Ichthyol Res
DOI 10.1007/s10228-013-0352-0

FULL PAPER

Two new species of deepwater cardinalfish from the Indo-Pacific,
with a definition of the Epigonus pandionis group (Perciformes:
Epigonidae)
Makoto Okamoto • Hiroyuki Motomura

Received: 29 March 2013 / Revised: 9 May 2013 / Accepted: 9 May 2013
 The Ichthyological Society of Japan 2013

Abstract Two new Indo-Pacific species of deepwater
cardinalfish, Epigonus lifouensis and E. tuberculatus aredescribed based on the specimens collected from the
Loyalty Islands and Cocos-Keeling Islands, respectively.
These species belong to the Epigonus pandionis groupdefined as lacking an opercular spine, having more than 43
pored lateral-line scales to the end of the hypural and dorsalfin rays VII-I, 9–11. Epigonus lifouensis is distinguishedfrom other members of the group by a combination of thefollowing characters: ribs present on the last abdominal
vertebra; tongue toothless; tubercle absent on inner symphysis of lower jaw; eye elliptical; total gill rakers 24–25;
pectoral-fin rays 18–19; pyloric caeca 10–13; body depth
17.0–17.1 % SL; and posterior half of oral cavity and tongue black. Epigonus tuberculatus is distinguished fromother members of the group by a combination of the following characters: ribs on the last abdominal vertebra
reduced or absent; tongue toothless; tubercle present on
inner symphysis of lower jaw; total gill rakers 21–22;
pectoral-fin rays 19–20; pyloric caeca 8–10; orbital

This article was registered in the Official Register of Zoological
Nomenclature (ZooBank) as
A987BB74-B30C-4A79-9E77-D4E1AC200436.
This article was published as an Online First article on the online
publication date shown on this page. The article should be cited byusing the doi number.
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

diameter 14.5–15.4 % SL; and lower-jaw length
16.0–17.6 % SL. A key to the species and some commentson the group are provided based on examination of all
members (nine species, including two new species) of thegroup.
Keywords Deepwater cardinal fish  Epigonus 
New Caledonia  Australia

Introduction
Fishes of the genus Epigonus Rafinesque 1810 (deepwater
cardinalfish) are distributed from temperate to tropical waters
in the world (Mayer 1974; Okamoto 2012; Okamoto andMotomura 2012). In a recent taxonomic study of the genus
Epigonus, Okamoto (2012) proposed four species groups:
the Epigonus constanciae group (Okamoto 2012), theEpigonus oligolepis group (Okamoto and Motomura 2011),
the Epigonus telescopus group (Abramov 1992), and an
unnamed group (Okamoto 2012). In the last group, Okamoto(2012) indicated that the group consists of seven species, viz.,
Epigonus denticulatus Dieuzeide 1950; Epigonus elongatus
Parin and Abramov 1986; Epigonus cavaticus Ida, Okamotoand Sakaue 2007; Epigonus fragilis (Jordan and Jordan
1922); Epigonus marisrubri Krupp, Zajonz and Khalaf 2009;
Epigonus pandionis (Goode and Bean 1881); and Epigonus
parini Abramov 1987, and is characterized by the following
combination of characters: opercular spine absent, more than
43 pored lateral-line scales to end of hypural, and VII-I, 9–11
dorsal-fin rays.
During an investigation of material of the genus fromthe Indo-Pacific, two undescribed species having theabove-mentioned characters collected from Lifou Island,
the Loyalty Islands and the Cocos-Keeling Islands were123

M. Okamoto, H. Motomura

found. Herein, these species are described as two new
species, and the species group is defined as ‘‘the Epigonus
pandionis group’’. Also, a key to the species and somecomments of the group are provided based on examinationof all members of the group.

Materials and methods
Meristic and morphometric methods followed Mayer (1974)
and Okamoto (2011). Missing lateral-line scales were estimated by counting scale pockets. The number of pored lateralline scales on the caudal fin is represented as ‘‘?n’’. The firstcaudal vertebra is defined as the first vertebra bearing a definite hemal spine. Measurements were made with calipers to
the nearest 0.1 mm. The formula for the configuration of thesupraneural bones, anterior neural spines, and anterior dorsal
fin pterygiophores follows Ahlstrom et al. (1976). Counts ofsupraneurals, vertebrae, and ribs were taken from radiographs.
The term ‘‘maxillary mustache-like process’’ is used for aprocess on the maxillary head (see Okamoto 2011: fig. 3). The
number of pyloric caeca and sex were established by dissection of the abdomen on the right side. Standard length and headlength are abbreviated as SL and HL, respectively. Institutional codes follow Sabaj Pérez (2012).
Epigonus lifouensis sp. nov.
(New English name: Loyalty Deepwater Cardinalfish)(Figs. 1, 2a)Holotype. MNHN 2000-0018, 139.7 mm SL, male,
21400 5900 S, 167310 5900 E, south of Lifou Island, Loyalty
Islands, New Caledonia, western South Pacific, 575 m
depth, beam trawl, 21 February 1989.
Paratype. MNHN 2003-1826, 202.5 mm SL, female,21230 3100 S, 167460 1000 E, south of Lifou Island, Loyalty
Islands, New Caledonia, western South Pacific, 500 m
depth, Waren dredge, 23 February 1989.
Diagnosis. A species of Epigonus with the following
combination of characters: opercular spine absent; ribspresent on the last abdominal vertebra; tongue toothless;maxillary mustache-like process absent; tubercle absent on
Fig. 1 Epigonus lifouensis sp.
nov., MNHN 2000-0018,
holotype, 139.7 mm SL

