Species Diversity 20: 1–12
25 May 2015
DOI: 10.12782/sd.20.1.001

Enneapterygius phoenicosoma, a New Species of Triplefin(Tripterygiidae) from the Western Pacific Ocean
Hiroyuki Motomura1,5, Ryohei Ota2, Masatoshi Meguro3, and Satokuni Tashiro4
1 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp

2 Numazu Deepsea Aquarium, 83 Senbonminatomachi, Numazu, Shizuoka 410-0845, Japan

3 Kagoshima Environmental Research and Service, 1-1-10 Nanatsujima, Kagoshima 891-0132, Japan

4 The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-24 Korimoto, Kagoshima 890-0065, Japan
5 Corresponding author(Received 19 November 2014; Accepted 13 February 2015)
A new species of triplefin (Tripterygiidae), Enneapterygius phoenicosoma, is described on the basis of 71 type specimens (17.2–37.2 mm standard length: SL) from southern Japan. An additional five specimens from the Caroline Islands
and Vanuatu are also identified as belonging to this species. Males of the new species have the same distinct nuptial coloration as males of the Hawaiian endemic species E. atriceps (Jenkins, 1903). Comparisons of the present material with the
holotype and 46 non-type specimens of E. atriceps show, however, that E. phoenicosoma differs from the latter in having asimple nasal tentacle (vs a bilobed tentacle in E. atriceps); lower counts of the anal-fin soft rays (16–18, mode 17 vs 19–21,
19); second dorsal-fin spines (11–13, 12 vs 12–15, 14); scale rows in longitudinal series (32–35, 33 vs 35–37, 36); and pored
lateral-line scales (16–19, 17 vs 17–20, 19); a higher count of the pectoral-fin rays (15–17, 16 vs 14–17, 15); and several
morphometric features, including the lengths of the dorsal-fin spines and rays, the second dorsal-fin base, and the anal-finbase. The new species grows larger than E. atriceps (largest recorded size, 37.2 mm SL vs 26.9 mm SL in E. atriceps).
Key Words: Actinopterygii, Teleostei, Perciformes, Enneapterygius atriceps, Japan, morphology.

Introduction

Materials and Methods

The triplefin genus Enneapterygius Rüppell, 1835, thelargest genus of tripterygiids, has been diagnosed by a discontinuous lateral line with an anterior series of 6–22 pored
scales and a posterior series of 13–27 notched scales, a firstdorsal fin with three spines, an anal fin with one spine, pelvic fins with one spine and two soft rays, and a naked head,
opercle, pectoral-fin base, and abdomen (Fricke 1997).
However, synapomorphies for the genus have not beenfound, and further investigation at the generic level is therefore necessary.
During surveys of the ichthyofauna of southern Japan
from southern Kyushu to the Ryukyu Islands (e.g., Motomura and Matsuura 2010, 2014; Motomura et al. 2013), 71
specimens of a species attributed to Enneapterygius werecollected in depths of less than 25 m. The specimens weresimilar to the Hawaiian endemic species Enneapterygius
atriceps (Jenkins, 1903) in their overall appearance and thecoloration of nuptial males, but differed in several aspects,
including the shape of the tentacle on the anterior nostril,
scale and fin-ray counts, and various morphometrics. The
species from Japan is herein described as new, and five specimens from the Caroline Islands and Vanuatu are also identified as belonging to it.

Counts and measurements follow Fricke (1997) and Holleman and Bogorodsky (2012), with the mandibular pore
formula following Hansen (1986). Measurements weremade to the nearest 0.1 mm with needle-point calipers
under a dissecting microscope. Standard length is abbreviated as SL. Nuptial coloration is defined as the colorationof adult males during the courtship display and after death;
all collected adult males retained the coloration (except for
red colors) after preservation. Curatorial procedures for
newly collected specimens followed Motomura and Ishikawa (2013). The specimens examined in this study are deposited in the Academy of Natural Sciences of Drexel University, Philadelphia (ANSP); Bishop Museum, Honolulu(BPBM); Laboratory of Marine Biology, Faculty of Science,
Kochi University, Kochi (BSKU); Coastal Branch of Natural
History Museum and Institute, Chiba, Katsuura (CMNH);
Field Museum of Natural History, Chicago (FMNH); theKagoshima University Museum, Kagoshima (KAUM); Muséum National d’Histoire Naturelle, Paris (MNHN); National Museum of Nature and Science, Tsukuba (NSMT);
Royal Ontario Museum, Toronto (ROM); and National Museum of Natural History, Smithsonian Institution, Suitland(USNM).

© 2015 The Japanese Society of Systematic Zoology

2

Hiroyuki Motomura et al.

Fig. 1. Holotype of Enneapterygius phoenicosoma n. sp. (KAUM–I. 21960, male, 35.1 mm SL, Ichiki-kushikino, Kagoshima, Japan). A, fresh
specimen; B, preserved specimen.

Enneapterygius phoenicosoma Motomura, Ota, andMeguro, n. sp.
[New English name: Red-bodied Triplefin; new Japanesename: Aka-hebigimpo](Figs 1–6, 8A–B, 9; Tables 1–3)
Enneapterygius sp. 1: Ogaya in Takagi et al. 2010: 115, unnumbered 5 figs (Morode, Ehime, Shikoku, Japan).
Enneapterygius sp. 2: Meguro in Motomura and Matsuura
2014: 457, unnumbered 2 figs (Yoron-jima island, Amami
Islands, Kagoshima, Japan).
Enneapterygius sp.: Iwatsubo et al. 2014: 89, fig. 11 (off Bandokorobana Nature Park, Ei, Minami-kyushu, Kagoshima, Japan).
Holotype. KAUM–I. 21960, male, 35.1 mm SL, off Nagasaki-bana, Kushikino-kose, Ichiki-kushikino, Kagoshima,
Japan, 31°42′24″N, 130°15′44″E, 0.5–1 m, 4 August 2009, T.
Yoshida, G. Ogihara, and M. Yamashita.
Paratypes. 70 specimens, 17.2–37.2 mm SL, allfrom Japan. IZU PENINSULA: NSMT-P 62088, female,27.2 mm SL, off Ryugu-jima island, Shimoda, Shizuoka,

