesthes gemmifer is reported here based on one specimen collected off Fernando de Noronha Archipelago, at 430 m depth (Fig. 7B).

Specimen examined. 1: NPM 4799, 1 (146 mm), AB2#41A.

Astronesthes gudrunae Parin & Borodulina, 2002

(Fig. 8C)

Diagnosis. Astronesthes gudrunae differs from congeners by the AC with 1–2 elevated photophores; IV forming an arch below the pectoral-fin base; the presence of an aggregation of luminous tissue above the pectoral-fin, larger than eye and longitudinally oriented; three luminous spots on upper jaw, the first largest and anterior to orbit; IC 51–53; OA 32–33; PV 2 and 3 at the same level; chin barbel without a terminal bulb (Parin, Borodulina, 2002; Sutton et al., 2020).

Geographical distribution. Previous records ofAstronesthes gudrunae are mostly restricted tothe North Atlantic, south to 40º N, including the Caribbean Sea, with a single record in the eastern North Atlantic, at 22°1’N, 32°8’W (Parin, Borodulina, 2002, 2003; Judkins, Haedrich, 2018; Sutton et al., 2020). Astronesthes gudrunae is reported for the first time in the Brazilian EEZ based on one specimen collected off Rocas Atoll, at 610 m depth (Fig. 7B), which also represents the southernmost record of the species known so far (about 3º S).

Specimen examined. 1: NPM 4802, 1 (111 mm), AB2#53A.

Astronesthes richardsoni (Poey, 1852)

(Fig. 8D)

Diagnosis. Astronesthes richardsoni differs from congeners by the pelvic-fin origin well in advance of the dorsal-fin origin; anal-fin base approximately equal to dorsal-fin base; IV series in a linear row; IC 55–60; PV 15–17; OV 14–16; OA 33–37; 2–3 palatine teeth; postorbital luminous organ small; a ring of luminous spots on the cheek; chin barbel without terminal bulb (Parin, Borodulina, 2000, 2003; Sutton et al., 2020).

Geographical distribution. Atlantic Ocean, between 60ºN and 30ºS (Gibbs, 1990a; Parin, Borodulina, 2000; Afonso et al., 2018; Sutton et al., 2020). In the western Atlantic, the species was reported from off Canada to Brazil, including the Gulf of Mexico and the Caribbean Sea (Gibbs, 1990a; McEachran, Fechhelm, 1998; Parin, Borodulina, 2000; Harold, 2003; Coad, 2018b). In Brazil, A. richardsoni was reported from off Maranhão and Rio Grande do Norte States, and off Fernando de Noronha and Saint Peter and Saint Paul archipelagos. Additional records in the western South Atlantic are known outside the Brazilian EEZ (Parin et al., 1974; Gibbs, 1990a; Parin, Borodulina, 2000, 2003; Judkins, Haedrich, 2018; Sutton et al., 2020). Astronesthes richardsoni is reported here based on specimens collected off Rio Grande do Norte State, the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 25 and 800 m (Fig. 7B).

Specimens examined. 10 (22–132 mm SL): NPM 3194, 2 (49–71 mm), AB1#9; NPM 4800, 1 (132 mm), AB2#39; NPM 4801, 1 (122 mm), AB2#53A; NPM 4805, 1 (31 mm), AB2#41B; NPM 4812, 2 (91–111 mm), AB2#42A; NPM 4813, 1 (22 mm), AB2#39; NPM 4814, 1 (31 mm), AB2#40A.

Astronesthes similus Parr, 1927

(Fig. 8E)

Diagnosis. Astronesthes similus differs from congeners by the anal-fin base longer than the dorsal-fin base; 11–12 dorsal-fin rays; 6 pectoral-fin rays; BR 20–21; OA 39–41; IC 59–63; AC in a straight line; only two filaments on the terminal bulb of the chin barbel (Gibbs et al.,1984; Parin, Borodulina, 2003).

Geographical distribution. Western Atlantic, between 43ºN and 27ºS, including the Gulf of Mexico and the Caribbean Sea (Parr, 1927; Gibbs et al., 1984; McEachran, Fechhelm, 1998; Harold, 2003; Menezes et al., 2003; Moore et al., 2003; Ramírez et al., 2019). In Brazil, the species is reported off the northeastern region (Eduardo et al., 2020c), from off Bahia to Rio de Janeiro States (Braga et al., 2007), and in the Vitória-Trindade Chain (Davis Seamount; Pinheiro et al., 2015). Records outside the Brazilian EEZ in the vicinities of the Saint Peter and Saint Paul and Trindade and Martin Vaz archipelagos are also known (Gibbs et al., 1984; Judkins, Haedrich, 2018). Astronesthes similus is reported here based on specimens collected between depths of 90 and 1,113 m off Paraíba and Rio Grande do Norte States (Fig. 7B).

Specimens examined. 10 (36–75 mm SL): NPM 3423, 1 (40 mm), AB1#34; NPM 4811, 2 (53–75 mm), AB2#59A; NPM 4822, 7 (36–48 mm), AB2#59B.

Borostomias Brauer, 1906

Diagnosis. Borostomias can be distinguished in the Astronesthinae by the presence of 8 or more short and widely spaced maxillary teeth; gill teeth in groups of 2–4; ceratohyal teeth absent; dorsal-fin origin close to pelvic-fin origin; photophore series in continuous rows (Gibbs, 1964). Six valid species (Fricke et al., 2022).

Borostomias elucens (Brauer, 1906)

(Fig. 9A)

FIGURE 9 | A. Borostomias elucens, NPM 4299, 196 mm SL; B. Heterophotus ophistoma, NPM 4400, 223 mm SL. Scale bars = 10 mm.

Diagnosis. Borostomias elucens can be distinguished from congeners by the fang-like premaxillary teeth; 13–16 anal-fin rays; ventral adipose fin absent; chin barbel with spherical terminal bulb, without filaments; AC 12–14, forming and arch behind anal-fin base; VAV 14–15; VAL 18 or less (Gibbs, 1964; Sutton et al., 2020).

Geographical distribution. Cincumglobal in tropical and subtropical waters (Gibbs, 1964; Bekker et al., 1975; Yamamoto, 1982; Gibbs, 1990a; Aizawa, 2002; Shinohara et al., 2005; Liao et al., 2006; Tatsuta et al., 2014; Wang et al., 2019; Sutton et al., 2020). In the western Atlantic, the species is reported from the Gulf of Mexico, Caribbean Sea, off French Guiana, and off northeastern Brazil outside the EEZ (Parin et al., 1974; Regan, Trewavas, 1929; Fujii, 1983b; Harold, 2003; Olivar et al., 2017; Judkins, Haedrich, 2018). Borostomias elucens is therefore reported for the first time in the Brazilian EEZ based on specimens collected off Rio Grande do Norte State (including the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 650 and 1,113 m (Fig. 7C).

Specimens examined. 45 (46–299 mm SL): NPM 4299, 6 (96–196 mm), AB2#44A; NPM 4300, 8 (54–288 mm), AB2#42A; NPM 4302, 4 (91–299 mm), AB2#59A; NPM 4303, 3 (72–164 mm), AB2#49A; NPM 4391, 14 (46–284 mm), AB2#39; NPM 4399, 13 (47–278 mm), AB2#53A.

Heterophotus Regan & Trewavas, 1929

Diagnosis. Heterophotus is a monotypic genus that can be distinguished in the Astronesthinae by the postorbital organ as long as the eye; ventral photophores in irregular groups of 1–5; IP 1–11; PV 32–35; VAV 13–14; OV 33–36; VAL 16–20; AC 13–15; 31–35 closely spaced and erect (not slanting) maxillary teeth; gill arch teeth in groups of 2; ceratohyal teeth absent; 14–20 palatine teeth; 12–17 anal-fin rays; 11–13 dorsal-fin rays; ventral adipose fin absent; chin barbel with flattened white tip without terminal filaments; tip of snout not upturned (Gibbs, 1964; Sutton et al., 2020).

Heterophotus ophistoma Regan & Trewavas, 1929

(Fig. 9B)

Diagnosis. Same as for the genus.

Geographical distribution. Tropical and subtropical waters of the Atlantic (Gibbs, 1964, 1990a; Harold, 2003; Czudaj et al., 2020; Sutton et al., 2020), Indian (Hutchins, 2001, Shcherbachev et al., 1986), and central and western Pacific oceans (Clarke, 1974; Aizawa, 2002; Mundy, 2005; Liao et al., 2006; Shinohara et al., 2009; Fricke et al., 2011). In the western Atlantic, the species is known from off United States to northeastern Brazil, including the Gulf of Mexico and the Caribbean Sea (Regan, Trewavas, 1929; Bekker et al., 1975; Fujii, 1983b; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Judkins, Haedrich, 2018). In Brazil the species was previously reported from off Maranhão State, and the Fernando de Noronha and Saint Peter and Saint Paul archipelagos (Judkins, Haedrich, 2018; Klautau et al., 2020; Marceniuk et al., 2021). Heterophotus ophistoma is reported here based on specimens collected between depths of 430 and 1,030 m off Rio Grande do Norte State (including the seamounts), and the Fernando de Noronha Archipelago (Fig. 7C).

Specimens examined. 8 (96–253 mm SL): NPM 4304, 1 (211 mm), AB2#39; NPM 4326, 1 (207 mm), AB2#41A; NPM 4398, 2 (219–253 mm), AB2#50A; NPM 4400, 2 (218–223 mm), AB2#54B; NPM 4401, 1 (216 mm), AB2#44A.

Chauliodontinae

Members of the Chauliodontinae can be recognized in the Stomiidae by the dorsal-fin origin well in advance of the pelvic-fin origin, shortly behind head; very long fang-like teeth on both jaws extending well beyond opposite jaw when mouth is closed; dorsal and ventral adipose fin present, dorsal adipose fin large, angular, distinctly placed posteriorly on body; body long, slender, compressed, marked with scale-like hexagonal patterns, each hexagonal cell containing 1 or 2 photophores; 5–7 dorsal-fin rays, the first one elongated, second one with a red photophore on tip; anal fin short, 10–13 rays, below dorsal adipose fin; chin barbel short to absent (Morrow, 1964b; Nelson et al., 2016; Stewart, 2015). A single genus (Fricke et al., 2022).

Chauliodus Bloch & Schneider, 1801

Diagnosis. Same as for the subfamily. Nine valid species (Fricke et al., 2022).

Chauliodus sloani Bloch & Schneider, 1801

(Fig. 10)

FIGURE 10 | Chauliodus sloani, NPM 3287, 167 mm SL. Scale bar = 10 mm.

Diagnosis. Chauliodus sloani differs from congeners by the presence of an approximately round postorbital organ; dorsal-fin origin over 4–10 OV; fourth premaxillary tooth longer than the third; predorsal length 17–28% SL; PV 17–23; VAV 22–28; AC 9–13; IC 62–69; OV 17–22; VAL 22–28; OA 41–48 (Morrow, 1964b; Parin, Novikova, 1974; Gibbs, 1986a; Stewart, 2015; Sutton et al., 2020).

