Taxonomy and Distribution of Deep-Sea Bigscales and Whalefishes (Teleostei: Stephanoberycoidei) Collected off Northeastern Brazil, Including Seamounts and Oceanic Islands

Abstract

Despite the increasing number of studies on the systematics of the Stephanoberycoidei (bigscales, pricklefishes, gibberfishes, hispidoberycids, and whalefishes) globally, knowledge about the diversity and distribution of the group in the western South Atlantic still remains fragmentary. In this study, we present new anatomical (meristic and morphometric) and distributional data for 18 species of the Stephanoberycoidei based on the examination of 150 specimens recently collected during the ABRACOS (Acoustics along the BRAzilian COaSt) expeditions off northeastern Brazil, including the Rocas Atoll, Fernando de Noronha Archipelago, and the seamounts off Rio Grande do Norte State, and additional museum specimens. In the Melamphaidae, remarks on the taxonomy of Melamphaes polylepis and of the Poromitra crassiceps and P. megalops species groups are made based on specimens examined. In addition, Scopeloberyx opercularis, currently considered as a junior synonym of Scopeloberyx robustus, is recognized as a valid species. Among the species identified, nine have their distributions extended in the western South Atlantic based on confirmed records: Melamphaes polylepis, M. typhlops, Poromitra megalops, Poromitra sp., Scopeloberyx opercularis, Scopeloberyx opisthopterus, Scopelogadus mizolepis, Cetostoma regani, and Rondeletia loricata. Eight further species are reported for the first time in Brazilian waters: Cetomimus sp. 1, Cetomimus sp. 2, Ditropichthys storeri, Gyrinomimus bruuni, Melamphaes eulepis, M. leprus, M. longivelis, and Melamphaes sp. Additional remarks on the taxonomy and distribution of the Stephanoberycoidei in the western South Atlantic are also provided.

Full text

Taxonomy and Distribution of Deep-Sea Bigscales and Whalefishes (Teleostei:
Stephanoberycoidei) Collected off Northeastern Brazil, Including Seamounts
and Oceanic Islands
Gabriel Vin´
ıcius Felix Afonso
1
, Fabio Di Dario
1
, Leandro Nol´
e Eduardo
2,3
, Fla´via
Lucena-Fr´
edou
2
, Arnaud Bertrand
2,3
, and Michael Maia Mincarone
1
Despite the increasing number of studies on the systematics of the Stephanoberycoidei (bigscales, pricklefishes,
gibberfishes, hispidoberycids, and whalefishes) globally, knowledge about the diversity and distribution of the group in
the western South Atlantic still remains fragmentary. In this study, we present new anatomical (meristic and
morphometric) and distributional data for 18 species of the Stephanoberycoidei based on the examination of 150
specimens recently collected during the ABRACOS (Acoustics along the BRAzilian COaSt) expeditions off northeastern
Brazil, including the Rocas Atoll, Fernando de Noronha Archipelago, and the seamounts off Rio Grande do Norte State,
and additional museum specimens. In the Melamphaidae, remarks on the taxonomy of Melamphaes polylepis and of the
Poromitra crassiceps and P. megalops species groups are made based on specimens examined. In addition, Scopeloberyx
opercularis, currently considered as a junior synonym of Scopeloberyx robustus, is recognized as a valid species. Among the
species identified, nine have their distributions extended in the western South Atlantic based on confirmed records:
Melamphaes polylepis,M. typhlops,Poromitra megalops,Poromitra sp., Scopeloberyx opercularis,Scopeloberyx opisthopterus,
Scopelogadus mizolepis,Cetostoma regani, and Rondeletia loricata. Eight further species are reported for the first time in
Brazilian waters: Cetomimus sp. 1, Cetomimus sp. 2, Ditropichthys storeri,Gyrinomimus bruuni,Melamphaes eulepis,M. leprus,
M. longivelis, and Melamphaes sp. Additional remarks on the taxonomy and distribution of the Stephanoberycoidei in the
western South Atlantic are also provided.
THE Stephanoberycoidei comprises 23 genera and
about 94 species of mostly meso- and bathypelagic
teleosts commonly known as bigscales, pricklefishes,
gibberfishes, hispidoberycids, and whalefishes (Nelson et al.,
2016; Fricke et al., 2020a). Fishes of this suborder usually
have a short to moderately long and somewhat compressed
body, the subocular shelf and orbitosphenoid are absent,
basibranchial tooth plates are also absent, with the exception
of the copular tooth plate in the Cetomimidae, ossification is
reduced to thin laminar bones on the surface of an
extensively cartilaginous neurocranium, the supramaxillae
are absent or reduced, and the extrascapular, when present, is
greatly enlarged, partially or entirely covering the parietal
bone (Johnson and Patterson, 1993; Moore, 1993; Wiley and
Johnson, 2010; Nelson et al., 2016).
Until recently, the Stephanoberycoidei was recognized as
an order (Stephanoberyciformes), distinct from the Beryci-
formes (e.g., Johnson and Patterson, 1993; Nelson, 2006;
Wiley and Johnson, 2010), but there is growing evidence
indicating that the former is a subgroup of the latter based
on both morphological (Stiassny and Moore, 1992; Moore,
1993) and molecular data (Miya et al., 2005; Near et al.,
2012; Betancur-R. et al., 2013; Dornburg et al., 2017).
Relationships within the Stephanoberycoidei are also
contentious: the Melamphaidae, traditionally recognized
as a family of the Stephanoberycoidei or Stephanoberyci-
formes (e.g., Johnson and Patterson, 1993; Moore, 1993;
Wiley and Johnson, 2010), has been proposed as the sister
group of the Berycidae in the Berycoidei (Miya et al., 2005;
Near et al., 2012; Betancur-R. et al., 2013; Dornburg et al.,
2017).However,forpurposesofthisstudy,wefollow
Moore’s (1993) traditional arrangement of the Stephanober-
ycoidei, including the families Melamphaidae, Stephano-
berycidae, Hispidoberycidae, Gibberichthyidae,
Rondeletiidae, Barbourisiidae, and Cetomimidae. The Mir-
apinnidae and Megalomycteridae, also traditionally includ-
ed in the Stephanoberycoidei (e.g., Moore, 1993; Nelson,
2006), are no longer recognized as valid since members of
those families are now regarded as larvae and males,
respectively, of the Cetomimidae (Johnson et al., 2009;
Nelson et al., 2016).
Several contributions have been made on the systematics
and biogeography of the Stephanoberycoidei in the last
decades (e.g., Ebeling, 1962; Ebeling and Weed, 1973; Keene,
1973, 1987; Keene et al., 1987; Paxton, 1989; Kotlyar, 1996,
1999, 2004a, 2004b, 2008a, 2011a, 2013, 2014, 2019; Merrett
and Moore, 2005; Bartow, 2010; Mincarone et al., 2014).
However, knowledge of the diversity and distribution of the
group in the western South Atlantic remains fragmentary,
despite the apparent relative abundance of some stephano-
berycoids in deep-sea environments (G¨
unther, 1887; Keene,
1987; Campos et al., 2008; Costa and Mincarone, 2010;
Mincarone et al., 2014; Judkins and Haedrich, 2018). This
study reports on new anatomical and taxonomic data of
mostly rare species of the Stephanoberycoidei recently
collected off northeastern Brazil. The extensively long
Brazilian coastline (c. 7,500 km; e.g., Reis et al., 2016) and
associated Exclusive Economic Zone (EEZ) encompasses a
substantial portion of the Tropical western South Atlantic.
Therefore, a review of the distribution of the species recorded
1
Instituto de Biodiversidade e Sustentabilidade, Universidade Federal do Rio de Janeiro, Av. Sa
˜o Jos´
e do Barreto, 764, Maca´
e, RJ, 27965-045,
Brazil; Email: (MMM) mincarone@macae.ufrj.br. Send reprint requests to MMM.
2
Departamento de Pesca e Aquicultura, Universidade Federal Rural de Pernambuco, Recife, Brazil.
3
Institut de Recherche pour le D´
eveloppement (IRD), MARBEC, Univ Montpellier, CNRS, Ifremer, IRD, Montpellier, France.
Submitted: 7 May 2020. Accepted: 11 November 2020. Associate Editor: M. P. Davis.
Ó2021 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/i2020069 Published online: 9 June 2021
Ichthyology & Herpetology 109, No. 2, 2021, 467–488

in this study in the Brazilian EEZ is also provided, with
references to additional records in the western South
Atlantic.
MATERIALS AND METHODS
Most specimens examined in this study are part of a large
collection of mesopelagic invertebrates and fishes sampled
during the ABRACOS expeditions (Acoustics along the
Brazilian Coast), carried out between 30 September and 20
October 2015 (ABRACOS 1–AB1; Bertrand, 2015), and
between 9 April and 6 May 2017 (ABRACOS 2–AB2;
Bertrand, 2017). Both expeditions were conducted by the
French RV Antea off northeastern Brazil and included
collectionsalongtheFernandodeNoronhaRidge(Rocas
Atoll, Fernando de Noronha Archipelago, and seamounts off
Rio Grande do Norte State). The survey comprised 82 fishing
stations, between the surface and 1,113 m depth, and
resulted in the collection of more than 9,000 fish speci-
mens. Sampling was conducted using micronekton (body
mesh 40–80 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 on the upper part of the trawl mouth. An open-
mouth net was employed, but pre-established target
(maximum) depths were defined for each trawl according
to the presence of an acoustic scattering layer or patches
detected with a Simrad EK60 split-beam scientific echo
sounder. At the target depths, trawling activity lasted for
about 30 minutes. Therefore, collection of specimens most
likely occurred at target depths, which are indicated as
capture depths in the species accounts.
Measurements and counts followed Hubbs and Lagler
(1947) with adjustments by Ebeling (1962) for the Melam-
phaidae and Paxton (1989) for the Cetomimidae. Cranial
bone nomenclature followed Kotlyar (1991). Vertebrae and
dorsal- and anal-fin ray counts were obtained through a
Faxitron LX-60 Cabinet X-ray System. Unless stated other-
wise, gill raker number refers to the total number of rakers in
the first gill arch. Species identifications were based on
descriptions and taxonomic keys provided by Goode and
Bean (1895), Parr (1934, 1946), Harry (1952), Rofen (1959),
Bigelow (1961), Ebeling (1962), Abe and Hotta (1963),
Richardson and Garrick (1964), Abe et al. (1965), Maul
(1969), Ebeling and Weed (1973), Fedorov et al. (1987),
Paxton (1989), McEachran and Fechhelm (1998), Moore
(2003), Paxton and Trnski (2003), Kotlyar (2004b, 2004c,
2005, 2008a, 2008b, 2009a, 2009b, 2009c, 2010, 2011a,
2011b, 2012a, 2012b, 2013, 2014, 2015a, 2015b, 2015c,
2016a, 2016b, 2016c, 2019, 2020), Iwasaki (2009), and
Mincarone et al. (2014). Institutional abbreviations follow
Sabaj (2020).
RESULTS
Melamphaidae
The Melamphaidae (bigscales) is the largest family in the
Stephanoberycoidei, comprising five genera and about 72
species of meso- and bathypelagic fishes (Ebeling and Weed,
1973; Kotlyar, 2004a, 2005, 2010, 2012b, 2013, 2016c).
Species of the group are reported from all oceans except the
Arctic and the Mediterranean Sea (Ebeling, 1962; Kotlyar,
2004b, 2004c, 2005, 2008a, 2008b, 2009b, 2009c, 2010,
2011a, 2011b, 2012a, 2012b, 2013, 2019; Moore, 2016;
Sutton et al., 2020). Juveniles occur in shallow oceanic
waters, whereas adults occur below 100–200 m to depths
greater than 3,000 m (Ebeling, 1962; Keene, 1987; Kotlyar,
2004a; Mincarone et al., 2014). In the Brazilian EEZ, ten
species of the Melamphaidae were previously recorded:
Melamphaes hubbsi,M. polylepis,M. typhlops,Poromitra
crassiceps,P. megalops,Poromitra sp., Scopeloberyx opisthopterus,
Scopeloberyx robustus,Scopelogadus beanii, and Scopelogadus
mizolepis (G¨
unther, 1887; Keene, 1987; Campos et al., 2008;
Costa and Mincarone, 2010; Mincarone et al., 2014; Judkins
and Haedrich, 2018). Records of 11 species of the Melam-
phaidae in the western South Atlantic are provided, four of
them new in the Brazilian EEZ.
Melamphaes eulepis Ebeling, 1962
Figure 1A, Table 1
Melamphaes eulepis Ebeling, 1962: 70 (type locality: off
Ghana, 008310S, 118020W, about 200 m depth; holotype:
ZMUC P41141).
Specimens examined.—NPM 5007, 3, 35.2–45.6 mm, RV
Antea,sta.AB2/41A,03819059.1 00 S, 32824042.1 00 Wto
03819031.8 00 S, 32825004.600 W, 430 m, 26 April 2017, 2144–
2206 h; NPM 5008, 3, 42.0–45.0 mm, sta. AB2/49A,
04810038.1 00 S, 33816007.400 Wto04810058.0 00 S, 33815003.800 W,
770–1020 m, 30 April 2017, 2117–2152 h; NPM 5009, 2,
41.5–45.1 mm (Fig. 1A), sta. AB2/53A, 03848058.700 S,
33859017.100 Wto03850005.8 00 S, 33858046.500 W, 610 m, 2
May 2017, 2208–2240 h; NPM 5224, 2, 45.5–46.8 mm, sta.
AB2/54B, 03845017.2 00 S, 34841004.000 Wto03844039.200 S,
34840004.5 00 W, 830–1030 m, 3 May 2017, 1311–1347 h.
Diagnosis.—Melamphaes eulepis differs from all congeners by
the presence of bone expansions on the head ridges. It also
differs from all congeners, except M. spinifer, by having
almost all body scales intact in preserved specimens.
Melamphaes eulepis differs from M. spinifer by the number of
pores in the angular portion of the cheek (3–4, usually 3 vs.
4–5, usually 5) and total number of vertebrae (28–30 vs. 26–
29, usually 27; Kotlyar, 2016c).
Distribution.—Melamphaes eulepis has a circumtropical distri-
bution, except for the eastern Pacific (Kotlyar, 2014). In the
Atlantic Ocean, the species is reported between 278N and 78S,
including the Gulf of Mexico and the Caribbean Sea (Ebeling,
1962; Keene, 1987; Kotlyar, 2014; Moore, 2016; Sutton et al.,
2020). In the western South Atlantic, the species was
previously known based on a single record made southeast
of Sa
˜o Pedro e Sa
˜o Paulo Archipelago (ISH 606/66—018240S,
268W; Keene, 1987). The ten specimens reported herein
therefore represent further confirmation of the occurrence of
the species in the western South Atlantic and the first records
in Brazilian waters. They were collected off the Fernando de
Noronha Archipelago, Rocas Atoll, and near the seamounts
off Rio Grande do Norte State, at depths ranging between 430
and 1,030 m (Fig. 2).
Habitat.—Melamphaes eulepis is a mesopelagic species, with
adults and juveniles occurring at a minimum depth of 150
and 200 m, whereas adults probably occur below 700 m at
night (Ebeling, 1962; Keene, 1987).
468 Ichthyology & Herpetology 109, No. 2, 2021