3–5 m, 20 December 2001, G. Shinohara and M. Yanagida.
SHIKOKU: BSKU 75367, male, 28.6 mm SL, BSKU 75368,
female, 30.0 mm SL, Murote Bay, Ainan, Ehime, 10 July
2005, E. Katayama; BSKU 101171, male, 23.7 mm SL, south
side of Murote Bay, Ainan, Ehime, 21 April 2005, T. Ogaya;
BSKU 101172, male, 22.8 mm SL, south side of Murote Bay,
Ainan, Ehime, 18 May 2005, T. Ogaya; BSKU 114638, female, 26.3 mm SL, BSKU 114639, male, 25.9 mm SL, off Tei
Beach, Kounan, Kochi, 23 June 2007. EASTERN KYUSHU(PACIFIC SIDE): KAUM–I. 6122, male, 33.0 mm SL,
KAUM–I. 9623, male, 28.1 mm SL, KAUM–I. 9624, female,
27.8 mm SL, KAUM–I. 9629, female, 25.9 mm SL, KAUM–I.
9630, male, 27.1 mm SL, tidepool at Shirahama, Miyazaki,
31°79′N, 131°48′E, 0.1–0.5 m, 26 April 2008, M. Meguroand H. Iwatsubo. WESTERN KYUSHU (EAST CHINASEA SIDE): KAUM–I. 4249, male, 29.8 mm SL, off Kurose
Beach, Kasasa, Minami-satsuma, Kagoshima, 31°22′29″N,
130°10′09″E, 2 m, 28 May 2007, M. Ito; KAUM–I. 16206,
male, 33.4 mm SL, KAUM–I. 16207, male, 32.9 mm SL,
KAUM–I. 16208, male, 32.9 mm SL, KAUM–I. 16232,
male, 27.6 mm SL, MNHN 2015–0122, female, 33.5 mm SL,
MNHN 2015–0123, male, 29.2 mm SL, off Nagasaki-bana,

New triplefin from the western Pacific

3Fig. 2. Males of Enneapterygius phoenicosoma n. sp. from Ichiki-kushikino, Kagoshima, Japan. A, KAUM–I. 40972, paratype, 31.0 mm SL;
B, KAUM–I. 40973, paratype, 31.7 mm SL. In each case, the preserved specimen and the underwater photograph represent the same individual. Underwater photographs taken at depths of 0.1–0.5 m show the normal coloration of males, and the coloration changes to the nuptial
coloration immediately after death.

Fig. 3. Female of Enneapterygius phoenicosoma n. sp. KAUM–I.
40970, paratype, 28.6 mm SL, Ichiki-kushikino, Kagoshima, Japan.
The two photographs represent the same individual.

Kushikino-kose, Ichiki-kushikino, Kagoshima, 31°42′24″N,
130°15′44″E, 0.5 m, 15 March 2009, M. Meguro and T.
Yoshida; KAUM–I. 21990, male, 33.9 mm SL, KAUM–I.
21991, male, 32.8 mm SL, KAUM–I. 21994, male, 33.7 mm
SL, KAUM–I. 25833, female, 34.5 mm SL, KAUM–I. 26185,
male, 33.7 mm SL, KAUM–I. 26186, male, 33.8 mm SL,
KAUM–I. 26187, male, 32.1 mm SL, KAUM–I. 26188,
male, 36.1 mm SL, KAUM–I. 26189, male, 37.2 mm SL,
KAUM–I. 26190, female, 33.9 mm SL, KAUM–I. 26191, female, 31.9 mm SL, KAUM–I. 26192, female, 35.0 mm SL,

KAUM–I. 26206, male, 35.1 mm SL, KAUM–I. 26207, male,28.6 mm SL, KAUM–I. 26208, male, 30.4 mm SL, KAUM–I.
26209, male, 29.6 mm SL, KAUM–I. 26210, male, 28.5 mm
SL, KAUM–I. 26211, male, 27.2 mm SL, KAUM–I. 26250,
male, 28.8 mm SL, KAUM–I. 26251, male, 27.5 mm SL,
KAUM–I. 26252, female, 32.0 mm SL, KAUM–I. 26253, female, 32.0 mm SL, KAUM–I. 26254, female, 31.8 mm SL,
KAUM–I. 26255, female, 31.3 mm SL, KAUM–I. 26256, female, 28.7 mm SL, same data as holotype; KAUM–I. 40970,
female, 28.6 mm SL, KAUM–I. 40971, female, 35.3 mm SL,
KAUM–I. 40972, male, 31.0 mm SL, KAUM–I. 40973, male,
31.7 mm SL, off Nagasaki-bana, Kushikino-kose, Ichiki-kushikino, Kagoshima, 31°42′24″N, 130°15′44″E, 0.1–0.5 m, 1
August 2011, M. Meguro; KAUM–I. 45375, male, 26.4 mm
SL, off Nagasaki-bana, Kushikino-kose, Ichiki-kushikino,
Kagoshima, 31°42′24″N, 130°15′44″E, 0.1–0.5 m, 1 August
2011, M. Meguro et al.; KAUM–I. 47301, male, 33.5 mm
SL, off Nagasaki-bana, Kushikino-kose, Ichiki-kushikino, Kagoshima, 31°42′24″N, 130°15′44″E, 0.1–0.5 m, 1
August 2011, H. Motomura et al. SOUTHERN KYUSHU:
KAUM–I. 32119, male, 30.2 mm SL, tidepool at Marukihama beach, Kushi, Bonotsu, Minami-satsuma, Kagoshima, 31°17′22″N, 130°12′45″E, 0.5 m, 10 September 2010,
H. Motomura et al.; KAUM–I. 38722, female, 27.4 mm SL,
off fishing port at Kushi, Bonotsu, Minami-satsuma, Kagoshima, 31°18′37″N, 130°13′19″E, 2–25 m, 14 June 2011,
A. Simons et al.; KAUM–I. 47259, male, 30.0 mm SL, east
coast of Chiringa-jima island, Ibusuki, Kagoshima Bay, Kagoshima, 31°16′37″N, 130°40′20″E, 0.3–1 m, 25 May 2012,
H. Motomura et al.; KAUM–I. 55381, male, 28.3 mm SL,
KAUM–I. 55382, male, 31.1 mm SL, KAUM–I. 55383, female, 30.8 mm SL, KAUM–I. 55384, male, 34.1 mm SL,
KAUM–I. 55385, male, 31.2 mm SL, off Bandokoro-bana4Hiroyuki Motomura et al.

