Species Diversity 18: 9–14
25 May 2013
DOI: 10.12782/sd.18.1.009

Japanese records of Helcogramma aquila9First Records of the Blacktail Triplefin(Perciformes: Tripterygiidae), Helcogramma aquila, fromJapan, with Notes on its Fresh Coloration
Satokuni Tashiro1 and Hiroyuki Motomura2,3
1 Faculty of Fisheries, Kagoshima University, 4–50–20 Shimoarata, Kagoshima 890–0056, Japan
2 The Kagoshima University Museum, 1–21–30 Korimoto, Kagoshima 890–0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp
3 Corresponding author(Received 19 November 2012; Accepted 9 January 2013)
Eight specimens (33.2–40.8 mm standard length) of Helcogramma aquila Williams and McCormick, 1990 (Tripterygiidae), previously known from the Philippines and the Mariana Islands, were collected from Okinoerabu-jima and Okinawajima islands in the Ryukyu Islands, Japan. These specimens represent the first records of this species from Japan, and thespecimen from Okinoerabu-jima island is the northernmost record for the species. Underwater photographs of male andfemale individuals and color photographs of a fresh male specimen are given for the first time for H. aquila.
Key Words: Ryukyu Islands, Helcogramma fuscopinna species group, distribution, fresh coloration.

Introduction
The original description of Helcogramma aquila Williams
and McCormick, 1990 was based on 13 specimens collectedfrom the Batanes Islands, Philippines. Subsequently, Fricke(1997) redescribed the species on the basis of the type series,
four new specimens from the Philippines, and five specimens from Guam. Recently, Myers and Donaldson (2003)
listed H. aquila from Saipan.
Field and museum collection surveys for tripterygiids inthe Ryukyu Islands, Japan, have yielded eight specimens of
H. aquila from Okinoerabu-jima and Okinawa-jima islands.
These specimens are described herein as the first recordsof H. aquila from Japan, and the specimen collected from
Okinoerabu-jima island represents the northernmost record
for this species. Williams and McCormick (1990) and Fricke(1997) provided morphological descriptions of the species
based only on preserved specimens; therefore, no information on the fresh coloration of this species has been available. Underwater photographs of male and female individuals taken at Okinawa-jima island and a color photograph ofa fresh male specimen are given here for the first time for H.
aquila.

Materials and Methods
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. The description is based on specimens from Japan.
The specimens of H. aquila examined in this study are deposited at the Australian Museum, Sydney, Australia (AMS);
the Kagoshima University Museum, Kagoshima, Japan(KAUM); the Hiwa Museum for Natural History, Hiwa, Hiroshima, Japan (HMNH); and the Museum Support Centerof the National Museum of Natural History, Smithsonian
Institution, Suitland, MD, USA (USNM).
Helcogramma aquila Williams and McCormick, 1990
[New standard Japanese name: Momiji-hebigimpo](Figs 1–4; Table 1)
Helcogramma aquila Williams and McCormick, 1990: 1021,
fig. 3A–B (type locality: Batan Island, Batanes Islands,
Philippines); Williams and Howe 2003: 158, figs 3–4(Batanes Islands, Philippines); Allen and Erdmann 2012:
772, unnumbered fig. (color photograph of preserved holotype).
Helcogramma aquilum: Fricke 1997: 409, fig. 81 (Guam,
Mariana Islands; Batanes Islands, Philippines); Fricke
and Williams 2000: 632 (listed name only); Williams andFricke 2001: 3534 (listed name only); Myers and Donaldson 2003: 634 (name listed from Saipan, Mariana Islands).
Helcogramma inclinata (not of Fowler 1946): Yoshigou andNakamura 2008: 70 (in part; listed as HMNH-P 4409,
9143–9148 from Okinawa-jima island, Ryukyu Islands).
Helcogramma sp.; Yoshigou and Nakamura 2008: 71 (inpart; listed as HMNH-P 8003 from Okinoerabu-jima island, Ryukyu Islands).
Material examined. 8 specimens from the Ryukyu
Islands, Japan: HMNH-P 8003, male, 37.3 mm SL, Nishihara, Okinoerabu-jima island, Amami Islands, Kagoshima© 2013 The Japanese Society of Systematic Zoology

10

S. Tashiro and H. Motomura

Fig. 1. Color photographs of fresh (A) and preserved (B–D) specimens of Helcogramma aquila. A–B, HMNH-P 9144, male, 35.0 mm
SL, Okinawa-jima island, Japan; C, HMNH-P 4409, female, 32.5 mm SL, Okinawa-jima island, Japan; D, USNM 298405, holotype, male,39.7 mm SL, Batan Island, Philippines.