123

inner symphysis of lower jaw; eye elliptical; total gillrakers 24–25; vertebrae 10 ? 15; pored lateral-line scales
48–49 ? 3–4; dorsal-fin rays VII-I, 10; pectoral-fin rays
18–19; pyloric caeca 10–13; body depth 17.0–17.1 % SL;
and posterior half of oral cavity and tongue black.
Description. Counts and proportional measurements are
given in Table 1. Data for the holotype are presented first,
followed by data of paratype (if different) in parentheses.
Body slender, laterally compressed, nape not humped,
deepest at pectoral-fin base. Head large and thick. Maxillarymustache-like process absent. Snout short and round; twonostrils close-set without dermal flap, elliptical, posterior
one slightly larger. Eye large, elliptical, diameter greaterthan postorbital length; bony rim of orbit slightly raised
above dorsal profile; interorbital region flat. Mouth large,
terminal, gape oblique; posterior margin of maxilla
extending to below anterior margin of pupil; lower jaw notprojecting when mouth closed; tubercle absent on innersymphysis of lower jaw. Teeth minute, two or three rows onanterior half of upper jaw, toothless at symphysis; one or
two rows on all of lower jaw but in three rows at symphyseal part. Villiform teeth on vomer and palatines. Tongue
toothless. Opercular spine absent; preopercle without
spines, ventral and posterior margins smooth. Origin of firstdorsal fin vertically above anterior portion of pelvic fin; firstdorsal-fin spine minute; two supernumerary spines on firstdorsal-fin pterygiophore. Spine of second dorsal fin weak
and short. Gap between dorsal fins longer than snout length.
Origin of anal fin vertically below posterior portion of
second dorsal-fin base; first anal-fin spine minute; second
spine long, length longer than length of first spine on firstdorsal fin; two supernumerary spines on first anal-fin
pterygiophore. Posterior tip of pectoral fin not reaching tovertical line from anus (tip broken). Pelvic fin short, posterior tip not reaching to anus. Caudal fin deeply forked.
Anus located anteriorly vertically below second dorsal-fin
origin. Ribs present on last abdominal vertebra. Supraneuralbones 3; formula for configuration of supraneural bones,
anterior neural spines, and anterior dorsal fin pterygiophores /0 ? 0/0 ? 2/1 ? 1/1/. Scales deciduous, cycloid,
covering the whole body except snout tip, anterior to rim oforbit, and surface of jaws; scales also present on basesof second dorsal, anal, and caudal fins; series of pored

Two new species of Epigonus

Fig. 2 Lateral view of vertebrae of abdominal region of a Epigonus
lifouensis sp. nov., MNHN 2000-0018, holotype, 139.7 mm SL,b Epigonus tuberculatus sp. nov., CSIRO H 5677-08, holotype,154.0 mm SL, and c Epigonus tuberculatus sp. nov., CSIRO H
5677-09, paratype, 132.3 mm SL