Fig. 1. Species of Melamphaidae reported in this study: (A) Melamphaes eulepis, NPM 5009, 45.1 mm SL; (B) Melamphaes leprus, NPM 5227, 90
mm SL; (C) Melamphaes longivelis, NPM 5229, 75.2 mm SL; (D) Melamphaes polylepis, NPM 5228, 49.0 mm SL; (E) Melamphaes typhlops, NPM
5225, 60.3 mm SL; (F) Melamphaes sp., NPM 5826, 61.9 mm SL; (G) Poromitra megalops, NPM 5632, 57.0 mm SL; (H) Poromitra sp., NPM 5331,
120.0 mm SL; (I) Scopeloberyx opercularis, NPM 5987, 32.0 mm SL; (J) Scopeloberyx opisthopterus, NPM 5985, 25.0 mm SL; (K) Scopelogadus
mizolepis, NPM 5990, 49.0 mm SL. Scale bar ¼1 cm.
Afonso et al.—Stephanoberycoidei off northeastern Brazil 469

Melamphaes leprus Ebeling, 1962
Figure 1B, Table 1
Melamphaes leprus Ebeling, 1962: 60 (type locality: north of
Ascension Island, 038450S, 108000W, about 350 m depth;
holotype: ZMUC P41172).
Specimen examined.—NPM 5227, 1, 90 mm (Fig. 1B), RV
Antea,sta.AB2/41A,03819059.1 00 S, 32824042.1 00 Wto
03819031.8 00 S, 32825004.600 W, 430 m, 26 April 2017, 2144–
2206 h.
Diagnosis.—Melamphaes leprus differs from all congeners,
except M. falsidicus,M. macrocephalus,M. pachystomus, and
M. polylepis, by having 20 or more gill rakers (rarely 19),
width near the median region of the larger gill rakers
approximately equal to the space between rakers, I, 7
pelvic-fin rays, posttemporal spine absent, and presence
simultaneously of 14–15 rays in the pectoral fin and 11
precaudal vertebrae. Melamphaes leprus differs from M.
macrocephalus by the number of transverse series of scales
(31–33 vs. 25–28) and by the relative position of the pelvic
and pectoral fins (pelvic fin originates after pectoral-fin
origin vs. pelvic-fin origin is anterior to pectoral-fin origin). It
differs from M. falsidicus and M. polylepis by the absence of
spurs on the haemal arch of the first caudal vertebra (vs.
spurs present), and from M. pachystomus by the anal-fin
origin (in line with the third or fourth dorsal-fin ray,
counting from the last ray vs. posterior to the last dorsal-fin
Table 1. Morphometric and meristic data of species of Melamphaes reported in this study.
Species M. eulepis M. leprus M. longivelis M. polylepis M. typhlops Melamphaes sp.
n10 1 2 37 7 1
Standard length (SL, mm) 35.2–46.8 90.0 73.8–75.2 36–70.2 37.3–71.0 61.9
Measurements in % SL
Head length 36.9–40.0 35.6 36.0–36.7 31.6–39.9 33.7–35.1 34.9
Head width 18.3–21.1 17.1 16.9 14.5–17.5 15.3–17.5 16.2
Eye diameter 5.3–6.0 4.8 6.2–6.3 3.9–5.6 4.1–5.5 4.8
Postorbital length 23.2–26.7 22.3 23.5–24.7 20.0–25.1 22.4–22.8 22.9
Snout length 9.1–10.3 8.6 8.4–9.4 6.9–9.3 7.5–8.6 8.1
Upper jaw length 17.2–18.8 17.0 18.0–18.1 14.5–17.6 16.6–18.2 16.5
Body depth 27.7–31.5 24.4 25.9 21.3–27.6 23.0–26.1 25.8
Prepectoral length 34.7–40.7 36.8 37.2–38.3 32.0–36.9 33.6–35.7 35.1
Prepelvic length 34.4–43.1 37.7 38.6–39.9 34.7–39.7 36.6–39.5 —
Predorsal length 43.7–49.9 43.1 41.3–42.8 40.6–45.4 39.9–42.6 43.3
Preanal length 62.5–73.8 63.9 71.8–74.8 60.6–71.1 70.0–72.6 67.9
Dorsal-fin base length 25.6–31.6 30.6 29.4–30.6 22.9–29.0 23.5–28.7 26.2
Anal-fin base length 7.7–11.1 11.1 8.9–10.2 8.0–11.4 7.3–10.0 9.7
Caudal peduncle length 22.7–25.5 23.1 21.4–23.0 21.3–30.6 17.9–24.1 26.3
Caudal peduncle depth 9.3–10.5 9.6 9.3–9.5 7.8–10.1 7.9–9.7 9.7
Counts
Gill rakers (upper þangle and lower) 4þ13–14 6þ16 4þ14 5–6þ15–17 4þ11–13 5þ15
Gill rakers (total) 17–18 22 18 20–23 16–17 20
Dorsal-fin rays III,14–16 III,15 III,17–18 III,13–15 III,15 III,14
Anal-fin rays I,8 I,8 I,8 I,7–8 I,8 I,8
Pectoral-fin rays 15 15 15 14–15 15 15
Pelvic-fin rays I,7 I,7 I,7 I,7 I,7 I,8
Principal caudal rays (upper/lower) 8–10/8–9 10/9 9/8–9 8–11/7–10 8–10/8–9 10/9
Procurrent caudal rays (upper/lower) 3–4/3 4/4 5/3 — — —
Vertebrae (precaudal þcaudal) 12–13þ16–18 11þ16 12þ17 11þ17–19 12–13þ14–16 11þ18
Vertebrae (total) 28–30 27 29 28–30 26–28 29
Fig. 2. Records of Melamphaes eulepis (triangle), M. leprus (circle), M.
longivelis (pentagon), M. polylepis (star), M. typhlops (square), and
Melamphaes sp. (diamond) off northeastern Brazil collected during the
ABRACOS surveys. Tip of arrow indicates same collection locality for
different species. FN–Fernando de Noronha Archipelago; PB–Para´
ıba;
PE–Pernambuco; RA–Rocas Atoll; RN–Rio Grande do Norte.
470 Ichthyology & Herpetology 109, No. 2, 2021

ray) and by the number of vertebrae (27 vs. 28–29; Kotlyar,
2011b, 2012b).
Distribution.—Ebeling (1962) described Melamphaes leprus
based on ten specimens from the eastern Tropical Atlantic,
collected between 118N and 48S. Subsequently, Keene (1987)
reported a wider distribution for the species in the Atlantic,
from 178Nto138S, and from 298Wto118E, with only three
records in the western South Atlantic. Kotlyar (2011b) also
reported on a single specimen from the Gulf of Guinea,
eastern Atlantic. The single specimen reported herein was
collected off the Fernando de Noronha Archipelago, at 430 m
depth, and represents the fourth record of the species in the
western South Atlantic and the first record in Brazilian waters
(Fig. 2).
Habitat.—Melamphaes leprus is a meso- to bathypelagic
species, with juveniles and half-grown specimens captured
at the upper limit of the mesopelagic zone, between 150 and
300 m at night (Ebeling, 1962). One adult specimen was
captured in bottom trawling at 1,550 m depth (Kotlyar,
2011b).
Melamphaes longivelis Parr, 1933
Figure 1C, Table 1
Melamphaes microps longivelis Parr, 1933: 16 (type locality: off
Acklins Island, Bahama, western Atlantic, 228310N,
748260W, 10,000 feet [3048 m] wire out; holotype: YPM
2833).
Specimens examined.—NPM 5229, 1, 75.2 mm (Fig. 1C), RV
Antea,sta.AB2/42A,03815028.100 S, 31848029.1 00 Wto
03815027.8 00 S, 31850040.600 W, 780 m, 27 April 2017, 1223–
1226 h; NPM 5230, 1, 73.8 mm, sta. AB2/35, 04819036.6 00 S,
35829051.6 00 Wto04818 032.400 S, 35832019.8 00 W, 630 m, 20
April 2017, 2235–2315 h.
Diagnosis.—Melamphaes longivelis differs from all congeners,
except M. eulepis and M. spinifer, by having 17–19 (more often
18) gill rakers, width near the median region of the larger gill
rakers less than three-quarters of the space between the
rakers, eye diameter equal to or larger than the suborbital
bone width, anal-fin origin in line with or posterior to the
last dorsal-fin ray origin (less than the width of one scale
pocket), caudal peduncle depth substantially more than two
times in the caudal peduncle length, and III,17–18 dorsal-fin
rays. Melamphaes longivelis differs from M. eulepis and M.
spinifer by having less than half (rarely more) body scales
present in preserved specimens (vs. all, or almost all, body
scales present in preserved specimens; Kotlyar, 2015a,
2016c).
Distribution.—Melamphaes longivelis occurs in the Atlantic
Ocean, with confirmed records restricted to the eastern and
western North Atlantic (Kotlyar, 2015a; Sutton et al., 2020)
and the eastern South Atlantic (Sutton et al., 2020). Keene
(1987) reported two specimens of M. longivelis in the western
South Atlantic off southeastern Sa
˜o Pedro e Sa
˜oPaulo
Archipelago, but his report was made before Kotlyar’s
(2015a) revision of the species complex. Therefore, the two
specimens collected off the Fernando de Noronha Archipel-
ago and off Rio Grande do Norte State, between depths of
630 and 780 m (Fig. 2) represent the first confirmed records
of M. longivelis in the western South Atlantic. Records of the
species in the region prior to Kotlyar’s (2015a) revision (e.g.,
Ebeling, 1962; Moore, 2003) require confirmation.
Habitat.—Melamphaes longivelis is a mesopelagic species, with
juveniles recorded in depths shallower than 150 m and
adults below 500 m (Ebeling, 1962; Keene, 1987; Kotlyar,
2015a).
Melamphaes polylepis Ebeling, 1962
Figure 1D, Table 1
Melamphaes polylepis Ebeling, 1962: 43 (type locality: South
of Sri Lanka, 058210N, 808380E, about 2250 m depth;
holotype: ZMUC P41178).
Specimens examined.—NPM 5228, 2, 49.0–60.5 mm (Fig. 1D),
RV Antea,sta.AB2/42A,03815028.100 S, 31848029.1 00 Wto
03815027.8 00 S, 31850040.600 W, 780 m, 27 April 2017, 1223–
1226 h; NPM 5231, 2, 46.3–57.6 mm, sta. AB2/60B,
03831043.0 00 S, 36821019.800 Wto03831046.8 00 S, 36822025.700 W,
670–700 m, 6 May 2017, 1249–1319 h; NPM 5233, 4, 59.0–
67.3 mm, sta. AB2/52A, 03843016.200 S, 33825009.8 00 Wto
03842014.200 S, 33824036.2 00 W, 822–984 m, 2 May 2017,
1147–1218 h; NPM 5234, 2, 52.6–56.8 mm, sta. AB2/16,
07836015.0 00 S, 33859030.000 Wto07836049.3 00 S, 33857018.700 W,
680 m, 14 April 2017, 2153–2239 h; NPM 5237, 8, 60.5–69.0
mm,sta.AB2/44A,03852052.5 00 S, 32817033.300 Wto
03852013.4 00 S, 32816028.000 W, 850 m, 28 April 2017, 1244–
1317 h, 1244–1317 h; NPM 5238, 4, 64.1–66.4 mm, sta. AB2/
53A, 03848058.7 00 S, 33859017.1 00 Wto03850005.8 00 S,
33858046.5 00 W, 610 m, 2 May 2017, 2208–2240 h; NPM
5239, 2, 36.0–66.0 mm, sta. AB2/49A, 04810038.100 S,
33816007.4 00 Wto04810058.000 S, 33815003.8 00 W, 770–1020 m,
30 April 2017, 2117–2152 h; NPM 5241, 2, 37.0–60.6 mm,
sta. AB2/54B, 03845017.2 00 S, 34841004.000 Wto03844039.2 00 S,
34840004.5 00 W, 830–1030 m, 3 May 2017, 1311–1347 h; NPM
5242, 11, 58.2–70.2 mm, sta. AB2/39, 04852026.9 00 S,
34835022.9 00 Wto04850052.800 S, 34851004.700 W, 650–800 m,
24 April 2017, 2149–2237 h.
Diagnosis.—Melamphaes polylepis differs from all congeners,
except M. falsidicus,M. macrocephalus,M. pachystomus, and
M. leprus, by having 20 or more (rarely 19) gill rakers, width
near the median region of the larger gill rakers approximately
equal to the space between rakers, I,7 pelvic-fin rays,
posttemporal spine absent, and presence simultaneously of
14–15 pectoral-fin rays and 11 precaudal vertebrae. Melam-
phaes polylepis differs from M. falsidicus by the number of
transverse series of scales (30–36 vs. 29–30) and the eye
diameter (10.0–16.3% HL vs. 16.4–19.2% HL). It differs from
M. leprus,M. macrocephalus, and M. pachystomus by the
presence of spurs on the haemal arch of the first caudal
vertebra (vs. spurs absent; Kotlyar, 2011b, 2012b).
Distribution.—Melamphaes polylepis has a circumglobal distri-
bution, except for the eastern Pacific Ocean (Kotlyar, 2011b).
It was originally reported from the North Atlantic between
the equator and 208N, the Indian Ocean and Indonesia
between 158N and 158S, and the North Pacific between 348N
and 68N (Ebeling, 1962). Ebeling (1962) also reported the
species from the South Pacific, at 308560S, 109817 0W, based
on a single specimen that might actually represent M.
pachystomus (Kotlyar, 2011b). Keene (1987) recorded M.
Afonso et al.—Stephanoberycoidei off northeastern Brazil 471