Prefecture, 0.5 m depth, 4 May 2004; HMNH-P 9143, male,
40.8 mm SL, HMNH-P 9144, male, 35.0 mm SL, HMNH-P
9145, male, 36.5 mm SL, HMNH-P 9146, male, 36.4 mm SL,
HMNH-P 9147, male, 36.5 mm SL, Ou-jima island, Nanjo,
Okinawa-jima island, Okinawa Prefecture, 1.0 m depth, 2
January 2006; HMNH-P 9148, male, 34.4 mm SL; HMNH-P

4409, female, 32.5 mm SL, Giizabanda, Asato, Yaese, Okinawa-jima island, Okinawa Prefecture, 1.0 m depth, 4 January
2002.
Diagnosis. A species of Helcogramma with the following combination of characters: 13 (rarely 14) second-dorsalfin spines; 24–31 pored lateral-line scales; 5–7 symphyseal

Japanese records of Helcogramma aquila11

Fig. 2. Cephalic sensory pore system of Helcogramma aquila. Dorsal (A), lateral (B), and ventral (C) views of head (HMNH-P 9148, male,
34.4 mm SL). Scale bar 2 mm.

Fig. 3. Schematic depiction of melanophore distribution patternof mature males of Helcogramma aquila, based on seven male specimens from the Ryukyu Islands.

mandibular pores; nape in front of dorsal fin without scales;
lower half of head (area bounded by snout, ventral marginof orbit, and preopercular margin) black; bright blue stripe
extending from anterior part of upper lip to preopercularmargin through ventral margin of orbit; dorsal fins black
distally; central part of pectoral fin base black; and entirecaudal fin black (all color diagnosis applying only to mature
males).
Description. Counts and measurements are given in
Table 1. Japanese specimens are shown in Fig. 1A–C and
Fig. 4. Cephalic sensory pore systems are illustrated in Fig.
2.
General morphology. Body moderately elongate, slightly
laterally compressed anteriorly, progressively more compressed posteriorly. Dorsal profile of snout moderately
steep. Anterior nostril a short membranous tube with an unbranched, thin tentacle; anterior nostril located at mid-levelof eye; posterior nostril opening elliptical. Eyes oriented
dorsolaterally; one minute, simple tentacle on upper posterior part of each eye. Interorbital space very narrow, width
less than pupil diameter. Mouth slightly oblique; posterior margin of maxilla extending beyond a vertical throughmiddle of pupil. Uppermost anterior margin of upper jaw
approximately level with lowermost margin of orbit. Lateralline continuous, with pored scales; pored scale series ending
below base of fourth to sixth ray of third dorsal fin. Body
covered with ctenoid scales; sizes of scales above and below
lateral line subequal. No scales on head (including maxilla, interorbital space, preopercle, and opercle), pectoral finbase, or ventral surface of body. No scales on nape in frontof first spine of first dorsal fin. No scales on fin membranes,
except basally on caudal fin. Origin of first dorsal fin at vertical between preopercular and opercular margins; firstspine longest, third spine shortest. Origin of second dorsal
fin above fourth or fifth pored lateral-line scale; third to fifth