Geographical distribution. Circumglobal, between 65ºN and 45ºS, including the Mediterranean Sea (Parin, Novikova, 1974; Gibbs, 1986a; Bilecenoglu et al.,2002; Vélez, 2009; Møller et al., 2010; Lipej, Dulčić, 2010; Duhamel et al., 2014; Stewart, 2015; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species is reported from off Canada to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Morrow, 1964b; Parin, Novikova, 1974; McAllister, 1990; McEachran, Fechhelm, 1998; Nión et al., 2016; Mecklenburg et al., 2018; Coad, 2018b; Ramírez et al., 2019). In Brazil, C. sloani was previously recorded off Amapá, Pará, Maranhão, Rio Grande do Norte, Bahia, Espírito Santo, Rio de Janeiro, and Rio Grande do Sul States, in addition to the Saint Peter and Saint Paul Archipelago (Parin, Novikova, 1974; Figueiredo et al., 2002; Menezes et al., 2003; Bernardes et al., 2005; Braga et al., 2007; Haimovici et al., 2008; Lins Oliveira et al., 2015; Judkins, Haedrich, 2018; Eduardo et al., 2020b). Larvae were recorded off Rio de Janeiro State (Bonecker, Castro, 2006; Bonecker et al., 2014). Chauliodus sloani is reported here based on specimens collected off Rio Grande do Norte, Paraíba, Pernambuco, the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 90 and 1,030 m (Fig. 7D).

Remarks. In addition to the 57 specimens reported here, 247 specimens of C. sloani collected in the ABRACOS expeditions were dissected in a study addressing the trophic ecology, vertical migration, and functional role of the species (Eduardo et al., 2020b).

Specimens examined. 57 (55–241 mm SL): NPM 3286, 1 (184 mm), AB1#22; NPM 3287, 1 (167 mm), AB1#14; NPM 4386, 11 (154–238 mm), AB2#41A; NPM 4389, 5 (84–153 mm), AB2#44A; NPM 4396, 2 (140–146 mm), AB2#21; NPM 4397, 2 (96–114 mm), AB2#39; NPM 4420, 5 (96–196 mm), AB2#16; NPM 4421, 9 (131–210 mm), AB2#35; NPM 4422, 15 (83–241 mm), AB2#42A; NPM 4447, 2 (90–219 mm), AB2#54B; NPM 4448, 3 (144–184 mm), AB2#49A; NPM 4921, 1 (55 mm), AB2#40A.

Stomiinae

Members of the Stomiinae can be recognized in the Stomiidae by an elongated to very elongated body; snout short; both the dorsal- and anal-fins about opposite to each other, located far behind the pelvic-fin; a long chin barbel, ending in a terminal swelling with either small filaments or appendages; 5–6 rows of closely spaced, scale-like, hexagonal cells; photophores present on each hexagonal cell; adipose fins absent (Gibbs, 1969; Harold, 2003; Nelson et al., 2016; Kenaley, Stewart, 2015f). A single genus (Fricke et al., 2022).

Stomias Cuvier, 1816

Diagnosis. Same as for the subfamily. A total of 11 valid species (Fricke et al., 2022).

Stomias danae Ege, 1933

(Fig. 11A)

FIGURE 11 | A. Stomias danae, NPM 4768, 95 mm SL; B. Stomias longibarbatus, NPM 4598, 173 mm SL. Scale bars = 10 mm.

Diagnosis. Stomias danae differs from congeners by the chin barbel stem with 2 filaments at base; terminal bulb of chin barbel without filaments; palatine with 2 teeth; 16–18 dorsal-fin rays; 20–21 anal-fin rays; six rows of hexagonal cells dorsal to lateral series of photophores; one photophore per hexagon cell in the upper two rows, usually 2–3 in the third row, and usually 3–5 in the fourth and fifth rows; total vertebrae 60–62 (Morrow, 1964d; Gibbs, 1969; Shcherbachev, Novikova, 1976).

Geographical distribution. Known from a few records in the western South Atlantic, western Indian and central Pacific oceans (Gibbs, 1969; Clarke, 1974; Shcherbachev, Novikova, 1976; Mundy, 2005). In the western South Atlantic, the species is known from one record off Trindade and Martin Vaz Archipelago (Brazil), and two records off northeastern Brazil, outside the country’s EEZ (Shcherbachev, Novikova, 1976; Gibbs, 1969; Menezes et al.,2003; Judkins, Haedrich, 2018). Stomias danae is reported here based on one specimen collected off Paraíba State, at 800 m depth (Fig. 7E).

Specimen examined. 1. NPM 4768, 1 (95 mm), AB2#21.

Stomias longibarbatus (Brauer, 1902)

(Fig. 11B)

Diagnosis. Stomias longibarbatus differs from congeners by an extremely elongated chin barbel, 6–11 times longer than head length and up to 75% SL; body length 20-35 times body depth; IC>170; OA>135; VAV 58–73; total vertebrae 164–172 (Morrow, 1964d; Shcherbachev, Novikova, 1976; Kenaley, Stewart, 2015f; Sutton et al., 2020).

Geographical distribution. Circumglobal, between 43ºN and 38ºS (Shcherbachev, Novikova, 1976; Gibbs, 1986b, 1990b; Mundy, 2005; Kenaley, Stewart, 2015f; Cherel et al., 2020; Sutton et al., 2020). In the western Atlantic, the species is reported from off United States, the Gulf of Mexico, Brazil, and Uruguay (McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Nión et al., 2016; Judkins, Haedrich, 2018). In Brazil, S. longibarbatus was reported from off the Fernando de Noronha Archipelago (Judkins, Haedrich, 2018). It is reported here based on specimens collected off Rio Grande do Norte State (near the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 260 and 1,020 m (Fig. 7E).

Specimens examined. 5 (173–390 mm SL): NPM 4390, 1 (390 mm), AB2#49A; NPM 4598, 1 (173 mm), AB2#56B; NPM 4599, 1 (272 mm), AB2#50A; NPM 4600, 1 (304 mm), AB2#58A; NPM 4601, 1 (267 mm), AB2#52B.

Malacosteinae

Members of the Malacosteinae can be recognized in the Stomiidae by the body moderately elongate; scales absent; snout short; mouth large to extremely large, jaws longer than skull; skin between mandibles absent, except in Pachystomias Günther, 1887; dorsal-fin origin over anal-fin origin and located far behind pelvic-fin origin; adipose fins absent; chin barbel present in most species, simple, without a complex distal tip; pectoral fin present or absent; 14–28 dorsal-fin rays; 17–32 anal-fin rays; caudal fin small, forked; one or more accessory orbital photophores anterior to orbit that often produces red light. Three genera (Nelson et al., 2016; Kenaley, Stewart, 2015g; Fricke et al., 2022).

Aristostomias Zugmayer, 1913

Diagnosis. Aristostomias can be distinguished in the Malacosteinae by the chin barbel present; pectoral-fin present; snout longer than eye diameter; 2 pairs of nostrils on either side of head; two rows of conspicuous photophores (lateral and ventral series) separated in groups (Morrow, 1964c; Goodyear, Gibbs, 1986; McEachran, Fechhelm, 1998; Kenaley, Stewart, 2015g; Sutton et al., 2020). Six valid species (Fricke et al., 2022).

Aristostomias grimaldii Zugmayer, 1913

(Fig. 12A)

FIGURE 12 | A. Aristostomias grimaldii, NPM 4659, 79 mm SL; B. Aristostomias tittmanni, NPM 4662, 76 mm SL; C. Malacosteus niger, NPM 4331, 124 mm SL; D. Pachystomias microdon, NPM 4592, 150 mm SL; E. Photostomias atrox, NPM 4575, 67 mm SL; F. Photostomias goodyeari, NPM 4590, 64 mm SL. Scale bars = 10 mm.

Diagnosis. Aristostomias grimaldii differs from congeners by the presence of 7–10 pectoral-fin rays; 14–16 PV photophores in close-set groups (5 groups of 2–4 photophores each); luminous dots between eye and postorbital organ; barbel ending in a thickened terminal region with a poorly defined bulb (Morrow, 1964c; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic, eastern Indian, and central Pacific oceans, between 43ºN and 35ºS (Porteiro et al., 1999; Moore et al., 2003; Mundy, 2005; Goodyear, 1990; Nión et al., 2016; Sutton et al., 2020). In the western Atlantic, the species was reported from off the United States, Suriname, French Guiana, and Uruguay, including the Gulf of Mexico and Caribbean Sea (Bekker et al., 1975; Fujii, 1983c; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Nión et al., 2016; Judkins, Haedrich, 2018). Aristostomias grimaldii is reported for the first time in the Brazilian EEZ based on five specimens collected between depths of 260 and 850 m off Rio Grande do Norte State, near the seamounts, and the Fernando de Noronha Archipelago (Fig. 7F).

Specimens examined. 5 (65–86 mm SL): NPM 4622, 4 (65–86 mm), AB2#44A; NPM 4659, 1 (79 mm), AB2#56C

Aristostomias tittmanni Welsh, 1923

(Fig. 12B)

Diagnosis. Aristostomias tittmanni differs from congeners by the presence of 6–7 pectoral-fin rays; 15–19 PV photophores in 5–6 close-set groups of 2–4 photophores each; luminous dots between eye and postorbital organ; barbel with a well-defined bulb (Morrow, 1964c; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic (Morrow, 1964c; McAllister, 1990; Porteiro et al., 1999; Harold, 2003; Béarez et al., 2017; Carneiro et al., 2019; Sutton et al., 2020) and Pacific oceans (Pequeño, 1989; Mundy, 2005). In the western Atlantic, the species is reported from off Canada, United States, French Guiana, and northeastern Brazil (off Maranhão State) (McEachran, Fechhelm, 1998; McAllister, 1990; Judkins, Haedrich, 2018; Ramírez et al., 2019). Aristostomias tittmanni is reported here based on four specimens collected off Paraíba State, the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 50 and 800 m (Fig. 7F).

Specimens examined. 4 (32–76 mm SL): NPM 3278, 1 (39 mm), AB1#12; NPM 4595, 1 (32 mm), AB2#45B; NPM 4596, 1 (37 mm), AB2#21; NPM 4662, 1 (76 mm), AB2#42A.

Malacosteus Ayres, 1848

Diagnosis. Malacosteus can be distinguished in the Malacosteinae by the presence of the pectoral fin; a single nostril on either side of head; snout shorter than eye diameter; chin barbel absent; serial photophores reduced in size and arranged in groups; a small postorbital organ behind eye; suborbital organ large and comma-shaped (Morrow, 1964c; Kenaley, 2007; Kenaley, Stewart, 2015g; Sutton et al., 2020). Two valid species (Fricke et al., 2022).