lateral-line scales complete, 4 (3) pored scales on caudal fin.
No trace of luminous organ around belly or visceral organs.
Color in alcohol (Fig. 1). Body, head, and all fins uniformly light brown, except for the posterior half of oralcavity and tongue which are black. All fins with manymelanophores on the soft rays and membranes.
Distribution. The holotype and paratype were collectedfrom south of Lifou Island, Loyalty Islands, New Caledonia, at depths of 500–575 m (Fig. 3).
Etymology. The specific name lifouensis refers to the
locality of the type specimens.
Comparison. Epigonus lifouensis belongs to a group ofsimilar species in the Epigonus pandionis group (present
study). Epigonus lifouensis is most similar to E. elongatus;
however, E. lifouensis is distinguished from E. elongatusin having ribs on the last abdominal vertebra (Fig. 2a)(vs. lacking in E. elongatus), and higher gill rakers andpyloric caeca counts (24–25 and 10–13 vs. 21–22 and 7–9
in E. elongatus, respectively). Epigonus cavaticus andE. fragilis also have 24–27 total gill rakers but differ fromE. lifouensis in having lower pectoral-fin ray and pyloric
caeca counts (16–17 and 7–8 vs. 18–19 and 10–13 in
E. lifouensis, respectively). Furthermore, E. lifouensis differs from E. denticulatus, E. marisrubri, E. pandionis, andE. parini in having fewer gill rakers (24–25 vs. 28–34 in
E. denticulatus, 29–30 in E. marisrubri, 26–30 in
E. pandionis, and 35–39 in E. parini), elliptic eye (vs.
round), and black tongue and oral cavity (vs. tan or lightbrown). In addition to the above-mentioned differences,
E. marisrubri has a tooth patch on the tongue (vs. absent in
E. lifouensis), and E. pandionis has a greater body
depth (24.0–28.6 % SL in E. pandionis vs. 17.0–17.1 %SL in E. lifouensis). Epigonus lifouensis is distinguishedfrom the second new species in the present study,
E. tuberculatus (Fig. 4), in having developed ribs on thelast abdominal vertebra (vs. ribs absent or reduced on
this vertebra in E. tuberculatus; Fig. 2b, c), lacking
tubercle on inner symphysis of lower jaw (vs. having in
E. tuberculatus; Fig. 5), and higher gill rakers count(24–25 vs. 21–22 in E. tuberculatus).
Remarks. Epigonus lifouensis had ribs on the last
abdominal vertebra (Fig. 2a). Mochizuki and Shirakihara(1983) regarded the presence or absence of a pair of ribs on
the last abdominal vertebra as an important diagnostic
character in this genus. However, in reviewing Epigonus,
Abramov (1992) proposed the taxonomy of the genus
without the character. In the present study, it is clear thatthe condition of ribs on the last abdominal vertebra in theE. pandionis group is important based on examination of
all members (Table 2).

123

M. Okamoto, H. Motomura
Table 1 Counts and measurements of Epigonus lifouensis sp. nov. and Epigonus tuberculatus sp. nov.
Epigonus lifouensis

Epigonus tuberculatus

Holotype MNHN
2000-0018

Paratype MNHN
2003-1826

Holotype CSIRO H
5677-08

Paratype CSIRO H
5677-09

139.7

202.5

154.0

132.3

Dorsal-fin rays
Anal-fin rays

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

18

19

19

20
48 ? 3Standard length (mm)CountsPored lateral-line scales

48 ? 349 ? 446 ? 4

Scales above lateral line

3

3

3

3

Scales below lateral line

9

9

9

9
5 ? 17 = 22

Gill rakers

7 ? 18 = 25

6 ? 18 = 24

5 ? 16 = 21

Pyloric caeca

13

10

8

10

Vertebrae

10 ? 15

10 ? 15

10 ? 15

10 ? 15

Head length

32.4

32.1

34.4

32.7

Head width

18.0

18.7

17.8

16.3

Head height

14.7

15.4

16.5

15.6

Body depth

17.0

17.1

18.8

17.8

Measurements (% standard length)

Body width

16.0

18.0

17.8

16.8

Caudal-peduncle depth
Caudal-peduncle length

7.1
24.3

6.6
24.2

7.7
25.2

7.9
27.1

Orbital diameter

14.5

13.8

15.4

14.5

Interorbital width

8.4

7.8

8.4

9.1

Postorbital length

10.1

11.5

10.4

10.4

Upper-jaw length

14.4

14.1

13.9

12.7

Lower-jaw length

17.6

17.1

17.6

16.0

Snout length

8.2

8.4

10.8

7.5

Pre-1st dorsal-fin length

39.2

38.7

40.1

38.1

Pre-2nd dorsal-fin length

62.6

62.5

61.9

60.9

Prepectoral fin length

32.4

32.8

34.8

31.9

Prepelvic fin length

36.9

39.1

36.7

35.5

Preanus length

55.2

59.5

57.8

57.6

Preanal fin length

66.5

68.2

68.7

66.7

1st spine length on 1st dorsal fin

3.7

Broken

3.3

Broken

2nd spine length on 1st dorsal fin

BrokenBroken8.8

Broken

3rd spine length on 1st dorsal fin
2nd dorsal-fin spine length

BrokenBrokenBroken
4.2

Broken
4.5

BrokenBroken

1st anal-fin spine length

1.9

2.9

1.4

2.2

2nd anal-fin spine length

Broken

4.1

4.4

5.3

Pelvic-fin spine length

Broken7.7

7.5

Broken

1st dorsal-fin base

12.2

15.3

11.2

10.8

2nd dorsal-fin base

9.4

9.4

9.5

10.4

Anal-fin base

7.9

7.6

8.9

8.8

Pectoral-fin length

Broken

12.2

16.6

BrokenPelvic-fin length

Broken9.8

BrokenBroken123

Two new species of Epigonus

Fig. 3 Collection localities of Epigonus lifouensis sp. nov. (circles)
and E. tuberculatus sp. nov. (star)