polylepis from 328Nto188S in the Atlantic Ocean, including
the southeastern Caribbean Sea and the western South
Atlantic, from eastern Sa
˜o Pedro e Sa
˜o Paulo Archipelago to
northern Trindade Island (ISH 484/66, 178360S, 288530W), off
the central Brazilian coast. Further records of the species in
Brazil were also made off Bahia and Esp´
ırito Santo States
based on two specimens collected between depths of 837 and
1,051 m (Mincarone et al., 2014), and off northeastern Brazil
(Eduardo et al., 2020a). The 37 specimens examined here
were collected off Pernambuco State, the Fernando de
Noronha Archipelago, the Rocas Atoll, and the seamounts
off Rio Grande do Norte State, between depths of 610 and
1,030 m (Fig. 2).
Habitat.—Melamphaes polylepis is a meso- and bathypelagic
species, with specimens captured by open-mouth nets at
depths down to 4,228 m (Ebeling, 1962; Keene, 1987; Keene
et al., 1987; Kotlyar, 2011b). According to Ebeling (1962),
juveniles and subadults probably occur at depths below 200–
300 m. Keene (1987) reported on vertical migration of
juveniles between 500 and 800 m during the day, and
mainly between 100 and 400 m at night.
Remarks.—In his unpublished Ph.D. dissertation, Keene
(1987) described ‘‘Melamphaes indicoides’’ based on specimens
collected in the Atlantic. The species, which was never
formally described and is, therefore, not valid, is morpho-
logically similar to M. polylepis, differing by the number of
diagonal series of scales (8 vs. 9–10, respectively). Subse-
quently, Bartow (2010), based on the examination of only
five specimens, proposed that ‘‘Melamphaes indicoides’’ and
M. polylepis might also differ by the following characters
(numbers in brackets refer to mode values for counts or mean
values for measurements): number of dorsal- (III,15–16
[III,15] vs. III,13–15 [III,14]) and caudal-fin rays (25–27 [25]
vs. 27–29 [28]), number of gill rakers on the first gill arch (20
vs. 20–23 [21]), number of scale in horizontal series (25–31
[28] vs. 33–35 [34]), number of scales in diagonal row (5–8 [7]
vs. 8–10 [9]), HL (29.76–32.44% [30.92%] SL vs. 35.0–41.4%
[37.5%] SL), distance between the end of dorsal fin to caudal-
fin origin (27.38–32.83% [30.37%] SL vs. 33.1–36.6% [35.0%]
SL), postanal length (25.56–30.77% [28.43%] SL vs. 35.6–
41.3% [38.1%] SL), orbit to cheek angle length (10.77–
13.75% [12.68%] SL vs. 9.6–11.7% [10.8%] SL), and caudal
peduncle length (19.72–23.21% [21.21%] SL vs. 26.6–30.7%
[28.3%] SL). The specimens recognized as M. polylepis in the
current study are partially damaged but might represent the
same species provisionally named by Keene (1987) as
‘‘Melamphaes indicoides.’’ More in-depth taxonomic studies
in the M. polylepis species complex are necessary.
Melamphaes typhlops (Lowe, 1843)
Figure 1E, Table 1
Metopias typhlops Lowe, 1843: 90 (type locality: off Madeira;
neotype: BMNH 1864.11.8.1 [designated by Ebeling,
1962]).
Specimens examined.—NPM 5225, 1, 60.3 mm (Fig. 1E), RV
Antea,sta.AB2/54B,03845017.200 S, 34841 004.000 Wto
03844039.200 S, 34840004.5 00 W, 830–1030 m, 3 May 2017,
1311–1347 h; NPM 5226, 1, 37.3 mm, sta. AB2/41A,
03819059.1 00 S, 32824042.100 Wto03819031.8 00 S, 32825004.600 W,
430 m, 26 April 2017, 2144–2206 h; NPM 5232, 1, 65.7 mm,
sta. AB2/52A, 03843016.2 00 S, 33825009.800 Wto03842014.200 S,
33824036.2 00 W, 822–984 m, 2 May 2017, 1147–1218 h; NPM
5235, 1, 61.6 mm, sta. AB2/16, 07836015.0 00 S, 33859030.000 W
to 07836049.3 00 S, 33857018.700 W, 680 m, 14 April 2017, 2153–
2239 h; NPM 5236, 1, 68.2 mm, sta. AB2/44A, 03852052.500 S,
32817033.3 00 Wto03852013.400 S, 32816028.000 W, 850 m, 28
April 2017, 1244–1317 h; NPM 5240, 2, 67.5–71.0 mm, sta.
AB2/39, 04852026.900 S, 34803032.3 00 Wto04850052.800 S,
34805006.5 00 W, 650–800 m, 24 April 2017, 2149–2237 h.
Diagnosis.—Melamphaes typhlops differs from congeners,
except M. contradictorius,M. eurous,M. inconspicuus,M.
indicus,M. janae,M. kobylyanskyi,M. longivelis,M. parvus,M.
proximus, and M. succedaneus, by having 19 or fewer (rarely
20) gill rakers, width near the median region of the larger gill
rakers less than three-quarters of the space between rakers,
eye diameter equal to or larger than suborbital bone width,
anal-fin origin posterior to the vertical through the last
dorsal-fin ray origin, and less than half of body with scales
present in preserved specimens (rarely more). Melamphaes
typhlops differs from M. contradictorius,M. inconspicuus,M.
janae,M. kobylyanskyi,M. longivelis,M. parvus,M. proximus,
and M. succedaneus by the distance between anal-fin origin
and the vertical through the last dorsal-fin ray origin (usually
equal to the width of one to one and a half scale pocket vs.
less than the width of one scale pocket) and number of gill
rakers on the lower portion of the first gill arch (10–11 vs. 12–
14 [a single specimen with 12 in the present study]). It differs
from M. eurous and M. indicus by having gill rakers of the
fourth branchial arch present as reduced, flat or slightly
convex, patches (vs. gill rakers of the fourth branchial arch in
the shape of short knobs; Kotlyar, 2016a, 2016c; this study).
Distribution.—Melamphaes typhlops occurs in the Atlantic
Ocean, from 418Nto288S, including the Gulf of Mexico
(Ebeling, 1962; Keene, 1987; Keene et al., 1987; Kotlyar,
2016a; Moore, 2016; Sutton et al., 2020). In the western
South Atlantic, the species was previously known from 22
specimens collected off southern Sa
˜o Pedro e Sa
˜o Paulo
Archipelago and off the central to southeastern Brazilian
coast, with some records inside the country’s EEZ (Keene,
1987; Mincarone et al., 2014). Seven specimens collected
between depths of 430 and 1,030 m off the Fernando de
Noronha Archipelago, the Rocas Atoll, the seamounts off Rio
Grande do Norte State, and off Rio Grande do Norte and
Pernambuco States are reported here (Fig. 2).
Habitat.—Melamphaes typhlops is a meso- to bathypelagic
species, with post-larvae captured between the surface and
down to 100 m, juveniles between depths of 150 and 1,000
m, and adults below 500 m (Ebeling, 1962; Keene et al. 1987;
Kotlyar, 2016a).
Melamphaes sp.
Figure 1F, Table 1
Specimen examined.—NPM 5826, 1, 61.9 mm (Fig. 1F), RV
Antea,sta.AB2/54B,03845017.200 S, 34841004.0 00 Wto
03844039.200 S, 34840004.5 00 W, 830–1030 m, 3 May 2017,
1311–1347 h.
Diagnosis.—The only specimen identified here as Melamphaes
sp. differs from all other species of the genus, except M.
ebelingi,M. nikolayi, and M. occlusus, by the number of pelvic-
472 Ichthyology & Herpetology 109, No. 2, 2021