spine longest, longer than first spine of first dorsal fin. Origin of third dorsal fin above 23rd or 24th pored lateral-line
scale; first soft ray longest. Origin of pelvic fin slightly anterior to vertical from origin of first dorsal fin. Base of uppermost pectoral fin ray below third spine of first dorsal fin;pectoral fin pointed posteriorly, posterior tip of longest ray
below base of 13th or 14th spine of second dorsal fin. Originof anal fin below seventh spine of second dorsal fin; posterior margin of caudal fin slightly rounded.
Coloration of mature males when alive (Figs 3, 4A).
Upper half of head reddish with irregular bluish stripes andblotches. Lower half of head black with bright blue stripe, its
width equal to or slightly narrower than pupil diameter, extending from anterior part of upper lip through ventral margin of orbit to preopercular margin; and blue blotches scattered on lower and under sides of head. Body reddish with
six poorly defined dark red saddles and irregular verticalblue bands between saddles. Pectoral fin red with distinct
black blotch, subequal in size to orbit diameter, on central
part of fin base and bright blue blotches in and around black
blotch. First dorsal fin black except for reddish basal membranes. Second and third dorsal fins red with black margins.
Anal fin red with dusky margin. Caudal fin black.
Coloration of females when alive (Fig. 4A, B). Headand body whitish, mottled with dark red, yellow, and blue
streaks, bands, spots, blotches, and saddles; indistinct blue
stripe extending from snout to preopercular margin; twobright yellow saddles behind second and third dorsal fins,respectively. Dorsal fin membranes transparent with reddish
and bluish rays. Pectoral fin membranes transparent; rays
yellowish basally and remaining parts pale red; central partof fin base with indistinct black blotch. Anal fin membranes
transparent; rays reddish with white marginally. Caudal fintransparent with three poorly defined vertical, narrow, reddish-brown bands.
Coloration of mature males when fresh (based on color
photographs of HMNH-P 8003 and 9144; Fig. 1A). Similar to coloration when alive, but blue stripes, blotches, andsaddles lost.
Coloration of preserved male specimens (Fig. 1B, D).
Lower half of head, including lips, blackish. First dorsal fin
black anteriorly. Second and third dorsal fins black distally.
Anal fin dusky distally. Caudal fin dusky to black. Melanophores scattered on body.
Coloration of preserved female specimen (Fig. 1C).

12

S. Tashiro and H. Motomura

Fig. 4. Underwater photographs of Helcogramma aquila. A, female (left) and male (right); B, female (same female individual as in A),
Maeda, Onna, Okinawa-jima island, Japan, 3–5 m, 13 May 2012. Photos by T. Katano.

Melanophores scattered on head and body, including baseof pectoral fin; body with four distinct saddles. Dorsal finswithout black margin. Anal fin with dusky margin. Caudal
fin mottled with dusky blotches.
Distribution. This species is currently known from the
Batanes Islands in the Philippines (Williams and McCormick 1990), Guam (Fricke 1997) and Saipan (Myers andDonaldson 2003) in the Mariana Islands, and Okinawa-jimaand Okinoerabu-jima islands in the Ryukyu Islands (thisstudy). All the Japanese specimens were collected from tidepools outside the barrier reef edges in depths of 0.5–1.0 m.
Remarks. The features of the eight specimens from
Okinoerabu-jima and Okinawa-jima islands agree well withthose of H. aquila given by Williams and McCormick (1990)
and Fricke (1997); i.e., the Japanese specimens have 13(rarely 14) second dorsal-fin spines; 24–31 pored lateral-line
scales; 5–7 symphyseal mandibular pores; a pre-first-dorsalfin region without scales; the lower half of the head (thearea bounded by the snout, ventral margin of the orbit, andthe preopercular margin) black in mature males; the dorsal
fins black distally in mature males; and the central part ofpectoral fin base and the entire caudal fin black in mature
males. There are some differences between the meristics for
H. aquila given here and these in previous studies (Williams
and McCormick 1990; Fricke 1997): 10 or 11 soft rays in thethird dorsal fin (7 rays in one deformed specimen; see Table
1) vs 11 rays in the latter; and i+8+vii–viii pectoral fin rays

vs 0–i+8–9+vii. The slight differences in ray counts probably reflect the limited number of specimens available in theprevious studies.
Some morphometric data for H. aquila given herein differ
from those of Fricke (1997); no morphometric data for thisspecies were given by Williams and McCormick (1990) andWilliams and Howe (2003). Fricke (1997) gave the snout
length as 5.5–6.4% of SL (vs 9.5–10.4% herein), and the upper-jaw length as 10.0–12.8% of SL (vs 13.2–15.0%) (Table1). The holotype and one paratype of the species (bothmales) from the Philippines were examined by both Fricke(1997) and this study, and our data for these two type specimens were consistent with those of the specimens of H. aquila from Japan reported herein. We consider Fricke’s (1997)
morphometric data to be inaccurate.
Helcogramma aquila is a member of the H. fuscopinna
species group, which is characterized by the presence of abright blue stripe extending from the anterior part of theupper lip to the preopercular margin through the ventral
margin of the orbit in mature males (Williams and Howe
2003). This group comprises 11 species (Williams and Howe
2003): H. albimacula Williams and Howe, 2003; H. aquila;
H. cerasina Williams and Howe, 2003; H. desa Williams andHowe, 2003; H. fuscopinna Holleman, 1982; H. inclinata(Fowler, 1946); H. lacuna Williams and Howe, 2003; H. nesion Williams and Howe, 2003; H. nigra Williams and Howe,
2003; H. randalli Williams and Howe, 2003; and H. vulcana

Japanese records of Helcogramma aquila13

Table 1.