Malacosteus niger Ayres, 1848

(Fig. 12C)

Diagnosis. Malacosteus niger can be distinguished from M. australis Kenaley, 2007 by IP 3–5 in 3–5 groups (vs. IP 4–8 in 3–7 groups); PO 5.8–11.9% upper jaw length in males (vs. PO 3.7–6.4% upper jaw length in males); PO 2.8–8.1% upper jaw length in females (vs. PO 2.5–4.4% upper jaw length in females) (Kenaley, 2007; Kenaley, Stewart, 2015g).

Geographical distribution. Circumglobal, between 66ºN and 40ºS (Goodyear, 1990; Harold, 2003; Kenaley, 2007; Kenaley, Stewart, 2015g; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species is reported from off Greenland to Brazil, including the Gulf of Mexico and the Caribbean Sea (Parin et al., 1974; Bekker et al., 1975; Fujii, 1983c; McAllister, 1990; McEachran, Fechhelm, 1998; Kenaley, 2007; Coad, 2018b; Afonso et al.,2018; Judkins, Haedrich, 2018). In Brazil the species was previously reported from off the northeastern region and from Bahia to Rio de Janeiro States, in addition to the Saint Peter and Saint Paul Archipelago (Braga et al., 2007; Costa, Mincarone, 2010; Olivar et al., 2017; Eduardo et al., 2018a; Judkins, Haedrich, 2018). Records outside the Brazilian EEZ are also known off the Fernando de Noronha Archipelago (Kenaley, 2007). Malacosteus niger is reported here based on 43 specimens collected off Rio Grande do Norte State (including the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago, between depths of 550 and 1,113 m (Fig. 7F).

Specimens examined. 43 (63–181 mm SL): NPM 4328, 10 (76–181 mm), AB2#44A; NPM 4331, 2 (106–124 mm), AB2#49A; NPM 4332, 1 (102 mm), AB2#42A; NPM 4333, 15 (71–111 mm), AB2#53A; NPM 4336, 2 (128–134 mm), AB2#54B; NPM 4337, 9 (65–181 mm), AB2#39; NPM 4338, 3 (63–125 mm), AB2#59A; NPM 4442, 1 (67 mm), AB2#52A.

Pachystomias Günther, 1887

Diagnosis. Pachystomias can be distinguished in the Malacosteinae by the very large and crescent-shaped suborbital organ, red or orange in fresh specimens; body depth about 13.8-20.0% SL; two nostrils on either side of head; a small postorbital organ below eye; mouth floor, i.e., the membrane between the mandibular symphysis and the hyoid, present; chin barbel without terminal bulb, its length shorter than head length in adults but about equal to 50% SL in juveniles; teeth uniserial in jaws, curved; 5–6 pectoral-fin rays, reduced in length; dorsal- and anal-fin bases approximately equal in length, anal-fin origin just posterior to dorsal-fin origin; photophores in both ventral and lateral series with long spaces separating them into clearly distinct groups (Morrow, Gibbs, 1964; Gibbs, 1986c; McEachran, Fechhelm, 1998; Kenaley, Stewart, 2015g; Sutton et al., 2020). A single valid species (Fricke et al., 2022).

Pachystomias microdon (Günther, 1878)

(Fig. 12D)

Diagnosis. Same as for the genus.

Geographical distribution. Circumglobal, from 58ºN to 40ºS (Gibbs, Barnett, 1990; Harold, 2003; Stevenson et al., 2009; Kenaley, Stewart, 2015g; Orlov, Tokranov, 2019; Sutton et al., 2020; Love et al., 2021). In the western Atlantic, the species was reported from off Canada to the Caribbean Sea (Morrow, Gibbs, 1964; McAllister, 1990; McEachran, Fechhelm, 1998; Moore et al., 2003). Pachystomias microdon is reported for the first time in the western South Atlantic based on nine specimens collected off Rio Grande do Norte State (including the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago in the Brazilian EEZ, between depths of 610 and 1,030 m (Fig. 7F).

Specimens examined. 9 (39–181 mm SL): NPM 4587, 1 (168 mm), AB2#39; NPM 4588, 2 (173–176 mm), AB2#54B; NPM 4589, 1 (181 mm), AB2#44A; NPM 4591, 1 (85 mm), AB2#49A; NPM 4592, 1 (150 mm), AB2#52A; NPM 4594, 2 (39–109 mm), AB2#53A; NPM 4625, 1 (155 mm), AB2#41A.

Photostomias Collett, 1889

Diagnosis. Photostomias can be distinguished in the Malacosteinae by the pectoral fin present only in larvae and transformed juveniles up to about 27 mm SL, absent in large juveniles and adults; pelvic-fin origin well before midlength of body, greatly elongate, 36.8–52.3% SL; 6 pelvic-fin rays, proximal ray flattened and separate from remaining five elongate rays; halves of each pelvic-fin ray fused together along their anterior margins for most of their length; pelvic fins attached near base to pelvic girdle by a thin membrane; chin barbel absent; one row of photophores on each side of isthmus (Morrow, 1964c; Fink, 1985; Kenaley, Hartel, 2005; Sutton et al., 2020). Seven valid species (Fricke et al., 2022).

Photostomias atrox (Alcock, 1890)

(Fig. 12E)

Diagnosis. Photostomias atrox differs from congeners by the presence of three pairs of basibranchial tooth patches; 22–26 dorsal-fin rays; 26–30 anal-fin rays; sum of dorsal- and anal-fin rays 47–55, usually 50-54; male PO moderate to large in specimens greater than 60 mm, 72.1–197.2% fleshy orbit, 16.7–46.6% jaw length; AO large in specimens greater than 60 mm, 9.2–37.8% fleshy orbit, 2.1–9.6% jaw length (Kenaley, Hartel, 2005; Sutton et al., 2020).

Geographical distribution. Circumglobal in tropical and subtropical waters, except the eastern Pacific Ocean (Kenaley, Hartel, 2005; Kenaley, 2009; Sutton et al., 2020). In the western Atlantic, the species is reported from off the United States to Brazil (Kaneley, Hartel, 2005; Hartel et al., 2008). In Brazil the species was recorded off the Trindade and Martin Vaz Archipelago, with additional records outside the EEZ off the north portion of the country and the Saint Peter and Saint Paul Archipelago (Kenaley, Hartel, 2005; Sutton et al., 2020). Photostomias atrox is reported here based on one specimen collected off Pernambuco, at 680 m depth (Fig. 7F).

Specimen examined. 1: NPM 4575, 1 (67 mm), AB2#16.

Photostomias goodyeari Kenaley & Hartel, 2005

(Fig. 12F)

Diagnosis. Photostomias goodyeari differs from congeners by the presence of three pairs of basibranchial tooth patches; 26–31 dorsal-fin rays, 30–33 anal-fin rays, sum of dorsal- and anal-fin rays 56–63, usually 57–61; male PO small to moderate in specimens larger than 60 mm, 54.5–103.2% fleshy orbit, 9.6–24.8% jaw length; male AO moderate in specimens larger than 60 mm, 11.2–25.1% fleshy orbit, 2.3–4.4% jaw length (Kenaley, Hartel, 2005; Sutton et al., 2020).

Geographical distribution. Atlantic Ocean, between 40ºN and 5ºS (Kenaley, Hartel, 2005; Hartel et al., 2008; Sutton et al., 2020). In the western Atlantic, the species was reported from off the United States, the Gulf of Mexico, the Caribbean Sea, Suriname, and northern Brazil, outside the EEZ (Kenaley, Hartel, 2005; Melo et al., 2020). Photostomias goodyeari was included in a list of Brazilian deep-sea fishes by Melo et al. (2020), without reference to voucher specimens. The occurrence of the species is therefore confirmed in the Brazilian EEZ based on one specimen collected off Rio Grande do Norte State, between depths of 600 and 860 m (Fig. 7F).

Specimen examined. 1: NPM 4590, 1 (64 mm), AB2#39.

Melanostomiinae

Members of the Melanostomiinae can be recognized in the Stomiidae by the body relatively elongate; snout short; mouth large to extremely large; chin barbel usually present, ranging from very short to longer than body, barbel simple to complex; skin between lower jaws present in all species; dorsal-fin origin usually opposite to anal-fin origin (except in Eustomias and Flagellostomias), both fins located far behind the pelvic fin; dorsal adipose fin absent, except in Chirostomias Regan & Trewavas, 1930; scales absent, body without hexagonal patterns; suborbital and postorbital light organs usually present, preorbital luminous organ absent; photophores in 2 continuous rows along flanks and ventral portion of the body (Gibbs, 1986c; Kenaley, Stewart, 2015h; Nelson et al., 2016). Sixteen genera (Fricke et al., 2022).

Bathophilus Giglioli, 1882

Diagnosis. Bathophilus can be distinguished in the Melanostomiinae by the absence of teeth on vomer; premaxilla with fixed teeth followed by a series of depressible teeth; suborbital organs reduced or absent; base of pelvic fin high on body, about near midlateral portion of body; dorsal- and anal-fin bases similar in length; presence of small photophores in the entire body (Morrow, Gibbs, 1964; Barnett, Gibbs Jr., 1968; Gibbs, 1986c; Kenaley, Stewart, 2015h; Sutton et al., 2020). A total of 19 valid species (Fricke et al., 2022).

Bathophilus nigerrimus Giglioli, 1882

(Fig. 13A)

FIGURE 13 | A. Bathophilus nigerrimus, NPM 4593, 84 mm SL; B. Bathophilus pawneei, NPM 4572, 124 mm SL. Scale bars = 10 mm.

Diagnosis. Bathophilus nigerrimus differs from congeners by the pelvic-fin origin below the body midline; 16–26 free pelvic-fin rays; a single group of 31–57 pectoral-fin rays; 13–15 dorsal- and anal-fin rays; about 22–24 photophores in lateral series (OV 12–13, VAL 9–12); about 33-35 photophores in ventral series (IP 4–5, PV 12–13, VAV 11–13, AC 3–6) (Morrow, Gibbs, 1964; Barnett, Gibbs Jr., 1968; Gibbs, 1986c; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic Ocean and the Mediterranean Sea (Gibbs, Barnett, 1990; Santos et al.,1997; Harold, 2003; Judkins, Haedrich, 2018; Sutton et al., 2020; Tsagarakis et al., 2021), western Pacific (Aizawa, 2002) and Indian oceans (Hutchins, 2001; Fricke et al., 2018). In the western Atlantic, the species is reported from off United States to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Morrow, Gibbs, 1964; Parin et al., 1974; McEachran, Fechhelm, 1998; Harold, 2003; Nión et al., 2016). Batophilus nigerrimus is reported here for the first time in the Brazilian EEZ based on two specimens collected off Rocas Atoll, between depths of 90 and 610 m (Fig. 14A). The likely occurrence of the species in Brazilian waters was previously mentioned by Bonecker, Castro (2006: 90) based on its wide distribution in the western Atlantic.

FIGURE 14 | Records of species of the Melanostomiinae (AF) collected during the ABRACOS expeditions off northeastern Brazil. Oceanic islands: RA, Rocas Atoll; FN, Fernando de Noronha Archipelago.