fin rays (I,8 vs. I,7). Melamphaes sp. differs from M. ebelingi by
the number of vertebrae (29 vs. 26–27) and by the presence
of spurs on the haemal arch of the first caudal vertebra (vs.
spurs absent). It differs from M. occlusus by the number of gill
rakers (20 vs. 22) and by the number of dorsal-fin rays (III,14
vs. III,16), and from M. nikolayi by the presence of spurs on
the haemal arch of the first caudal vertebra (vs. spurs absent)
and by the number of precaudal vertebrae (11 vs. 12; Ebeling,
1962; Keene, 1973; Bartow, 2010; Kotlyar, 2012b). Kotlyar
(2015c) reported on the occurrence of one spine and eight
soft rays in one side of the pelvic fin of a single specimen of
M. lentiginosus (typical condition: I,7 pelvic-fin rays), but the
specimen reported in the present study differs from M.
lentiginosus by the number of gill rakers (20 vs. 15–17; Table
1).
Distribution.—Known only from a single specimen, collected
near the seamounts off Rio Grande do Norte State, between
depths of 830 and 1,030 m (Fig. 2).
Remarks.—This specimen most likely belongs to a new
species, which will be described in a forthcoming study.
Poromitra megalops (L¨
utken, 1878)
Figure 1G, Table 2
Melamphaes megalops L¨
utken, 1878: 176 (type-locality: south
of Azores, eastern North Atlantic [stomach content];
holotype: ZMUC 84).
Specimens examined.—NPM 5632, 9, 32.5–57.0 mm (Fig. 1G),
RV Antea,sta.AB2/53A,03848058.700 S, 33859017.1 00 Wto
03850005.8 00 S, 33858046.500 W, 610 m, 2 May 2017, 2208–
2240 h; NPM 5927, 2, 46.5–53.0 mm, sta. AB2/52A,
03843016.2 00 S, 33825009.800 Wto03842014.2 00 S, 33824036.200 W,
822–984 m, 2 May 2017, 1147–1218 h; NPM 5928, 1, 53.0
mm,sta.AB2/49A,04810038.1 00 S, 33816007.400 Wto
04810058.0 00 S, 33815003.800 W, 770–1020 m, 30 April 2017,
2117–2152 h; NPM 5929, 1, 54.5 mm, sta. AB2/39,
04852026.9 00 S, 34803032.300 Wto04850052.8 00 S, 34805006.500 W,
650–800 m, 24 April 2017, 2149–2237 h; NPM 5931, 2, 33.0–
42.0 mm, sta. AB2/35, 04819036.600 S, 35829051.6 00 Wto
04818032.4 00 S, 35832019.800 W, 630 m, 20 April 2017, 2235–
2315 h; NPM 5933, 5, 35.0–59.0 mm, sta. AB2/44A,
03852052.5 00 S, 32817033.300 Wto03851043.6 00 S, 32816020.000 W,
850 m, 28 April 2017, 1244–1317 h; NPM 5937, 1, 34.0 mm,
sta. AB2/59A, 03838001.6 00 S, 36803010.600 Wto03838007.900 S,
36802022.6 00 W, 700–1113 m, 5 May 2017, 2157–2237 h; NPM
5938, 6, 47.0–59.0 mm, sta. AB2/54B, 03845017.2 00 S,
34841004.0 00 Wto03844039.200 S, 34840004.5 00 W, 830–1030 m,
3 May 2017, 1311–1347 h; NPM 6089, 1, 25.0 mm, sta. AB1/
22, 04807044.8 00 S, 33847024.500 Wto04807 000.700 S,
33848057.9 00 W, 525 m, 8 October 2015, 2132–2212 h.
Diagnosis.—Poromitra megalops differs from all congeners,
except P. jucunda and P. macrophthalma, by the eye diameter
(2.9–3.2 in HL vs. 4.0–17.2 in HL; Kotlyar, 2010). According
to Kotlyar (2010), P. megalops differs from P. macrophthalma
by the number of gill rakers (26–28 vs. 21–24), in addition to
Table 2. Morphometric and meristic data of species of Poromitra,Scopeloberyx, and Scopelogadus reported in this study.
Species
Poromitra
megalops
Poromitra
sp.
Scopeloberyx
opercularis
Scopeloberyx
opisthopterus
Scopelogadus
mizolepis
n28 27 1 4 19
Standard length (SL, mm) 25.0–59.0 48.0–121.0 32.0 25.0–32.0 37.0–70.0
Measurements in % SL
Head length 32.9–39.7 39.3–44.8 39.0 29.0 34.8–38.6
Head width 11.7–14.5 12.8–16.1 — — —
Eye diameter 8.8–12.7 4.3–6.8 6.3 — —
Postorbital length 17.3–23.0 23.7–28.3 — — —
Snout length 4.5–6.9 8.8–11.3 — — 8.0–10.7
Upper jaw length 13.2–18.3 16.4–19.2 — — —
Body depth 22.3–26.7 20.7–26.3 — — 20.4–22.9
Prepectoral length 35.6–41.3 39.6–47.2 43.8 — 35.2–43.0
Prepelvic length 32.0–36.3 39.8–44.5 — — 36.9–39.8
Predorsal length 42.0–46.3 44.7–50.8 — — 41.4–44.8
Preanal length 55.1–62.7 61.9–72.8 — — 57.6–60.2
Dorsal-fin base length 20.8–25.0 23.6–28.9 — — 17.6–20.2
Anal-fin base length 8.4–11.8 8.7–12.2 — — 9.0–11.1
Caudal peduncle length 28.9–34.4 19.4–25.3 22.8 — 29.1–34.9
Caudal peduncle depth 5.8–7.6 7.7–10.0 — — 8.1–9.4
Counts
Gill rakers (upper þangle and lower) 6–8þ16–20 9–11þ21–23 18þ8 3–4þ11–13 7–8þ15–17
Gill rakers (total) 23–27 30–34 26 14–17 22–25
Dorsal-fin rays II–III,11–12 III,11–12 — — II,11
Anal-fin rays I,9 I,8 I,7 — I,8
Pectoral-fin rays 12–14 14–15 — — 13–14
Pelvic-fin rays I,7 I,7 — I,7–8 —
Principal caudal rays (upper/lower) 10/8–10 9–10/9–10 — — —
Procurrent caudal rays (upper/lower) 3/3 3/3 — — —
Vertebrae (precaudal þcaudal) 10þ19–20 10–11þ15–17 — — —
Vertebrae (total) 29–30 26–27 26 25 24–25
Afonso et al.—Stephanoberycoidei off northeastern Brazil 473

number of pyloric caeca (6 vs. 4–5), number of spines on the
posterior margin of the preopercle (2–15 vs. 0–1), and width
of the angular region of the preopercle (11.6–13.4% HL vs. 7–
12% HL), and from P. jucunda also by the number of gill
rakers (26–28 vs. 23–25; but see Remarks, below).
Distribution.—Poromitra megalops has a circumtropical distri-
bution, being more common in the eastern Atlantic, Indo-
Pacific and eastern Central Pacific (Ebeling and Weed, 1973;
Keene, 1987; Moore, 2016; Sutton et al., 2020). Previous
records in the western South Atlantic were restricted to nine
specimens collected off southeastern Sa
˜o Pedro e Sa
˜o Paulo
Archipelago and by one further isolated record made at
328490S, 268260W (Keene, 1987; Judkins and Haedrich, 2018).
The species was also recorded off Ascension Island, middle
Atlantic, based on two specimens (Keene, 1987). Ebeling and
Weed (1973) reported intraspecific variation between popu-
lations from the Atlantic and eastern Central Pacific and
those of the Indo-Pacific. Kotlyar (2010), however, proposed
that those different populations should be recognized as
distinct species, restricting P. megalops to the Atlantic, except
the western South Atlantic (see Remarks, below). In Brazilian
waters, the species was previously reported off Sa
˜o Pedro e
Sa
˜o Paulo Archipelago (018200S, 278370W; 018440S, 278440W;
Keene, 1987; Judkins and Haedrich, 2018). In the current
study, P. megalops is reported from 28 specimens collected off
the Fernando de Noronha Archipelago, the Rocas Atoll, and
from the seamounts off Rio Grande do Norte State, between
depths of 525 and 1,113 m. This also represents the largest
single collection of P. megalops in the western South Atlantic
made to date (Fig. 3).
Habitat.—The species is meso- to bathypelagic, with adults
usually occurring below depths of 400–500 m (maximum
depth 1,113 m; this study). Juveniles and post-larvae occur in
shallow waters down to 150–200 m (Ebeling and Weed, 1973;
Keene, 1987; Keene et al., 1987).
Remarks.—Poromitra macrophthalma was recognized as valid
until recently, when Ebeling and Weed (1973) proposed that
the species is a junior synonym of P. megalops. According to
them, specimens previously assigned to P. macrophthalma
would represent a different morphotype of P. megalops
restricted to the Indo-Pacific, only slightly distinct from the
Atlantic and middle to eastern Pacific remaining population
of the species in the number of anal-fin rays (I,8–10 vs. I,9–
10), number of gill rakers in the lower portion of the first gill
arch (14–18 vs. 16–20), number of vertebrae (26–28 vs. 28–
30), and number of dorsal-fin spines (usually II vs. usually
III). Subsequently, Kotlyar (2010) revalidated P. macrophthal-
ma, restricting its distribution to the Indo-Pacific. He
furthermore restricted the distribution of P. megalops to the
Atlantic and described P. jucunda from the Central and
eastern Pacific. According to Kotlyar (2010), P. megalops
differs from P. jucunda by the eye diameter (31.3–34.0% HL
vs. 24.2–32.6% HL) and number of gill rakers (26–28 vs. 23–
25). However, eye diameter of some specimens of P. megalops
from the Atlantic examined by Keene (1987) is also around
20% HL, with number of gill rakers ranging from 22 to 28.
The eye diameter and gill raker counts of some specimens
identified herein as P. megalops also fall within the range
proposed by Kotlyar (2010) for P. jucunda (eye diameter 24.3–
37.5% HL and number of gill rakers on the first branchial
arch 23–27; Table 2). Separation between P. megalops and P.
macrophthalma sensu Kotlyar (2010) is also problematic.
According to Kotlyar (2010), P. megalops differs from P.
macrophthalma by the following characters: number of gill
rakers (26–28 vs. 21–24), number of pyloric caeca (6 vs. 4–5),
number of spines on the posterior margin of preopercle (2–15
vs. 0–1), and width of the angular region of the preopercle
(11.6–13.4% HL vs. 7–12% HL). However, specimens identi-
fied here as P. m e g a l o p s have 23–27 gill rakers, 1–5
inconspicuous spines on the posterior margin of the
preopercle, and width of the angular region of the preopercle
from 10 to 16.5% HL (Table 2). Summing up, data available
from specimens identified here as P. megalops, in association
with information provided by Keene (1987) and Kotlyar
(2010), suggest that P. macrophthalma and P. jucunda might
not be valid. If correct, the situation would be similar to the
one proposed by Ebeling and Weed (1973), in which a single
species of the group (P. megalops, the senior synonym), with a
circumglobal distribution, should be recognized. However, a
more thorough taxonomic study of this group of species is
necessary, based on examination of more specimens from the
Atlantic and Indo-Pacific Oceans.
Poromitra sp.
Figure 1H, Table 2
Specimens examined.—NPM 3189, 1, 110.0 mm, RVAntea, sta.
AB1/14, 03858057.400 S, 34803023.1 00 Wto03857043.500 S,
34804050.5 00 W, 510 m, 6 October 2015, 2140–2226 h; NPM
3190, 4, 59.5–72.0 mm, sta. AB1/22, 04807044.800 S,
33847024.500 Wto04807000.7 00 S, 33848057.900 W, 525 m, 8
October 2015, 2132–2212 h; NPM 3198, 1, 48.0 mm, sta.
AB1/51, 08856029.500 S, 34829003.5 00 Wto08859005.600 S,
34828035.2 00 W, 45–200 m, 19 October 2015, 2209–2335 h;
Fig. 3. Records of Poromitra megalops (circle) and Poromitra sp.
(square) off northeastern Brazil collected during the ABRACOS surveys.
Tip of arrow indicates same collection locality for both species. FN–
Fernando de Noronha Archipelago; PB–Para´
ıba; PE–Pernambuco; RA–
Rocas Atoll; RN–Rio Grande do Norte.
474 Ichthyology & Herpetology 109, No. 2, 2021