Fig. 4. Underwater photographs of nuptial males and females of Enneapterygius phoenicosoma n. sp. A, nuptial male, off Nobeoka, Miyazaki Prefecture, east coast of Kyushu, Japan, 0–2 m depth, 5 August 2007 (photo by K. Ota); B, nuptial male, off east coast of Izu Peninsula,
Japan, 6–10 m depth (photo by S. Yamamoto); C, nuptial male, off Kii Peninsula, Japan, 11–15 m depth, 12 October 2008 (photo by D. Itou);
D, nuptial male and two females, off Otsuki, Kochi Prefecture, Shikoku, Japan, 5 m depth, 29 October 2014 (photo by K. Matsuno). Live nuptial males have a reddish body with a black head and pectoral-fin base, a distinct white band along the posterior margin of the black area on
the pectoral-fin base, and no dense black fins.

National Park, Beppu, Ei, Minami-kyushu, Kagoshima,
31°14′N, 130°26′E, 0.3 m, 23 June 2013, H. Iwatsubo;
KAUM–I. 56701, female, 30.3 mm SL, off Bandokoro-banaNational Park, Beppu, Ei, Minami-kyushu, Kagoshima,
31°14′N, 130°25′E, 4 m, 29 September 2013, H. Iwatsubo. OSUMI ISLANDS: KAUM–I. 29443, male, 31.2 mm
SL, south coast of Iou-jima island, Mishima, Kagoshima,
30°46′32″N, 130°16′43″E, 10–20 m, 25 May 2010, KAUM
Fish Team; KAUM–I. 53997, male, 28.2 mm SL, off Harutahama, Ambo, east coast of Yaku-shima island, Kagoshima, 30°18′03″N, 130°39′18″E, 8–10 m, 10 April 2013,
S. Tashiro; KAUM–I. 56567, male, 19.0 mm SL, off Urata,
Kunigami, Nishinoomote, Tanegashima island, Kagoshima,
30°49′36″N, 131°02′11″E, 4–8 m, 19 September 2013, S.
Tashiro et al. AMAMI ISLANDS: CMNH-ZF 11219, male,25.3 mm SL, Kurasaki, Tatsugo, Amami-oshima island,
Kagoshima, 28°25′55″N, 129°37′57″E, 10 m, 1 July 2004,
Y. Ikeda and M. Aizawa; KAUM–I. 17513, male, 26.2 mm
SL, tidepool at Tsuchihama, Kasari, Amami-oshima island, Kagoshima, 28°24′34″N, 129°40′31″E, 0.5–1.0 m, 27
March 2009, M. Meguro and T. Yoshida; KAUM–I. 40407,
male, 21.3 mm SL, KAUM–I. 40408, male, 18.9 mm SL, off
Maehama, Yoron-jima island, Kagoshima, 27°01′13″N,128°26′26″E, 2–10 m, 17 August 2011, M. Meguro and E.
Katayama; KAUM–I. 51390, male, 22.4 mm SL, off Chabana,Yoron-jima island, Kagoshima, 27°03′40″N, 128°25′02″E,

8 m, 26 October 2012, KAUM Fish Team; KAUM–I.
55016, male, 17.2 mm SL, off Yoron Port, Yoron-jima island, Kagoshima, 27°03′40″N, 128°25′02″E, 8 m, 29 June
2013, T. Trnski et al.; NSMT-P 31249, male, 23.6 mm SL,
off Sakinome Beach, Amami-oshima island, Kagoshima,
28°11′N, 128°16′E, 2–3 m, 12 September 1989, M. Aizawa.
OKINAWA ISLANDS: KAUM–I. 41905, male, 27.6 mm
SL, off Nago, Okinawa-jima island, 26°30′N, 128°02′E, 24
March 2011, Y. Sakurai.
Non-type material examined. 5 specimens, 20.4–
25.3 mm SL. CAROLINE ISLANDS: NSMT-P 22918, maleand female, 20.4–22.0 mm SL, south coast of Eten Island,
Chuuk Islands, 0.5 m, 6 July 1982, K. Matsuura. VANUATU:
USNM 362385, male and female, 22.1–23.1 mm SL, Fisheries Wharf, Santo Island, 15°31′30″S, 167°09′32″E, 5 m, 12
May 1997, J. Williams et al.; USNM 344010, male, 25.3 mm
SL, Port Narevin, Erromango, 18°44′25″S, 169°12′41″E,
5.5 m, 28 May 1996; J. Williams et al.
Diagnosis. A small- to medium-sized species of Enneapterygius with the following combination of characters:
11–13 (mode 12) second dorsal-fin spines; 8–10 (9) third
dorsal-fin soft rays; 15–17 (16) pectoral-fin rays; 16–18 (17)anal-fin soft rays; 32–35 (33) scale rows in longitudinal series; 16–19 (17) pored lateral-line scales; 16–19 (18) notchedlateral-line scales; 1 scale row between last pored lateral-line
scale and first notched lateral-line scale; mandibular pore

New triplefin from the western Pacific

5Fig. 6. Distributional records of Enneapterygius phoenicosoma n.
sp. and E. atriceps. Stars, localities of specimens of E. phoenicosoma;
closed circles, localities of specimens of E. atriceps examined in thisstudy; open circles, localities of specimens of E. atriceps examined
by Fricke (1997).