Specimens examined. 2 (84–95 mm SL): NPM 4593, 1 (84 mm), AB2#53A; NPM 4628, 1 (95 mm), AB2#49B.

Bathophilus pawneei Parr, 1927

(Fig. 13B)

Diagnosis. Bathophilus pawneei differs from congeners by the pelvic-fin origin close to, but slightly below, the body midline; 12–15 pelvic-fin rays; 2 pectoral-fin rays (rarely 1) in a single group; 14–17 dorsal-fin rays; 15–18 anal-fin rays; usually a small luminous spot behind the pelvic fin, and sometimes another luminous spot on the shoulder region; about 24–29 photophores in the lateral series (OV 11–14, VAL 13–15); about 36–42 photophores in the ventral series (IP 5, PV 12–15, VAV 13–15, AC 6–7) (Morrow, Gibbs, 1964; Barnett, Gibbs Jr., 1968; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic, Indian, and Pacific oceans, from 45ºN to 27ºS (Morrow, Gibbs, 1964; Clarke, 1974; Gibbs, Barnett, 1990; Aizawa, 2002; Shinohara et al., 2009; Sutton et al., 2020). In the western Atlantic, the species is reported from off the United States to Brazil, including the Gulf of Mexico and the Caribbean Sea (Parr, 1927; Morrow, Gibbs, 1964; Parin et al., 1974; Gibbs, Barnett, 1990; McEachran, Fechhelm, 1998; Moore et al., 2003; Judkins, Haedrich, 2018). In Brazil the species was reported from off Fernando de Noronha and Saint Peter and Saint Paul archipelagos (Judkins, Haedrich, 2018; Sutton et al., 2020), with additional records outside the Brazilian EEZ off the Rio de Janeiro State (Gibbs, Barnett, 1990). Bathophilus pawneei is reported here based on specimens collected off the Rocas Atoll and Fernando de Noronha Archipelago, between depths of 430 and 610 m (Fig. 14A).

Specimens examined. 4 (30–124 mm SL): NPM 4571, 1 (81 mm), AB2#41A; NPM 4572, 1 (124 mm), AB2#40A; NPM 4928, 2 (30–32 mm), AB2#53B.

Eustomias Vaillant, 1884

Diagnosis. Eustomias can be distinguished in the Melanostomiinae by the trunk slender, mouth large, snout tapering and longer than eye, upper jaw protrusible; three pairs of basibranchial teeth; no teeth on vomer or palatine; anal-fin origin well in advance of the dorsal-fin origin; anal-fin base distinctly longer than the dorsal-fin base; 0–13 pectoral-fin rays, lacking free rays; notochord forming a U-shaped or S-shaped bend behind head, with anterior 6 or 7 vertebrae incomplete; chin barbel with great anatomical variation (Morrow, Gibbs, 1964; Gibbs, 1986c; Kenaley, Stewart, 2015h; Sutton et al., 2020). A total of 128 valid species in 11 subgenera (Fricke et al., 2022).

Eustomias bibulbosus Parr, 1927

(Fig. 15A)

FIGURE 15 | A. Eustomias bibulbosus, NPM 4642, 87 mm SL; B. Eustomias braueri, NPM 4816, 56 mm SL; C. Eustomias brevibarbatus, NPM 4646, 95 mm SL; D. Eustomias enbarbatus, NPM 4819, 60 mm SL; E. Eustomias minimus, NPM 4881, 69 mm SL; F. Eustomias schmidti, NPM 4882, 68 mm SL. Scale bars = 10 mm.

Diagnosis. Eustomias bibulbosus differs from congeners by the presence of 3 well developed pectoral-fin rays; long chin barbel (60–85% SL), with two terminal bulbs separated by a long interspace; distal bulb usually 0.9–1.6% SL; terminal filament long (22–33% SL), rarely with 1 or 2 branches; chin barbel stem darkly pigmented; lower jaw about as long as upper jaw and not upturned at tip; deep groove on belly absent; preorbital and suborbital organs absent (Gibbs et al., 1983; McEachran, Fechhelm, 1998).

Geographical distribution. Previous records are restricted to the western North Atlantic, between 40ºN and 13ºN (Vinnichenko, 1997; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003), with records off United States, Bermudas and Suriname (Parr, 1927; Bekker et al., 1975; Gibbs et al., 1983; Judkins, Haedrich, 2018). Eustomias bibulbosus is therefore reported for the first time in the South Atlantic based on one specimen collected off Pernambuco State in the Brazilian EEZ, at 680 m depth (Fig. 14B).

Specimen examined. 1: NPM 4642, 1 (87 mm), AB2#16.

Eustomias braueri Zugmayer, 1911

(Fig. 15B)

Diagnosis. Eustomias braueri differs from congeners by the presence of 9–16 pectoral-fin rays; 8 pelvic-fin rays; chin barbel length approximately equal to head length, 12–17% SL; one ovoid terminal bulb at the chin barbel with two anterior filaments; chin barbel stem darkly pigmented; a series of luminous spots on lower side of body that resembles large photophores; lower jaw about as long as upper jaw and not upturned at tip; wide ventral groove extending from isthmus to behind pectoral-fin base present; preorbital and suborbital organs absent (Morrow, Gibbs, 1964; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic and western North Pacific oceans, between 40ºN and 30ºS (Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1974; Bekker et al., 1975; Vinnichenko, 1997; McEachran, Fechhelm, 1998; Tatsuta et al., 2014; Sutton et al., 2020). Menezes et al. (2003) included the species in a list of Brazilian marine fishes based on Morrow, Gibbs (1964), but that record is outside the country’s EEZ, off the northeastern region. Melo et al. (2020: 208), in turn, regarded the species as “not confirmed” in Brazil. Therefore, occurrence of E. braueri is confirmed in the Brazilian EEZ based on one specimen collected off Pernambuco State, at 680 m depth (Fig. 14B).

Specimen examined. 1: NPM 4816, 1 (56 mm), AB2#16.

Eustomias brevibarbatus Parr, 1927

(Fig. 15C)

Diagnosis. Eustomias brevibarbatus differs from congeners by the presence of two long and free pectoral-fin rays; barbel length 13–33% SL; chin barbel usually with two terminal bulbs separated by a short interspace (0.1–1.5% SL); anterior portion of distal bulb with distinctive black pigment; short terminal filament (0.3–2.5% SL) bifurcated or trifurcated near its base; barbel stem lightly to moderately pigmented; deep ventral groove on belly absent; preorbital and suborbital organs absent (Gomon, Gibbs, 1985; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Western Atlantic, between 35ºN and 25ºS, reported from off the United States to southeastern Brazil, including the Gulf of Mexico and the Caribbean Sea (Morrow, Gibbs, 1964; Bekker et al., 1975; Gomon, Gibbs, 1985; McEachran, Fechhelm, 1998; Harold, 2003). In Brazil the species was reported from off Fernando de Noronha Archipelago and Rio de Janeiro State, with additional records in the western South Atlantic outside the country’s EEZ (Gomon, Gibbs, 1985; Menezes et al., 2003; Costa, Mincarone, 2010; Judkins, Haedrich, 2018; Sutton et al., 2020). Eustomias brevibarbatus is reported here based on specimens collected off Rio Grande do Norte State (near the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago between depths of 90 and 1,030 m (Fig. 14B).

Specimens examined. 6 (85–128 mm SL): NPM 4643, 1 (87 mm), AB2#49A; NPM 4644, 1 (85 mm), AB2#41A; NPM 4645, 1 (128 mm), AB2#50A; NPM 4646, 1 (95 mm), AB2#49B; NPM 4689, 1 (85 mm), AB2#59A; NPM 5013, 1 (102 mm), AB2#54B.

Eustomias enbarbatus Welsh, 1923

(Fig. 15D)

Diagnosis. Eustomias enbarbatus differs from congeners by the presence of 3–4 (usually 3) well developed pectoral-fin rays; chin barbel 27.5–34.9% SL, with one terminal bulb; one filament before the terminal bulb, which ends in a branched stem or in a group of short filaments; deep groove on belly absent (Morrow, Gibbs, 1964; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. Atlantic, Indian, and Pacific oceans (Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1974; Gibbs, Barnett, 1990; Aizawa, 2002; Harold, 2003; Mundy, 2005; Kenaley, Stewart, 2015h; Carneiro et al., 2019; Sutton et al., 2020). In the western Atlantic, the species is reported from off the United States to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Parin et al., 1974; Morrow, Gibbs, 1964; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Nión et al., 2016). The species was recorded in Brazilian waters off Saint Peter and Saint Paul and Trindade and Martin Vaz archipelagos (Judkins, Haedrich, 2018; Sutton et al., 2020). Eustomias enbarbatus is reported here based on specimens collected between depths of 680 and 780 m off the Pernambuco State and the Fernando de Noronha Archipelago (Fig. 14C).

Specimens examined. 3 (54–60 mm SL): NPM 4818, 1 (54 mm), AB2#16; NPM 4819, 2 (55–60 mm), AB2#42A.

Eustomias minimus Clarke, 1999

(Fig. 15E)

Diagnosis. Eustomias minimus differs from congeners by the presence of 2 pectoral-fin rays; chin barbel with a single, simple and thin branch arising from the stem, branch length barely half that of distal barbel; terminal filaments of the chin barbel simple and short, not or barely reaching beyond bulb tip (Clarke, 1999).

Geographical distribution. Previously known from four poorly preserved specimens, including the holotype, collected in the central and western equatorial Pacific (Clarke, 1999). Eustomias minimus is reported for the first time in the Atlantic Ocean based on one specimen collected off the Fernando de Noronha Archipelago in the Brazilian EEZ, at 780 m depth (Fig. 14C).

Remarks. Morphometric and meristic data of the specimen examined are within the ranges reported for the species (Clarke, 1999), except for the barbel length, which is slightly shorter (7.5% SL) than in the original description (11–12.7% SL).

Specimen examined. 1: NPM 4881, 1 (69 mm), AB2#42A.

Eustomias schmidti Regan & Trewavas, 1930

(Fig. 15F)

Diagnosis. Eustomias schmidti differs from congeners by the presence of two pectoral-fin rays bound together in black tissue; chin barbel short, about 7–17% of SL; a distinct chin barbel anatomy, with three branches arising from the stem: the medial branch is stout, bulbous and with distal filaments, barely reaching to barbel tip, the lateral branches taper distally, extending well beyond end of barbel; terminal bulb of chin barbel unpigmented and long, occupying most of distal barbel, tightly constricted distally into a spheroidal terminal section, and with a terminal filament or single-based group of filaments arising ventrally from this constriction; deep ventral groove on belly relatively short, extending to PV 4–6; anal-fin base approximately twice as long as dorsal-fin base; postorbital photophore and barbel photophore probably glows green or blue (Morrow, Gibbs, 1964; Gibbs, 1986c; McEachran, Fechhelm, 1998; Clarke, 2000; Kenaley, Stewart, 2015h; Sutton et al., 2020).

