H. Motomura and K. Matsuura (eds) Fishes of Yaku-shima Island
National Museum of Nature and Science, Tokyo, 10 March 2010

Apogonid fishes (Teleostei: Perciformes)of Yaku-shima Island, Kagoshima Prefecture,southern Japan
Tomohiro Yoshida1*, Shigeru Harazaki2 and Hiroyuki Motomura1
1

Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan
Diving Service Mori to Umi, 1559-1 Miyanoura, Yakushima, Kumage, Kagoshima 891-4205, Japan*
Corresponding author: e-mail: k5299534@kadai.jp
2

Abstract Forty five species of apogonid fishes were confirmed to occur at Yaku-shima Island, Kagoshima Prefecture, Kyushu, southern Japan. Of these, 27 species represent the first records from theisland, including the first Japanese record of Apogon chrysotaenia. A new Japanese name, Akafujitenjikudai, is proposed for Apogon crassiceps. Each species account provides a synonym list, material examined, descriptions based on collected specimens, distribution in Japanese waters, taxonomic
and nomenclatural remarks, and ecological notes from Yaku-shima Island.
Key words: Apogonidae, cardinalfishes, distribution, Yaku-shima Island, Japan.

Introduction
The family Apogonidae, characterized by having two separated dorsal fins, the first with 6–8
spines and the second with 1 spine and 8–14 softrays, anal fin with 2 spines and 8–18 soft rays, andusually 24 vertebrae, consists of about 23 genera and roughly 273 species (Nelson, 2006). Of
these, about 15 genera and 94 species have beenrecorded from Japanese waters (Matsuura and Tachikawa, 1994; Hayashi, 2002, 2004; Mabuchi etal., 2003, 2004; Yoshigou and Yoshino, 2004; Miyahara et al., 2005; Shibukawa et al., 2007; Fraserand Allen, 2010).
The first apogonid fish reported from Yakushima Island was Apogon notatus in 1906. Jordan
and Starks (1906) gave an illustration of this species (as Amia notata) by Mr. W. S. Atkinson on
the basis of a single specimen from the island. The
specimen is now missing from collections of the
California Academy of Sciences, San Francisco,or the Museum Support Center of the Smithsonian
Institution National Museum of Natural History,
Suitland. Subsequently, Arai and Ida (1975) reported two genera with four species of the family

from Kusugawa on the northeast coast of Yakushima Island on the basis of collected specimens,
which are deposited at the National Museum of
Nature and Science, Tokyo. Ichikawa et al. (1992)
listed three genera with 10 species from the island,
and Kuniyasu (1999) listed two genera with 12
species from Kurio on the southwest coast, bothbeing based only on underwater observations.
In addition to the previous records, our Yakushima Island ichthyofaunal surveys in 2008–2009(see Motomura et al., 2010) and the second author’s underwater observations over the last six
years resulted in 11 genera with 45 species of
apogonid fishes occurring off Yaku-shima Island.
This paper provides an annotated checklist ofthese species, along with color photographs if
available.
Materials and methods
Counts and measurements generally follow
Fraser (2005). Counts of longitudinal stripes on
the lateral surface of the body begin with the uppermost stripe, including a stripe running along
the dorsal-fin bases. Measurements were made to27

T. Yoshida et al.

the nearest 0.1 mm. Unless specified otherwise,
the length of specimens listed throughout thispaper is the standard length (abbreviated as SL).
Identifications are generally based on Allen et al.
(2005), Greenfield, (2001, 2007a, b), Gon (1993),
Gon and Randall (2003), Fraser (2005, 2008),
Fraser and Allen (2010), Hayashi (2000, 2002),
Mabuchi et al. (2003, 2004), Randall (2005),
and Randall et al. (1997a). Randall (2005) reclassified the genus Apogon and divided it into
four genera, Apogon, Ostorhinchus, Pristiapogon,
and Zoramia, but we use Apogon rather than thesegeneric names. Scientific names are arrangedalphabetically for each genus. The synonym lists
contain only the original description citation andpapers related to Yaku-shima Island, viz., Jordan
and Starks (1906), Arai and Ida (1975), Ichikawa
et al. (1992), Kuniyasu (1999), and Yoshida andMotomura (2009). Descriptions are based on
specimens collected from Yaku-shima Island during ichthyofaunal surveys of the island (Motomura
et al., 2010) and four specimens of A. taeniophorus reported by Arai and Ida (1975).
A photograph of A. indicus is based on a preserved specimen; other specimen photographs
were taken when fresh. Underwater photographs
were taken by the second author (SH) and I.
Takaku (Yakushima-Diving-Life) around the island. Underwater observations have been made by
SH for more than six years, and ecological noteson each species at Yaku-shima Island are given on
the basis of his observations.
Acronyms of the following institutions areused in the text: Australian Museum, Sydney(AMS); Bernice Pauahi Bishop Museum, Honolulu (BPBM); Laboratory of Marine Biology, Kochi
University, Kochi (BSKU); Fisheries ResearchLaboratory, Mie University, Shima (FRLM);
Institute of Oceanic Research and Development,
Tokai University, Shimizu (IORD); Kagoshima
University Museum, Kagoshima (KAUM); Kanagawa Prefectural Museum of Natural History,
Odawara (KPM); Department of Plant and Animal
Sciences, Faculty of Agriculture, University ofMiyazaki, Miyazaki (MUFS); and National Museum of Nature and Science, Tokyo (NSMT). Colorphotographs of fresh specimens are deposited at
BSKU, KAUM, and KPM.
28

List of species of Apogonidae
Apogon amboinensis Bleeker, 1853
[Japanese name: Amami-ishimochi](Figs. 1A, B)
Apogon amboinensis Bleeker, 1853: 329 (type locality: Ambon, Moluccas, Indonesia); Yoshida
and Motomura, 2009: 96, fig. 1 (mouth ofAmbo River, Yaku-shima Island).
Material examined. KAUM–I. 21588, 53.5mm SL, mouth of Ambo River; KAUM–I. 21589,
49.0 mm SL, mouth of Ambo River.
Description. Dorsal-fin rays VI-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
7; circumpeduncular scales 12; total gill rakers
21–22, developed gill rakers 19–20; body lightbrown, with two black stripes, one from snout to
opercular margin through eye, and the other onanterior portion of lateral line; anterior part of firstdorsal fin blackish; and a distinct black spot on
caudal-fin base.
Remarks. Yoshida and Motomura (2009) reported two specimens (KAUM–I. 21588, 21589)
of A. amboinensis as the first records from Yakushima Island and the northernmost records of thespecies. They considered A. amboinensis to reproduce in the mouth of the Ambo River on Yakushima Island because one of the two specimens(KAUM–I. 21588) had about 3,000 eggs in his
mouth. Subsequently, this species was also confirmed by the second author to occur in the mouthof the Isso River on the northern coast of the island on 24 Oct. 2009 and the mouth of the Ambo
River on 8 Jan. 2010 (Fig. 1B).
Apogon amboinensis is distributed in thewestern Pacific, from Papua New Guinea to Japan(Allen et al., 2005). In Japanese waters, this species has been reported from Yaku-shima Island(Yoshida and Motomura, 2009; this study) and the
Ryukyu Islands [Amami-oshima Island (Schmidt,
1930; Okada, 1938; Matsubara, 1963; Yoshigou
et al., 2001), Okinawa Island (Yoshigou et al.,2005), Kume-jima Island (Yoshigou and Nakamura, 2003; Yoshigou, 2007), Miyako-jima Island(Yoshigou et al., 2005), Ishigaki-jima Island (Sakai

Apogonid fishes in Yaku-shima Island

et al., 2001), Iriomote-jima Island (Hayashi andKishimoto, 1983; Yoshigou et al., 2001), and Yonaguni-jima Island (Sakai et al., 2001)]. At Yakushima Island, the species inhabits river mouths on
substrates mixed with mud and deciduous leaves
in depths of 1–2 m.