Fig. 5. Cephalic sensory pore system of Enneapterygius phoenicosoma n. sp. Dorsal (A), lateral (B), and ventral (C) views of headof holotype (KAUM–I. 21960, male, 35.1 mm SL, Ichiki-kushikino,
Kagoshima, Japan); dorsal view (D) of head of paratype (KAUM–I.
26186, male, 33.8 mm SL, Ichiki-kushikino, Kagoshima, Japan).
Nostrils and tentacles not illustrated; scale bar 2 mm.

formula 4–6+1+4–6 (4+1+4); long dorsal-fin spines andrays [e.g., lengths of second spine 8.6–12.6% of SL (mean10.1%) and third spine 7.2–9.8% of SL (8.5%) in first dorsal fin; first spine 12.8–17.2% of SL (15.2%), second spine
13.8–18.2% of SL (15.9%), and third spine 14.2–18.4% of SL(15.8%) in second dorsal fin; and first soft ray 14.8–19.5%
of SL (17.3%), second soft ray 14.7–18.9% of SL (16.9%),and third soft ray 13.9–19.1% of SL (15.9%) in third dorsal fin]; short second dorsal-fin base [25.0–32.9% of SL(mean 29.2%)]; short anal-fin base [36.9–43.5% of SL (mean40.5%)]; nasal tentacle unbranched, flat, broadened distally(Fig. 8A, B); lower two-thirds of head, including lower halfof eye, snout, lips, cheek, and opercle, and pectoral-fin base,dense black in nuptial males; upper head and body reddishbrown in males and females, reddish in nuptial males, with
H-shaped and/or upside-down Y-shaped oblique brownbars on lateral surface of body in both sexes; caudal-fin basewith vertical brownish band, its width subequal to pupil
diameter, and narrow vertical white band behind brownish band in both sexes; narrow, indistinct, white line alongposterior margin of black area on pectoral-fin base in nuptial males; pectoral, pelvic, and anal fins reddish in nuptial
males; no fins black.
Description. In following text, data for the holotype arepresented first, followed by paratype data (if different) in parentheses. Counts and measurements of the holotype, paratypes, and non-types are given in Tables 1–3. The cephalicsensory pore systems of the holotype are illustrated in Fig.
5A–C; major variation of sensory pores on the occipital region is shown in Fig. 5D.
Dorsal fin III, XII, 9 (III, XI–XIII, 8–10); anal fin I, 17 (I,
16–18); 16 (15–17) pectoral-fin rays; 34 (32–35) scale rows
in longitudinal series; 17 (16–19) pored lateral-line scales;
17 and 18 (16–19) notched lateral-line scales; 4 (3 or 4) scalerows above first pored lateral-line scale; 3½ and 4 (2½–4)
scale rows between first spine base of second dorsal fin andpored lateral-line scale; 3½ (2½–4) scale rows below firstnotched lateral-line scale; 2½ and 3 (2–3) scale rows above
last pored lateral-line scale; 12 (11 or 12) circumpeduncularCounts
1st, 2nd, and 3rd dorsal-fin rays
Anal-fin rays
Pectoral-fin rays
(unbranched+branched+unbranched rays)
Scale rows in longitudinal series
Pored lateral-line scales
Notched lateral-line scales
Scale rows above 1st pored lateral-line scale
Scale rows between 1st spine base of second
dorsal fin and pored lateral-line scale
Scale rows below 1st notched lateral-line scale
Scale rows above last pore lateral-line scale
Circumpeduncular scales
Mandibular pore formula
Measurements
Body depth
Body width
Head length
Snout length
Orbit diameter
Interorbital width
Upper-jaw length
Postorbital lengthPre-1st dorsal-fin lengthPre-2nd dorsal-fin lengthPre-3rd dorsal-fin lengthPre-anal-fin length
Pre-pectoral-fin length
Pre-pelvic-fin length
Caudal-peduncle length
Caudal-peduncle depth
1st spine length of 1st dorsal fin
2nd spine length of 1st dorsal fin
3rd spine length of 1st dorsal fin

Standard length (mm)III, XI–XIII, 8–10
I, 16–18
15–17(ii–vi+3–7+vi–viii)
33–35
16–191
16–191
3–4
2½–4
2½–4
2–3
11–12 (usually 12)
4–6+1 (rarely 3)+4–6
17.3–21.0 (18.9)
15.7–20.6 (17.9)
28.0–31.4 (29.5)
7.8–10.5 (9.1)
9.2–11.7 (10.1)
2.2–3.3 (2.7)
10.5–13.1 (11.8)
11.5–14.1 (13.0)
24.5–28.1 (26.2)
34.7–39.0 (36.6)
68.0–74.1 (70.6)
45.3–53.1 (49.0)
29.5–34.2 (31.7)
18.3–24.8 (22.0)
8.8–12.4 (10.9)
7.3–9.9 (8.5)
8.8–13.0 (10.7)
8.6–12.6 (10.0)
7.2–9.7 (8.5)

3½
2½–3
12
4+1+4
18.8
17.4
29.0
8.5
9.5
3.1
12.1
13.2
24.5
36.5
69.9
46.7
31.1
21.7
11.9
8.5
11.3
11.0
—

17.2–37.2

17.8–22.0 (19.5)
16.7–20.7 (18.8)
29.8–32.1 (30.5)
7.9–10.9 (9.1)
10.2–11.0 (10.5)
2.4–3.1 (2.6)
11.8–13.4 (12.4)
13.9–14.4 (14.1)
25.1–27.1 (26.2)
36.9–39.1 (38.0)
69.4–72.6 (71.0)
49.5–53.9 (51.4)
31.2–34.0 (32.1)
19.0–24.7 (20.9)
10.8–12.1 (11.4)
7.8–8.9 (8.2)
12.0–13.7 (13.1)
10.7–12.1 (11.4)
8.0–9.8 (8.6)

3–3½
2½
11–12
4+1+4

III, XII–XIII, 8–9
I, 16–17
15–17(ii–vi+5–8+viii)
32–34
16–17
16–18
4
3½–4

20.4–25.3

Non-types
n=5

Enneapterygius phoenicosoma n. sp.
Paratypes
n=70

III, XII, 9I, 1716(iii+6+vii)
34
17
17–181
4
3½–4

35.1

Holotype
KAUM–I. 21960

Means2
19.0
17.9
29.6
9.1
10.2
2.7
11.8
13.1
26.2
36.7
70.6
49.1
36.8
22.0
11.0
8.5
10.9
10.1
8.5

Table 1. Counts and measurements, expressed as percentages of standard length, of Enneapterygius phoenicosoma n. sp. and E. atriceps. Means in parentheses.