Geographical distribution. Atlantic and Pacific oceans, between 40ºN and 40ºS (Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1974; Bekker et al., 1975; Gibbs, 1986c; Gibbs, Barnett, 1990; Vinnichenko, 1997; Aizawa, 2002; Harold, 2003; Moore et al., 2003; Mundy, 2005; Kenaley, Stewart, 2015h; Carneiro et al., 2019; Sutton et al., 2020). In the western Atlantic the species is reported from off the United States to the Caribbean Sea, with a single record off southeastern Brazil, outside the country’s EEZ (Clarke, 2000; Menezes et al., 2003; Judkins, Haedrich, 2018). Melo et al. (2020) also included E. schmidti in a list of deep-sea fishes of Brazil, without providing information on voucher specimens or locality data. Therefore, the occurrence of E. schmidti is confirmed in Brazilian waters based on one specimen collected off the Fernando de Noronha Archipelago, at 850 m depth (Fig. 14C).

Specimen examined. 1: NPM 4882, 1 (68 mm), AB2#42A.

Grammatostomias Goode & Bean, 1896

Diagnosis. Grammatostomias can be distinguished in the Melanostomiinae by the anal-fin origin below the dorsal-fin origin; adipose fin absent; 4–11 pectoral-fin rays, free from membrane, some of them bearing a mass of luminous tissue; premaxillary teeth with 2 large fangs followed by 7–16 smaller teeth; teeth on vomer absent; 2–4 palatine teeth in a single row; conspicuous line or loop of blue luminous tissue on flank (Morrow, Gibbs, 1964; Sutton et al., 2020). Four valid species (Fricke et al., 2022).

Grammatostomias dentatus Goode & Bean, 1896

(Fig. 16A)

FIGURE 16 | A. Grammatostomias dentatus, NPM 3184, 114 mm SL; B. Grammatostomias ovatus, NPM 3191, 67 mm SL; C. Leptostomias gladiator, NPM 4782, 83 mm SL. Scale bars = 10 mm.

Diagnosis. Grammatostomias dentatus differs from congeners by the luminous tissue on both sides of body forming a distinct line from the operculum to beyond the pelvic-fin base; 5 pectoral-fin rays; PV 15–16 (Morrow, Gibbs, 1964; Sutton et al., 2020).

Geographical distribution. North Atlantic, Indian, and western Pacific oceans (Gibbs, Barnett, 1990; Vinnichenko, 1997; Harold, 2003; Fricke, 2011; Carneiro et al., 2019; Sutton et al., 2020). In the western Atlantic the species is reported from off the United States and Bermudas, between 43ºN and 28ºN (Morrow, Gibbs, 1964; Harold, 2003; Moore et al., 2003; Judkins, Haedrich, 2018). Melo et al. (2020) included G. dentatus in a list of deep-sea fishes of Brazil, without providing information on voucher specimens or locality data. Previously known records of the species in the South Atlantic are actually outside the Brazilian EEZ (ISH 1364-1968; ISH 762-1966). Therefore, the occurrence of G. dentatus is confirmed in Brazilian waters based on one specimen collected off Pernambuco State, at depths between 45 and 200 m (Fig. 14D).

Specimen examined. 1: NPM 3184, 1 (114 mm), AB1#51.

Grammatostomias ovatus Prokofiev, 2014

(Fig. 16B)

Diagnosis. Grammatostomias ovatus differs from congeners by the luminous tissue on flank always longer than deep, forming an elongate circle with a wavy outline, ending far in advance of the pelvic-fin origin; 6–9 pectoral-fin rays; first pectoral-fin ray isolated and heavily pigmented, remaining rays unpigmented, with masses of luminescent tissue, enormously developed on the second ray; PV 17–18 (Prokofiev, 2014; Prokofiev, unpubl. data).

Geographical distribution. The species was recently described based only on the holotype from the Canary Basin (Prokofiev, 2014), but additional specimens are also known from the tropical Atlantic Ocean, between 20ºN and 4ºN (Prokofiev, unpub. data). Grammatostomias ovatus is therefore reported for the first time in the South Atlantic based on one specimen collected off Pernambuco State in the Brazilian EEZ, between depths of 45 and 200 m (Fig. 14D).

Remarks. Menezes et al. (2003) included a morphologically similar species, G. circularis Morrow, 1959, in a checklist of Brazilian marine fishes based on Morrow, Gibbs (1964), who actually did not record the species in the Brazilian EEZ. Melo et al. (2020: 208), in turn, considered the occurrence of G. circularis as not confirmed in Brazilian waters.In the description of G. ovatus, Prokofiev (2014) indicated that the species could be distinguished from G. circularis by the number of pectoral-fin rays (6 vs. 9). Examination of additional specimens of G. ovatus indicates that the number of pectoral-fin rays is variable in the species (6–9). However, G. circularis is still distinct from G. ovatus in the shape of the luminous loop on the flank, which is nearly circular in the former (e.g., Morrow, Gibbs, 1964). The degree of elongation of the luminous loop in G. ovatus is also somewhat variable, but it is always longer than deep in that species instead of almost as long as deep in the case of G. circularis. The luminous tissue associated with the pectoral-fin rays of G. ovatus is also conspicuously more developed than in G. circularis, even though some ontogenetic variation in this character might be expected. In summary, the additional specimens of G. ovatus examined after its description in 2014 supports the distinction between that species and G. circularis; the occurrence of the later in Brazilian waters has not yet been confirmed.

Specimen examined. 1: NPM 3191, 1 (67 mm), AB1#51.

Leptostomias Gilbert, 1905

Diagnosis. Leptostomias can be distinguished in the Melanostomiinae by a very long and slender body, maximum depth around 10% SL or less; anal-fin origin below the dorsal-fin origin, adipose fin absent; 10–12 pectoral-fin rays; 20–29 anal-fin rays; 16–22 dorsal-fin rays; one tooth on each side of vomer; one or two pairs of teeth on basibranchials; PV 36 or more; VAV 20–24 (Morrow, Gibbs, 1964; Gibbs, 1986c; Kenaley, Stewart, 2015h; Sutton et al., 2020). A total of 12 valid species (Fricke et al., 2022).

Leptostomias gladiator (Zugmayer, 1911)

(Fig. 16C)

Diagnosis. Leptostomias gladiator differs from congeners by the long chin barbel, 30–75% SL, the stem with 1–2 filaments near its base; terminal bulb of chin barbel not split, bearing one or more minute filaments or tubercles at tip; 19–22 dorsal-fin rays; 23–29 anal-fin rays; head small, 10.0–13.4% SL; PV 39–44; VAV 20–22; AC 11–13 (Morrow, Gibbs, 1964; Kenaley, Stewart, 2015h; Sutton et al., 2020).

Geographical distribution. Circumglobal, between 56ºN and 45ºS, except the western Indian and western Pacific oceans (Gibbs, Barnett, 1990; Porteiro et al., 1999; Bañón et al., 2010; Fricke et al., 2014; Kenaley, Stewart, 2015h; Orlov, Tokranov, 2019; Sutton et al., 2020). In the western Atlantic, the species is reported from off United States to Uruguay, including the Gulf of Mexico and the Caribbean Sea (Morrow, Gibbs, 1964; McEachran, Fechhelm, 1998; Harold, 2003; Moore et al., 2003; Nión et al., 2016; Judkins, Haedrich, 2018). Menezes et al. (2003) included L. gladiator in a list of Brazilian marine fishes based on records outside the country’s EEZ (Morrow, Gibbs, 1964; Gibbs, 1986c). The occurrence of the species in Brazilian waters was later considered as “not confirmed” by Melo et al. (2020: 208). Therefore, occurrence of L. gladiator is confirmed in the Brazilian EEZ based on one specimen collected off Fernando de Noronha Archipelago, at 780 m depth (Fig. 14D). One additional specimen, collected off Rio de Janeiro State in Brazilian waters (MNRJ 30806), is also known.

Specimen examined. 1: NPM 4782, 1 (83 mm), AB2#42A.

Melanostomias Brauer, 1902

Diagnosis. Melanostomias can be distinguished in the Melanostomiinae by the dorsal and anal fins placed posteriorly in the body, near the caudal-fin base; anal-fin origin below dorsal-fin origin; pectoral fin without isolated rays; 3–7 (usually 5) pectoral-fin rays; 13–18 dorsal-fin rays; 16–21 anal-fin rays; jaw teeth long and depressible; 8 teeth on premaxilla; 4–5 teeth on maxilla; two pairs of teeth on basibranchials; palatine with 5 teeth; vomer with 1 tooth on each side; postorbital organ large; chin barbel present; body depth larger than 10% SL (Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1978; McEachran, Fechhelm, 1998; Sutton et al., 2020). A total of 18 valid species (Fricke et al., 2022).

Melanostomias bartonbeani Parr, 1927

(Fig. 17A)

FIGURE 17 | A. Melanostomias bartonbeani, NPM 4609, 185 mm SL; B. Melanostomias biseriatus,NPM 4603, 177 mm SL; C. Melanostomias tentaculatus, NPM 3181, 201 mm SL. Scale bars = 10 mm.

Diagnosis. Melanostomias bartonbeani differs from congeners by the chin barbel length 17–33% SL; terminal end of chin barbel with masses of opaque material filling much of the membrane on both sides of the axis; barbel tip unpigmented, about 5 times longer than wide; relatively large, white spots on head; 12–16 dorsal-fin rays; 16–20 anal-fin rays; PV 22–26; VAV 12–16; AC 8–11; IC 55–60; OV 22–25; VAL 11–14 (Parr, 1927; Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1978; Gibbs, 1986c; Sutton et al., 2020; specimens examined herein).

Geographical distribution. Atlantic and Indian oceans, between 70ºN and 40ºS (Morrow, Gibbs, 1964; De Groot, Nijssen, 1971; Gibbs, 1986c; Vinnichenko, 1997; Møller et al., 2010; Carneiro et al.,2019; Sutton et al., 2020). In the western Atlantic, the species was previously reported off Greenland, Canada, United States, Bermudas and Uruguay (Morrow, Gibbs, 1964; McAllister, 1990; Møller et al., 2010; Nión et al., 2016). The species was reported in Brazilian waters by Melo et al. (2020) likely based on one specimen collected off Espírito Santo State (MNRJ 30905). The occurrence of M. bartonbeani is therefore corfirmed in the Brazilian EEZ based on two specimens collected off Paraíba State and the Fernando de Noronha Archipelago, between depths of 90 and 780 m (Fig. 14E).

Remarks. Morphometric and meristic data of the two specimens examined are within the ranges reported for the species (Morrow, Gibbs, 1964; Gibbs, 1986c), except for the barbel length, which is longer (26–33% SL) than previously indicated (17–24% SL).

Specimens examined. 2 (50–185 mm SL): NPM 3422, 1 (50 mm), AB1#34; NPM 4609, 1 (185 mm), AB2#42A.