AB

stripe) reaching a black spot on caudal-fin base;and caudal-fin base spot subequal in size to pupil.
Remarks. Apogon angustatus is widely distributed in the Indo-West Pacific where it ranges
from the Red Sea to Australia and Japan (Allenet al., 2005; Randall, 2005). In Japanese waters,
this species has been reported from Hachijojima Island (Senou et al., 2002) and the Ryukyu
Islands [Amami-oshima Island (Hayashi, 2002),
Okinoerabu-jima Island (Yoshigou et al., 2005),
Ie-jima Island (Senou et al., 2006a), Okinawajima Island (Yoshigou et al., 2005), Sesoko-jimaIsland (Hayashi, 2002), Tokashiki-jima Island(Watai et al., 2009), Miyako Group (Senou et al.,2007), and Iriomote-jima Island (Yoshigou et al.,
2001; Yoshigou and Nakamura, 2002)].
At Yaku-shima Island, specimens of A. angustatus were collected only from Kurio, representing the first reliable records of A. angustatusfrom the island. This solitary species (not forming
schools) lives in the shade of rocky reefs at depthsof less than 15 m at Yaku-shima Island; it can beobserved in coastal areas of the island throughout
the year.

Fig. 1. Apogon amboinensis. A, KAUM–I. 21589, 49.0 mm
SL, mouth of Ambo River; B, mouth of Ambo River, 1
m, 8 Jan. 2010, S. Harazaki.

Apogon angustatus (Smith and Radcliffe, 1911)[Japanese name: Usujima-ishimochi](Fig. 2)
Amia angustata Smith and Radcliffe in Radcliffe,
1911: 253, fig.1 (type locality: Malanipa Island, east of Zamboanga in Mindanao, Philippines).
Material examined. KAUM–I. 20268, 17.8mm SL, Kurio; KAUM–I. 21721, 14.3 mm SL,
Kurio.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
4; circumpeduncular scales 12; total gill rakers
19, developed gill rakers 14; white body, with fivepoorly defined dark brown stripes on lateral surface of body, posterior end of third stripe (middle

Fig. 2. Apogon angustatus. Off Isso, 15 m, 16 Oct. 2009, S.
Harazaki.

Apogon apogonides (Bleeker, 1856)[Japanese name: Aohana-tenjikudai](Fig. 3)Cheilodipterus apogonides Bleeker, 1856a: 37[type locality: Manado, Sulawesi (Celebes),
Indonesia].
29

T. Yoshida et al.

Remarks. The photographed individual (Fig.
3) is herein identified as Apogon apogonides on
the basis of the following combination of characters: pinkish orange body, with brassy yellow
abdomen; two blue lines from front of snout to
opercular margin; bluish spots in three rows on
mid-lateral surface of body; and no black markings on caudal peduncle. Specimens have notbeen collected from Yaku-shima Island.
Apogon apogonides is widely distributed in
the Indo–Pacific where it ranges from the Red
Sea east to the Society Islands, and north to Japan(Allen et al., 2005; Randall, 2005). In Japanesewaters, this species has been reported from the
Izu Islands [Miyake-jima Island (Hayashi, 2002)
and Hachijo-jima Island (Senou et al., 2002) ],the Sagami Sea (Senou et al., 2006b), Shikoku[Oshima Island (Aizawa and Senou, 1991) andKashiwa-jima Island (Hirata et al., 1996)], andthe Ryukyu Islands [Ie-jima Island (Senou et al.,2006a), Okinawa-jima Island (Hayashi, 2002),
and Miyako Group (Senou et al., 2007)].
Figure 3 represents the first confirmed recordof A. apogonides from Yaku-shima Island. This
species is always observed in aggregations with
Apogon notatus throughout the year around a
wreck at a depth of 15 m off Isso, Yaku-shimaIsland; it was not observed elsewhere around theisland. Off Isso, the spawning period of the species is from July to August.

Fig. 3. Apogon apogonides. Off Isso, 18 m, 15 Sept. 2009,
S. Harazaki.

Apogon aureus (Lacepède, 1802)[Japanese name: Aosuji-tenjikudai](Figs. 4A, B)
30

Centropomus aureus Lacepède, 1802: 253, 273(type locality: Mauritius).
Apogon aureus; Ichikawa et al., 1992: 9 (Yakushima Island, Japan).
Material examined. BSKU 96634, 63.3 mm
SL, Isso.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 25; predorsal scales
5; circumpeduncular scales 12; total gill rakers
24, developed gill rakers 21; no oblique dark bars
on cheek; yellowish orange body, with two blue
stripes from snout to opercle through eye; and abroad, black band, its width subequal to pupil diameter, on caudal-fin base.
Remarks. Apogon aureus is widely distributed in the Indo-West Pacific where it ranges fromthe east coast of Africa east to Tonga, and northto Japan (Allen et al., 2005; Randall, 2005). In
Japanese waters, this species has been reported
from the Izu Islands [Miyake-jima Island (Ida andMoyer, 1974; Hayashi, 2002) and Hachijo-jima Island (Senou et al., 2002)], the Sagami Sea (Senou
et al., 2006b), Shikoku [Oshima Island (Aizawa
and Senou, 1991) and Kashiwa-jima Island (Hirata
et al., 1996)], Yaku-shima Island (Ichikawa et al.,1992), and the Ryukyu Islands [Ie-jima Island(Senou et al., 2006a), Miyako Group (Senou etal., 2007), and Iriomote-jima Island (Hayashi andKishimoto, 1983)].
At Yaku-shima Island, only a single specimenof A. aureus was collected off Isso, but it has also
been observed at depths of 6–30 m off the island
throughout the year, sheltering among rocky reefs.
At Yaku-shima Island, spawning of A. aureus begins in early June and ends in July. Although juveniles aggregate, adults usually behave as a pairat the island.
Apogon caudicinctus Randall and Smith, 1988
[Japanese name: Koyari-ishimochi](Fig. 5)
Apogon caudicinctus Randall and Smith, 1988: 2,
fig.1 (type locality: south side of Mei Point,Rapa).

Apogonid fishes in Yaku-shima IslandA

information on A. caudicinctus at Yaku-shima Island is unknown since it is difficult to distinguishthe species from other small reddish species of
Apogon during underwater observation.

B
Fig. 5. Apogon caudicinctus. KAUM–I. 21765, 33.8 mm
SL, Kurio.

Fig. 4. Apogon aureus. A, BSKU 96634, 63.3 mm SL,
Isso; B, off Isso, 18 m, 22 Apr. 2005, S. Harazaki.