20.4
15.7
30.2
9.0
10.6
3.1
11.8
13.8
27.2
39.6
74.6
48.1
31.3
20.3
9.5
8.8
11.2
—
8.4

—
—
—
4+1+4

35
—
—
—
—

III, XIV, 9
I, 20
15

22.1

16.8–21.0 (19.1)
15.1–18.9 (17.4)
26.0–30.2 (28.3)
7.3–9.1 (8.0)
8.8–11.8 (10.5)
2.4–3.2 (2.8)
10.3–12.4 (11.2)
11.6–13.6 (12.6)
23.5–28.9 (26.0)
32.1–39.4 (35.9)
69.5–76.7 (71.3)
44.3–52.3 (48.0)
28.4–33.6 (31.1)
19.1–24.4 (22.2)
9.4–13.0 (10.8)
7.7–10.1 (8.8)
8.4–13.0 (10.8)
8.0–11.3 (9.2)
6.5–9.1 (7.8)

3–5
2–4
12
4–6+1 (rarely 2)+4–5

III, XII–XV, 9–11
I, 19–21
14–17(ii–iv+5–7+vi–viii)
35–37
17–20
16–20
2–5
2–5

14.9–26.9

Non-types
n=46

Enneapterygius atriceps
Holotype
USNM 50719

6
Hiroyuki Motomura et al.

15.0
17.9
—
17.4
18.0
17.3
34.9
15.8
25.0
5.5
30.0
16.8
41.0

Holotype
13.7–17.2 (15.3)
14.2–18.2 (15.9)
14.2–18.4 (15.8)
14.8–19.5 (17.3)
14.7–18.9 (17.0)
13.9–19.1 (15.9)
30.6–37.4 (33.6)
14.2–19.1 (16.4)
19.8–26.2 (23.0)
4.1–6.2 (5.2)
25.0–32.9 (29.3)
13.9–19.2 (17.1)
38.1–43.5 (40.6)

Paratypes

n=44
n=76

n=46
n=76

E. atriceps
E. phoenicosoma n. sp.

E. atriceps
E. phoenicosoma n. sp.

1 Includes count of holotype.

n=45
n=75

2
—

14

—
10

11

—
4

16

4
6

13

281
8

15

—
651

17

13
621

16

Pectoral-fin rays (left)

1
591

12

Second dorsal-fin spines

1
6

17

—
7

n=41
n=64

n=46
n=76

Anal-fin soft rays

1
—

391
—

15

14

18

Frequency distribution of selected fin ray counts of Enneapterygius phoenicosoma n. sp. and E. atriceps.

E. atriceps
E. phoenicosoma n. sp.

Table 2.

12.8–14.2 (13.6)
13.8–15.1 (14.3)
14.2–15.3 (14.9)
14.9–15.9 (15.5)
15.3–15.4 (15.3)
13.9–15.0 (14.4)
28.9–34.6 (32.6)
15.6–17.2 (16.6)
21.3–23.3 (22.4)
5.3–6.4 (5.7)
26.5–29.8 (28.1)
15.8–16.9 (16.2)
36.9–40.4 (38.3)

Non-types

Enneapterygius phoenicosoma n. sp.

1 Including scale counts of right side of body; 2 including data for holotype, paratypes, and non-types; — damaged.

1st spine length of 2nd dorsal fin
2nd spine length of 2nd dorsal fin
3rd spine length of 2nd dorsal fin
1st soft ray length of 3rd dorsal fin
2nd soft ray length of 3rd dorsal fin
3rd soft ray length of 3rd dorsal fin
Pectoral-fin length
Pelvic-fin spine length
Pelvic-fin soft-ray length
1st dorsal-fin base length
2nd dorsal-fin base length
3rd dorsal-fin base length
Anal-fin base length

Table 1. Continued.

27
—

19

15.2
15.9
15.8
17.3
16.9
15.9
33.5
16.5
23.2
5.2
29.2
17.1
40.5

3
—

14

—
5

8

18
11

10

1
—
—

21

1
7

17

3
—

11

12.9–15.3 (14.2)
13.6–15.8 (14.8)
14.2–17.3 (15.9)
13.6–16.7 (15.6)
13.4–16.2 (14.8)
29.4–35.5 (32.9)
13.9–18.9 (17.3)
19.6–26.9 (23.5)
4.2–6.8 (5.1)
27.4–38.1 (31.8)
14.4–19.0 (16.6)
39.5–47.5 (43.1)

181

15
491

16

Non-types
12.5–14.2 (13.4)

20

221
8

15

Pectoral-fin rays (right)

601

251

9

Third dorsal-fin soft rays

—
13.7
14.7
—
16.3
15.4
—
18.9
24.7
6.0
33.6
16.6
46.0

Holotype

Enneapterygius atriceps

New triplefin from the western Pacific
7

Hiroyuki Motomura et al.

1
—
6
2
12
32
8
201
1
6

19
18
17
16

Notched lateral-line scales (right side)

—
12

16

20

6
—
12
1
9
9

19
18
17

Pored lateral-line scales (right side)

1
—

20

6
—

20

10
3
10
321
6
20
1 Includes count of holotype.

n=28
n=59
E. atriceps
E. phoenicosoma n. sp.

1
4

19
18
17
16

Notched lateral-line scales (left side)

11
1
9
3
2
461
n=28
n=61
E. atriceps
E. phoenicosoma n. sp.

—
11

19
18
17
16

Pored lateral-line scales (left side)

13
—
26
—
51
—
—
391

37
36
35

Scale rows in longitudinal series (left)

34
33

—
37
n=44
n=76
E. atriceps
E. phoenicosoma n. sp.

Table 3.

