Ichthyol Res (2011) 58:155–160
DOI 10.1007/s10228-011-0205-7

FULL PAPER

Epigonus carbonarius, a new species of deepwater cardinalfish(Perciformes: Epigonidae) from the Marquesas Islands,
with a redefinition of the Epigonus oligolepis group
Makoto Okamoto • Hiroyuki Motomura

Received: 14 November 2010 / Revised: 13 January 2011 / Accepted: 13 January 2011Ó The Ichthyological Society of Japan 2011

Abstract A new epigonid fish, Epigonus carbonarius, isdescribed on the basis of four specimens (50.8–95.3 mmstandard length) collected from off Nuku Hiva Island,
Marquesas Islands. This species belongs to the Epigonus
oligolepis group, redefined in this study. It is distinguishedfrom the three recognized species of the group by thefollowing combination of characters: a small number of
total gill rakers (21–23) and pyloric caeca (6), presence of
lingual teeth, and absence of teeth from posterior partof vomer. A key to the species in the E. oligolepis group isprovided.
Keywords Epigonus carbonarius  Lingual teeth  Rib 
Nuku Hiva Island

Introduction
The genus Epigonus Rafinesque 1810 is distributed fromtemperate to tropical waters in the world, with 29 valid
species (Mayer 1974; Okamoto 2011). In a worldwide
taxonomic review of the deepwater cardinalfish genus
Epigonus, Abramov (1992) proposed four species groups,
the Epigonus denticulatus group, the Epigonus oligolepis
group, the Epigonus robustus group, and the Epigonus
telescopus group. Of these, the E. oligolepis group was the
M. Okamoto (&)
Seikai National Fisheries Research Institute,1551-8 Taira-machi, Nagasaki 851-2213, Japane-mail: okamako@affrc.go.jp
H. Motomura
The Kagoshima University Museum, 1-21-30 Korimoto,Kagoshima 890-0065, Japane-mail: motomura@kaum.kagoshima-u.ac.jp

smallest species group, including Epigonus devaneyi Gon
1985, Epigonus glossodontus Gon 1985, and Epigonus
oligolepis Mayer 1974. Abramov (1992) defined this groupby the following characters: ‘‘small size (up to 130 mm),large scales (33–39 in lateral line), the occurrence of 7
spines on the 1st dorsal fin and 10 soft rays in the 2nd
dorsal fin as well as by the absence of an opercular spine.’’However, he did not examine the specimens of the members of the group in the revision, and proposed the taxonomy of the genus without an important diagnostic
character, viz. presence or absence of a rib on the last
abdominal vertebra (see Mochizuki and Shirakihara 1983).
While investigating epigonid fish specimens for taxonomic study (Ida et al. 2007; Okamoto 2011), the firstauthor found four specimens of a new species from theMarquesas Islands. This species is described herein, andthe E. oligolepis group is redefined based on examinationof all members of the group. A key to the species of theE. oligolepis group is provided.

Materials and methods
Meristic and morphometric methods generally followed
Mayer (1974). Body depth and body width were measured
at the pectoral-fin base. Lateral-line scales were estimatedby counting scale pockets when specimens had lost theirscales. The number of pored lateral-line scales on the caudal
fin is represented as ‘‘? n’’. The term ‘‘mustache-like process’’ is used for a process on the maxillary head (see Mayer
1974). The first caudal vertebra is defined as the first vertebra bearing a definite hemal spine. Measurements weremade with digital calipers to the nearest 0.1 mm. Terminology and formula of the supraneural bones followed
Mabee (1988) and Ahlstrom et al. (1976), respectively.

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Counts of supraneurals, vertebrae, and ribs were taken fromradiographs. Sex was determined by dissection of theabdomen on the right side. Standard length is abbreviated as
SL. Institutional codes follow Leviton et al. (1985).