Melanostomias biseriatus Regan & Trewavas, 1930

(Fig. 17B)

Diagnosis. Melanostomias biseriatus differs from congeners by the chin barbel length, about 20–40% SL; barbel with a slender stem and a flattened distal tip terminating in a prominent ovoid bulb; length of the expanded distal part of the chin barbel about 10 times longer than wide; the distal tip of chin barbel bears two narrow flattened wings with a row of small and discrete opaque luminous bodies on each side; PV 27–30; OV 26–28 (Morrow, Gibbs, 1964; McEachran, Fechhelm, 1998; Sutton et al., 2020).

Geographical distribution. North Atlantic, between 35º and 5º N, and a single record in the North Pacific Ocean (Gibbs, 1960; Morrow, Gibbs, 1964; Moore et al., 2003; Sutton et al., 2020). In the western Atlantic, the species is reported from off the United States to French Guiana, including the Gulf of Mexico and the Caribbean Sea (Morrow, Gibbs, 1964; Bekkker et al., 1975; Fujii, 1983d; Judkins, Haedrich, 2018). Melanostomias biseariatus is therefore reported for the first time in the western South Atlantic based on two specimens collected between depths of 610 and 680 m off Pernambuco State and the Rocas Atoll, in the Brazilian EEZ (Fig. 14E). Two additional specimens collected outside the Brazilian EEZ off north South America (MCZ 132252) and off northeastern Brazil (ISH 665-1966) likely represent further records of the species in the western South Atlantic.

Specimens examined. 2 (29–177 mm SL): NPM 4603, 1 (177 mm), AB2#53A; NPM 4815, 1 (29 mm), AB2#16.

Melanostomias tentaculatus (Regan & Trewavas, 1930)

(Fig. 17C)

Diagnosis. Melanostomias tentaculatus differs from congeners by the chin barbel not flattened, slender, its length about 2–3 times in HL or 25% to 50% SL; barbel with one or two pale elongated terminal bulbs, with very small terminal filaments; main stem of chin barbel extended as a free tentacle along the bulb; 5–7 pectoral-fin rays; anal-fin origin just posterior to dorsal-fin origin; PV 25–27; OV 24–26 (Morrow, Gibbs, 1964; Parin, Pokhilskaya, 1978; McEachran, Fechhelm, 1998; Kenaley, Stewart, 2015h; Sutton et al., 2020).

Geographical distribution. Atlantic, Indian, and westernPacific oceans (Parin, Pokhilskaya, 1978; Gibbs, Barnett, 1990; Vinnichenko, 1997; Bordes et al., 1999; Aizawa, 2002; Harold, 2003; Tatsuta et al., 2014; Kenaley, Stewart, 2015h; Carneiro et al., 2019; Sutton et al., 2020). In the western Atlantic, the species is reported from off the United States to Argentina, including the Gulf of Mexico and the Caribbean Sea (Parin et al., 1974; Morrow, Gibbs, 1964; Moore et al.,2003; McEachran, Fechhelm, 1998; Judkins, Haedrich, 2018). In Brazil the species was previously reported off Maranhão State and the east coast to about 8ºS (McEachran, Fechhelm, 1998; Judkins, Haedrich, 2018). Melanostomias tentaculatus is reported here based on specimens collected off the Pernambuco State and Fernando de Noronha Archipelago, between depths of 45 and 800 (Fig. 14E).

Specimens examined. 4 (182–201 mm SL): NPM 3181, 1 (201 mm), AB1#51; NPM 4604, 1 (182 mm), AB2#40A; NPM 4605, 1 (194 mm), AB2#41A; NPM 4607, 1 (188 mm), AB2#21.

Photonectes Günther, 1887

Diagnosis. Photonectes can be distinguished in the Melanostomiinae by the dorsal and anal fins opposed, rays covered with black skin in some species, leaving only tips of rays visible; 0–3 pectoral-fin rays; lower jaw longer than upper jaw and strongly curved upwards; lower jaw teeth uniserial; retroarticular with a process directed backwards; vomer with one or two pairs of teeth, rarely toothless; postorbital organ well-developed; some species with white luminous tissue on top of head and/or in shoulder region, others with blue luminous tissue in spots on head, inside mouth or in bands or stripes along body; 48–67 vertebrae (Morrow, Gibbs, 1964; Gibbs, 1986c; Klepadlo, 2011; Flynn, Klepadlo, 2012; Kenaley, Stewart, 2015h; Prokofiev, 2016, 2019; Prokofiev, Klepadlo, 2019; Prokofiev, Frable, 2021). A total of 29 valid species (Fricke et al., 2022).

Photonectes achirus Regan & Trewavas, 1930

(Fig. 18A)

FIGURE 18 | A. Photonectes achirus, NPM 4629, 79 mm SL; B. Thysanactis dentex, NPM 4639, 138 mm SL. Scale bars = 10 mm.

Diagnosis. Photonectes achirus differs from congeners by the absence of the pectoral fin; IV 31–36; one pair of luminous organs inside mouth; chin barbel length 70–80% HL, with a small terminal bulb and a terminal appendage with a secondary, small, bulb at its tip; presence of blue luminous tissue in a midventral band extending from the pectoral region to the pelvic fin, with short transverse streaks alternating with serial photophores, and in patches under OV-1 and OV-2 photophores, on sides of isthmus, under the lower jaw, above the posterior end of the maxilla, between the eyes, and above the occipito-vertical articulation; BR 6–8 (Morrow, Gibbs, 1964; Flynn, Klepadlo, 2012; Prokofiev, Klepadlo, 2019; Prokofiev, Frable, 2021).

Geographical distribution. Known from a few records in the western and central Pacific and western North Atlantic oceans, off United States and in the Caribbean Sea (McEachran, Fechhelm, 1998; Harold, 2003; Mundy, 2005; Klepadlo, 2011; Flynn, Klepadlo, 2012). Photonectes achirus is reported for the first time in the South Atlantic based on three specimens collected off Rio Grande do Norte (including the seamounts) and Paraíba States in the Brazilian EEZ, between depths of 70 and 800 m (Fig. 14F).

Remarks. Two specimens listed as Photonectes sp. (NPM 6867, 26–29 mm; AB2#42A) in the Supplementary Material S1 likely represent P. achirus, but their identification could not be confirmed at species level due to their extremely small sizes.

Specimens examined. 3 (33–79 mm SL): NPM 4629, 1 (79 mm), AB2#54B; NPM 4695, 1 (33 mm), AB2#21; NPM 4924, 1 (damaged), AB2#28.

Thysanactis Regan & Trewavas, 1930

Diagnosis. Thysanactis can be distinguished in the Melanostomiinae by the dorsal and anal fins placed posteriorly on body, anal-fin origin slightly anterior to dorsal-fin origin; first pectoral-fin ray long and isolated, ending in 5–6 filaments, followed by 10–11 additional pectoral-fin rays; chin barbel short, similar in length to head length, ending in a round terminal bulb covered with filaments; IP 20; PV 31–32; VAV 14–16; OV 30–32; VAL 14–16; AC 11–12 (Morrow, Gibbs, 1964; Sutton et al., 2020). A single valid species (Fricke et al., 2022).

Thysanactis dentex Regan & Trewavas, 1930

(Fig. 18B)

Diagnosis. Same as for the genus.

Geographical distribution. Atlantic, eastern Indian, and western and central Pacific oceans (Morrow, Gibbs, 1964; Clarke, 1974; Gibbs, Barnett, 1990; Hutchins, 2001; Aizawa, 2002; Harold, 2003; Fricke et al., 2011; Sutton et al., 2020). In the western Atlantic, the species is reported between 18ºN and 7ºS, from the Caribbean Sea to Brazil, where it was recorded off the northeastern region (Eduardo et al., 2018a; Judkins, Haedrich, 2018). Thysanactis dentex is reported here based on specimens collected from off Rio Grande do Norte State (including the seamounts), the Rocas Atoll, and the Fernando de Noronha Archipelago, at depths between 90 and 1,113 m (Fig. 14F).

Specimens examined. 40 (43–150 mm SL): NPM 3281, 1 (52 mm), AB1#12; NPM 3282, 1 (50 mm), AB1#14; NPM 3421, 1 (damaged), AB1#22; NPM 4626, 2 (68–92 mm), AB2#41A; NPM 4627, 8 (67–131 mm), AB2#44A; NPM 4632, 3 (137–145 mm), AB2#52A; NPM 4633, 1 (135 mm), AB2#39; NPM 4635, 1 (94 mm), AB2#59B; NPM 4636, 1 (81 mm), AB2#54B; NPM 4637, 3 (132–145 mm), AB2#28; NPM 4638, 2 (113–145 mm), AB2#59A; NPM 4639, 2 (46–138 mm), AB2#52B; NPM 4640, 1 (51 mm), AB2#44B; NPM 4641, 1 (90 mm), AB2#42A; NPM 4649, 9 (43–55 mm), AB2#49B; NPM 4694, 2 (115–150 mm), AB2#50A.

Discussion​


Scientific expeditions conducted off the Brazilian coast in recent decades contributed significantly to the understanding of the deep-sea, especially the taxonomic composition and geographic distribution of different groups of fishes (Séret, Andreata, 1992; Figueiredo et al., 2002; Bernardes et al., 2005; Braga et al., 2007; Mincarone et al., 2008; Costa, Mincarone, 2010; Lima et al., 2011; Lins Oliveira et al., 2015; Melo et al., 2020). With the total number of 108 species of the Stomiiformes confirmed in the Brazilian EEZ, including 90 species previously recorded (Tab. 2), the order might be regarded as one of the most well-known groups of deep-sea fishes in the western South Atlantic. However, knowledge on deep-sea fishes in general remains highly unsatisfactory in the region (Irigoien et al., 2014; Reis et al., 2016; St. Jonh et al., 2016). One example of this is that, among the 12 new records for the Brazilian EEZ presented here, seven are also new for the western South Atlantic, the South Atlantic, or even the Atlantic.

Phosichthyidae is the least diverse family of the Stomiiformes in the Brazilian EEZ. Of the eigth species of the family confirmed for the region, four were identified here (Tab. 2). The second most well represented family of the Stomiiformes in the Brazilian EEZ is the Sternoptychidae, with 15 confirmed species, followed by the Gonostomatidae, with 14 species. Nine species of the Gonostomatidae were recorded here, including Gonostoma denudatum, a new record for the region. In the Sternoptychidae, there is also a likely new species of Polyipnus from off the central coast of Brazil that still lacks formal description (Lima et al., 2011). However, of the 18 new or confirmed records of stomiiforms presented here for Brazilian waters based on specimens we examined, 16 are members of the highly diverse Stomiidae. When the 50 additional species of the Stomiidae previously reported for the Brazilian EEZ are also considered, a total of 66 species of the family is confirmed for the region (Tab. 2). Our collections also include at least five species of Eustomias and one species of Melanostomias that likely represent new species awaiting formal description.

TABLE 2 | Confirmed records of Stomiiformes in the Brazilian Exclusive Economic Zone.