Counts and measurements, expressed as percentages of standard length, of Helcogramma aquila. Means in parentheses.
Williams andMcCormick(1990)

Fricke (1997)

Batan Island, Philippines

Philippines

Philippines andGuamThis study
Okinoerabu-jima andOkinawa-jima islands

Standard length (mm)CountsSecond and third dorsal-fin rays
Pectoral fin rays
Anal fin rays
Pored lateral-line scales
Mandibular pore formulaPre-dorsal scales
Measurements (% of SL)
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
Longest spine length of 2nd dorsal fin
Longest ray length of 3rd dorsal fin
Pectoral fin length
2nd ray length of pelvic fin

Non-types

Non-types

Holotype

Paratype

Holotype andparatypes

Holotype,paratypes, andnon-types

Males

FemaleMale

Male

Males andfemales

Males andfemales

n=7

n=1

n=1

n=1

n=13

n=22

34.4–40.8

33.2

39.7

37.0

26.6–40.1

13.1–40.5

XIII–XIV+10–11
i+8+vii–viii
I, 19–20
24–29
6–8+5–7+6–8
absentXIII+71
i+8+vii
I, 20
31
5+6+7
absentXIII+11
i+8+vii
I, 20
25
8+6+7
absentXIII+11
i+8+vii
I, 19
31
8+6+8
absentXIII–XIV+11
0–i+8–9+vii
I, 19–20
24–31
5–8+5–8+6–8
absentXIII–XV+11
0–i+8–9+vii
I, 19–20
24–31
5–8+5–8+5–8
absent

21.1–23.2 (21.9)
20.0–22.8 (21.6)
29.5–32.5 (31.1)
9.5–10.4 (9.9)
9.6–10.4 (10.0)
2.9–3.6 (3.1)
13.2–14.7 (13.9)
12.5–13.6 (13.2)
24.9–26.7 (25.9)
35.7–36.9 (36.5)
69.2–73.4 (71.5)
48.9–52.6 (50.6)
30.1–32.5 (31.6)
22.3–26.7 (23.9)
9.5–11.8 (10.7)
7.0–8.0 (7.6)
10.6–12.7 (11.7)
8.8–10.4 (9.6)
7.4–8.6 (8.0)
13.4–15.4 (14.4)
15.2–16.9 (16.3)
27.9–30.3 (28.9)
19.7–21.6 (20.6)

23.0
22.2
32.7
10.4
10.5
3.2
15.0
13.3
28.0
39.0
73.6
53.4
34.2
27.0
10.5
7.8
10.5
9.1
7.9
15.0
15.8
30.5
21.9

19.9
20.6
29.5
9.3
10.8
3.6
14.7
13.0
23.4
34.8
68.7
47.1
31.3
22.0
9.7
7.9
11.8
10.7
9.9
14.2
17.8
30.1
21.1

21.1
20.5
30.6
9.3
11.1
3.0
15.0
13.1
25.6
35.8
71.3
50.6
32.7
22.7
8.8
7.9
12.2
10.8
8.4
14.4
16.7
28.8
22.1

—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—

21.4–22.0
19.4–20.1
26.1–28.0
5.5–6.4
9.8–11.7
2.5–3.5
10.0–12.8—
23.3–24.8
34.3–36.1
68.1–70.9
48.0–51.0
31.8–33.8
20.0–22.3
12.1–13.8
7.7–8.2
10.2–11.1
8.3–9.5
7.5–9.0
13.1–15.8
15.8–17.7
23.5–36.8
16.7–18.8