Epigonus carbonarius sp. nov. (New English name:Charcoal Deepwater Cardinalfish) (Figs. 1, 2a, e, 3;
Table 1).
Holotype. MNHN 2010-983: 95.3 mm SL, female,
08°580 S, 140°040 W, off Nuku Hiva Island, Marquesas
Islands, 391–408 m depth, 24 August 1997.
Paratypes. Three specimens collected with the holotype: MNHN 2010-984, 89.7 mm SL, female; MNHN
2010-985, 67.7 mm SL, sex unknown; MNHN 2010-986,
50.8 mm SL, sex unknown.
Diagnosis. A species of Epigonus with the following
combination of characters: opercular spine absent; ribabsent on the last abdominal vertebra; lingual teeth present;
vomerine teeth patch on anterior part, not extending posteriorly along midline of palate; mustache-like processabsent; gill rakers 5–6 ? 16–17 = 21–23; vertebrae
10 ? 15; pored lateral-line scales 35–36 to end of hypural

M. Okamoto, H. Motomura

plus 4–5 on caudal fin; dorsal fin VII-I, 10; anal fin II, 9;pyloric caeca 6.
Description. Counts and proportional measurements are
given in Table 1. Body moderately slender, compressed,
nape not humped, deepest at pectoral-fin base. Head large,slightly compressed. Mustache-like process absent. Snout
short and round, length shorter than interorbital width; twonostrils closely set in front of upper edge of pupil, anterior
nostril short and with membranous tube, posterior nostril
elliptical without dermal flap. Eye large, round, orbital
diameter slightly shorter than postorbital length; bony rimof orbit raised above dorsal profile; interorbital region flat.
Mouth large, gape oblique; posterior margin of maxilla
reaching to below anterior margin of pupil; lower jaw notprojecting when mouth is closed, anterior projecting teeth
or nub-like structure absent on symphysis of lower jaw.
Small conical teeth on both jaws and palatines, upper jaw
and palatines teeth arranged in a single row, lower jaw
teeth arranged in a single row but in two or three rows at
symphyseal part. Vomerine teeth minute, 2–5 teeth
restricted to anterior part (Fig. 2a). Lingual teeth present,
villiform, forming v-shaped tooth patch (Fig. 2e). Opercular spine absent, not forming median ridges; preopercularedges smooth. Origin of first dorsal fin vertically above

Fig. 1 Epigonus carbonarius sp. nov., MNHN 2010-983, holotype, 95.3 mm SL

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New epigonid from Marquesas Islands

Fig. 2 Dentitions of vomer and palatine (upper part) and basihyal(lower part) of Epigonus oligolepis group. a, e Epigonus carbonarius
sp. nov., MNHN 2010-983, 95.3 mm SL, holotype; b, f E. devaneyi,

Fig. 3 X-ray photograph of abdominal region of Epigonus carbonarius sp. nov., MNHN 2010-983, holotype. Arrow shows the 10th
abdominal vertebra. Bar 5 mm

anterior portion of pectoral fin; first dorsal-fin spine minute; third dorsal-fin spine the longest. Spine of second
dorsal fin long, thicker than first dorsal-fin spines. First andsecond dorsal fins widely separated by a distance longer

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BPBM 30277, 68.5 mm SL, paratype; c, g E. glossodontus, BPBM
30278, 54.3 mm SL, paratype; d, h E. oligolepis, USNM 207719,
77.6 mm SL, paratype. Bars 1 mm

than snout length. Origin of anal fin vertically below posterior portion of second dorsal-fin base; first anal spine
minute; second spine fairly thick. Posterior tips of pectoraland pelvic fins not reaching vertical line from anus. Caudal
fin deeply forked. Anus located vertically below first soft
ray of second dorsal fin. Abdominal vertebrae 10, rib absent
on last vertebra (Fig. 3); caudal vertebrae 15. Supraneuralbones three (0?0/0?2/1?1/1). Scales deciduous, weakly
ctenoid, covering the whole body except snout tip, anterior
to rim of orbit and surface of jaws; scales also present on
bases of second dorsal, anal, and caudal fins; series of pored
lateral-line scales complete, 4–5 pored scales on caudal fin.
No trace of luminous organ around belly or visceral organ.
Color in alcohol. Body and head uniformly dark brown.
All fins black, except for pectoral fin (light brown). Mouth
cavity pale with scattered small melanophores.
Distribution. All specimens were collected from off
Nuku Hiva Island, Marquesas Islands, at a depth of
391–408 m.
Etymology. The specific name, carbonarius, is a Latinnoun (meaning ‘‘charcoal’’) referring to the blackish coloration of the body and fins of the new species.
Comparison. Epigonus carbonarius sp. nov. belongs
to a group of three similar species in the E. oligolepis
group (see ‘‘Remarks’’); E. devaneyi, E. glossodontus, andE. oligolepis (Fig. 4). Although E. carbonarius is similar
to E. glossodontus and E. oligolepis in having lingual teeth

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M. Okamoto, H. Motomura

Table 1 Counts and measurements of Epigonus carbonarius sp. nov.
Holotype
MNHN
2010-983