Material examined. KAUM–I. 21765, 33.8mm SL, Kurio.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 12; pelvic-fin rays I,
5; pored lateral-line scales 24; predorsal scales 7;
circumpeduncular scales 12; total gill rakers 16,
developed gill rakers 10; second spine length offirst dorsal fin 1.8 in head length; reddish body,with a broad, black circumpeduncular bar, its
width greater than orbit diameter; and edges ofscales on dorsal body dark.
Remarks. Apogon caudicinctus has been
known from Réunion, the Marquesas Islands,
the Pitcairn Islands, Rapa, Fiji, and the Ryukyu
Islands (Randall, 2005). In Japanese waters, thisspecies has been reported from the Ogasawara
Islands (Randall et al., 1997b; Hayashi, 2002)
and the Ryukyu Islands [Amami-oshima Island(Hayashi, 2002), Ie-jima Island (Senou et al.,2006a), Kume-jima Island (Yoshigou and Nakamura, 2003), and Ishigaki-jima Island (Hayashi,
2002)].
At Yaku-shima Island, a single specimen of
A. caudicinctus was collected at a depth of lessthan 3 m in Kurio, representing the first reliable
records of A. caudicinctus from the island and thenorthernmost record for the species. Ecological

Apogon chrysotaenia Bleeker, 1851
[Japanese name: None](Fig. 6)
Apogon chrysotaenia Bleeker 1851: 168 [type locality: Jakarta (Batavia), Java, Indonesia].
Remarks. The photographed individual (taken
off Isso) is identified as Apogon chrysotaenia on
the basis of the following characters reported for
adults of the species: a yellowish orange body,with several faint brownish stripes dorsally; 6 fluorescent blue stripes radiating from front of head;
and tip of second dorsal fin elongated.
Apogon chrysotaenia has been reported only
from the South China Sea, Indonesia, Papua New
Guinea, and Australia (Allen, 2000; Allen et al.,2005), and no records of the species have been
published from Japanese waters. Thus, the photographed individual represents the first record of
the species from Japan. Specimens of the species
from Yaku-shima Island are required to confirm
this new record.
The second author (SH) first observed a single
individual of the species at a depth of 20 m off
Isso, Yaku-shima Island, on 5 July 2004, and hephotographed that individual on 11 July 2004 (Fig.
6). Subsequently, SH found a pair of adult andsubadult individuals at a depth of 30 m off Isso
on 21 November 2009. These individuals usually
shelter under or behind large rocks but are often
forced into open water by strong unidirectional
currents around the rocks.
31

T. Yoshida et al.

Fig. 6. Apogon chrysotaenia. Off Isso, 20 m, 11 July 2004,
S. Harazaki.

Apogon cookii Macleay, 1881
[Japanese name: Suji-ishimochi](Figs. 7A–C)
Apogon cookii Macleay, 1881: 344 (type locality:
Endeavour River and Darnley Island, Australia).
Material examined. KAUM–I. 11207, 27.3mm SL, Kurio; KAUM–I. 21714, 31.0 mm SL,
Kurio; KAUM–I. 21715, 28.0 mm SL, Kurio;
KAUM–I. 21716, 22.0 mm SL, Kurio; KAUM–I.
21717, 18.0 mm SL, Kurio; NSMT–P 91664, 76.0mm SL, Kurio.
Description. Dorsal-fin rays VII-I, 9; anal-fin
rays II, 8; pectoral-fin rays 14; pelvic-fin rays I, 5;
pored lateral-line scales 24; predorsal scales 3–4;
circumpeduncular scales 12; total gill rakers 18–
20, developed gill rakers 12–13; body white, with
six dark brown stripes on lateral surface of body;third stripe beginning from upper eye, ending at
middle of body (below second dorsal-fin origin);
fourth stripe (middle stripe) posteriorly reaching
to a black spot on caudal-fin base; and caudal-finbase spot subequal in size to pupil diameter.
Remarks. Apogon cookii is similar to A.
nigrofasciatus, but the former differs in having
12–13 (vs. 15–19 in the latter) developed gill rakers, 15 (vs. usually 14) pectoral-fin rays, and a
postocular stripe (vs. no stripe) (Gon and Randall,2003).
Apogon cookii is widely distributed in the
Indo-West Pacific where it ranges from the Red
Sea and east coast of Africa east to New Caledonia and Japan (Allen et al., 2005; Randall, 2005).
32

In Japanese waters, this species has been reported
from the Sagami Sea (Senou et al., 2006b) and the
Ryukyu Islands [Amami-oshima Island (Yoshigou
et al., 2001), Ie-jima Island (Senou et al., 2006a),
Okinawa-jima Island (Yoshigou and Nakamura,
2002, 2003; Yoshigou et al., 2005), Sesoko-jimaIsland (Yoshigou and Nakamura, 2003), Ou-jimaIsland (Yoshigou et al., 2005), Kume-jima Island(Yoshigou and Nakamura, 2003), Tokashiki-jimaIsland (Watai et al., 2009), Minamidaitou-jimaIsland (Yoshigou et al., 2005), Shimoji-shimaIsland (Yoshigou et al., 2005), Miyako Group(Senou et al., 2007), Iriomote-jima Island (Hayashi
and Kishimoto, 1983), and Yonaguni-jima Island(Yoshigou et al., 2001)].
At Yaku-shima Island, specimens of A. cookii
were collected only from Kurio and these specimens represent the first reliable records of A.
cookii from the island. This solitary species (notforming schools) lives in the shade of rocky reefs
at depths of less than 5 m at Yaku-shima Island;
it can be observed in coastal areas of the island
throughout the year.

A

B

C

Fig. 7. Apogon cookii. A, KAUM–I. 11207, 27.3 mm SL,
Kurio; B, off Isso, 5 m, 22 May 2007, S. Harazaki; C,
off Isso, 3 m, 14 Apr. 2005, S. Harazaki.

Apogonid fishes in Yaku-shima Island

Apogon crassiceps Garman, 1903
[New Japanese name: Akafuji-tenjikudai](Figs. 8A, B)
Apogon crassiceps Garman, 1903: 230 (type locality: Suva Reef, Viti Levu, Fiji).
Material examined. BSKU 96552, 27.2mm SL, Yudomari; KAUM–I. 11445, 37.4mm SL, Isso; KAUM–I. 20159, 20.3 mm SL,
Yudomari; KAUM–I. 20196, 42.5 mm SL, Yudomari; KAUM–I. 20205, 22.8 mm SL, Yudomari; KAUM–I. 20206, 17.0 mm SL, Yudomari;
KAUM–I. 20249, 24.9 mm SL, Kurio; KAUM–I.
20331, 34.3 mm SL, Isso; KAUM–I. 21767, 29.2mm SL, Kurio; KAUM–I. 21768, 27.2 mm SL,
Kurio; KPM–NI 22513, 24.4 mm SL, Yudomari;
KPM–NI 22546, 28.4 mm SL, Kurio.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 13; pelvic-fin rays I,
5; pored lateral-line scales 24 ; predorsal scales
5–6; large scale between first dorsal fin and lateralline 1 (often with a smaller second scale at base of
dorsal fin); circumpeduncular scales 12; total gillrakers 15–16, developed gill rakers 8–9; full scale
between the first dorsal fin and the lateral line 1,
often with a smaller second scale; a free edge ofskin near the anterior nasal opening; and body
semitransparent red, with a thin, dark red stripe on
mid-lateral surface of body when alive.
Remarks. The Apogon coccineus complex,
comprising A. campbelli Smith 1949, A. coccineus Rüppell 1838, A. crassiceps Garman 1903,
A. kautamea Greenfield and Randall 2004, andA. kominatoensis Ebina, 1935 (see Greenfield,
2001; Greenfield and Randall, 2004), was definedby Greenfield (2001) as having a free edge ofskin near the anterior nasal opening, and a single,
large, full scale (often a smaller second scale) between the lateral line and the dorsal fin. Our specimens from Yaku-shima Island is herein identified
as A. crassiceps of the A. coccineus complex anddiffer from A. campbelli and A. kautamea in having 13 pectoral-fin rays (vs. 14 rays in the lattertwo species; Greenfield and Randall, 2004) andfrom A. kominatoensis in lacking numerous small
brownish spots scattered on the body (vs. having the spots in A. kominatoensis; Ebina, 1935).