Frequency distribution of selected scale counts of Enneapterygius phoenicosoma n. sp. and E. atriceps.

n=40
n=76

n=27
n=65

n=28
n=60

—
1

32

—
431

20

11
—
26
—
31
1
—
351
—
39

35
34
33

Scale rows in longitudinal series (right)

36

37

8

scales; 4+1+4 (4–6+1+4–6) mandibular pores.
Body moderately elongate, slightly compressed anteriorly,
progressively more compressed posteriorly; dorsal profile ofsnout not strongly steep. Mouth slightly oblique; posterior
margin of maxilla extending slightly beyond vertical drawn
through anterior margin of pupil; anterior tip of upper jaw
almost reaching as high as ventral margin of orbit. Anterior
nostril membranous tube with thin, but distally broadened,unbranched tentacle (Fig. 8A, B); anterior nostril located at
level of middle of eye, slightly closer to eye than to upper
lip; posterior nostril opening elliptical. Eye oriented dorsolaterally; simple tentacle on posterodorsal margin of eye,its length subequal to length of nostril tentacle. Interorbital
space narrow, its width narrower than pupil diameter. Opercular margin extending beyond vertical drawn through baseof third spine of first dorsal fin.
Lateral line discontinuous, with anterior series of pored
scales and posterior series of notched scales; pored scaleseries ending below membrane between last spine of second dorsal fin and body; notched scale series beginning
below last pored scale (or below third or second scale fromlast pored scale or below a point just posterior to last pored
scale) and ending at caudal-fin base; one scale row between
posteriormost pored scale and anteriormost notched scale;one additional notched scale present (or scale absent) anterior to 2 (1–4) normal scales followed by series of notchedscales. Body scales ctenoid; no scales on head, including
maxilla, interorbital space, preopercle, and opercle, pectoral-fin base, pre- and inter-pelvic-fin region, or pre-dorsalfin region; no scales on fin membranes except for basal partof caudal fin.
Origin of first dorsal fin at vertical drawn between preopercular and opercular margins; first spine of this fin longest, third spine shortest. Origin of second dorsal fin above
fifth (third to fifth) pored lateral-line scale, second (rarely
third) spine of this fin longest. Origin of third dorsal fin
above 20th (19th to 21st) longitudinal scale; second (orfirst) ray of this fin longest. Origin of pelvic fin anterior tovertical drawn through base of first spine of first dorsal fin,
below posterior edge of preopercle. Base of uppermost pectoral-fin ray below second pored lateral-line scale. Pectoral
fin pointing posteriorly, with posterior tip of its longest ray
not reaching vertical drawn through origin of third dorsal
fin (usually just reaching vertical drawn through base of lastspine of second dorsal fin). Origin of anal fin below sixth(fifth to seventh) spine base of second dorsal fin.
Color of nuptial male specimens when fresh—Based oncolor photographs of holotype (Fig. 1A) and 22 paratypes(Fig. 2): lower two-thirds of head, including lower half ofeye, snout, lips, cheek, and opercle, and pectoral-fin base,dense black (dark grey with scattered large gray blotches);remaining part of head and body reddish with 11 (9–12)
H-shaped and/or upside-down Y-shaped oblique brownbars on lateral surface of body. Narrow, indistinct, whiteline along posterior margin of black area on pectoral-finbase. Caudal-fin base with vertical brownish band, its width
subequal to pupil diameter, and narrow vertical white band
behind brownish band. First dorsal fin semi-transparent,New triplefin from the western Pacific

9Fig. 8. Tentacle on anterior nostril. A–B, holotype (KAUM–I.
21960, 35.1 mm SL) of Enneapterygius phoenicosoma n. sp.; C–D,
non-type (ANSP 28022, 18.6 mm SL) of E. atriceps. Upper (A, C)
and lower (B, D) represent lateral and posterior views respectively.
Scale bars 0.1 mm.

Fig. 7. Cephalic sensory pore system of Enneapterygius atriceps.
Dorsal (A), lateral (B), and ventral (C) views of head (ANSP 95035,
male, 20.8 mm SL, Waikiki, Oahu, Hawaiian Islands). Nostrils andtentacles not illustrated; scale bar 1 mm.

yellowish (or reddish) basally and distally; spines reddish.
Second dorsal fin semi-transparent with melanophores scattered on basal one-third of fin, forming broad, pale grayband; distal margin of fin grayish. Third dorsal fin semitransparent without distinct bands or stripes. Pectoral fin
semi-transparent with red rays. Pelvic fin rays red (or pink).
Anal fin red with reddish (or whitish) rays. Caudal fin semitransparent with whitish or reddish rays and without distinct bands or stripes, with reddish ventral margin.
Color in life of nuptial males—Based on several underwater photographs (see Fig. 4): similar to the above, but narrow white line along posterior margin of black area on pectoral-fin base more distinct.

Color in life of pale males and females—Based on several
underwater photographs (see Figs 2–3): head mottled withbrown, white, blue, and green blotches, spots, and lines.
Body whitish with 9–12 H-shaped and/or upside-down Yshaped oblique brown bars on lateral surface of body; caudal-fin base with vertical brownish band and narrow verticalwhite band behind brownish band. First dorsal fin whitish.
Second dorsal fin semi-transparent with pale gray band basally. Third dorsal, anal, and caudal fins semi-transparent.
Pectoral fin semi-transparent; narrow white line along posterior margin of pectoral-fin base. Color of female specimens when fresh (Fig. 3) similar to their color in life.
Color in preservative of nuptial males—Lower two-thirdsof head, including lower half of eye, snout, lips, cheek, andopercle, and pectoral-fin base black; remaining part of headand body whitish with oblique grayish bars on lateral surface of body. Caudal-fin base with vertical grayish band.
Second dorsal fin semi-transparent, grayish basally. Third
dorsal, pectoral, anal, and caudal fins semi-transparent withwhite rays. Pelvic-fin rays white.
Color in preservative of females—Similar to coloration ofpreserved male specimens, but head and pectoral-fin basepale and mottled with scattered melanophores.
Distribution and ecological notes. Currently known
from the western Pacific Ocean: Japan, the Caroline Islands,
and Vanuatu. In Japanese waters, E. phoenicosoma has beenrecorded from the Pacific coast of southern Japan, including the Izu Peninsula, Shikoku, and southern Kyushu, andthe Ryukyu Islands. This species usually occurs in open
tide-pools and areas with strong regular surges generated bywave swells. Individuals usually inhabit the surface of large

10

Hiroyuki Motomura et al.