Paratypes(n = 3)

95.3

50.8–89.7

Dorsal-fin rays
Anal-fin rays

VII-I, 10
II, 9VII-I, 10II, 9Pectoral-fin rays

17

17

Pored lateral-line scales

36 ? 5

35 ? 4Standard length (mm)CountsScales above lateral line3

3

Scales below lateral line

78Gill rakers

5 ? 16 = 21

5–6 ? 16–17= 21–23

Pyloric caeca

66Vertebrae

10 ? 15

10 ? 15

Head length

34.7

35.8–38.2

Head width
Head height

14.4
16.1

13.0–15.1
16.6–17.1

Body depth

21.6

21.9–23.1

Body width

12.8

11.7–14.6

Caudal-peduncle depth

9.9

9.6–10.6

Caudal-peduncle length

29.1

25.0–28.4

Orbital diameter

13.6

13.4–15.2

Interorbital width

9.1

8.9–9.3

Postorbital length

14.1

14.2–15.9

Upper-jaw length

13.7

13.6–14.6

Lower-jaw length

14.8

15.5–17.0

Snout length

8.0

8.1–8.7

Pre-1st dorsal-fin length

37.4

36.4–39.7

Pre-2nd dorsal-fin length

57.7

56.1–60.7

Measurements (% standard length)

Pre-pectoral-fin length

34.4

34.8–35.8

Pre-pelvic-fin lengthPre-anus-length

36.5
55.3

36.6–39.0
56.9–58.0

Pre-anal-fin length

65.6

64.6–66.3

1st spine length on 1st dorsal fin

2.4

2.1–3.9

2nd spine length on 1st dorsal fin

14.4

14.3–17.7

3rd spine length on 1st dorsal fin

15.7

14.5–16.8

2nd dorsal-fin spine length

14.1

14.0–15.0

1st anal-fin spine length

2.2

2.5–3.3

2nd anal-fin spine length

13.1

13.9–14.4

Pelvic-fin spine length

13.2

13.0–13.3

1st dorsal-fin base

15.8

13.2–15.9

2nd dorsal-fin base

13.0

12.0–12.7

Anal-fin base

11.1

10.2–10.8

Pectoral-fin length

18.8

18.6–22.2

Pelvic-fin length

18.8

18.1–18.5

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Fig. 4 Other three species of Epigonus oligolepis group. a Epigonus
devaneyi, BPBM 30277, 68.5 mm SL, paratype; b E. glossodontus,
BPBM 40445, 78.0 mm SL; c E. oligolepis, USNM 207719, 77.6 mm
SL, paratype

Fig. 5 Distribution records of the Epigonus oligolepis group. Epigonus carbonarius sp. nov. (star); E. devaneyi (triangle); E. glossodontus (circle); E. oligolepis (squares)(Fig. 2), the new species differs in having 21–23 total gillrakers (vs. 24–27 and 29–32 in E. glossodontus andE. oligolepis, respectively) and vomerine teeth not extending posteriorly (vs. teeth present posteriorly along midlineof palate in the latter two species; Fig. 2). The new species
also clearly differs from E. devaneyi in having lingual teeth(vs. lingual teeth absent in E. devaneyi, Fig. 2) and fewer
gill rakers and pyloric caeca (21–23 and 6 in E. carbonarius vs. 26–29 and 8–10 in E. devaneyi). In addition, the
new species is distributed in the South Pacific, although allother members of the species group have been reported
only from the Northern Hemisphere (Fig. 5). Selectedcharacters of the four species are compared in Table 2.

New epigonid from Marquesas Islands

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Table 2 Comparison of the selected characters and distribution among four species of the Epigonus oligolepis group
Species
Sources

E. carbonarius sp. nov.
Present study

E. devaneyi
Present study,
Gon (1985)

E. glossodontus
Present study,
Gon (1985)

E. oligolepis
Present study,
Mayer (1974)

Lingual teeth

Present

AbsentPresent

Present

Vomerine teeth on posterior part

AbsentAbsentPresent

Present

Anterior projecting teeth
on lower jaw

AbsentAbsentPresent

AbsentGill rakers (total)