Apogon coccineus is an Indian Ocean species(Randall, 2005).
All previous records of A. crassiceps “Akanetenjikudai” in Japanese waters (e.g., Masuda et al.,
1984; Hayashi, 2000, 2002) were based on misidentifications of a species of the A. talboti group(sensu Greenfield, 2007b). Comparisons of A.
crassiceps from Yaku-shima Island with “Akanetenjikudai” (= A. crassiceps in Japanese literature)from Ie-jima and Iriomote-jima Islands revealed
that the former differs from the latter in having asemitransparent red body (vs. opaque dark red),a thin, dark red stripe on the mid-lateral surfaceof the body (vs. stripe absent), 15 or 16 total gillrakers (vs. 18 or 19), and a single full scale (oftenwith a smaller second scale) between the first dorsal fin and lateral line (vs. 2 large full scales, with
1 smaller third scale) (this study; Hayashi, 2000,
2002). In addition, A. crassiceps attains 5 cm SL(Randall, 2005), whereas “Akane-tenjikudai”
reaches 12 cm SL (Hayashi, 2000, 2002).
“Akane-tenjikudai” is most similar to A.
talboti Smith, 1961 in overall body appearance
and sharing the following characters (“Akanetenjikudai” vs. A. talboti): dorsal-fin rays VI-I, 9(vs. VI-I, 9); anal-fin rays II, 8 (II, 8); pectoral-fin
rays 13 or 14 (13); pored lateral-line scales 24 (24);total gill rakers 19 (20 or 21), developed rakers
18 (16–19); and supraneural bones 3 (3) [data for“Akane-tenjikudai” were based on KPM–NI 2524
(79.9 mm SL, Ie-jima Island, Japan, 16 m depth,
coll. by H. Senou et al., 17 Jan. 1996), Masuda
et al. (1984), and/or Hayashi (2000, 2002), anddata for A. talboti were based on Gon and Randall (2003), Greenfield and Randall (2004), andGreenfield (2007b)]. Accordingly, we tentatively
identify “Akane-tenjikudai” as A. talboti, andpropose a new standard Japanese name, Akafujitenjikudai, for A. crassiceps.
Apogon crassiceps is widely distributed in
the western Pacific where it ranges from the Line
Islands and Tauamotu east to Australia, and northto Japan (Allen et al., 2005; Randall, 2005; thisstudy). At Yaku-shima Island, specimens of A.
crassiceps were collected from Isso, Kurio, andYudomari in depths of less than 5 m, representing the first reliable records of A. crassiceps fromthe island. The photographed individual (Fig. 8B)
33

T. Yoshida et al.

is herein identified as A. crassiceps by having abody semitransparent red, with a thin, dark red
stripe on the mid-lateral surface of the body.

A

B

Fig. 8. Apogon crassiceps. A, KAUM–I. 20331, 34.3 mm
SL, Isso; B, off Isso, 8 m, 22 Oct. 2009, S. Harazaki.

Apogon doederleini Jordan and Snyder, 1901
[Japanese name: Osuji-ishimochi](Figs. 9A, B)
Apogon doederleini Jordan and Snyder, 1901:
901, fig. 6 (type locality: Nagasaki, Japan);Ichikawa et al., 1992: 9 (Yaku-shima Island,
Japan); Kuniyasu, 1999: 12 (Kurio, Yakushima Island, Japan).
Material examined. FRLM 34706, 103.7mm SL, Yudomari; KAUM–I. 11124, 99.2 mm
SL, Ambo; KAUM–I. 11440, 46.5 mm SL, Isso;
KAUM–I. 11441, 47.5 mm SL, Isso; KAUM–
I. 11442, 46.5 mm SL, Isso; KAUM–I. 11443,
52.7 mm SL, Isso; KAUM–I. 11444, 44.5 mm
SL, Isso; KAUM–I. 11446, 48.7 mm SL, Isso;
KAUM–I. 11447, 43.9 mm SL, Isso; KAUM–
I. 11448, 41.2 mm SL, Isso; KAUM–I. 11449,
41.5 mm SL, Isso; KAUM–I. 11450, 51.6 mm
SL, Isso; KAUM–I. 11451, 52.1 mm SL, Isso;
KAUM–I. 11452, 51.9 mm SL, Isso; KAUM–I.
20071, 85.7 mm SL, Yudomari; KAUM–I. 20072,
62 mm SL, Yudomari; KAUM–I. 20343, 56 mm
SL, Isso; KAUM–I. 21751, 91.9 mm SL, Ambo;
34

KAUM–I. 25234, 78.1 mm SL, Ambo; MUFS
25446, 92.7 mm SL, Ambo; MUFS 25447, 85.4mm SL, Ambo; MUFS 25448, 70.9 mm SL,
Ambo; MUFS 25449, 69.2 mm SL, Ambo; MUFS
25450, 69.4 mm SL, Ambo.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 15; pelvic-fin raysI, 5; pored lateral-line scales 24 ; predorsal scales
3; circumpeduncular scales 12; total gill rakers
18, developed gill rakers 12; body pinkish brown,with four dark brown stripes on lateral surface ofbody; third stripe (middle stripe) posteriorly notreaching to a black spot on caudal-fin base; andcaudal-fin base spot subequal in size to pupil diameter.
Remarks. Apogon doederleini is distributedin the Indo-West Pacific where it ranges from
Western Australia east to the Kermadec Islands,
and north to Japan, but no records from Indonesia
or New Guinea (Randall et al., 1997a; Randall,2005). In Japanese waters, this species has beenreported from the Izu Islands [Hachijo-jima Island(Senou et al., 2002)], the Ogasawara Islands (Kuwamura et al., 1983; Sato, 1991; Randall et al.,1997b), Tateyama Bay (Hagiwara and Kimura,2005), the Sagami Sea (Senou et al., 2006b),
Shikoku [Oshima Island (Aizawa and Senou,
1991), Kochi (Kamohara, 1964), and Kashiwajima Island (Hirata et al., 1996)], Nagasaki (Jordan
and Snyder, 1901), Yaku-shima Island (Ichikawa
et al., 1992; Kuniyasu, 1999), and the Ryukyu Islands [Amami-oshima Island (Yamakawa, 1969),
Yagaji-shima Island (Yoshigou and Nakamura,
2002; Yoshigou et al., 2005), Ie-jima Island (Senou
et al., 2006a), Okinawa-jima Island (Yoshigou etal., 2001; Yoshigou et al., 2005), and Iriomotejima Island (Hayashi and Kishimoto, 1983)].
At Yaku-shima Island, specimens of A. doederleini were collected from Ambo, Isso, Kurio
and Yudomari. The species has been observed
at depths of less than 10 m in coastal areas of
the island throughout the year, usually sheltering among rocky reefs in the daytime. Althoughjuveniles form small aggregations, the adults areusually solitary.

Apogonid fishes in Yaku-shima Island

A

B

Fig. 9. Apogon doederleini. A, KAUM–I. 11124, 99.2 mm
SL, Ambo; B, off Isso, 5 m, 26 Jan. 2005, S. Harazaki.

species has been reported from the Ogasawara
Islands (Randall et al., 1997b), Tateyama Bay(Hagiwara and Kimura, 2005), Sagami Bay (Senou, 1999), the Sagami Sea (Senou et al., 2006b),
Shikoku [Kochi (Kamohara, 1964) and Kashiwajima Island (Hirata et al., 1996)], Yaku-shimaIsland (Arai and Ida, 1975; Ichikawa et al., 1992;
Kuniyasu, 1999), and the Ryukyu Islands [Amami-oshima Island (Kamohara and Yamakawa,1965) and the Kerama Islands (Hayashi, 2002)].
At Yaku-shima Island, specimens of A.
endekataenia were collected only from Isso. This
species is always observed in aggregations ofabout 10 individuals in depths of 25–30 m off
Isso throughout the year; not observed elsewherearound the island.