The holotype of E. lifouensis was a male with a fullyripe testis. The paratype was a female with a great numberof eggs in several developmental stages. The most developed eggs were ca. 0.4 mm in diameter, and were roundwith a single oil globule.
Epigonus tuberculatus sp. nov.
(New English name: Keeling Deepwater Cardinalfish)(Figs. 2b, c, 4, 5)
Holotype. CSIRO H 5677-08, 154.0 mm SL, female,
14450 S, 95240 E–14450 S, 95220 E, south of Cocos-Keeling Islands, Indian Ocean, 473–570 m depth, 11 April
2001, trawl, collected by M. Scott.
Paratype. CSIRO H 5677-09, 132.3 mm SL, female,same data as holotype.
Diagnosis. A species of Epigonus with the following
combination of characters: opercular spine absent; ribs on
the last abdominal vertebra reduced or absent; tongue
toothless; maxillary mustache-like process absent; tubercle
present on inner symphysis of lower jaw; total gill rakers
21–22; vertebrae 10 ? 15; pored lateral-line scales
46–48 ? 3–4; dorsal-fin rays VII-I, 10; pectoral-fin rays
19–20; pyloric caeca 8–10; orbital diameter 14.5–15.4 %SL; and lower-jaw length 16.0–17.6 % SL.
Description. Counts and proportional measurements are
given in Table 1. Data for the holotype are presented first,
followed by data of paratype (if different) in parentheses.

Body slender, laterally compressed, nape not humped,
deepest at pectoral-fin base. Head large and thick. Maxillary mustache-like process absent. Snout short and round;
two nostrils close-set without dermal flap, elliptical, posterior one slightly larger. Eye large, round, diameter greaterthan postorbital length; bony rim of orbit slightly raised
above dorsal profile; interorbital region flat. Mouth large,
terminal, gape oblique; posterior margin of maxilla
extending to below anterior margin of pupil; lower jaw notprojecting when mouth closed; developed tubercle present
on inner symphysis of lower jaw (smaller tubercle). Teeth
minute, in one or two rows on anterior portion of upper
jaw, toothless at symphysis; one or two rows on all of
lower jaw with a pair of small conical teeth at symphyseal
part. Small conical teeth on vomer and palatines. Tongue
toothless. Opercular spine absent, not forming medianridges; preopercle without spines, ventral and posterior
margins smooth. Origin of first dorsal fin vertically above
anterior portion of pelvic fin; first dorsal-fin spine minute;two supernumerary spines on first dorsal-fin pterygiophore.
Spine of second dorsal fin short, slightly thicker than firstdorsal-fin spines (broken). Gap between dorsal fins longer
than snout length. Origin of anal fin vertically below posterior portion of second dorsal-fin base; first anal-fin spine
minute; second spine short, length subequal to spine ofsecond dorsal fin; two supernumerary spines on first analfin pterygiophore. Posterior tip of pectoral fin not reaching
to vertical line from anus. Pelvic fin short, not reaching toanus. Caudal fin deeply forked. Anus located anteriorly vertically below second dorsal-fin origin. Reduced ribs present onlast abdominal vertebra (ribs absent). Supraneural bones 3;
formula for configuration of supraneural bones, anterior
neural spines, and anterior dorsal fin pterygiophores /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 (3) pored scales on caudal fin. No trace of luminous organ around belly or visceral organs.
Color in alcohol (Fig. 4). Body, head, and caudal fin
uniformly dark brown, except for the lips and oral cavity
which are tan. All fins except caudal fin light brown, withmany melanophores on the soft rays and membranes.

Fig. 4 Epigonus tuberculatus
sp. nov., CSIRO H 5677-08,
holotype, 154.0 mm SL (taken
by T. Carter)123

M. Okamoto, H. Motomura

Fig. 5 Tip of mouth of Epigonus tuberculatus sp. nov., CSIRO H
5677-08, holotype, 154.0 mm SL. a Front view, b lateral view.
Arrows show the tubercle of lower jaw. Bars 10 mm

Distribution. The holotype and paratype were collectedfrom south of the Cocos-Keeling Islands, Indian Ocean, at
a depth of 473–570 m (Fig. 3).
Etymology. The specific name tuberculatus is from theLatin tuber meaning swelling, in reference to the tubercle
found on the inner symphysis of the lower jaw (Fig. 5).
Comparison. Epigonus tuberculatus belongs to a groupof similar species in the Epigonus pandionis group (present
study). The holotype of Epigonus tuberculatus has atubercle on inner symphysis of lower jaw and can be distinguished from other members by that character. On theother hand, the smaller paratype of this species (132.3 mmSL) had an undeveloped tubercle compared with holotype(154.0 mm SL). Thus, there may be an ontogenetic change
in this character. However, E. tuberculatus can be distinguished from E. elongatus by having a greater orbital
diameter and lower-jaw length (14.5–15.4 % SL and123