NPM 5331, 3, 100.0–120.0 mm (Fig. 1H), sta. AB2/53A,
03848058.7 00 S, 33859017.100 Wto3850005.800 S, 33858046.500 W,
610 m, 2 May 2017, 2208–2240 h; NPM 5926, 1, 59.0 mm,
sta. AB2/16, 07836014.4 00 S, 33859033.800 Wto07836049.300 S,
33857018.7 00 W, 680 m, 14 April 2017, 2153–2239 h; NPM
5930, 2, 65.0–111.0 mm, sta. AB2/35, 04819036.600 S,
35829051.6 00 Wto04818 032.400 S, 35832019.8 00 W, 630 m, 20
April 2017, 2235–2315 h; NPM 5932, 2, 51.5–66.5 mm, sta.
AB2/39, 04852026.900 S, 34803032.300 Wto04850052.800 S,
34805006.5 00 W, 650–800 m, 24 April 2017, 2149–2237 h;
NPM 5934, 6, 81.0–121.0 mm, sta. AB2/41A, 03819059.1 00 S,
32824042.1 00 Wto03819 031.800 S, 32825004.6 00 W, 430 m, 26
April 2017, 2144–2206 h; NPM 5935, 1, 62.0 mm, sta. AB2/
44A, 03852052.500 S, 32817033.300 Wto03851043.600 S,
32816020.0 00 W, 850 m, 28 April 2017, 1244–1317 h; NPM
5936, 1, 62.0 mm, sta. AB2/59A, 03838001.6 00 S, 36803010.600 W
to 03838007.9 00 S, 36802022.600 W, 700–1113 m, 5 May 2017,
2157–2237 h; NPM 5939, 5, 62.0–85.0 mm, sta. AB2/54B,
03845017.2 00 S, 34841004.000 Wto03844039.2 00 S, 34840004.5 00 W,
830–1030 m, 3 May 2017, 1311–1347 h.
Diagnosis.—Kotlyar (2008a) defined five species groups of
Poromitra based on the anatomy of the preopercle. The
specimens of Poromitra sp. examined here are more similar to
those of the P. crassa and P. crassiceps species groups (Kotlyar,
2008a). However, Poromitra sp. differs from P. crassa (the
single species in the P. crassa species group) by the number of
dorsal-fin rays (III,11–12 vs. III,10), number of gill rakers (30–
34 vs. 23–25), and body depth (20.7–26.3% SL vs. 31.5–
34.2% SL; Kotlyar, 2008a). In the P. crassiceps species group,
Poromitra sp. differs from P. coronata by the number of dorsal-
fin rays (III,11–12 vs. III,10), from P. rugosa and P. decipiens by
the relative position of pelvic and pectoral fins (pelvic-fin
origin is beneath or slightly anterior to pectoral-fin origin vs.
pelvic fin originates after pectoral-fin origin), and from P.
curilensis by the number of dorsal-fin rays (III,11–12 vs.
III,12–14), anal-fin origin (in line with the second to fifth
dorsal-fin ray origins vs. in line with the sixth to seventh
dorsal-fin ray origins, counting from the last ray), and
extension of the posterior margin of the upper jaw
(extending beyond the posterior margin of the eye vs. in
line with the posterior margin of the eye). See Remarks for
further details.
Distribution.—The 27 specimens identified in this study as
Poromitra sp. were collected off the Fernando de Noronha
Archipelago, Rocas Atoll, seamounts off Rio Grande do Norte
State, and off Rio Grande do Norte and Pernambuco States,
between depths of 45 and 1,113 m (Fig. 3). The species was
previously reported off Esp´
ırito Santo and Rio de Janeiro
States, southeastern Brazil, at depths between 837 and 1,762
m (Mincarone et al., 2014).
Remarks.—The specimens identified here as Poromitra sp.
represent the same species also recognized as Poromitra sp. by
Mincarone et al. (2014). Those authors concluded that
Poromitra sp. belongs to the P. crassiceps group of Kotlyar
(2008a, 2008b), and this conclusion is supported in the
present study based on the new specimens examined.
According to Kotlyar (2008a, 2008b), only two species of
the Poromitra crassiceps group occur in the western Atlantic
Ocean, P. crassiceps and P. kukuevi.Poromitra crassiceps differs
from Poromitra sp. by the number of dorsal- (III,12–13 vs.
III,11–12 [a single specimen with 12]) and anal-fin rays (I,9–
10 vs. I,8), number of vertebrae (27–29 vs. 26–27), and
relative position of pelvic and pectoral fins (pelvic fin
originates after pectoral-fin origin vs. pelvic-fin origin is
beneath or slightly anterior to pectoral-fin origin). Despite
similarities in terms of shape of the preopercle, number and
presence of spines in the preopercle, and counts of pectoral-,
pelvic-, dorsal-, and anal-fin rays, P. kukuevi (which was
described based on a single and possibly juvenile specimen)
differs substantially from Poromitra sp. in the number of gill
rakers (26 vs. 30–34, respectively). Measures and counts of 12
specimens identified as Poromitra sp. by Mincarone et al.
(2014) are in accordance with those presented here, except
by the number of vertebrae (26 vs. 26–27, respectively; Table
2). This variation is interesting, since the single known
specimen of P. kukuevi also has 27 vertebrae (Kotlyar, 2008b).
Mincarone et al. (2014) indicated that the taxonomic
situation of at least part of the Poromitra crassiceps group is
complex, concluding, among other things, that P. kukuevi
might be a junior synonym of P. indooceanica, which has
priority over the former by six printed pages (Kotlyar, 2008b).
This situation renders the proper identification of Poromitra
sp. as even more problematic. Bartow (2010) also noted the
current taxonomic complexity of the genus Poromitra in the
Atlantic, especially of the species included in the Poromitra
crassiceps group. Keene (1987), in his unpublished Ph.D.
dissertation, informally described a distinct Atlantic species
of the genus and provisionally named it as ‘‘Poromitra gibbsi’’
(not Poromitra gibbsi Parin and Borodulina, 1989, which is a
valid and distinct species), with records along the western
South Atlantic, including off Brazil. Measurements and
counts of ‘‘Poromitra gibbsi’’ sensu Keene (1987), such as
number of vertebrae (25–27) and dorsal-fin rays (III,10–12),
are similar to those reported for P. glochidiata,P. indooceanica,
P. kukuevi, and P. unicornis, all of them included in the
Poromitra crassiceps species group of Kotlyar (2008a). Meristic
and morphometric data of the specimens identified here as
Poromitra sp. also fall within the ranges and description
provided by Keene (1987) for his ‘‘Poromitra gibbsi.’’ There-
fore, it is possible that the species recognized by Kotlyar
(2008a) as P. kukuevi, in addition to ‘‘Poromitra gibbsi’’ sensu
Keene (1987) and Poromitra sp. sensu Mincarone et al. (2014)
and this paper, might represent the same Atlantic species.
The taxonomic problems of the Poromitra crassiceps group
can be properly addressed only with a major global review of
the group, including the examination of a substantial
number of specimens and the type material of all nominal
species included in the complex.
Scopeloberyx opercularis Zugmayer, 1911
Figure 1I, Table 2
Scopeloberyx opercularis Zugmayer, 1911: 8 (type-locality: off
Portugal, 368070N, 108180W, 0–4740 m depth; holotype:
MOM 0091-1179).
Specimen examined.—NPM 5987, 1, 32.0 mm (Fig. 1I), RV
Antea,sta.AB2/42A,03815028.1 00 S, 31848029.100 Wto
03815026.4 00 S, 31848022.900 W, 780 m, 27 April 2017, 1223–
1226 h.
Diagnosis.—According to Keene (1987), Scopeloberyx opercu-
laris differs from all congeners, except S. rubriventer, by the
horizontal distance between the pelvic- and pectoral-fin
Afonso et al.—Stephanoberycoidei off northeastern Brazil 475

origins (5% SL or less vs. 7.5% SL or more) and number of gill
rakers (23 or more [rarely 22] vs. 21 or fewer [rarely 22]).
Scopeloberyx opercularis differs from S. rubriventer by the HL
(39.0–44.9% SL vs. 46.1–47.8% SL; Keene, 1987; this study).
In a recent review, Kotlyar (2004b, 2004c, 2005) described
three species of Scopeloberyx:S. bannikovi,S. pequenoi, and S.
rossicus.Scopeloberyx opercularis differs from S. bannikovi and
S. pequenoi by the number of gill rakers (23–26 vs. 15–16) and
from S. rossicus by the number of vertebrae (25–28 [usually
26] vs. 23–25 [usually 24–25]; Keene, 1987; Kotlyar, 2004b,
2004c, 2005; this study). Kotlyar (2004a, 2004b) also
recognized S. opercularis as a junior synonym of S. robustus,
and this conclusion is generally accepted (e.g., Kotlyar,
2004a, 2004b; Mincarone et al., 2014). However, the present
study follows Keene (1987) and Moore (2003, 2016), which
consider S. opercularis as a valid species (see Remarks). In
addition to characters indicated previously, S. opercularis also
differs from S. robustus by the number of gill rakers (23–25 vs.
19–22; Keene, 1987; this study).
Distribution.—Scopeloberyx opercularis occurs in the western
Tropical Atlantic between 288N and 58S, including the Gulf of
Mexico and south of Caribbean Sea, and in the eastern
Atlantic between 398N and 168S (Keene, 1987). Keene (1987)
indicated records of the species in Brazilian waters off
southeastern Sa
˜o Pedro e Sa
˜o Paulo Archipelago (018200S,
278370W; 018440S, 278440W), in addition to records outside
the Brazilian EEZ. The species was also recorded in Brazilian
waters off northern Bahia State (Mincarone et al., 2014; as
Scopeloberyx robustus). In the present study, a single juvenile
specimen was collected off the Fernando de Noronha
Archipelago, at 780 m depth (Fig. 4).
Habitat.—According to Keene (1987), most specimens of S.
opercularis were captured below 700 m depth, and there is no
evidence of migratory behavior. The species is, therefore,
apparently meso- to bathypelagic.
Remarks.—Scopeloberyx opercularis is currently considered as a
junior synonym of S. robustus (e.g., Ebeling and Weed, 1973;
Maul, 1973; Fricke et al., 2020a), but there is still some
controversy in the literature about the validity of the species.
The synonymy of the two species was followed in the recent
revision of Kotlyar (2004b), who concluded that variations in
the anal-fin origin in relation to the dorsal-fin origin, number
of transverse series of scales, and number of pyloric caeca
indicated ‘‘a sub-species level of difference between the fishes
from the Atlantic Ocean and the Indo-Pacific.’’ However,
Keene (1987), in his unpublished dissertation, recognized S.
opercularis as a distinct and valid species based on the
examination of 162 specimens distributed throughout the
Tropical Atlantic. Moore (2003, 2016), probably following
Keene (1987), also considered the species as valid. Mincarone
et al. (2014) accepted that synonymy but indicated that most
characters of the single specimen identified by them as S.
robustus collected off Bahia State, central coast of Brazil, were
in accordance with the description of S. opercularis provided
by Keene (1987). Identification of the specimen examined
here is also in accordance with the diagnosis of S. opercularis
provided by Keene (1987) based on the number of gill rakers:
6–7þ16–18 ¼23–25 (8þ18 in our specimen, including one
rudimentary raker in the upper branch) vs. 5–6þ14–17 ¼19–
22 in S. robustus according to Kotlyar (2004b, 2004c, 2005).
In addition, the eye diameter of the juvenile specimen
examined herein (6.3% SL) is within the range for S.
opercularis (4.2–4.9% in adults and 5.7–6.4% in juveniles)
and differs from values proposed for S. robustus by Keene
(1987; 2.5–3.5% in adults and 4.2–5.8% in juveniles).
Therefore, and following Keene’s (1987) diagnosis and
taxonomic conclusion, we propose that S. opercularis is a
valid species. With the revalidation of S. opercularis, and
following Keene (1987), distribution of S. robustus is restricted
to the North Atlantic between about 30–408N, and the
Tropical and Subtropical Atlantic, east of 308W, including one
record in Brazilian waters, off southeastern Sa
˜o Pedro e Sa
˜o
Paulo Archipelago (018440S, 278440W).
Scopeloberyx opisthopterus (Parr, 1933)
Figure 1J, Table 2
Melamphaes opisthopterus Parr, 1933: 18 (type-locality: off Cat
Island, Bahamas, 248290N, 758530W, 7000 feet [2134 m];
holotype: YPM 2816).
Specimens examined.—NPM 5985, 1, 25.0 mm (Fig. 1J), RV
Antea,sta.AB2/49A,04810038.1 00 S, 33816007.4 00 Wto
04810058.0 00 S, 33815003.800 W, 770–1020 m, 30 April 2017,
2117–2152 h; NPM 5988, 1, 32.0 mm, sta. AB2/39,
04852026.9 00 S, 34803032.300 Wto04850052.8 00 S, 34805006.500 W,
650–800 m, 24 April 2017, 2149–2237 h; NPM 5989, 2, 29.0–
30.0 mm, sta. AB2/44A, 03852052.500 S, 32817033.3 00 Wto
03851043.6 00 S, 32816020.000 W, 850 m, 28 April 2017, 1244–
1317 h.
Diagnosis.—Scopeloberyx opisthopterus differs from all conge-
ners, except S. microlepis, by the horizontal distance between
the verticals through the ventral margin of the pectoral fin
Fig. 4. Records of Scopeloberyx opercularis (circle), Scopeloberyx
opisthopterus (square), and Scopelogadus mizolepis (triangle) off
northeastern Brazil collected during the ABRACOS surveys. Tip of arrow
indicates same collection locality for different species. FN–Fernando de
Noronha Archipelago; PB–Para´
ıba; PE–Pernambuco; RA–Rocas Atoll;
RN–Rio Grande do Norte.
476 Ichthyology & Herpetology 109, No. 2, 2021