Fig. 9. Comparisons of lengths of (A) second spine of first dorsal fin, (B) first spine of second dorsal fin, (C) second spine of second dorsal
fin, (D) third spine of second dorsal fin, (E) second dorsal-fin base, and (F) anal-fin base (% standard length) with standard length in Enneapterygius phoenicosoma n. sp. (stars) and E. atriceps (circles).

rocks, but also slits and holes in large rocks, at depths of lessthan 25 m (usually less than 15 m). Spawning of E. phoenicosoma at Shikoku occurs from early morning until about 10
am starting in June (Ogaya in Takagi et al. 2010; reported as
Enneapterygius sp. 1).
Etymology. From the Greek “phoenico” meaning red
and “soma” body, in reference to the red body of nuptial
males. The name is used as a noun in apposition.
Remarks. The nuptial coloration of males of E. phoenicosoma n. sp., including their black head and pectoralfin base and reddish body, is distinct from the colorationof females; however, detailed analyses of 32 morphometricfeatures (listed in Table 1) revealed no other sexual dimorphism.
Most specimens of the new species were collected fromshallow waters in southern Japan, where it is relatively common. We also found a few specimens from the Caroline Islands and Vanuatu, which we have tentatively identified as
belonging to this species. More specimens are required to
fully assess the taxonomic status of these specimens fromthe central western Pacific Ocean, so they are excluded fromthe type series of E. phoenicosoma.

Enneapterygius atriceps (Jenkins, 1903)[English name: Hawaiian Blackhead Triplefin](Figs 6–7, 8C–D, 9; Tables 1–3)
Tripterygium atriceps Jenkins, 1903: 505, fig. 46 (type locality: Honolulu, Oahu, Hawaiian Islands).
Enneapterygius atriceps: Fricke 1997: 161, 566, figs 25, 26(Hawaiian Archipelago, including French Frigate Shoals,
Laysan Island, and Midway Atoll); Williams and Fricke
2001: 3534 (name only); Fricke 2009: 38 (Hawaiian Islands, north to Midway Atoll); Randall 2007: 373, unnumbered figs (Hawaiian Islands); Meguro and Motomura in Motomura and Matsuura 2010: 6 (name only).
Holotype. USNM 50719, male, 22.1 mm SL, Honolulu,
Oahu, Hawaiian Islands, 1889, O. Jenkins.
Non-type material examined. 46 specimens, 14.9–
26.9 mm SL, all from the Hawaiian Archipelago. ANSP
28022, 3 males, 15.7–19.4 mm SL, Honolulu, Oahu, Hawaiian Islands, 1900, O. Jenkins; ANSP 95035, male and female,
14.9–20.8 mm SL, Waikiki, Oahu, Hawaiian Islands, 1992, C.
Edmondson; BPBM 3465, male, 25.8 mm SL, Waikiki Reef,
Honolulu, Oahu, 1923, V. Pietschmann; BPBM 19658, male

New triplefin from the western Pacific

and female, 19.0–20.9 mm SL, Kaneohe Bay, Oahu, 2 m, 13
September 1975, B. Carlson et al.; BPBM 34846, 13 males
and 5 females, 19.5–23.9 mm SL, southeast side of Sand Island, Midway Atoll, 8 m, 16 September 1991, J. Randall etal.; FMNH 47658, male, 22.1 mm SL, Kaneohe Bay, Oahu,
April–May 1935, Shedd Aquarium; FMNH 63605, 9 males
and 5 females, 17.0–26.9 mm SL, Kahuku, Oahu, 21 April
1961, L. Woods et al.; ROM 28022, 3 males (one in poor
condition, its SL not included), 16.0–18.5 mm SL, Honolulu,
Oahu, US Fish. Commission; ROM 95035, 2 males, 15.6–
20.4 mm SL, Waikiki, Honolulu, Oahu, 1922, C. Edmondson.
Diagnosis. A small-sized species of Enneapterygius
with the following combination of characters: 12–15 (mode14) second dorsal-fin spines; 9–11 (9) third dorsal-fin softrays; 14–17 (15) pectoral-fin rays; 19–21 (19) anal-fin softrays; 35–37 (36) scale rows in longitudinal series; 17–20(19) pored lateral-line scales; 16–20 (18) notched lateral-line
scales; 1 or 2 scale rows between last pored lateral-line scale
and first notched lateral-line scale; mandibular pore formula 4–6+1+4–5 (4+1+4); short dorsal-fin spines and rays[e.g., lengths of second spine 8.0–11.3% of SL (mean 9.2%)
and third spine 6.5–9.1% of SL (7.8%) in first dorsal fin; firstspine 12.5–14.2% of SL (13.4%), second spine 12.9–15.3%
of SL (14.2%), and third spine 13.6–15.8% of SL (14.8%) in
second dorsal fin; first soft ray 14.2–17.3% of SL (15.9%),
second soft ray 13.6–16.7% of SL (15.6%), and third soft
ray 13.4–16.2% of SL (14.8%) in third dorsal fin]; long second dorsal-fin base [27.4–38.1% of SL (mean 31.8%)]; longanal-fin base [39.5–47.5% of SL (mean 43.1%)]; nasal tentacle bilobed (Fig. 8C, D; rarely unbranched; 2 of 47 specimens with unbranched tentacle); lower two-thirds of head,
including lower half of eye, snout, lips, cheek, and opercle,
and pectoral-fin base dense black in nuptial males; upper
head and body reddish-brown in females and males, reddishin nuptial males, with 4–6 H-shaped and/or upside-down
Y-shaped oblique brown bars on lateral surface of body;
caudal-fin base with vertical brownish band, its width subequal to pupil diameter, narrow vertical white band behindbrownish band; narrow, indistinct, white line along posterior margin of black area on pectoral-fin base; no fins black.
Distribution. Known only from the Hawaiian Archipelago (Fricke 1997; this study).
Remarks. Counts and measurements of the above-listed holotype and non-types of E. atriceps are given in Tables
1–3. The cephalic sensory pore system of the species is illustrated in Fig. 7. Color photographs of male and female
specimens were published by Randall (2007: 373).
Comparisons. Enneapterygius phoenicosoma n. sp. issimilar to the Hawaiian endemic species E. atriceps in sharing the following features: 16–20 pored lateral-line scales;a single scale row between the last pored lateral-line scale
and the first notched lateral-line scale; the lower two-thirdsof the head, including the lower half of the eye, snout, lips,
cheek, and opercle, and the pectoral-fin base dense black in
nuptial males; the upper head and body reddish in nuptial
males but reddish-brown in females and non-nuptial males,with H-shaped or upside-down Y-shaped oblique brown