Species

References

DIPLOPHIDAE

 

Diplophos australis Ozawa, Oda & Ida, 1990

Figueiredo et al. (2002); Menezes et al. (2003); Melo et al. (2020); this study

Diplophos taenia Günther, 1873

Figueiredo et al. (2002); Menezes et al. (2003); Bonecker, Castro (2006); Braga et al. (2007); Bonecker et al. (2014); Pinheiro et al. (2015); Stocco, Joyeux (2015); Eduardo et al. (2020c); Melo et al. (2020); this study

Manducus maderensis (Johnson, 1890)

Mukhacheva (1978); Smith et al. (1991); Figueiredo et al. (2002); Menezes et al. (2003); Braga et al. (2007); Costa, Mincarone (2010); Pinheiro et al. (2015); Judkins, Haedrich (2018); Melo et al. (2020); this study

Triplophos hemingi (McArdle, 1901)

This study

GONOSTOMATIDAE

 

Cyclothone acclinidens Garman, 1899

Bonecker et al. (2012, 2014); Stocco, Joyeux (2015); Olivar et al. (2017); Melo et al. (2020); Pinheiro et al. (2020)

Cyclothone alba Brauer, 1906

Séret, Andreata (1992); Bonecker et al. (2012; 2014); Olivar et al. (2017); Judkins, Haedrich (2018); Melo et al. (2020); Pinheiro et al. (2020)

Cyclothone braueri Jespersen & Tåning, 1926

Séret, Andreata (1992); Bonecker et al. (2012, 2014); Stocco, Joyeux (2015); Melo et al. (2020)

Cyclothone microdon (Günther, 1878)

Judkins, Haedrich (2018); Melo et al. (2020)

Cyclothone obscura Brauer, 1902

Mukhacheva (1974)

Cyclothone pallida Brauer, 1902

Olivar et al. (2017); Judkins, Haedrich (2018); Melo et al. (2020); Pinheiro et al. (2020)

Cyclothone parapallida Badcock, 1982

Olivar et al. (2017); Melo et al. (2020)

Cyclothone pseudopallida Mukhacheva, 1964

Mukhacheva (1974); Bonecker et al. (2012, 2014); Stocco, Joyeux (2015); Olivar et al. (2017); Judkins, Haedrich (2018); Melo et al. (2020)

Gonostoma atlanticum Norman, 1930

Mukhacheva (1972); Judkins, Haedrich (2018); Melo et al. (2020); this study

Gonostoma denudatum Rafinesque, 1810

This study

Margrethia obtusirostra Jespersen & Tåning, 1919

Figueiredo et al. (2002); Menezes et al. (2003); Braga et al. (2007); Bonecker et al. (2012, 2014); Melo et al. (2020); this study

Sigmops bathyphilus (Vaillant, 1884)

Costa, Mincarone (2010); Melo et al. (2020); this study

Sigmops elongatus (Günther, 1878)

Mukhacheva (1972); Figueiredo et al. (2002); Menezes et al. (2003); Bernardes et al. (2005); Braga et al. (2007); Haimovici et al. (2008); Bonecker et al. (2012, 2014); Mincarone et al. (2017); Eduardo et al. (2018a); Judkins, Haedrich (2018); Melo et al. (2020); this study

Zaphotias pedaliotus (Goode & Bean, 1896)

Mukhacheva (1976); Eduardo et al. (2018a); Judkins, Haedrich (2018); Melo et al. (2020); this study

STERNOPTYCHIDAE

 

Maurolicinae

 

Argyripnus atlanticus Maul, 1952

Braga et al. (2007); Lima et al. (2011); Melo et al. (2020)

Maurolicus stehmanni Parin & Kobyliansky, 1993

Figueiredo et al. (2002); Bernardes et al. (2005); Braga et al. (2007); Haimovici et al. (2008); Lima et al. (2011); Bonecker et al. (2012, 2014); Pinheiro et al. (2015); Stocco, Joyeux (2015); Melo et al. (2020), Rees et al. (2020)

Maurolicus weitzmani Parin & Kobyliansky, 1993

Judkins, Haedrich (2018); Rees et al. (2020)

Valenciennellus tripunctulatus (Esmark, 1871)

Figueiredo et al. (2002); Bonecker et al. (2014); Olivar et al. (2017); Judkins, Haedrich (2018); Melo et al. (2020); this study

Sternoptychinae

 

Argyropelecus aculeatus Valenciennes, 1850

Borodulina (1978); Figueiredo et al. (2002); Menezes et al. (2003); Bernardes et al. (2005); Braga et al. (2007); Haimovici et al. (2008); Costa, Mincarone (2010); Lima et al. (2011); Bonecker et al. (2014); Eduardo et al. (2018a); Judkins, Haedrich (2018); Eduardo et al. (2020a); Melo et al. (2020); this study

Argyropelecus affinis Garman, 1899

Borodulina (1978); Eduardo et al. (2018a); Eduardo et al. (2020a); Melo et al. (2020); this study

Argyropelecus gigas Norman, 1930

Borodulina (1978); Bernardes et al. (2005); Haimovici et al. (2008); Melo et al. (2020); this study

Argyropelecus hemigymnus Cocco, 1829

Borodulina (1978); Figueiredo et al. (2002); Menezes et al. (2003); Bernardes et al. (2005); Haimovici et al. (2008); Costa, Mincarone (2010); Lima et al. (2011); Bonecker et al. (2014); Olivar et al. (2017); Judkins, Haedrich (2018); Eduardo et al. (2020a); Melo et al. (2020); this study

Argyropelecus lychnus Garman, 1899

Judkins, Haedrich (2018)

Argyropelecus sladeni Regan, 1908

Baird (1971); Borodulina (1978); Menezes et al. (2003); Lima et al. (2011); Bonecker et al. (2014); Olivar et al. (2017); Eduardo et al. (2018a); Judkins, Haedrich (2018); Melo et al. (2020); this study

Polyipnus clarus Harold, 1994

Lins Oliveira et al. (2015); Melo et al. (2020)

Polyipnus laternatus Garman, 1899

Lins Oliveira et al. (2015); Melo et al. (2020)

Polyipnus sp.

Lima et al. (2011)

Sternoptyx diaphana Hermann, 1781

Baird (1971); Borodulina (1978); Séret, Andreata (1992); Figueiredo et al. (2002); Menezes et al. (2003); Braga et al. (2007); Costa, Mincarone (2010); Lima et al. (2011); Bonecker et al. (2012, 2014); Lins Oliveira et al. (2015); Olivar et al. (2017); Eduardo et al. (2018a); Judkins, Haedrich (2018); Eduardo et al. (2020a); Melo et al. (2020); Pinheiro et al. (2020); this study

Sternoptyx pseudobscura Baird, 1971

Baird (1971); Menezes et al. (2003); Braga et al. (2007); Costa, Mincarone (2010); Lima et al. (2011); Eduardo et al. (2018a); Judkins, Haedrich (2018); Eduardo et al. (2020a); Melo et al. (2020); this study

Sternoptyx pseudodiaphana Borodulina, 1977

Costa, Mincarone (2010); Lima et al. (2011); Lins Oliveira et al. (2015); Mincarone et al. (2017); Judkins, Haedrich (2018); Melo et al. (2020); this study

PHOSICHTHYIDAE

 

Ichthyococcus australis Mukhacheva, 1980

Krefft (1983); Menezes et al. (2003); Melo et al. (2020)

Ichthyococcus ovatus (Cocco, 1838)

Bonecker et al. (2014); Judkins, Haedrich (2018); Melo et al. (2020)

Ichthyococcus polli Blache, 1964

Krefft (1983); Melo et al. (2020); this study

Phosichthys argenteus Hutton, 1872

Figueiredo et al. (2002); Bernardes et al. (2005); Braga et al. (2007); Haimovici et al. (2008); Melo et al. (2020); this study

Pollichthys mauli (Poll, 1953)

Figueiredo et al. (2002); Bernardes et al. (2005); Bonecker, Castro (2006); Braga et al. (2007); Haimovici et al. (2008); Costa, Mincarone (2010); Goçalo et al. (2011); Bonecker et al. (2012, 2014); Pinheiro et al. (2015); Melo et al. (2020); this study

Polymetme thaeocoryla Parin & Borodulina, 1990

Haimovici et al. (1994); Mincarone et al. (2004); Bernardes et al. (2005); Braga et al. (2007); Haimovici et al. (2008); Lins Oliveira et al. (2015); Mincarone et al. (2017); Melo et al. (2020)

Vinciguerria nimbaria (Jordan & Williams, 1895)

Gorbunova (1972); Figueiredo et al. (2002); Bonecker, Castro (2006); Braga et al. (2007); Goçalo et al. (2011); Bonecker et al. (2012, 2014, 2019); Pinheiro et al. (2015, 2020); Stocco, Joyeux (2015); Judkins, Haedrich (2018); Melo et al. (2020); this study

Vinciguerria poweriae (Cocco, 1838)

Figueiredo et al. (2002); Stocco, Joyeux (2015); Melo et al. (2020)

STOMIIDAE

 

Astronesthinae

 

Astronesthes atlantica Parin & Borodulina, 1996

Parin, Borodulina (1996); Judkins, Haedrich (2018); this study

Atronesthes barbata Kner, 1860

Kner (1860); Parin, Borodulina (2002); Melo et al. (2020)

Astronesthes boulengeri Gilchrist, 1902

Parin et al. (1999)

Astronesthes gemmifer Goode & Bean, 1896

Judkins, Haedrich (2018); Melo et al. (2020); this study

Astronesthes gudrunae Parin & Borodulina, 2002

This study

Astronesthes haplophos Parin & Borodulina, 2002

Parin, Borodulina (2003); Melo et al. (2020)

Astronesthes illuminata Parin, Borodulina & Hulley, 1999

Parin et al. (1999)

Astronesthes macropogon Goodyear & Gibbs, 1970

Parin, Borodulina (1997); Figueiredo et al. (2002); Braga et al. (2007); Costa, Mincarone (2010); Mincarone et al. (2017); Melo et al. (2020)

Astronesthes micropogon Goodyear & Gibbs, 1970

Parin, Borodulina (2003); Judkins, Haedrich (2018)

Astronesthes richardsoni (Poey, 1852)

Parin, Borodulina (2000, 2003); Judkins, Haedrich (2018); this study

Astronesthes similus Parr, 1927

Braga et al. (2007); Pinheiro et al. (2015); Eduardo et al. (2020c); Melo et al. (2020); this study

Borostomias elucens (Brauer, 1906) 

This study

Heterophotus ophistoma Regan & Trewavas, 1929

Judkins, Haedrich (2018); Klautau et al. (2020); Marceniuk et al., (2021); this study

Chauliodontinae

 

Chauliodus danae Regan & Trewavas, 1929

Judkins, Haedrich (2018); Melo et al. (2020)

Chauliodus minimus Parin & Novikova, 1974

Parin, Novikova (1974); Bernardes et al. (2005); Haimovici et al. (2008); Melo et al. (2020)