16.0–17.6 % SL vs. 12.1–13.6 % SL and 13.4–15.1 % SL
in E. elongatus, respectively). Furthermore, E. tuberculatus
differs from E. denticulatus, E. marisrubri, E. parini, andE. pandionis in having fewer gill rakers (21–22 vs. 28–34 in
E. denticulatus, 29–30 in E. marisrubri, 35–39 in E. parini,and 26–30 in E. pandionis) and ribs absent or reduced on thelast abdominal vertebra (Fig. 2b, c) (vs. developed ribspresent in the latter species). In addition to the above-mentioned differences, E. tuberculatus is distinguished from
E. marisrubri and E. pandionis in having a toothless tongue(vs. tooth patch present on tongue in E. marisrubri) andshallower body depth (17.8–18.8 % SL vs. 24.0–28.6 %SL in E. pandionis), respectively. The other two speciesof the group, E. cavaticus and E. fragilis, differ fromE. tuberculatus in having developed ribs on the last
abdominal vertebra and 16–17 pectoral-fin rays (vs. ribs
absent or reduced on the last abdominal vertebra and 19–20
pectoral-fin rays in E. tuberculatus). The diagnostic characters between two new species in the present study aredescribed in the ‘‘Comparison’’ of E. lifousensis.
Remarks. The ribs on the last abdominal vertebra areusually present in most species of the genus Epigonus
including members of the E. pandionis group defined in thepresent study (21 species, Okamoto 2012; Okamoto et al.
2012). On the other hand, 10 species of the genus lack theribs on the last abdominal vertebra (Okamoto and Fukui
2011; Okamoto and Motomura 2011; Okamoto et al. 2011).
However, the holotype and paratype of E. lifouensis havethe reduced ribs and lack ribs on the last abdominal vertebra, respectively (Fig. 2b, c). This condition is also foundin Epigonus marimonticolus Parin and Abramov 1986 of
the E. constanciae group (see Okamoto 2012).
The holotype and a paratype of E. tuberculatus werefemales with a great number of eggs at several developmental stages. The most developed eggs were ca. 0.5 mm
in diameter, and were round with a single oil globule.

The Epigonus pandionis group
Included species. Nine species (Figs. 1, 4, 9): Epigonus
cavaticus, E. denticulatus, E. elongatus, E. fragilis,
E. lifouensis, E. marisrubri, E. pandionis, E. parini, andE. tuberculatus.
Diagnosis. Opercular spine absent; more than 43 pored
lateral-line scales to end of hypural; and dorsal fin rays VIII, 9–11.
Remarks. The E. pandionis group comprises nine species and is distinguished from the E. constanciae group (16species; see Okamoto 2012) in lacking an opercular spine(vs. opercular spine present in the E. constanciae group).
Furthermore, the E. pandionis group differs from theE. telescopus group (four species; see Okamoto et al. 2012)

Two new species of Epigonus
Table 2 Comparison of the selected characters among nine species of the Epigonus pandionis group
Species

Tooth patch
on tongue

Ribs onlast AVGill
rakersPyloriccaeca

P1 rays

1st spine
on D1

Sources

E. cavaticus

AbsentPresent

24–27

7–8

16

VisiblePresent study (n = 4); Ida et al. (2007)

E. denticulatus

AbsentPresent

28–34

10–14

18–21

VisiblePresent study (n = 24); Mayer (1974)

E. elongatus

AbsentAbsent

22–23

7–9

18–20

VisiblePresent study (n = 3); Parin and
Abramov (1986)
Present study (n = 5); Mayer (1974)

E. fragilis

AbsentPresent

25–26

7–8

16–17

VisibleE. lifouensis

AbsentPresent

24–25

10–13

18–19

VisiblePresent study (n = 2)

E. marisrubri

Present

Present

29–30

10

17–18

VisiblePresent study (n = 2); Krupp et al. (2009)

E. pandionis

AbsentPresent

26–30

10–13

17–19

VisiblePresent study (n = 44); Mayer (1974)

E. parini

AbsentPresent

35–39

8–11

18–20

Invisible

Present study (n = 1); Abramov (1987)

E. tuberculatus

AbsentAbsent or reduced

21–22

8–10

19–20

VisiblePresent study (n = 2)