A

Apogon endekataenia Bleeker, 1852
[Japanese name: Kosuji-ishimochi](Figs. 10A, B)
Apogon endekataenia Bleeker, 1852: 449 [typelocality: Bangka (Banka) or Lepar Island, Indonesia]; Arai and Ida, 1975: 192 (Kusugawa,Yaku-shima Island, Japan); Ichikawa et al.,
1992: 9 (Yaku-shima Island, Japan); Kuniyasu, 1999: 12 (Kurio, Yaku-shima Island,
Japan).
Material examined. KAUM–I. 21786, 26.8mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; anal-fin
rays II, 8; pectoral-fin rays 14; pelvic-fin rays I,
5; pored lateral-line scales 24 ; predorsal scales 3;
circumpeduncular scales 12; total gill rakers 17,
developed gill rakers 12; body white, with about
seven yellowish brown stripes on lateral surface ofbody; fifth stripe (middle stripe) posteriorly reaching to a black spot on caudal-fin base; caudal-finbase spot subequal in size to pupil diameter.
Remarks. Apogon endekataenia is widelydistributed in the western Pacific (Randall et al.,1997a; Hayashi, 2002). In Japanese waters, thisB

Fig. 10. Apogon endekataenia. A, KAUM–I. 21786, 26.8mm SL, Isso; B, off Isso, 30 m, 16 Sept. 2009, S.
Harazaki.

Apogon exostigma (Jordan and Starks, 1906)[Japanese name: Yukata-ishimochi](Figs. 11A, B)
Amia exostigma Jordan and Starks, 1906: 238, fig.
31 (type locality: Apia, Upolu Island, Western
Samoa).
Material examined. BSKU 96553, 45.8 mm
SL, Yudomari; KPM–NI 22551, 28.1 mm SL, Kurio.
35

T. Yoshida et al.

Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 23; predorsal scales
4; circumpeduncular scales 12; total gill rakers
16–17, developed gill rakers 10–12; no dark saddle on body below second dorsal-fin base; body
depth 3.0–3.4 in SL; body whitish silver, with a
middle stripe extending from front of snout to a
black spot, smaller than pupil diameter, at caudalfin base; and lower margin of caudal-fin base spot
on lateral line.
Remarks. Apogon exostigma is widely distributed in the Indo-West Pacific where it ranges
from the Red Sea east to the Line and Pitcairn
Islands, and north to Japan (Allen et al., 2005;
Randall, 2005). In Japanese waters, this species
has been reported only from the Ryukyu Islands[Amami-oshima Island (Kamohara and Yamakawa, 1968), Ie-jima Island (Senou et al., 2006a),
Okinawa-jima Island (Yoshigou and Nakamura,2002), Tokashiki-jima Island (Watai et al., 2009),
Miyako Group (Senou et al., 2007), Ishigaki-jimaIsland (Yoshigou et al., 2001), and Iriomote-jimaIsland (Hayashi and Kishimoto, 1983)].
Two specimens of A. exostigma, collectedfrom Kurio and Yudomari, represent the first reliable records of the species from Yaku-shima Island and the northernmost records for the species.
This solitary species lives in the shade of rocky
reefs in depths of less than 10 m at Yaku-shimaIsland; it can be observed in coastal areas of theisland throughout the year (relatively common).

A

B

Fig. 11. Apogon exostigma. A, KPM–NI 22551, 28.1 mm
SL, Kurio; B, off Isso, 12 m, 21 Sept. 2009, S. Harazaki.

36

Apogon fasciatus (White, 1790)[Japanese name: Furai-ishimochi](Fig. 12)
Mullus fasciatus White, 1790: 268, pl. 53, fig.
1 (type locality: Port Jackson, New South
Wales, Australia).
Remarks. Many Japanese authors (e.g.,
Hayashi, 2000, 2002; Senou et al., 2006b) haveused a name, Apogon quadrifasciatus Cuvier,
1828, for this species. However, Randall andLachner (1986) and Fraser (2005) regarded A.
quadrifasciatus as a junior synonym of A. fasciatus.
The photographed individual (Fig. 12) is
identified as Apogon fasciatus on the basis of thefollowing combination of characters: a semitransparent body, with two black stripes on the lateralsurface of the body, the upper stripe extending
from the snout to the dorsal portion of the caudalfin base via above the eye, the lower stripe fromthe snout to the caudal-fin margin via the middleof the eye; and two whitish stripes (or dotted
stripes) running above and below each black
stripe.
Apogon fasciatus is widely distributed in the
Indo-West Pacific where it ranges from the Red
Sea east to Australia, and north to Japan (Allenet al., 2005; Fraser, 2005). In Japanese waters,
this species has been reported as A. quadrifasciatus from the Sagami Sea (Senou et al., 2006b),
Nagasaki Prefecture and southward (Hayashi,
2002), including the Ryukyu Islands [Okinawajima Island (Yoshigou and Nakamura, 2003) and
Iriomote-jima Island (Hayashi and Kishimoto,1983)].
Figure 12 represents the first confirmed record of A. fasciatus from Yaku-shima Island. The
second author observed this species forming aschool with five or six individuals behind a big
piece of driftwood over a sandy bottom at a depthof 20–25 m off Isso, Yaku-shima Island. No otherindividuals have recently been confirmed aroundYaku-shima Island; it seems to be very rare in theisland. Specimens have not been collected from
Yaku-shima Island.

Apogonid fishes in Yaku-shima Island

Although juveniles ca. 3–4 cm SL are common in
the island, adults over ca. 10 cm SL are relatively
rare.

AFig. 12. Apogon fasciatus. Off Isso, 20 m, 10 Sept. 2005, S.
Harazaki.

Apogon fraenatus Valenciennes, 1832
[Japanese name: Hitosuji-ishimochi](Figs. 13A, B)

B

Apogon fraenatus Valenciennes, 1832: 57, pl. 4,
fig. 4 (type locality: New Guinea).
Remarks. Photographed individuals (Figs.
13A, B) are identified as A. fraenatus on the basis of the following combination of characters:a pinkish silver body with a dark black stripe on
lateral surface of body, middle stripe posteriorly
reaching to a black spot on caudal-fin base; andcaudal-fin base spot subequal in size to pupil diameter. Specimens have not been collected from
Yaku-shima Island.
Apogon fraenatus is widely distributed in the
Indo-West Pacific where it ranges from the Red
Sea and Gulf of Oman east to the Line Islands
and Tuamotu Archipelago, and north to Japan(Allen et al., 2005; Randall, 2005). In Japanesewaters, this species has been reported from the
Izu Islands [Miyake-jima Island (Hayashi, 2002)
and Hachijo-jima Island (Senou et al., 2002)],
Wakayama Prefecture (Hayashi, 2002), Kashiwajima Island (Hirata et al., 1996), and the Ryukyu
Islands [Amami-oshima Island (Kamohara andYamakawa, 1965), Ie-jima Island (Senou et al.,2006a), Okinawa-jima Island (Yoshigou et al.,2005), Miyako Group (Senou et al., 2007), andIriomote-jima Island (Hayashi and Kishimoto,
1983; Yoshigou et al., 2001; Yoshigou and Nakamura, 2002)].
Figure 13 represents the first confirmed recordof A. fraenatus from Yaku-shima Island. This species is always observed in aggregations, sheltering
among rocky reefs, throughout the year aroundYaku-shima Island at depths of less than 20 m.

Fig. 13. Apogon fraenatus. A, off Isso, 25 m, 15 Sept.
2009, S. Harazaki; B, off Isso, 10 m, 14 Jan. 2010, S.
Harazaki.