and the origin of the pelvic fin (4.1–9.9% SL vs. 0–5.9% SL).
Scopeloberyx opisthopterus differs from S. microlepis by the
number of pelvic-fin rays (6–8 [rarely 6] vs. 6) and number of
vertebrae (25–27 vs. 27–29; Kotlyar, 2005).
Distribution.—Scopeloberyx opisthopterus has a circumglobal
distribution, occurring in both sides of the Atlantic Ocean
from off the United Kingdom to approximately 108S off
Africa (Kotlyar, 2005; Sutton et al., 2020). The species was
previously reported in Brazilian waters off Sa
˜o Pedro e Sa
˜o
Paulo Archipelago (Keene, 1987; Judkins and Haedrich,
2018). Other records in the western South Atlantic were also
restricted to the region of Sa
˜o Pedro e Sa
˜o Paulo Archipelago,
but outside the Brazilian EEZ (Keene, 1987; Judkins and
Haedrich, 2018). The four specimens collected off the
Fernando de Noronha Archipelago, between depths of 650
and 1,020 m (Fig. 4), represent an extension of the
distribution of the species in the western South Atlantic.
Habitat.—Scopeloberyx opisthopterus is meso- to bathypelagic,
with larger post-larvae and all other stages inhabiting depths
between 800 and at least 1,550 m, whereas smaller post-
larvae occur between 50 and 300 m (Keene et al., 1987).
Scopelogadus mizolepis (G¨
unther, 1878)
Figure 1K, Table 2
Scopelus mizolepis G¨
unther, 1878: 185 (type-locality: south of
New Guinea, off Aru Island; Molucca Islands, Indonesia,
Arafura Sea, western Pacific, 058410S, 13480403000 E, 800
fathoms [1463 m] depth; holotype: BMNH 1887.12.7.9).
Specimens examined.—NPM 5990, 3, 37.0–49.0 mm (Fig. 1K),
RV Antea,sta.AB2/41A,03819059.100 S, 32824042.100 Wto
03819031.8 00 S, 32825004.600 W, 430 m, 26 April 2017, 2144–
2206 h; NPM 5991, 1, 54.5 mm, sta. AB2/52A, 03843016.200 S,
33825009.8 00 Wto03842014.200 S, 33824036.200 W, 822–984 m, 2
May 2017, 1147–1218 h; NPM 5992, 5, 42.0–58.0 mm, sta.
AB2/42A, 03815028.100 S, 31848029.100 Wto03815026.400 S,
31848022.9 00 W, 780 m, 27 April 2017, 1223–1226 h; NPM
5993, 1, 43.0 mm, sta. AB2/59A, 03838001.6 00 S, 36803010.600 W
to 03838007.9 00 S, 36802022.600 W, 700–1113 m, 5 May 2017,
2157–2237 h; NPM 5994, 3, 46.0–70.0 mm, sta. AB2/44A,
03852052.5 00 S, 32817033.300 Wto03851043.6 00 S, 32816020.0 00 W,
850 m, 28 April 2017, 1244–1317 h; NPM 5995, 2, damaged–
45.0 mm, sta. AB2/39, 04803032.3 00 S, 34835022.900 Wto
04850052.8 00 S, 34805006.500 W, 650–800 m, 24 April 2017,
2149–2237 h; NPM 5996, 3, 37.0–40.0 mm, sta. AB2/53A,
03848058.7 00 S, 33859017.100 Wto03850005.8 00 S, 33858046.5 00 W,
610 m, 2 May 2017, 2208–2240 h; NPM 6090, 1, 40.0 mm,
sta. AB1/22, 04807044.8 00 S, 33847024.500 Wto04807000.7 00 S,
33848057.9 00 W, 525 m, 8 October 2015, 2132–2212 h.
Diagnosis.—Four species of Scopelogadus are currently consid-
ered as valid, with only S. beanii and S. mizolepis occurring in
the Atlantic (Fricke et al., 2020a). Scopelogadus mizolepis
differs from S. beanii by the number of gill rakers 21–26 (6–
8þ15–18) vs. 26–32 (8–10þ18–22; but see Remarks), and by
the stomach noticeably darkened posteriorly (vs. stomach
not darkened posteriorly; Sutton et al., 2020).
Distribution.—Scopelogadus mizolepis occurs in all oceans
except the eastern Pacific (Kotlyar, 2020). In the Atlantic,
the species is reported between 438N and 308S (Ebeling and
Weed, 1973; Keene, 1987; McEachran and Fechhelm, 1998;
Moore, 2016; Sutton et al., 2020). Several records are known
in the western South Atlantic and also off Ascension Island
(Keene, 1987; Judkins and Haedrich, 2018). In Brazilian
waters, the species was previously reported off Sa
˜o Pedro e
Sa
˜o Paulo Archipelago, Vit ´
oria-Trindade Seamount Chain
(Keene, 1987), Rocas Atoll, Fernando de Noronha Archipel-
ago (Judkins and Haedrich, 2018), and off Rio de Janeiro
State, southeastern Brazil (Costa and Mincarone, 2010;
Mincarone et al., 2014). The 19 specimens identified here
as S. mizolepis were collected near the Fernando de Noronha
Archipelago, Rocas Atoll, and the seamounts off Rio Grande
do Norte State, between depths of 430 and 1,113 m (Fig. 4).
Habitat.—The species is meso- to bathypelagic, with adults
collected below 500 m and post-larvae and juveniles
collected between depths of 50 and 300 m (Ebeling and
Weed, 1973; McEachran and Fechhelm, 1998).
Remarks.—There is some inconsistency in values of the
number of gill rakers on the first gill arch presented by
Kotlyar (2020) in his recent review of S. mizopelis. In the
diagnosis (Kotlyar, 2020: 4), it is stated ‘‘on first branchial
arch, 16–24 (usually 19–21) rakers,’’ whereas in the descrip-
tion, presented a few lines later in the same page, it is
mentioned ‘‘(6–8)þ1þ(12–17) ¼19–25’’ as the number of gill
rakers on the first gill arch. Variation in the number of gill
rakers on the first gill arch of the 19 specimens of S. mizolepis
examined here (22–25; Table 2) is in accordance with values
presented by both Kotlyar (2020: description) and Sutton et
al. (2020).
Two subspecies of Scopelogadus mizolepis were recognized
by Ebeling and Weed (1973): Scopelogadus mizolepis bispinosus
(Gilbert 1915), from the eastern Tropical Pacific, and
Scopelogadus mizolepis mizolepis (G¨
unther 1878), from the
Tropical Atlantic and Central Pacific. These subspecies are
not considered herein, as in Iwasaki (2009) and Mincarone et
al. (2014). However, according to Kotlyar (2020), the two
subspecies of S. mizolepis proposed by Ebeling and Weed
(1973) actually represent species that await formal recogni-
tion at that level.
Cetomimidae
The Cetomimidae (whalefishes) includes meso- and bathy-
pelagic fishes occurring in all oceans between 528N and 728S
(Paxton, 1989). After Johnson et al.’s (2009) study, the
number of species in the family became rather uncertain, as
molecular data revealed that species belonging to the
Mirapinnidae and Megalomycteridae are, in fact, larvae and
males, respectively, of the Cetomimidae, which was previ-
ously known only by females. About 21 to 26 nominal
species belonging to nine genera have been recognized as
valid in the Cetomimidae (Paxton, 1989; Johnson et al.,
2009; Nelson et al., 2016; Fricke et al., 2020b), with several
species still lacking formal description (Paxton, 1989; Nelson
et al., 2016). The Cetomimidae comprises one of the most
species-rich groups in the bathypelagic zone (1,000–4,000 m)
and it is suspected that this is the most abundant fish family
below 1,800 m (Paxton, 1989; Nelson et al., 2016). However,
records of the family in the western South Atlantic are still
scarce (e.g., Paxton, 1989; Mincarone et al., 2014). Cetomi-
mid fishes are mainly recognized by an elongated body,
enormous mouth extending far behind eye, reduced or
Afonso et al.—Stephanoberycoidei off northeastern Brazil 477

rudimentary eyes, absence of scales, and pelvic fins totally
absent in females, usually absent in males, and jugular in
juveniles (Paxton, 1989; Johnson et al., 2009; Mincarone et
al., 2014; Nelson et al., 2016).
Cetomimus sp. 1
Figure 5A, Table 3
Specimens examined.—NPM 5004, 1, 65.1 mm, RV Antea, sta.
AB2/16, 07836014.4 00 S, 33859033.8 00 Wto07836049.300 S,
33857018.7 00 W, 680 m, 14 April 2017, 2153–2239 h; NPM
5005, 1, 62.5 mm (Fig. 5A), sta. AB2/42A, 03815028.1 00 S,
31848029.1 00 Wto03815026.400 S, 31848022.9 00 W, 780 m, 27
April 2017, 1223–1226 h.
Diagnosis.—Cetomimus has no single, derived character that
distinguishes it from other genera of the Cetomimidae
(Paxton, 1989). Three free branchial arches are present in
Cetomimus, and also in Rhamphocetichthys and Gyrinomimus.
Cetomimus differs from Rhamphocetichthys by having a
cavernous lateral-line system formed by large canals pierced
by wide pores on the head and body (vs. absence of these
canals), a much shorter, rounded snout (vs. elongated and
pointed snout), and by the absence of ventral pharyngeal
tooth plates (vs. presence of ventral pharyngeal tooth plates).
Cetomimus is most similar to Gyrinomimus, differing from this
genus by the shape of teeth (short, in indistinct diagonal
rows vs. long, in distinct, usually longitudinal, rows) and by
the shape of the vomerine tooth patch (round or elliptical
and dome shaped vs. rectangular or laterally elongate and
flat; Paxton, 1989).
Distribution.—Species of Cetomimus are reported from the
Atlantic and Indo-Pacific Oceans, ranging from 418Nto578S
in the Pacific and from 418Nto408S in the Atlantic (Paxton,
1989). Two specimens badly damaged during the trawl (NPM
5004 and 5005) were collected off Pernambuco State and the
Fernando de Noronha Archipelago, between depths of 680
and 780 m (Fig. 6). Those specimens represent the first
confirmed records of the genus Cetomimus in Brazilian waters
and some of the few records in the western South Atlantic.
Habitat.—Species of Cetomimus are meso- to bathypelagic,
ranging from depths between 500 and approximately 3,300
m (Paxton, 1989; Tolley et al., 1989; Angulo, 2015; Paxton et
al., 2016). Juvenile specimens have been recorded in shallow
waters (Paxton et al., 2016).
Remarks.—Cetomimus currently includes seven nominal
species, and at least five species await description (Paxton,
1989). The seven species considered as valid are: Cetomimus
compunctus, from the western North Pacific and the western
South and eastern Central Atlantic (Paxton et al., 2016);
Cetomimus craneae, from Bermuda (Harry, 1952); Cetomimus
gillii, from the Atlantic, eastern Pacific, and western Indian
Oceans (Angulo, 2015; Paxton et al., 2016); Cetomimus
hempeli, from the Atlantic Ocean and possibly the North
Pacific (Paxton et al., 2016); Cetomimus kerdops, from the
Bahamas (Parr, 1934; Moore and Boardman, 1991); Cetomi-
mus picklei, from the eastern South Atlantic (Paxton and Bray,
1986); and Cetomimus teevani, from the western Atlantic
(Harry, 1952). The identification at the species level of the
two specimens reported here based on the ABRACOS
collection (NPM 5004 and 5005; Cetomimus sp. 1) was not
possible due to their poor state of preservation. Both
specimens are somewhat distorted, and the skin is almost
completely unattached to the remaining integument. An-
other specimen of the genus (MNRJ 26794) was identified as
Cetomimus sp. 2 (Fig. 5B), collected off Esp´
ırito Santo State,
southeastern Brazil (19842034.100 S, 38832001.800 Wto
19842041.1 00 S, 38836057.700 W), between depths of 875 and
942 m. The skin of specimen MNRJ 26794 is also damaged,
compromising the observation of lateral-line pores and flaps
and of the cavernous tissue. Nevertheless, some measure-
ments were successfully obtained, in addition to the number
of vertebrae, and of the dorsal-, anal-, and pectoral-fin rays
(Table 3). Cetomimus sp. 1 differs from Cetomimus sp. 2 in the
shape of the vomerine tooth patch (oval vs. triangular, with
its anterior tip narrower) in addition to meristic and
morphometric data provided in Table 3. Therefore, at least
two species of the genus occur in Brazilian waters, one of
them recorded off northeastern Brazil (NPM 5004 and 5005,
Cetomimus sp. 1) and the other collected off Esp´
ırito Santo
State (MNRJ 26794, Cetomimus sp. 2).
Cetostoma regani Zugmayer, 1914
Figure 5C, Table 3
Cetostoma regani Zugmayer, 1914: 4 (type locality: eastern
Atlantic, 30845030 00 N, 258470W, 0–2000 m depth; holotype:
MOM 0091-1729).
Specimens examined.—NPM 3185, 1, 81.0 mm (Fig. 5C), RV
Antea, sta. AB1/22, 04807044.8 00 S, 33847024.5 00 Wto
04807000.7 00 S, 33848057.900 W, 525 m, 8 October 2015, 2132–
2212 h; NPM 5001, 1, 96.8 mm, sta. AB2/54B, 03845017.200 S,
34841004.0 00 Wto03844039.2 00 S, 34840004.500 W, 830–1030 m, 3
May 2017, 1311–1347 h; NPM 5002, 2, 85.2–113.7 mm, sta.
AB2/42A, 03815028.1 00 S, 31848029.100 Wto03815026.4 00 S,
3184802.9 00 W, 780 m, 27 April 2017, 1223–1226 h; NPM
5151, 1, 95.0 mm, sta. AB2/44A, 03852052.5 00 S, 32817033.300 W
to 03851043.6 00 S, 32816020.000 W, 850 m, 28 April 2017, 1244–
1317 h.
Diagnosis.—Cetostoma regani is the single species of the
genus, and it differs from other genera of the Cetomimidae
by the number of dorsal-fin rays (29–37 vs. 13–22), number
of anal-fin rays (26–34 vs. 13–20), dorsal- and anal-fin bases
elevated in relation to the body (vs. not elevated), predorsal
length (1.7–2.0 in SL vs. 1.3–1.6 in SL), a very long, narrow
copular tooth patch present as three separate dentigerous
plates (vs. one solid plate), the gill slit behind the angle of
fourth gill arch tiny and tubular (vs. gill slit behind the
ventral arm of fourth gill arch either elongate or absent),
numerous small skin ridges along the belly from the pectoral-
fin base to the anus (vs. absence of skin ridges), and the fin
membrane between last ten anal-fin rays voluminous and
curtain-like (vs. fin membrane between posterior anal-fin
rays not voluminous and not curtain-like; Paxton, 1989).
Distribution.—Cetostoma regani has the broadest distribution
of all cetomimids, occurring in the Atlantic and Indo-Pacific
Oceans (except the eastern South Pacific), from 508Nto408S
(Paxton et al., 2016). The species was previously recorded in
Brazilian waters off Sa
˜o Pedro e Sa
˜o Paulo Archipelago, North
Atlantic, at 028410N, 288560W, 0–ca. 1,100 m depth (MCZ
42844), and at 008170N, 278310W, 0–ca. 300 m depth (MCZ
42843; Paxton, 1989). In this study, five specimens were
478 Ichthyology & Herpetology 109, No. 2, 2021

Fig. 5. Species of Cetomimidae and
Rondeletiidae reported in this study:
(A) Cetomimus sp. 1, NPM 5005,
62.5 mm SL; (B) Cetomimus sp. 2,
MNRJ 26794, 92.0 mm SL; (C)
Cetostoma regani, NPM 3185, 81.0
mm SL; (D) Ditropichthys storeri,
NPM 5003, 49.0 mm SL; (E) Gyrino-
mimus bruuni, NPM 5000, 66.2 mm
SL; (F) Gyrinomimus cf. bruuni, MNRJ
26793, 305.0 mm SL; (G) Rondeletia
loricata, NPM 3197, 32.3 mm SL.
Scale bar ¼1 cm.
Afonso et al.—Stephanoberycoidei off northeastern Brazil 479