AV abdominal vertebra, P1 pectoral fin, D1 first dorsal finin lacking an isolated spine between the first and second
dorsal fins (vs. an isolated spine present in the E. telescopus
group; see Okamoto et al. 2012). The remaining species groupof the genus, viz., the E. oligolepis group (four species; see
Okamoto and Motomura 2011), is characterized by a small
number of pored lateral-line scales (33–40, mostly 35–38 in
the E. oligolepis group vs. more than 43 pored lateral-line
scales in the E. pandionis group).
Most species of the other three species groups ofthe genus Epigonus (the E. constanciae group, theE. oligolepis group, and the E. telescopus group) havebeen reported from continental slopes or seamounts, withrecords ranging from 200 m to depths exceeding 1,000 m(Mochizuki 1990; Gon 1985; Okamoto 2011, 2012;
Okamoto and Motomura 2011, 2012). Similarly, six species of the E. pandionis group, including two new species,
have been collected from deeper than 200 m depth (Mayer
1974; Mochizuki 1990; Abramov 1987, 1992; Mytilineou
et al. 2005; Ruiz-Pico et al. 2012), viz. E. denticulatus,
E. elogatus, E. lifouensis sp. nov., E. pandionis, E. parini,and E. tuberculatus sp. nov. On the other hand, of theremaining three species of the group, E. cavaticus wasreported from a cave facing the drop off of the fringing reefof Palau at a depth of about 20 m (Ida et al. 2007).
Epigonus fragilis is a species endemic to the Hawaiian
Island, which usually occurs at a depth of 120–125 m(Mayer 1974). Recently, a single specimen of the species
was collected by fishing at a depth of 3–4.5 m (BPBM
41101); this is the shallowest record for the genus.
Epigonus marisrubri has been reported from a coral reefarea at a depth of 52 m at the northern tip of the Gulf of
Aqaba, Red Sea (Krupp et al. 2009). In addition, the firstauthor found a single specimen of E. marisrubri collectedfrom the Gulf of Aden at depths of 100–200 m in the
research collection of the National Museum of Natural
History, Smithsonian Institution (USNM 305950). The
specimen represents the first record of the species from theGulf of Aden. Thus, these three species of the group inhabit
the coastal reef area and continental shelf.
In the present study, it is clear that ribs on the last
abdominal vertebrae are absent in E. elongatus (Fig. 6a).
Also, the presence of ribs on the last abdominal vertebra of
E. marisrubri and E. parini is reported for the first time in
the present study (Fig. 6b, c). These characters are thus
useful for distinguishing these species and other related
species (see below). Selected characters of the nine species
are compared in Table 2.
The presence or absence of a tooth patch on the tongue
is an important diagnostic character in the genus (Gon
1985; Okamoto and Motomura 2012). Krupp et al. (2009)
reported that E. marisrubri has a tooth patch on the tongue(Fig. 7), this is unique to E. marisrubri within theE. pandionis group. Epigonus denticulatus rarely present
1–4 min teeth on the tongue, but they are scattered and do
not form a dense tooth patch. According to Okamoto andMotomura (2011) and Okamoto (2012), in other species
groups of the genus, one species of the E. constanciae
group and three species of the E. oligolepis group have thetooth patch on the tongue.
Abramov (1987) described that E. parini has six spines
on the first dorsal fin, which is a unique character among
the genus. However, it is clear that this species has sevenspines on the first dorsal fin. Although the first spine is
obscure because the spine is hidden under the skin of thefirst dorsal fin base (Fig. 8a), it can be distinguished inradiographs of the fish (Fig. 8b).

Key to the species of the Epigonus pandionis group
1a. First spine on first dorsal fin hidden under skin, six
spines on first dorsal fin visible; second dorsal-fin rays I, 9;gill rakers 35–39 (east of New Zealand; Fig. 9g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. parini

123

M. Okamoto, H. Motomura

Fig. 7 Dorsal view of toothed tongue of Epigonus marisrubri,
USNM 305950, 139.8 mm SL

Fig. 6 X-ray photograph of abdomen of a Epigonus elongatus, ZIN
47138, holotype, 160.0 mm SL, b E. marisrubri, SMF 31636,
paratype, 134.0 mm SL, and c E. parini, NSMT-P 41252,
168.0 mm SL. Arrows show the ribs on the last abdominal vertebra,
bearing no ribs in a. AV abdominal vertebra, CV caudal vertebra. Bars
5 mm

1b. First spine on first dorsal fin not hidden under skin,seven spines on first dorsal fin visible; second dorsal-fin
rays I, 10–11; gill rakers 21–34 . . . . . . . . . . . . . . . … 2
2a. Tooth patch of 2–4 rows of dense teeth present on
tongue (Gulf of Aqaba, Red Sea; Gulf of Aden; Figs. 7, 9e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . … E. marisrubri

123

Fig. 8 Lateral view of anterior part of first dorsal fin of Epigonus
parini, NSMT-P 41252, 168.0 mm SL. a Photograph and b X-ray
photograph. Arrow shows the first spine. Bars 10 mm