Apogon indicus Greenfield, 2001
[Japanese name: Ryukyu-ishimochi](Fig. 14)
Apogon indicus Greenfield, 2001: 465, figs.
2B, C (type locality: off le Morne, passe del’Ambulante, southwestern coast of Mauritius).
Material examined. NSMT–P 95448, 32.9mm SL, Haruo.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 13; pelvic-fin rays I,
5; pored lateral-line scales 23; predorsal scales 6;
large scales between first dorsal fin and lateral line
2, with a smaller third scale at base of dorsal fin;circumpeduncular scales 14; total gill rakers 15,
developed gill rakers 7; no free edge of skin nearanterior nasal opening; and body whitish, withblack pigments scattered on anterodorsal portionof body and caudal-fin base when preserved.
Remarks. Many Japanese authors (e.g.,
Hayashi, 2000, 2002) have used a name, Apogon
erythrinus Snyder, 1904, for this species. However, Greenfield (2001) reviewed the Apogon
37

T. Yoshida et al.

erythrinus complex and reclassified “A. erythrinus” into four species: A. erythrinus, endemic
to the Hawaiian Islands and Johnston Atoll; A.
marquesensis Greenfield, 2001, endemic to theMarquesas Islands; A. indicus, widespread in the
Indo–Pacific; and A. susanae Greenfield, 2001,
restricted to the Pacific Ocean. The A. erythrinus
complex is characterized by the following two
characters: skin at the end of the snout covering
the nasal bones and over the ascending processesof the premaxilla smooth with no free edge near
the anterior nasal opening, and two large fullscales (often a smaller third scales at base of firstdorsal fin) between the lateral line and the third
spine of the first dorsal fin (Greenfield, 2001).
Apogon indicus is distinguished from othermembers of the complex by having 12 or 13
pectoral-fin rays (vs. usually 14 rays in the latter;
Greenfield, 2001). Yoshigou et al. (2005) reported
eight specimens of the species (2 specimens from
Okinoerabu-jima Island, 1 specimen from Irabujima Island, and 5 specimens from Okinawa-jimaIsland) from Japan as A. indicus (instead of A.
erythrinus) for the first time.
Apogon indicus has been known from Mauritius, Vietnam, Philippines, Palau, Mariana Islands,
New Caledonia, Tonga, and American Samoa(Randall, 2005). In Japanese waters, this species
has been reported as A. erythrinus [except for
Yoshigou et al. (2005, as A. indicus)] from Hachijo-jima Island (Senou et al., 2002), Kashiwajima Island (Hirata et al., 1996), and the Ryukyu
Islands [Okinoerabu-jima Island (Yoshigou etal., 2005), Ie-jima Island (Senou et al., 2006a),
Okinawa-jima Island (Yoshigou and Nakamura,
2002, 2003; Yoshigou et al., 2005), Tokashikijima Island (Watai et al., 2009), Irabu-jima Island(Yoshigou et al., 2005), Miyako Group (Senou etal., 2007), and Iriomote-jima Island (Hayashi andKishimoto, 1983; Yoshigou et al., 2001)].
At Yaku-shima Island, a single specimen of A.
indicus was collected at a depth of 4 m off Haruo,
representing the first reliable record of A. indicus
from the island. Ecological information on A. indicus at Yaku-shima Island is unknown since it isdifficult for us to distinguish it from other reddish
species of Apogon during underwater observation.

38

Fig. 14. Apogon indicus. NSMT–P 95448, 32.9 mm SL,
Haruo.

Apogon kallopterus Bleeker, 1856
[Japanese name: Kasuri-ishimochi](Fig. 15)
Apogon kallopterus Bleeker, 1856a: 33 [typelocality: Manado, Sulawesi (Celebes), Indonesia]; Kuniyasu, 1999: 12 (Kurio, Yaku-shimaIsland, Japan).
Remarks. The photographed individual (Fig.
15) is identified as Apogon kallopterus on thebasis of the following combination of characters:a pale pinkish brown body, with an indistinct,
broad, black stripe on lateral surface of body; anterior part of first dorsal fin yellowish; and poorly
defined black spot, its size greater than pupil diameter, above middle of caudal-fin base.
Apogon kallopterus is widely distributed in
the Indo–Pacific where it ranges from the Red
Sea and the east coast of Africa to the Hawaiian and Pitcairn Islands, including Japan (Allenet al., 2005; Randall, 2005). In Japanese waters,
this species has been reported from Hachijo-jimaIsland (Senou et al., 2002), the Ogasawara Islands(Randall et al., 1997b; Hayashi, 2002), Wakayama
Prefecture and southward (Hayashi, 2002), including Yaku-shima Island (Kuniyasu, 1999), andthe Ryukyu Islands [Ie-jima Island (Senou et al.,2006a), Okinawa-jima Island (Yoshigou and Nakamura, 2003; Yoshigou et al., 2005), Tokashikijima Island (Watai et al., 2009), Miyako Group(Senou et al., 2007), and Iriomote-jima Island(Hayashi and Kishimoto, 1983)].
Apogon kallopterus lives solitarily (not forming schools) in the shade of rocky reefs at depthsof less than 10 m at Yaku-shima Island; it can beApogonid fishes in Yaku-shima Island

observed in coastal areas of the island throughout
the year. Although it is common around the island,no specimens were collected during this study.

Fig. 15. Apogon kallopterus. Off Isso, 6 m, 11 Sept. 2009, S.
Harazaki.

Apogon kominatoensis Ebina, 1935
[Japanese name: Kominato-tenjikudai](Fig. 16)

ish spots scattered on head and body when alive,
and without a thin, dark red stripe on mid-lateralsurface of body; and no black mark on caudal peduncle or lower lobe of caudal fin.
Remarks. Many Japanese authors (e.g.,
Hayashi, 2000, 2002) have used a name, Apogoncoccineus Rüppell, 1838, for this species. However, Randall (2005) has recently regarded that A.
coccineus is distributed only in the western Indian
Ocean. Our specimens from Yaku-shima Island
appear to be identical with A. kominatoensis, a
species of the A. coccineus complex (see Remarksof A. crassiceps). Because previous Japanese records of “A. coccineus” may include more than
one species, the reliable distributional range of A.
kominatoensis in Japan is unknown.
At Yaku-shima Island, specimens of A. kominatoensis were collected from Isso, Ambo, Haruo,
Yudomari, and Kurio, at depths of less than 5 m.

Apogon kominatoensis Ebina, 1935: 212, fig.1[type locality: tidepools at Kominato, Chiba,(Toba-ken: probably Chiba-ken), Japan].
Material examined. BSKU 96655, 29.7 mm
SL, Isso; FRLM 34709, 38.3 mm SL, Yudomari;
KAUM–I. 11133, 29.0 mm SL, Kurio; KAUM–I.
11290, 37.7 mm SL, Yudomari; KAUM–I. 11291,
42.3 mm SL, Yudomari; KAUM–I. 11652, 27.9mm SL, Ambo; KAUM–I. 20041, 40.2 mm SL,
Yudomari; KAUM–I. 20248, 39.3 mm SL, Kurio;
KAUM–I. 20259, 41.3 mm SL, Kurio; KAUM–
I. 20260, 41.2 mm SL, Kurio; KAUM–I. 20261,
32.7 mm SL, Kurio; KAUM–I. 20332, 35.4 mm
SL, Isso; KAUM–I. 21766, 27.1 mm SL, Kurio;
NSMT–P 91356, 40 mm SL, Yudomari; NSMT–P
91357, 37 mm SL, Yudomari; NSMT–P 95461,
36.7 mm SL, Haruo.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 13; pelvic-fin rays I,
5; pored lateral-line scales 22 ; predorsal scales
5–6; large full scale between first dorsal fin andlateral line 1, often with a smaller second scale atbase of dorsal fin; circumpeduncular scales 12;
total gill rakers 15–16, developed gill rakers 8; afree edge of skin near the anterior nasal opening;
body semitransparent red, with numerous brown-

Fig. 16. Apogon kominatoensis. KAUM–I. 20332, 35.4mm SL, Isso.