Table 3. Morphometric and meristic data of species of Cetomimidae and Rondeletiidae reported in this study (d ¼damaged).
Species
Cetomimus
sp. 1
Cetomimus
sp. 2
Cetostoma
regani
Ditropichthys
storeri
Gyrinomimus
bruuni
Gyrinomimus
cf. bruuni
Gyrinomimus
sp.
Rondeletia
loricata
n2151211 3
Standard length (SL, mm) 62.5–65.1 92.0 81.0–113.7 49.0 60.0–66.2 305.0 95.0 32.3–78.4
Measurements in % SL
Head length d–30.1 26.0 21.4–28.0 33.7 25.5–26.9 27.9 34.8 44.2–45.9
Head width 7.2–10.7 — 6.4–7.6 16.5 — 16.4 19.5 17.2–19.8
Eye diameter — — — — — 1.0 1.7 6.5–7.4
Postorbital length — — — — — 22.0 24.2 16.4–20.0
Snout length — 9.5 11.9 — 10.6–10.7 8.4 11.6 18.6–20.0
Upper jaw length d–29.9 23.0 16.1–19.6 — 24.3–26.4 23.0 32.6 25.0–26.0
Body depth 12.9–13.3 — 6.9–9.0 21.8 — 23.3 — 32.1–37.5
Prepectoral length 36.2–38.7 — 23.2–29.5 36.1 27.7–30.7 30.0 — 43.3–48.5
Prepelvic length — — — — — — — 60.1–69.3
Predorsal length 72.6–75.0 66.0 54.5–58.8 69.4 74.0–75.5 71.5 67.4 66.6–72.8
Preanal length 73.6–76.0 66.0 55.8–60.0 70.4 74.6–76.7 72.1 — 68.1–74.0
Dorsal-fin base length 16.3–16.4 13.0 32.2–40.2 26.7 15.7–15.9 17.0 18.0 19.4–27.9
Anal-fin base length 16.0–16.9 13.9 26.3–32.7 20.4 13.6–13.8 16.6 19.0 16.6–27.9
Caudal peduncle length 9.9–11.5 10.8 9.5–12.7 15.3 8.3–9.0 10.5 12.0 7.7–10.8
Caudal peduncle depth 5.4–5.9 3.3 3.3–4.3 6.0 5.1–5.2 4.4 7.4 8.3–12.4
Head length (HL, mm) d–17.1 23.9 — — d–17.8 85.0 33.1 —
Measurements in % HL
Copular tooth plate length d–29.2 33.5 — — d–27.0 27.0 30.2 —
Copular tooth plate width max. d–7.6 6.7 — — d–19.1 10.6 13.6 —
Copular tooth plate width min. d–5.8 4.2 — — d–7.9 5.3 12.0 —
Copular tooth plate ratio (max/min) d–1.3 1.6 — — d–2.4 2.0 1.1 —
Counts
Dorsal-fin rays 15–16 18 30–34 21 20 19 16 11–12
Anal-fin rays 16 17 27–32 16 18–19 18 15 11–12
Pectoral-fin rays 15–20 16 17–20 18 16–18 16 18 7–8
Pelvic-fin rays — — — — — — — 4–6
Principal caudal rays (upper/lower) 7–8/9 — 7–9/7–10 6/7 8/9–10 8/9 8/9 10–11/10
Procurrent caudal rays (upper/lower) 2–3/2–3 — — 3/2 3/2 5/4 5/3 5/2–3
Vertebrae (precaudal þcaudal) 32þ17 33þ21 22–26þ23–26 22þ17 36–38þ19–21 38þ20 — 10–11þ15–16
Vertebrae (total) 48–49 54 48–50 39 57 58 48 25–27
Jaws teeth rows (upper/lower) — — — — d–3–4/3–4 6–9/7–10 3/3 —
Vomerine teeth rows — — — — 2 indistinct 3 —
480 Ichthyology & Herpetology 109, No. 2, 2021

collected off the Fernando de Noronha Archipelago, the
Rocas Atoll, and the seamounts off Rio Grande do Norte
State, between depths of 525 and 1,030 m (Fig. 6). In
addition, another specimen identified as C. regani (MNRJ
26795), collected off Esp´
ırito Santo State (21812017.600 S,
40800053.0 00 Wto21809034.600 S, 40800027.7 00 W) between
depths of 1,333 and 1,390 m, extends the occurrence of
the species further south in Brazilian waters. Specimens
reported here also represent some of the few confirmed
records of the species in the western South Atlantic (Paxton,
1989).
Habitat.—Females of Cetostoma regani are meso- to bathype-
lagic, with adults collected between 100 and 3,700 m,
whereas juveniles are reported from shallow waters (Paxton,
1989; Paxton et al., 2016). Males are bathypelagic (Paxton et
al., 2016).
Remarks.—In this study, only females were collected. The
absence of males might be related to the depth of collections
(maximum 1,113 m), which did not reach the bathypelagic
zone.
Ditropichthys storeri (Goode and Bean, 1895)
Figure 5D, Table 3
Cetomimus storeri Goode and Bean, 1895: 453 (type locality:
western North Atlantic, 39803015 00 N, 7085004500 W, 1535
fathoms [2807 m] depth; holotype: USNM 35634).
Specimen examined.—NPM 5003, 1, 49.0 mm (Fig. 5D), RV
Antea,sta.AB2/53A,03848058.700 S, 33859017.1 00 Wto
03850005.8 00 S, 33858046.500 W, 610 m, 2 May 2017, 2208–
2240 h.
Diagnosis.—Ditropichthys storeri is the single species of the
genus, differing from all other genera in the Cetomimidae by
having fully developed, club-shaped gill rakers, a pair of thin
dermal folds along the abdomen, and anal lappets connected
as an unbroken fold of skin containing lappet scales over the
anal-fin base (Paxton, 1989).
Distribution.—Ditropichthys storeri has a circumglobal distri-
bution between 458N and 458S (Paxton et al., 2016). The
absence of the species in some regions may be related to the
lack of collecting efforts (Paxton, 1989). The species was
previously recorded in the western South Atlantic in five
localities off Uruguay and Argentina (Paxton, 1989). A single
specimen of D. storeri was collected off the Rocas Atoll, at 610
m depth (Fig. 6). This represents the first record of the species
in Brazilian waters and one of the few confirmed records in
the western South Atlantic (Paxton, 1989).
Habitat.—Small specimens of D. storeri (,40 mm) are
mesopelagic, occurring from 650 to 1,000 m, whereas larger
specimens (.60 mm) are bathypelagic, occurring from 1,000
to approximately 5,000 m (Paxton et al., 2016).
Gyrinomimus bruuni Rofen, 1959
Figure 5E, Table 3
Gyrinomimus bruuni Rofen, 1959: 257 (type locality: off
Kenya, 058250S, 478090E, over 4820 m depth; holotype:
ZMUC P23452).
Specimens examined.—NPM 5000, 2, 60.0–66.2 mm (Fig. 5E),
RV Antea,sta.AB2/54B,03845017.2 00 S, 34841 004.000 Wto
03844039.200 S, 34840004.500 W, 830–1030 m, 3 May 2017,
1311–1347 h.
Diagnosis.—Gyrinomimus differs from other genera of the
Cetomimidae, except Rhamphocetichthys and Cetomimus,by
having three free branchial arches (vs. four). Gyrinomimus
differs from Rhamphocetichthys by having a round snout (vs.
beak-like snout), and by the presence of tooth plates on the
second and third branchial arches (vs. plates absent). It
differs from Cetomimus by having jaw teeth arranged in
distinct longitudinal rows (vs. jaw teeth arranged in
indistinct diagonal rows), by the length of the teeth (except
the newest teeth) more than three times the width of its base
(vs. less than two times the width of its base), and by the
shape of the vomerine tooth plate, which is flat and
rectangular or oval (vs. domed and round or rarely oval;
Paxton, 1989). Gyrinomimus bruuni differs from its congeners
by the following characters: number of dorsal-fin rays (19–20
vs. 14–17 in G. andriashevi,G. grahami,G. myersi, and G.
parri); number of anal-fin rays (18–20 vs. 14–17 in G.
andriashevi,G. grahami,G. myersi, and G. parri); number of
lateral-line pores (19 vs. 14–15 in G. myersi and G. parri, and
23 in G. andriashevi); and pectoral-fin length (6.2–9.8% SL vs.
2.9% SL in G. andriashevi; Parr, 1934; Richardson and Garrick,
1946; Rofen, 1959; Bigelow, 1961; Fedorov et al., 1987;
Paxton, 1989).
Distribution.—Gyrinomimus bruuni has a circumglobal distri-
bution between 308N and 108S (Paxton, 2003). This is the
first report of this species in Brazilian waters, and it is based
on two specimens collected off the Rocas Atoll, between
depths of 830 and 1,030 m (Fig. 6). Those specimens also
Fig. 6. Records of Cetomimus sp. 1 (circle), Cetostoma regani (square),
Ditropichthys storeri (triangle), Gyrinomimus bruuni (pentagon), and
Rondeletia loricata (diamond) off northeastern Brazil collected during
the ABRACOS surveys. Tip of arrow indicates same collection locality for
different species. FN–Fernando de Noronha Archipelago; PB–Para´
ıba;
PE–Pernambuco; RA–Rocas Atoll; RN–Rio Grande do Norte.
Afonso et al.—Stephanoberycoidei off northeastern Brazil 481