2b. Tooth patch not on tongue (rarely present in
E. denticulatus, but reduced, a small number of teeth patches of 1–4 teeth scattered on tongue). . . . . . . . . . . . . 3
3a. Gill rakers 21–23; ribs on last abdominal vertebra
reduced or absent (Figs. 2b, c, 6a) . . . . . . . . . . . . . … 4
3b. Gill rakers 24–34; ribs present on last abdominal
vertebra (Figs. 2a, 6b, c) . . . . . . . . . . . . . . . . . . . . . . 5
4a. Tubercle present on inner symphysis of lower jaw(Fig. 5); orbital diameter 14.5–15.4 % SL; lower-jaw

Two new species of Epigonus

5a. Pyloric caeca 7–8; pectoral-fin rays 16–17 . . . . . 6
5b. Pyloric caeca 10–14; pectoral-fin rays 18–21 (rarely
17 in E. pandionis) . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6a. Teeth on lower jaw minute; orbital diameter
44.4–47.5 % HL; upper jaw-length 40.2–42.5 % HL; adultnot exceeding 100 mm SL (Palau, western North Pacific;
Fig. 9a) . . . . . . . . . . . . . . . . . . . . . . . . . . E. cavaticus
6b. Teeth on lower jaw conical; orbital diameter
38.4–42.9 % HL; upper-jaw length 45.5–48.2 % HL; adultexceeding more than 150 mm SL (Hawaiian Islands;
Fig. 9d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fragilis
7a. Gill rakers 24–25; eye elliptical; posterior half oforal cavity and tongue black (Loyalty Islands, New Caledonia; Fig. 1) . . . . . . . . . . . . . . . . E. lifouensis sp. nov.
7b. Gill rakers 26–34; eye round; posterior half of oralcavity and tongue tan or light brown . . . . . . . . . . . . . 88a. Body moderately deep, body depth 22.4–29.6 %SL; caudal-peduncle length 22.0–26.3 % SL; gill rakers
26–30; 100–130 mm SL specimens with posterodorsallycanted ring encircling the caudal peduncle (Atlantic;
Fig. 9f) . . . . . . . . . . . . . . . . . . . . . . . . . . E. pandionis
8b. Body slender, body depth 15.8–23.6 % SL; caudalpeduncle length 25.9–33.1 % SL; gill rakers 28–34;
100–130 mm SL specimens without posterodorsally canted
ring on caudal peduncle (Indo-Pacific; Atlantic; Mediterranean; Fig. 9b) . . . . . . . . . . . . . . . . . . .E. denticulatus

Fig. 9 Other seven species of the Epigonus pandionis group. a Epigonus
cavaticus, NSMT-P 72637, holotype, 65.4 mm SL, b E. denticulatus,
USNM 269827, 88.4 mm SL, c E. elongatus, ZIN 47138, holotype,160.0 mm SL (photograph taken by M. Nazarkin), d E. fragilis, BPBM
41101, 111.0 mm SL, e E. marisrubri, SMF 31636, paratype, 134.0 mm
SL, f E. pandionis, CAS 223178, 97.0 mm SL, g E. parini, NSMT-P
41252, 168.0 mm SL

length 16.0–17.6 % SL (Cocos-Keeling Islands, Indian
Ocean; Fig. 4) . . . . . . . . . . . . . E. tuberculatus sp. nov.
4b. Tubercle absent on inner symphysis of lower
jaw; orbital diameter 12.1–13.6 % SL; lower-jaw
length 13.4–15.1 % SL (western Indian Ocean;
Fig. 9c) . . . . . . . . . . . . . . . . . . . . . . . . . . E. elongatus

Comparative materials. Epigonus cavaticus: NSMT-P
72637, holotype, 65.4 mm SL, southern fringing reef ofNgemelis Island, Palau, 20 m depth, 27 February 2003;
NSMT-P 72638–72640, 3 paratypes, 66.6–69.5 mm SL,
same data as holotype. Epigonus denticulatus: AMS
I.21669-005, 2 specimens, 124.0–132.3 mm SL, 33310 S,
152040 E, New South Wales, Australia, 28 September
1977; AMS I.29540-002, 125.3 mm SL, 35320 S,
150460 E, New South Wales, Australia, 10 August 1977;
CAS 223187, 2 specimens, 97.0–112.6 mm SL, 16370 N,
11180 E, off Angola, 590–608 m depth, 28 March 2005;
FAKU-S 1114, 5 specimens, 109.1–132.8 mm SL,
2116.80 N, 1729.00 W, North Atlantic, 19 December 1971;
FAKU 43989, 76.0 mm SL, 3846.60 S, 16846.60 E, New
Zealand, 510 m depth, 27 November 1970; FAKU 44154,
148.4 mm SL, 3840.60 S, 16745.50 E, New Zealand,
655 m depth, 26 November 1970; FAKU 104596,
118.4 mm SL, 2445.40 N, 1633.60 W, North Atlantic,
740 m depth, 6 January 1972; FAKU 108973–108975, 3
specimens, 121.6–125.8 mm SL, 3840.60 S, 16745.50 E,
New Zealand, 655 m depth, 26 November 1970; HUMZ
99934, 122.8 mm SL, 26090 S, 06210 E, eastern South
Pacific, 246 m depth, 9 March 1983; HUMZ 99937,
119.2 mm SL, same data as HUMZ 99934; HUMZ
148540, 123.0 mm SL, 19070 S, 11260 E, off Namibia,
445 m depth, 7 September 1995; UF 177077, 2 specimens,123