Chauliodus schmidti Ege, 1948

Judkins, Haedrich (2018)

Chauliodus sloani Bloch & Schneider, 1801

Parin, Novikova (1974); Figueiredo et al. (2002); Menezes et al. (2003); Bernardes et al. (2005); Bonecker, Castro (2006); Braga et al. (2007); Haimovici et al. (2008); Bonecker et al. (2014); Lins Oliveira et al. (2015); Judkins, Haedrich (2018); Eduardo et al. (2020b); Melo et al. (2020); this study

Stomiinae

 

Stomias affinis Günther, 1887

Haimovici et al. (1994); Figueiredo et al. (2002); Bernardes et al. (2005); Bonecker, Castro (2006); Braga et al. (2007); Haimovici et al. (2008); Costa, Mincarone (2010); Bonecker et al. (2014); Mincarone et al. (2017); Melo et al. (2020)

Stomias boa (Risso, 1810)

Shcherbachev, Novikova (1976); Gibbs (1969); Melo et al. (2020)

Stomias colubrinus Garman, 1899

Séret, Andreata (1992); Melo et al. (2020)

Stomias danae Ege, 1933

Gibbs (1969); Menezes et al. (2003); Judkins, Haedrich (2018); Melo et al. (2020); this study

Stomias longibarbatus (Brauer, 1902)

Judkins, Haedrich (2018); this study

Idiacanthinae

 

Idiacanthus atlanticus Brauer, 1906

Figueiredo et al. (2002); Mincarone et al. (2004); Bernardes et al. (2005); Haimovici et al. (2008); Melo et al. (2020)

Malacosteinae

 

Aristostomias grimaldii Zugmayer, 1913

This study

Aristostomias tittmanni Welsh, 1923

Judkins, Haedrich (2018); this study

Aristostomias xenostoma Regan & Trewavas, 1930

Judkins, Haedrich (2018)

Malacosteus niger Ayres, 1848

Braga et al. (2007); Costa, Mincarone (2010); Olivar et al. (2017); Mincarone et al. (2017); Eduardo et al. (2018a); Judkins, Haedrich (2018); Melo et al. (2020); this study

Pachystomias microdon (Günther, 1878)

This study

Photostomias atrox (Alcock, 1890)

Kenaley, Hartel (2005); Melo et al. (2020); this study

Photostomias goodyeari Kenaley & Hartel, 2005

Melo et al. (2020); this study

Photostomias guernei Collett, 1889

Bonecker, Castro (2006); Melo et al. (2020)

Melanostomiinae

 

Bathophilus brevis Regan & Trewavas, 1930

Judkins, Haedrich (2018); Sutton et al. (2020)

Bathophilus nigerrimus Giglioli, 1882

This study

Bathophilus pawneei Parr, 1927

Judkins, Haedrich (2018); Sutton et al. (2020); this study

Echiostoma barbatum Lowe, 1843

Menezes et al. (2003); Melo et al. (2020)

Eustomias arborifer Parr, 1927

Gibbs et al. (1983); Judkins, Haedrich (2018); Melo et al. (2020)

Eustomias bibulbosus Parr, 1927

This study

Eustomias bigelowi Welsh, 1923

Clarke (2000)

Eustomias braueri Zugmayer, 1911

Melo et al. (2020); this study

Eustomias brevibarbatus Parr, 1927

Gomon, Gibbs (1985); Menezes et al. (2003); Costa, Mincarone (2010); Melo et al. (2020); this study

Eustomias curtifilis Clarke, 2000

Clarke (2000); Melo et al. (2020)

Eustomias enbarbatus Welsh, 1923

Judkins, Haedrich (2018); this study

Eustomias filifer (Gilchrist, 1906)

Menezes et al. (2003); Melo et al. (2020)

Eustomias fissibarbis (Pappenheim, 1914)

Judkins, Haedrich (2018)

Eustomias ignotus Gomon & Gibbs, 1985

Gomon, Gibbs (1985); Melo et al. (2020)

Eustomias longibarba Parr, 1927

Menezes et al. (2003); Melo et al. (2020)

Eustomias macronema Regan & Trewavas, 1930

Clarke (2000)

Eustomias minimus Clarke, 1999

This study

Eustomias posti Gibbs, Clarke & Gomon, 1983

Gibbs et al. (1983); Judkins, Haedrich (2018); Melo et al. (2020)

Eustomias satterleei Beebe, 1933

Clarke (2000); Judkins, Haedrich (2018)

Eustomias schmidti Regan & Trewavas, 1930

Melo et al. (2020); this study

Eustomias spherulifer Gibbs, Clarke & Gomon, 1983

Gibbs et al. (1983); Melo et al. (2020)

Eustomias tetranema Zugmayer, 1913

Costa, Mincarone (2010); Melo et al. (2020)

Melanostomiinae

 

Flagellostomias boureei (Zugmayer, 1913)

Bernardes et al. (2005); Bonecker, Castro (2006); Haimovici et al. (2008); Melo et al. (2020)

Grammatostomias dentatus Goode & Bean, 1896

Melo et al. (2020); this study

Grammatostomias ovatus Prokofiev, 2014

This study

Leptostomias gladiator (Zugmayer, 1911)

Melo et al. (2020); this study

Leptostomias longibarba Regan & Trewavas, 1930

Judkins, Haedrich (2018); Melo et al. (2020)

Melanostomias bartonbeani Parr, 1927

Melo et al. (2020); this study

Melanostomias biseriatus Regan & Trewavas, 1930

This study

Melanostomias niger Gilchrist & von Bonde, 1924

Haimovici et al. (1994); Figueiredo et al. (2002); Bernardes et al. (2005); Haimovici et al. (2008); Melo et al. (2020)

Melanostomias tentaculatus (Regan & Trewavas, 1930)

Judkins, Haedrich (2018); Melo et al. (2020); this study

Melanostomias valdiviae Brauer, 1902

Menezes et al. (2003); Melo et al. (2020)

Photonectes achirus Regan & Trewavas, 1930

This study

Photonectes mirabilis Parr, 1927

Bonecker, Castro (2006); Judkins, Haedrich (2018); Melo et al. (2020)

Thysanactis dentex Regan & Trewavas, 1930

Eduardo et al. (2018a); Judkins, Haedrich (2018); this study

 

These results reinforce the vast amount of information still necessary for a more comprehensive knowledge of the deep-sea diversity of the western South Atlantic to emerge, even in relatively well-known families as the Stomiidae. The new records and geographic range extensions uncovered here based on just two collecting expeditions also highlight the need for additional mesopelagic surveys along the Brazilian coast.

Acknowledgments​


We thank the French oceanographic fleet for funding the at-sea survey ABRACOS and the officers and crew of the RV Antea for their contribution to the success of the operations. This study could not have been carried out without the support of the team members of the BIOIMPACT (UFRPE) and LIZ (NUPEM/UFRJ) labs. We thank Alexei Orlov (P.P. Shirshov Institute of Oceanology) and Sergei Evseenko (in memorian) for providing literature. Thanks, are also due to Alfredo Carvalho-Filho and an anonymous reviewer for insightful comments and suggestions to the manuscript. CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico – PROTAX 443302/2020) and CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) provided scholarships to Bárbara Villarins. CNPq Research Grants were provided to Flávia Lucena-Frédou, and Michael Mincarone (grants 308554/2019–1 and 314644/2020–2, respectively). Artem Prokofiev is supported by the Russian Science Foundation (grant 19-14-00026). The NPM Fish Collection is supported by FUNBIO (grant 104/2016 – “Pesquisa Marinha e Pesqueira”) and FAPERJ (grant E–26/210.290/2021). This study is a contribution to the LMI TAPIOCA, CAPES/COFECUB program (88881.142689/2017–01), and EU H2020 TRIATLAS project (grant agreement 817578). This is part of the study conducted by the first author to obtain the BSc Degree in Biological Sciences at the Universidade Federal do Rio de Janeiro (UFRJ). We are especially grateful to members of the examination board, Cristiano Moreira (MNRJ) and Luciano Fischer (NUPEM/UFRJ) for comments and suggestions.

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Authors


Bárbara T. Villarins1, Fabio Di Dario2, Leandro N. Eduardo3,4, Flávia Lucena-Frédou3, Arnaud Bertrand3,4, Artem M. Prokofiev5,6 and Michael M. Mincarone2

[1]    Programa de Pós-Graduação em Ciências Ambientais e Conservação, Instituto de Biodiversidade e Sustentabilidade,Universidade Federal do Rio de Janeiro, Av. São José do Barreto, 764, 27965-045 Macaé, RJ, Brazil. btvillarins@gmail.com.

[2]    Instituto de Biodiversidade e Sustentabilidade, Universidade Federal do Rio de Janeiro, Av. São José do Barreto, 764, 27965-045 Macaé, RJ, Brazil. (FDD) didario@gmail.com, (MMM) mincarone@macae.ufrj.br (corresponding author)

[3]    Departamento de Pesca e Aquicultura, Universidade Federal Rural de Pernambuco, Rua Dom Manuel de Medeiros, s/n, 52171- 900 Recife, PE, Brazil. (LNE) leandronole@hotmail.com, (FLF) flavialucena@hotmail.com.

[4]    MARBEC, Univ. Montpellier, CNRS, Ifremer, IRD, Sète, France. Arnaud.Bertrand@ird.fr.

[5]    A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninski prospekt 33, Moscow 119071, Russia. prokartster@gmail.com.

[6]    P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovski rospect 36, Moscow 117997, Russia.

Authors’ Contribution


Bárbara T. Villarins: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Fabio Di Dario: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Validation, Writing-original draft, Writing-review and editing.

Leandro N. Eduardo: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Flávia Lucena-Frédou: Data curation, Funding acquisition, Project administration, Supervision, Writing-review and editing.

Arnaud Bertrand: Data curation, Funding acquisition, Project administration, Supervision, Writing-review and editing.

Artem M. Prokofiev: Investigation, Methodology, Writing-original draft, Writing-review and editing.

Michael M. Mincarone: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Visualization, Writing-original draft, Writing-review and editing.

Ethical Statement​


The authors state that all methods were approved and conducted in accordance with guidelines and regulations of the Ministério do Meio Ambiente (SISBIO authorization number: 47270–5). Operations of the RV Antea were approved by the Brazilian Navy Authority (“Estado-Maior da Armada”) under the Ordinances 178 (08/09/2015) and 4 (24/01/2017).

Competing Interests


The authors declare no competing interests.

How to cite this article


Villarins BT, Di Dario F, Eduardo LN, Lucena-Frédou F, Bertrand A, Prokofiev AM, Mincarone MM. Deep-sea dragonfishes (Teleostei: Stomiiformes) collected from off northeastern Brazil, with a review of the species reported from the Brazilian Exclusive Economic Zone. Neotrop Ichthyol. 2022; 20(2):e220004. https://doi.org/10.1590/1982-0224-2022-0004


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Accepted May 16, 2022 by Luiz Osmar

Submitted January 17, 2022

Epub July 08, 2022