21–23

26–29

24–27

29–32

Pyloric caeca

6

8–10

6–8

8–10

Distribution

Marquesas Islands

Hawaiian Islands

Hawaiian Islands

Gulf of Mexico to
Caribbean region

Remarks. Although Abramov (1992) recognized four
species groups in the genus Epigonus, he did not examinematerial of all members of the E. oligolepis group. In
addition, he did not discuss the importance of the presenceor absence of a rib on the last abdominal vertebra, although
Mochizuki and Shirakihara (1983) considered it to be an
important diagnostic character in the genus. As a result,
there are species that do not fall under the definition of anyof the four species group of the genus (treated as exceptions; Abramov 1992), and the limits between the four
species groups are not clear. Our examination of thepresence or absence of the rib on the last abdominal vertebra in the E. oligolepis group revealed that all membersof the group lack the rib. On the other hand, presence orabsence of this rib is variable within the other three species
groups (Mochizuki and Shirakihara 1983; Okamoto 2011).
The holotype (95.3 mm SL) and one paratype (89.7 mm
SL, MNHN 2010-984) of E. carbonarius are females with
developed mature ova. Gon (1985) reported the largest sizeof E. devaneyi and E. glossodontus of the E. oligolepis
group as 78.3 mm SL and 100.0 mm SL, respectively.
Epigonus oligolepis was found in this study to reach atleast 150 mm SL (based on one specimen, UF 43252).
With the exception of a small species, Epigonus cavaticus
Ida, Okamoto and Sakaue 2007 (\100 mm SL), adultspecimens of the other species groups are known to attain
100–350 mm SL (Abramov 1992; Okamoto 2011). Thus,
the E. oligolepis group appears to be a group with relatively small-sized species.
Epigonus carbonarius was collected from a depth of
391–408 m in the Marquesas Islands. All other species of
the E. oligolepis group occur in depths of more than 300 m(Mayer 1974; Gon 1985).

The Epigonus oligolepis group
The Epigonus oligolepis group: Abramov 1992: 100

Diagnosis. Pungent opercular spine absent; rib absent onlast abdominal vertebra; dorsal fin VII-I, 10; and 33–40
pored lateral-line scales to end of hypural.
Key to the species of the Epigonus oligolepis group
1a. Lingual teeth absent (Hawaiian Islands)  E. devaneyi
1b. Lingual teeth present 2
2a. Total gill rakers 21–23; vomerine teeth patch on
anterior part, not extending posteriorly along midline ofpalate (Marquesas Islands)  E. carbonarius sp. nov.
2b. Total gill rakers 24–31; vomerine teeth patch on
anterior part and extending posteriorly along midline ofpalate 3
3a. Maxillary teeth small and conical, visible when mouth
is closed; two or three large anterior projecting teeth on theeach side of the symphysis of the lower jaw (individuals
larger than 50 mm SL); total gill rakes 24–27 (Hawaiian
Islands) E. glossodontus
3b. Maxillary teeth minute, not visible when mouth isclosed; large teeth absent on the symphysis of the lower
jaw; total gill rakes 29–32 (Gulf of Mexico to Caribbean
region)E. oligolepis
Comparative materials. Epigonus devaneyi: BPBM
30277, paratype, 1 specimen, 68.5 mm SL, off south end of
Necker Ridge, northwestern Hawaiian Islands, 311–347 m
depth, 15 October 1976; HUMZ 129680, 1 specimen,
40.7 mm SL, 24°540 N, 165°360 W, off Hawaii, February
1994. E. glossodontus: BPBM 30278, paratypes, 2 specimens, 39.6–54.3 mm SL, off Pearl Harbor, Mamala Bay,Oahu, Hawaii, 1 September 1982; BPBM 40445, 1 specimen, 78.0 mm SL, 20°570 N, 157°230 W, Penguin Bank, off
Molokai, Hawaii, 30 October 2002. E. oligolepis: UF
43252, 6 specimens, 104.5–150.3 mm SL, 26°110 N,
84°430 W, Gulf of Mexico, Florida, 300 m depth, 19 September 1985; USNM 207719, paratypes, 2 specimens,

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52.0–77.6 mm SL, 12°010 N, 61°530 W, Caribbean Sea,
384–457 m depth, 4 September 1974.
Acknowledgments We are especially grateful to G. Duhamel and
R. Causse (MNHN) for opportunity to examine the specimens of the
new species. We also thank A.Y. Suzumoto (BPBM), M. Yabe and
T. Kawai (HUMZ), L.M. Page and R.H. Robins (UF), and J.T. Williams
and E. Wilbur (USNM) for loans of comparative materials, and O. Gon(South African Institute for Aquatic Biodiversity) for his criticalreading of this manuscript.

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