M. Okamoto, H. Motomura

54.2–59.0 mm SL, 24200 N, 83140 W, Florida Keys, 23
April 2003; USNM 269827, 2 specimens, 84.3–88.4 mm
SL, 11240 N, 51350 E, Somalia, western Indian Ocean,
75–175 m depth, 17 December 1964; USNM 272063,
112.2 m SL, east of Brush Island, New South Wales,
Australia, 5 July 1976. Epigonus elongatus: ZIN 47138,
holotype (photograph and radiograph), 160.0 mm SL,
11000 S, 50420 E, western Indian Ocean, 320 m depth, 9April 1983; ZIN 47139, 2 paratypes (radiographs),
140.0–149.0 mm SL, same data as holotype. Epigonus
fragilis: BPBM 21175, 107.0 mm SL, northwestern
Hawaiian Islands, 30 June 1977; BPBM 24929, 2 specimens, 89.0–102.0 mm SL, Hawaiian Islands, 15 October
1975; BPBM 28853, 167.0 mm SL, Oahu, Hawaiian
Islands, 12 March 1982; BPBM 37063, 170.0 mm SL,
Oahu, Hawaiian Islands, 15 February 1993; BPBM
41101, 111.0 mm SL, Lighthouse area, Puna, Hawaiian
Islands, 3–4.5 m depth; December 2010; USNM 207704,
66.9 mm SL, Hawaiian Islands, 3 November 1968.
Epigonus marisrubri: SMF 31636, paratype, 134.0 mm
SL, 2927.30 N, 3458.30 E, Gulf of Aqaba, 52 m depth, 2
May 1990; USNM 305950, 139.8 mm SL, 12030 N,
50540 E, Gulf of Aden, Indian Ocean, 100–200 m depth,
7 November 1989. Epigonus pandionis: CAS 223178,
97.0 mm SL, 11150 S, 13420 E, eastern Atlantic, 20–21 m
depth (probably data an error), 4 April 2005; FAKU-S
1270, 100.0 mm SL, 17130 N, 16440 W, eastern North
Atlantic, 11 December 1971; MCZ 144904, 11 specimens,
64.7–74.3 mm SL, 36570 N, 74370 W, western North
Atlantic, 298 m depth, 7 September 1995; TCWC
7027.10, 5 specimens, 72.8–80.0 mm SL, 28470 N,
86040 W, Gulf of Mexico, 351–357 m depth, 14 May
1985; UF 221291, 4 specimens, 136.7–182.0 mm SL,
03450 N, 08030 E, Equatorial Guinea, western Atlantic,
409–485 m depth, 14 May 1965; UF 221434, 19 specimens,
51.2–61.3 mm SL, 03490 N, 07380 E, Equatorial Guinea,
western Atlantic, 264–269 m depth, 14 May 1965; UF
222947, 3 specimens, 162.9–180.0 mm SL, Caribbean Sea,
373–434 m depth, 14 July 1966. Epigonus parini: NSMT-P
41252, 168.0 mm SL, 43330 S, 161260 E, Louisville Ridge,
New Zealand, 623 m depth, 3 April 1985.
Acknowledgments We are most grateful to the following persons
and institutions for specimen loans: M. McGrouther, A. Hay and
S. Reader (AMS); A. Suzumoto (BPBM); A. Graham (CSIRO);
Y. Kai (FAKU); M. Yabe and T. Kawai (HUMZ); K. Hartel(MCZ); K. Matsuura, G. Shinohara and M. Nakae (NSMT);
F. Krupp (formerly SMF); K. Conway and H. Prestridge (TCWC);
L. Page, R. Robins and R. Singer (UF); and J. Williams and
E. Wilbur (USNM). We also thank A. Balushkin, M. Nazarkin and
V. Spodareva (ZIN) for providing data on the type specimens,
T. Carter (CSIRO) for photographing of specimens, and M. Love
(Marine Science Institute, University of California) for his criticalreading of the manuscript.

123

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