represent one of the few records of the genus in the South
Atlantic and apparently the first confirmed record of the
species in the region (see Remarks).
Habitat.—Maximum depth reported for the species is 1,805
m (MNRJ 26793, this study). Other species of the genus are
bathypelagic, captured between 1,594 and 2,350 m (Mincar-
one et al., 2014; Paxton et al., 2016).
Remarks.—In addition to G. bruuni, four other species of
Gyrinomimus are currently regarded as valid: G. andriashevi,
from the Antarctic Ocean; G. grahami, cosmopolitan in the
South Hemisphere; G. myersi, circumglobal; and G. parri,
from the western Atlantic and western Pacific (Paxton, 1989;
Paxton et al., 2016; Fricke et al., 2020a). Two species groups
of Gyrinomimus in the North Atlantic are recognized: the
bruuni species group, with G. bruuni and two undescribed
species, and the myersi species group, with G. myersi,G. parri,
and one undescribed species (Moore et al., 2003; Paxton et
al., 2016).
Mincarone et al. (2014) reported the first specimen of
Gyrinomimus in Brazilian waters (MNRJ 36421, 95 mm SL),
collected off Rio de Janeiro State, southeastern Brazil. This
specimen was highly damaged and its identification at the
species level was not possible. A comparison made between
the specimens reported here as G. bruuni with the one
reported by Mincarone et al. (2014) as Gyrinomimus sp.
clearly indicates that the latter belongs to a different species.
Gyrinomimus sp. differs from G. bruuni by the following
characters: three distinct gill arches bearing well-developed
holobranchs, a reduced gill slit behind the ventral arm of the
third arch, near the angle vs. four distinct gill arches bearing
well-developed holobranchs, a relatively well-developed gill
slit behind the ventral arm of the third arch; holobranchs on
fourth gill arch highly undeveloped, in the shape of tubercles
vs. holobranchs more developed (0.5 times length of
holobranchs on first gill arch) and with a regular shape;
number of dorsal-fin rays (16 vs. 20); number of anal-fin rays
(15 vs. 18–19); number of distinct teeth rows on vomer (3 vs.
2); number of vertebrae (48 vs. 57); middle portion of preural
centra distinctly constricted vs. centra only slightly con-
stricted; HL (34.8% SL vs. 25.5–26.9% SL); upper-jaw length
(32.6% SL vs. 24.3–26.4% SL); predorsal length (67.4% SL vs.
74.0–75.5% SL); dorsal-fin base length (18.0% SL vs. 15.7–
15.9% SL); anal-fin base length (19.0% SL vs. 13.6–13.8% SL);
caudal-peduncle length (12.0% SL vs. 8.3–9% SL); and caudal
peduncle depth (7.4% SL vs. 5.1–5.2% SL; Table 3).
In addition to G. bruuni and Gyrinomimus sp., another
specimen of the genus (MNRJ 26793, 305 mm SL) collected
off Rio de Janeiro State, from 21828036.7 00 S, 39840018.200 Wto
21825031.4 00 S, 39840026.600 W, between depths of 1,790 and
1,805 m, was tentatively identified as Gyrinomimus cf. bruuni,
but it may represent an undescribed species (Fig. 5F, Table 3;
Paxton, 1989; J. Paxton, pers. comm.). Morphometric and
meristic data comparing G. bruuni ‘‘stricto sensu’’ (NPM 5000,
2 specimens) and Gyrinomimus cf. bruuni (MNRJ 26793) are
presented in Table 3. Other characters that differ between G.
bruuni and Gyrinomimus cf. bruuni are: the shape of preural
centra (middle portion of the centra only slightly constricted
vs. distinctly constricted) and the number of teeth rows on
upper (3–4 vs. 6–9, increasing anteriorly) and lower jaws (3–4
vs. 7–10, increasing anteriorly). Gyrinomimus cf. bruuni also
has cavernous tissue up to above the third anal-fin ray, about
three anal lappets, about 21–22 lateral-line pores, and the
length of the holobranchs on the fourth arch is 0.54 the
length on those of the first arch (J. Paxton, pers. comm.;
present study). A further specimen of Gyrinomimus (MCZ
50688), collected off Rio Grande do Sul State, southern Brazil,
in 1967, referred to as Gyrinomimus sp. by Mincarone et al.
(2014), still seems to be lost.
Rondeletiidae
The Rondeletiidae includes only two species, Rondeletia
bicolor Goode and Bean 1895 and Rondeletia loricata Abe and
Hotta 1963 (Paxton et al., 2001). Both species are meso- and
bathypelagic in tropical and temperate waters, with R. bicolor
occurring in the Atlantic and Pacific Oceans, and R. loricata
having an almost circumglobal distribution (Paxton and
Trnski, 2003; Kobyliansky et al., 2020). Rondeletia is mainly
recognized among the Stephanoberycoidei by having the
following combination of characters: large mouth with jaws
not extending beyond the posterior margin of eye; pelvic fins
with five or six soft rays; lack of teeth on basibranchials;
lateral line as vertical rows of papillae without supporting
internal scales; and lack of external body scales (Paxton and
Trnski, 2003).
Rondeletia loricata Abe and Hotta, 1963
Figure 5G, Table 3
Rondeletia loricata Abe and Hotta, 1963: 43, Pls. 11 (figs. 1–7),
12 (figs. 8–9) (type locality: off Kesennuma, Miyagi
Prefecture, Japan, 750 m depth; holotype: ZUMT 52196).
Specimens examined.—NPM 3197, 1, 32.3 mm (Fig. 5G), RV
Antea, sta. AB1/22, 04807044.8 00 S, 33847024.5 00 Wto
04807000.7 00 S, 33848057.900 W, 525 m, 8 October 2015, 2132–
2212 h; NPM 4144, 1, 78.4 mm, sta. AB2/54B, 03845017.200 S,
34841004.0 00 Wto03844039.2 00 S, 34840004.500 W, 830–1030 m, 3
May 2017, 1311–1347 h; NPM 4228, 1, 46.4 mm, sta. AB2/
44A, 03852052.500 S, 32817033.3 00 Wto03851043.6 00 S,
32816020.0 00 W, 850 m, 28 April 2017, 1244–1317 h.
Diagnosis.—Rondeletia loricata differs from R. bicolor by the
number of vertical rows of lateral-line pores (14–19 vs. 24–
26), the lack of a bony sphenotic hook over the orbit (vs.
presence of a bony sphenotic hook), and supratemporal and
cleithrum with large posterior extensions (vs. absence of
large posterior extensions; Paxton, 1974; Paxton and Trnski,
2003).
Distribution.—Rondeletia loricata has an almost circumglobal
distribution, being reported from 608Nto508S in all oceans
(Paxton, 1974; Bast and Klinkhardt, 1990; Kotlyar, 1996;
Paxton et al., 2001; Paxton and Trnski, 2003; Kharin, 2006;
Balanov and Kharin, 2009; Møller et al., 2010; Mincarone et
al., 2014). Records of the species in the western South
Atlantic are restricted to off Argentina and Brazil (Figueroa et
al., 1998; Mincarone et al., 2014). The species was first
reported in Brazilian waters by Mincarone et al. (2014), based
on three specimens collected off Bahia and Esp´
ırito Santo
States, between depths of 837 and 1,049 m. Rondeletia loricata
is recorded here based on three specimens collected off Rocas
Atoll, the Fernando de Noronha Archipelago, and seamounts
off Rio Grande do Norte State, between depths of 525 and
1,030 m (Fig. 6).
482 Ichthyology & Herpetology 109, No. 2, 2021

Habitat.—The species is meso- to bathypelagic, with most
records of adults below 400 m, with a maximum record of
1,200 m depth (Paxton et al., 2001; Kharin, 2006; Balanov
and Kharin, 2009). Larvae (3.5–4.6 mm SL) are captured in
shallow waters, between 8 and 40 m, and juveniles (,20 mm
SL) are captured between 110 and 175 m (Paxton et al.,
2001).
DISCUSSION
Scientific expeditions conducted since the last decades of the
20
th
century resulted in new records and new species
descriptions of several groups of deep-sea fishes in Brazilian
waters, substantially contributing to the understanding of
this important component of the diversity in the western
South Atlantic (e.g., Figueiredo et al., 2002; Melo, 2008;
Santos and Figueiredo, 2008; Carvalho-Filho et al., 2010;
Melo et al., 2010; Lima et al., 2011; Braga et al., 2014;
Pinheiro et al., 2015; Eduardo et al., 2018, 2019a, 2019b,
2020a, 2020b; Mincarone et al., 2019, 2020). However,
knowledge on the deep-sea fish diversity of the western
South Atlantic is still insufficient (Paxton, 1989; Mincarone
et al., 2014; Nielsen et al., 2015; Reis et al., 2016). The eight
new records and nine range extensions of species of the
Stephanoberycoidei reported here for Brazil, for instance,
were based on two relatively short deep-sea collecting
campaigns, indicating that a substantial diversity of deep-
sea fishes is still waiting to be discovered and properly
studied in the region.
With the new records presented here, a total of 26 species
of the Stephanoberycoidei are reported from off Brazil
(Keene, 1987; Paxton, 1989; Mincarone et al., 2014; Judkins
and Haedrich, 2018; Table 4). Based on the distribution of
melamphaids reported by Ebeling (1962) and Keene (1987),
other species of the family that potentially occur in Brazilian
waters are: Melamphaes suborbitalis (recorded in the central
Atlantic and off Rio da Prata, between Uruguay and
Argentina); M. microps (south of 278S, near the Brazilian
EEZ off Rio Grande do Sul State); M. simus (central Atlantic);
and Sio nordenskjoldii (south of 328S, off Uruguay). Scopelober-
yx nigrescens, which was considered as a junior synonym of
Scopeloberyx robustus by Kotlyar (2004b), but considered as
Table 4. Species of the Stephanoberycoidei recorded in Brazilian waters. * The four specimens of B. rufa (and only known records of the species off
Brazil) reported as missing by Mincarone et al. (2014) have now been located, in lots MNRJ 42181, 42182, 42183, and 42184.
Species Distribution References
Barbourisiidae
Barbourisia rufa Circumglobal Mincarone et al. (2014)*
Cetomimidae
Cetostoma regani Circumglobal, except eastern
South Pacific
Paxton (1989), present study
Cetomimus sp. 1 off northeastern Brazil present study
Cetomimus sp. 2 off Esp´
ırito Santo, Brazil present study
Ditropichthys storeri Circumglobal present study
Gyrinomimus bruuni Circumglobal present study
Gyrinomimus cf. bruuni off Rio de Janeiro, Brazil present study
Gyrinomimus sp. off Rio de Janeiro, Brazil Mincarone et al. (2014); present study
Gibberichthyidae
Gibberichthys pumilus western Tropical Atlantic Asano Filho et al. (2005); Mincarone et al. (2014)
Melamphaidae
Melamphaes eulepis Circumtropical, except eastern Pacific present study
Melamphaes hubbsi Tropical South Atlantic Judkins and Haedrich (2018)
Melamphaes leprus Tropical Atlantic present study
Melamphaes longivelis Circumglobal, except eastern Pacific present study
Melamphaes polylepis Circumglobal Keene (1987); Mincarone et al. (2014); Eduardo et al. (2020a);
present study
Melamphaes typhlops Atlantic Keene (1987); Mincarone et al. (2014); present study
Melamphaes sp. off Rio Grande do Norte, Brazil present study
Poromitra megalops Circumtropical Keene (1987); Judkins and Haedrich (2018); present study
Poromitra sp. off Brazil Mincarone et al. (2014); present study
Scopeloberyx opercularis Tropical Atlantic Keene (1987); Mincarone et al. (2014, as S. robustus); present
study
Scopeloberyx opisthopterus Circumglobal Keene (1987); Judkins and Haedrich (2018); present study
Scopeloberyx robustus Circumglobal, except eastern Pacific Keene (1987); Judkins and Haedrich (2018)
Scopelogadus beanii Circumglobal Judkins and Haedrich (2018)
Scopelogadus mizolepis Circumglobal Keene (1987); Costa and Mincarone (2010); Mincarone et al.
(2014); Judkins and Haedrich (2018); present study
Stephanoberycidae
Acanthochaenus luetkenii Atlantic, Indian and South Pacific Mincarone et al. (2014)
Stephanoberyx monae western Atlantic Mincarone et al. (2014)
Rondeletiidae
Rondeletia bicolor Atlantic and Pacific Mincarone et al. (2014)
Rondeletia loricata Circumglobal Mincarone et al. (2014); present study
Afonso et al.—Stephanoberycoidei off northeastern Brazil 483

valid by Moore (2003, 2016), was also reported from the
central Atlantic and, if valid, also potentially occurs off Brazil
(Keene, 1987).
The complex taxonomic scenario revealed by the exami-
nation of relatively few specimens of Melamphaes,Poromitra,
and Scopeloberyx in this study indicates that the systematics
of certain components of those genera are still in need of
revision, despite Kotlyar’s extensive taxonomic work (e.g.,
Kotlyar, 2004b, 2004c, 2005, 2008a, 2008b, 2009a, 2009b,
2009c, 2010, 2011a, 2011b, 2012a, 2012b, 2013, 2014,
2015a, 2015b, 2015c, 2016a, 2016b, 2016c). Part of the
problem might be due to the fact that a substantial number
of species in the group have been described based on
relatively few specimens or even on a single specimen in
some cases, as previously noted by other authors (e.g.,
Ebeling, 1962; Bartow, 2010). This situation is quite common
for deep-sea organisms and might not necessarily represent a
problem in itself when species are unambiguously distinct
from congeners or are presumably rare or with relatively
restricted geographic ranges, for instance. However, some
species of the Melamphaidae, particularly those of the genera
Melamphaes,Poromitra,andScopeloberyx,areapparently
abundant and have presumably large geographic ranges.
Therefore, descriptions or taxonomic revisions of compo-
nents of those genera based on relatively few specimens
patchily distributed over large areas have a worrying
tendency of neglecting relevant anatomical variation. The
experience accumulated in the last decades indicate that
extensive taxonomic studies including proper examination
of the type series and a large, truly representative number of
specimens on a global scale are required for a more coherent
and realistic taxonomic scenario of the group to emerge. In
this context, and also based on the results presented by
Mincarone et al. (2014), more investments in deep-sea
collections in historically neglected regions such as the
South Atlantic are still necessary in order to properly assess
the diversity of the Stephanoberycoidei.
DATA ACCESSIBILITY
Unless otherwise indicated in the figure caption, the
published images and illustrations in this article are licensed
by the American Society of Ichthyologists and Herpetologists
for use if the use includes a citation to the original source in
accordance with the Creative Commons Attribution CC BY
License.
ACKNOWLEDGMENTS
We thank the French oceanographic fleet for funding the
ABRACOS at-sea survey and the officers and crew of the RV
Antea for their effort and technical skills during the
expeditions. Thanks are also due to James Maclaine (Natural
History Museum, London) for sharing data, illustrations, and
radiographs of the type specimens for the study of Mincar-
one et al. (2014), which were also relevant in the present
study. We sincerely thank Jon A. Moore and John Paxton for
providing insightful comments and for sharing unpublished
information on stephanoberycoids that substantially im-
proved the manuscript. We further extend our gratitude to
Marcelo Britto and Cristiano Moreira (MNRJ) for loan of
specimens, and Jørgen Nielsen (ZMUC) for providing
literature. The first author received a student scholarship
from the ‘‘Programa Institucional de Bolsas de Inicia¸ca
˜o
Cient´
ıfica–Universidade Federal do Rio de Janeiro’’ (PIBIC/
UFRJ) during the study. Leandro Eduardo was supported by
CAPES (grant 88882.436215/2019-01), CAPES-Print (grant
88887.364976/2019-00), and FUNBIO/HUMANIZE (‘Progra-
ma Bolsas Funbio–Conservando o Futuro’–011/2019). Fabio
Di Dario, F. Lucena-Fr´
edou, and M. Mincarone are supported
by CNPq (grants PROTAX 443302/2020, 308554/2019-1, and
314644/2020-2, respectively). This study is a contribution to
the LMI TAPIOCA, program CAPES/COFECUB (88881.
142689/2017-01), and EU H2020 TRIATLAS project under
Grant Agreement 817578. The NPM Fish Collection has been
supported by the project Multipesca (FUNBIO) under the
grant ‘‘Pesquisa Marinha e Pesqueira’’ (104/2016). This study
is part of the final paper developed by the first author to
obtain a bachelor’s degree in Biological Sciences at the
Universidade Federal do Rio de Janeiro. We are especially
grateful to members of the examination board, Marcelo
Britto (MNRJ), and Luciano Fischer (NUPEM/UFRJ) for their
valuable suggestions.
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