1 Probable deformity; — No data were reported.

Randall and Clark, 1993. Helcogramma aquila can be separated from all other members of the H. fuscopinna species
group, except for H. inclinata and H. lacuna, by having morethan five symphyseal mandibular pores (Williams and Howe
2003). Helcogramma aquila is distinguished from H. inclinata and H. lacuna by having 13 s-dorsal-fin spines (vs 14 in H.
lacuna, 15 in H. inclinata), no scales on the nape anterior tothe origin of the first dorsal fin (vs present in H. inclinata),
and 16 pectoral fin rays (vs 17 in H. lacuna) (Williams andHowe 2003).
Williams and McCormick (1990), Fricke (1997), andWilliams and Howe (2003) provided morphologicaldescriptions of H. aquila based on preserved specimens andtherefore lacked information on fresh coloration. In thisstudy, underwater photographs of mature male and female
individuals (Fig. 4) and color photographs of a fresh mature
male specimen (Fig. 1A) are given for the first time for H.
aquila. These photographs reveal that males of H. aquila
can be easily distinguished from males of all members ofthe H. fuscopinna species group, except H. lacuna, by having both the anal fin and the caudal peduncle red instead of
black (Figs 1A, 3, 4). Williams and Howe (2003) described
H. lacuna on the basis of the preserved holotype and fourparatypes and gave color photographs of male (fig. 13) and14

S. Tashiro and H. Motomura

female (fig. 14) non-type specimens (without indication ofregistration numbers) as H. lacuna. However, these maleand female specimens were not of H. lacuna, but were of H.
rosea Holleman, 2006 (Holleman 2006: 97). Thus, the coloration of H. lacuna when fresh or a live is still unknown(Holleman 2006; Allen and Erdmann 2012). Judging froma photograph of the preserved type specimen of H. lacunafrom the Similan Islands, Thailand, given by Allen and Erdmann (2012: unnumbered fig. on p. 775), H. aquila can be
distinguished from H. lacuna by having a black caudal fin[vs a pale caudal fin (=no melanophores) the in preserved
specimen of H. lacuna].
Helcogramma aquila has previously been recorded fromthe Batanes Islands in the Philippines and Guam in the
Mariana Islands on the basis of collected specimens (William and McCormick 1990; Fricke 1997). Williams andHowe (2003) overlooked the record from Guam by Fricke(1997) and stated that H. aquila was endemic to the Batanes
Islands. Myers and Donaldson (2003) listed the species
from Saipan in the Mariana Islands; this record was probably based on an unpublished underwater photographof a male individual taken by H. Kimura (R. Myers, pers.
comm.). Allen and Erdmann (2012) stated the distributionof H. aquila as “Philippines (Batanes Province) and Guam”,
overlooking the record from Saipan. The present specimensfrom the Ryukyu Islands represent the first records of H. aquila from Japan and, as well, the northernmost record of thespecies.
A standard Japanese name, Momiji-hebigimpo, is proposed here for H. aquila. English (common) names, Blacktail Triplefin and Darktail Triplefin, were proposed by Fricke(1997) and Allen and Erdmann (2012), respectively.
Comparative material examined. Helcogramma aquila:
USNM 298405, holotype, male, 39.7 mm SL, White Beach,past Mahatae, Batan Island, Batanes Islands, Philippines,
20°24′45″N, 121°55′02″E, 0–6 m depth, D. Johnson and W.
Smith-Vaniz, 22 April 1987; AMS I. 29427-001, paratype,male, 37.0 mm SL, same data as holotype.

Acknowledgements
We are especially grateful to M. Meguro (KAUM) for
examining the holotype and paratype of H. aquila during
his stay at USNM, T. Katano (Okinawa Diving Center) for
providing underwater photographs of H. aquila, and H. Yoshigou (HMNH) for the loan of tripterygiid specimens deposited at HMNH. We thank M. McGrouther (AMS) and J.
Williams (USNM) for opportunities to examine comparative materials, R. Myers (Florida, USA) for providing information concerning the Saipan record of H. aquila, andvolunteers of KAUM and members of the KAUM fish laboratory for their kind curatorial assistance. This study was
supported in part by Grants-in-Aid for Scientific Research(B, 24370041 and C, 23580259) from the Japan Society for
the Promotion of Science, Tokyo, Japan (JSPS), a Grantin-Aid for Young Scientists (B, 19770067) from the Ministry of Education, Science, Sports and Culture, Japan, theJSPS Asian Core Program – Establishment of Research and
Education Network on Coastal Marine Science in Southeast Asia, and the Coastal Area Capability Enhancement inSoutheast Asia Project of the Research Institute for Humanity and Nature, Kyoto.

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