Apogon moluccensis Valenciennes, 1832
[Japanese name: Sehoshi-tenjikudai](Figs. 17A, B)
Apogon moluccensis Valenciennes, 1832: 54 [typelocality: Ambon, Moluccas, Indonesia].
Material examined. BSKU 96640, 46.8 mm
SL, Isso; KAUM–I. 20378, 43.1 mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
3; circumpeduncular scales 12; total gill rakers
22–23, developed gill rakers 19–21; body yellowish brown, with 2 reddish stripes, upper narrower
stripe along anterior portion of lateral line, and39

T. Yoshida et al.

lower broader stripe from snout to opercular margin through eye; about 7 vertical, brownish bars
on lateral surface of lower body, a small whitespot at posterior end of second dorsal-fin base;and no black spot on caudal peduncle.
Remarks. Many Japanese authors (e.g.,
Hayashi, 2000, 2002) have used a name, Apogonventrifasciatus Allen, Kuiter and Randall, 1994,
for this species. However, Fraser et al. (2002) regarded A. ventrifasciatus as a junior synonym of A.
moluccensis.
Apogon moluccensis is widely distributedin the western Pacific, from Australia to Japan(Hayashi, 2002; Randall et al., 1997a). In Japanese waters, this species has been reported as A.
ventrifasciatus from the Ryukyu-Islands [Okinawa-jima Island (Hayashi, 2002) and Iriomote-jimaIsland (Hayashi and Yano, 1996; Hayashi, 2002)].
At Yaku-shima Island, two specimens of A.
moluccensis were collected from Isso, representing the first reliable records of A. moluccensis
from the island and the northernmost records forthe species. This species is always observed inaggregations (ca. 20 individuals) throughout theyear around a wreck at a depth of 4 m off Isso,
Yaku-shima Island; no other localities confirmed
from the island.

A

B

Fig. 17. Apogon moluccensis. A, KAUM–I. 20378, 43.1mm SL, Isso; B, off Isso, 20 m, 18 Sept. 2007, S.
Harazaki.

40

Apogon nigrofasciatus Lachner, 1953
[Japanese name: Minami-futosuji-ishimochi](Figs. 18A, B)
Apogon nigrofasciatus Lachner, 1953: 466, fig.
81, pl. 37C, D (type locality: Yuro Island, Bikini Atoll, Marshall Islands); Kuniyasu, 1999:
12 (Kurio, Yaku-shima, Japan).
Apogon aroubiensis (not of Hombron and Jacquinot); Ichikawa et al., 1992: 9 (Yaku-shimaIsland, Japan).
Material examined. BSKU 96605, 58.9 mm
SL, Kurio.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
4; circumpeduncular scales 12; total gill rakers
24, developed gill rakers 18; snout pointed; bodywhite, with 5 broad, dark reddish brown stripes
on lateral surface of body; middle stripes broader
than pale interspace; and stripes not extending
onto caudal fin.
Remarks. Apogon nigrofasciatus is widelydistributed in the Indo-West Pacific where it ranges from the Red Sea and Gulf of Aden east to the
Marshall Islands and Tuamotu Archipelago, andnorth to Japan (Allen et al., 2005; Randall, 2005).
In Japanese waters, this species (as A. aroubiensis or A. nigrofasciatus) has been reported from
Hachijo-jima Island (Senou et al., 2002), the Ogasawara Islands (Randall et al., 1997b), the Sagami
Sea (Senou et al., 2006b), Kashiwa-jima Island(Hirata et al., 1996), Yaku-shima Island (Ichikawa
et al., 1992; Kuniyasu, 1999), and the Ryukyu
Islands [Amami-oshima Island (Kamohara andYamakawa, 1965; Yoshigou et al., 2001), Okinoerabu-jima Island (Yoshigou et al., 2005), Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou and Nakamura, 2002, 2003; Yoshigou
et al., 2005), Tokashiki-jima Island (Watai et al.,2009), Miyako Group (Senou et al., 2007), andIriomote-jima Island (Hayashi and Kishimoto,1983)].
At Yaku-shima Island, only a single specimenof A. nigrofasciatus was collected off Kurio, but it
has been observed at depths of 10–30 m throughout the year, sheltering among rocky reefs. At

Apogonid fishes in Yaku-shima IslandYaku-shima Island, spawning of A. nigrofasciatus
begins in early April and ends in May.

AB

len et al., 2005; Randall et al., 1997a). In Japanese waters, this species has been reported from
Hachijo-jima Island (Senou et al., 2002), Sagami
Bay (Senou, 1999), the Sagami Sea (Senou et al.,2006b), Shikoku [Oshima Island, (Aizawa andSenou, 1991) and Kashiwa-jima Island (Hirata etal., 1996)], Yaku-shima Island (Jordan and Starks,
1906; Ichikawa et al., 1992), and the Ryukyu
Islands [Ie-jima Island (Senou et al., 2006a), Miyako Group (Senou et al., 2007), and Iriomotejima Island (Hayashi and Kishimoto, 1983)].
Apogon notatus has been observed only at
two diving spots at a depth of 20 m off Isso. A
single school usually comprises more than 1,000
individuals, the number of individuals decreasing
to ca. 100 in winter. At Yaku-shima Island, thespawning period of the species is from May to
July.

Fig. 18. Apogon nigrofasciatus. A, BSKU 96605, 58.9 mm
SL, Kurio; B, off Isso, 15 m, 11 Sept. 2009, S. Harazaki.

Apogon notatus (Houttuyn, 1782)[Japanese name: Kurohoshi-ishimochi](Fig. 19)
Sparus notatus Houttuyn, 1782: 320 (type locality: Japan)
Amia notata: Jordan and Starks, 1906: 698, fig. 4(Yaku-shima Island, Japan).
Apogon notatus: Ichikawa et al., 1992: 9 (Yakushima Island, Japan).
Remarks. Although the specimen’s whereabouts is unknown, Jordan and Starks’ figure (fig.
4 as Amia notata) clearly shows it to be identified
as Apogon notatus in having a black band fromthe snout to the eye, a distinct spot on the nape, a
black spot on the caudal-fin base, a black blotch
on the first dorsal fin distally, and blackish bands
along the bases of second dorsal and anal fins. The
photographed individual (Fig. 19) is also herein
identified as A. notatus.
Apogon notatus is widely distributed in the
Indo-West Pacific, from Australia to Japan (Al-Fig. 19. Apogon notatus. Off Isso, 20 m, 20 June 2005, S.
Harazaki.