11

bars on the lateral surface of the body; the caudal-fin basewith a vertical brownish band, its width subequal to thepupil diameter, and a vertical narrow white band behindthe brownish band; a narrow, indistinct, white line along theposterior margin of the pectoral-fin base; and no black fins.
With the exception of E. phoenicosoma and E. atriceps, nonominal species with combination of the above-mentioned
characters have been reported in the genus (Fricke 1997;
this study).
The new species is clearly distinguished from E. atriceps
by having a simple nasal tentacle (vs bilobed in the latter;
Fig. 8) and 16–18 (mode 17) anal-fin soft rays [vs 19–21(19); Tables 1–2]. Enneapterygius phoenicosoma further
differs from E. atriceps in having 11–13 (mode 12) second
dorsal-fin spines [vs 12–15 (14); Tables 1–2], 15–17 (16)pectoral-fin rays [vs 14–17 (15); Tables 1–2], 32–35 (33)
scale rows in the longitudinal series [vs 35–37 (36); Tables
1, 3], and 16–19 (17) pored lateral-line scales [vs 17–20 (19);Tables 1, 3].
In morphometrics, E. phoenicosoma tends to have slightly
longer spines and rays of the dorsal fins than E. atriceps: e.g.,lengths of the second spine (mean 10.1% of SL vs 9.2% in
E. atriceps; Table 1, Fig. 9A) and third spine (8.5% vs 7.8%;
Table 1) in the first dorsal fin; lengths of the first spine(15.2% vs 13.4%; Table 1, Fig. 9B), second spine (15.9% vs
14.2%; Table 1, Fig. 9C), and third spine (15.8% vs 14.8%;
Table 1, Fig. 9D) in the second dorsal fin; and the lengthsof the first soft ray (17.3% vs 15.9%; Table 1), second ray(16.9% vs 15.6%; Table 1), and third ray (15.9% vs 14.8%;
Table 1) in the third dorsal fin. In contrast, the lengths of thesecond dorsal-fin base and anal-fin base in E. phoenicosoma
tend to be slightly shorter than those of E. atriceps (mean29.2% of SL and 40.5%, respectively, vs 31.8% and 43.1%, respectively, in E. atriceps; Table 1, Fig. 9E, F).
Enneapterygius phoenicosoma grows larger than E. atriceps (largest recorded size, 37.2 mm SL vs 26.9 mm SL; thisstudy). Fricke (1997) examined 87 specimens of E. atriceps
ranging from 11 to 26 mm SL. On the basis of examinationof otoliths of E. atriceps, Longenecker and Langston (2005)
determined the age of the oldest individual of E. atriceps as
117 days, with sexual maturity occurring at about 55 days
and 15 mm SL. No biological information on E. phoenicosoma is known. These two species are allopatrically distributedin the Pacific Ocean (Fig. 6).
Enneapterygius etheostoma (Jordan and Snyder, 1902)
co-occurs with E. phoenicosoma through at the entire distributional range of the latter in Japanese waters. Colorationof females and pale males in the two species is very similar
to each other either in life or when fresh. Besides the coloration of nuptial males, however, E. phoenicosoma differsfrom E. etheostoma in the following features: nasal tentacle
simple (vs bilobed in the latter), 11–13 (mode 12) second
dorsal-fin spines [vs 12–16 (14)], 16–18 (17) anal-fin softrays [vs 17–20 (19)], 2½–4 (3½) scale rows between base of
first spine of second dorsal fin and pored lateral-line scale[vs 4–7½ (6)], and 2–3 (2½) scale rows above last pored
lateral-line scale [vs 3½–5½ (4)] (Fricke 1997; Meguro andMotomura unpub. data).

Hiroyuki Motomura et al.

12

Acknowledgments
We are especially grateful to M. Sabaj (ANSP), A. Suzumoto and L. O’Hara (BPBM), and J. Williams (USNM) for
opportunities to examine specimens deposited in their museums. We thank H. Endo (BSKU), H. Kawase (CMNH),
S. Mochel and K. Swagel (FMNH), G. Shinohara and M.
Nakae (NSMT), and R. Winterbottom, E. Holm, D. Stacey,and M. Zur (ROM) for loans of specimens, R. Causse and
P. Pruvost (MNHN) for registering type specimens, and S.
Harazaki, S. Yamamoto, D. Itou, K. Ota, and K. Matsuno forproviding underwater photographs. We thank Y. Haraguchi,
volunteers, and students of KAUM for their kind assistancewith collecting and curating specimens, and G. Yearsley (Ellipsis Editing) for checking an early version of this manuscript. This study was supported in part by Grants-in-Aid
for Scientific Research (A: 26241027, B: 24370041 and C:
23580259 and 26450265) from the Japan Society for the
Promotion of Science, Tokyo, Japan (JSPS); the JSPS AsianCore Program, “Establishment of Research and Education
Network on Coastal Marine Science in Southeast Asia”; theJSPS International Training Program “Protect Diversity of
Bioresources in the Tropical Area”; the “Coastal Area Capability Enhancement in Southeast Asia Project” of the
Research Institute for Humanity and Nature, Kyoto, Japan;
the “Establishment of Research and Education Network on
Biodiversity and Its Conservation in the Satsunan Islands”
project of Kagoshima University adopted by the Ministry of
Education, Culture, Sports, Science and Technology, Japan;
and the “Biological Properties of Biodiversity Hotspots inJapan” project of the National Museum of Nature and Science, Tsukuba, Japan.

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