Apogon novemfasciatus Cuvier, 1828
[Japanese name: Tasuji-ishimochi](Fig. 20)
Apogon novemfasciatus Cuvier in Cuvier andValenciennes, 1828: 154 (type locality: Timorand Guam); Kuniyasu, 1999: 12 (Kurio, Yakushima Island, Japan).
Material examined. KAUM–I. 21759, 33.7mm SL, Ambo.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
4; circumpeduncular scales 12; total gill rakers
41

T. Yoshida et al.

20, developed gill rakers 16; snout pointed; bodywhite, with five poorly defined dark brown to
black stripes on lateral surface of body; and third
stripe (middle stripe) uneven in width, reachingcaudal fin, but not ending a spot.
Remarks. Apogon novemfasciatus is widelydistributed in the Indo-West Pacific where it
ranges from the Cocos Keeling Islands east to
the Line Islands, and the islands of Micronesia,
and north to Japan (Allen et al., 2005; Randall,2005). In Japanese waters, this species has beenreported from the Ogasawara Islands (Randall
et al., 1997b), the Sagami Sea (Senou et al.,2006b), Shikoku [Kochi (Kamohara, 1964) andKashiwa-jima Island (Hirata et al., 1996)], Yakushima Island (Kuniyasu, 1999), and the Ryukyu
Islands [Amami-oshima Island (Kamohara andYamakawa, 1965; Yoshigou et al., 2001), Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou et al., 2001; Yoshigou and Nakamura,
2002, 2003), Sesoko-jima Island (Yoshigou andNakamura, 2003), Tokashiki-jima Island (Wataiet al., 2009), Miyako Group (Senou et al., 2007),
Ishigaki-jima Island (Yoshigou et al., 2001),
Iriomote-jima Island (Hayashi and Kishimoto,1983), and Yonaguni-jima Island (Yoshigou et al.,2001)].
At Yaku-shima Island, a single specimen of
A. novemfasciatus was collected off Ambo. This
species lives solitarily in the shade of rocky reefs(not forming schools) at depths of less than 1 m at
Yaku-shima Island; it can be observed in coastal
areas of the island throughout the year.

Fig. 20. Apogon novemfasciatus. KAUM–I. 21759, 33.7mm SL, Ambo.

42

Apogon parvulus (Smith and Radcliffe, 1912)[Japanese name: Neon-tenjikudai]
Amia parvula Smith and Radcliffe in Radcliffe,
1912: 432, pl. 34, fig. 2 (type locality: Tataan
Pass, Tawi Tawi Group, Philippines).
Apogon parvulus: Ichikawa et al., 1992: 9 (Yakushima Island, Japan).
Remarks. The identification of Ichikawa etal.’s (1992) Apogon parvulus is uncertain. The
second author has dived to observe fishes aroundYaku-shima Island almost everyday for six years,
but has never seen this species. Apogon parvulus
is distributed in the western Pacific, from Kalimantan to Japan (Allen et al., 2005); known only
from Iriomote-jima Island (Hayashi and Kishimoto, 1983) in Japanese waters.
Apogon cf properuptus (Whitley, 1964), dotted type
[Japanese name: “kinsen-ishimochi, dotto-gata”](Figs. 21A, B)
Lovamia properupta Whitley, 1964: 167, pl. 10(type locality: Frederick Reef, Coral Sea).
Material examined. KAUM–I. 21789,
37.8 mm SL, Isso; KAUM–I. 21790, 40.2 mm
SL, Isso; KAUM–I. 21791, 36.6 mm SL, Isso;
KAUM–I. 21792, 35.1 mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
3; circumpeduncular scales 12; total gill rakers
21, developed gill rakers 18; white body, with
6 yellowish brown stripes on lateral surface ofbody, posterior end of fourth stripe (middle stripe)
reaching to caudal-fin margin; and a whitish interspace, between fifth and sixth stripes, on cheekand flank, with a series of white spots.
Remarks. In Japanese waters, “Kinsen-ishimochi, A. properuptus” has long been regarded
to exhibit two color forms (e.g., Masuda and Kobayashi, 1996; Hayashi, 1997). Recently, Mabuchi et al. (2003) verified that “Kinsen-ishimochi,
A. properuptus” in Japanese waters contained
two distinct species, named as dotted and lined
types, on the basis of examination of nucleotide

Apogonid fishes in Yaku-shima Island

sequence data of approximately 650 bp from theanterior half of the mitochondrial 16S rRNA gene.
Their “dotted type” is characterized by a seriesof white spots on a whitish interspace, between
the fifth and sixth body stripes, and on the cheekand flank, whereas the “lined type” has a narrow stripe on a whitish interspace on the cheekand flank. Twelve specimens of “dotted type ofKinsen-ishimochi, A. properuptus” were reportedby Mabuchi et al. (2003) from Japan: 3 specimensfrom Ajiro, Kochi; 3 from Okinoshima Island,
Fukuoka; 3 from Tatsunokuchi, Nagasaki; and 3
from Bonotsu, Kagoshima). Senou (1999) reported the dotted type as A. properuptus from Sagami
Bay. Kuiter and Kozawa (1999) published underwater photographs of this species as Apogon sp. 9
from Shizuoka, Mie, Kochi, and Kagoshima Prefectures. Hagiwara and Kimura (2005) listed thisspecies as “A. properuptus (dotted type)” from
Tateyama Bay. Although Ichikawa et al. (1992)
and Arai and Ida (1975) listed “Kinsen-ishimochi”
as A. cyanosoma from Yaku-shima Island, andKuniyasu (1999) listed it as A. properuptus fromthe island, the taxonomic status of each of their“Kinsen-ishimochi” is unknown.
Mabuchi et al. (2003) suggested that the name
A. properuptus cannot be applied to either dottedor lined types because A. properuptus (type locality: Frederick Reef, Coral Sea) is most likely to be
restricted to the southwestern Pacific, as Randall
et al. (1997a) stated. We tentatively identified here
the “dotted type of Kinsen-ishimochi” in Japanesewaters as “A. cf properuptus, dotted type”.
Comparisons of underwater photographs of“A. cf properuptus, dotted type” from Yaku-shimaIsland and published underwater photographs of A.
holotaenia Regan, 1905 (e.g., Kuiter and Kozawa,
1999) showed that their overall body appearance
and color pattern, including lines and spots, wereindistinguishable. However, comparison of thepreserved specimens from Yaku-shima Island witha specimen of A. holotaenia from its type locality(BPBM34498, 37.6 mm SL, Muscat, Oman, coll.
by J. E. Randall, 13 May 1990) shows that thepigmentation of the fourth stripe on the caudalfin membrane is different (barely remained faintstripe in the former vs. distinct black stripe in thelatter), suggesting that the two may be distinct

species.
At Yaku-shima Island, only four specimens of“A. cf properuptus, dotted type” were collected off
Isso, but it has been observed in depths of morethan 25 m throughout the year, usually sheltering
among rocky reefs. At Yaku-shima Island, spawning of this species begins in early summer.

A

B

Fig. 21. Apogon cf properuptus, dotted type. A, KAUM–I.
21790, 40.2 mm SL, Isso; B, off Isso, 30 m, 13 Dec.
2004, S. Harazaki.

Apogon cf properuptus (Whitley, 1964), lined type
[Japanese name: “kinsen-ishimochi, rain-gata”](Figs. 22A, B)
Lovamia properupta Whitley, 1964: 167, pl. 10(type locality: Frederick Reef, Coral Sea).
Material examined. KAUM–I. 20112, 32.3mm SL, Yudomari; KAUM–I. 20207, 31.7 mm
SL, Yudomari; KPM–NI 22506, 41.1 mm SL, Yudomari.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24 ; predorsal scales
3; circumpeduncular scales 12; total gill rakers
22–23, developed gill rakers 19; white body with
6 yellow stripes on lateral surface of body, posterior end of forth stripe (middle stripe) reachingon caudal fin margin; a narrow stripe on whitish
interspace on cheek and flank.
43

T. Yoshida et al.

Remarks. Taxonomic remarks are given under species account of “A. cf properuptus, dottedtype”. Nine specimens of “lined type of Kinsenishimochi, A. properuptus,” were reported by
Mabuchi et al. (2003) from Japan: 3 specimensfrom Ajiro, Kochi; 3 from Kuchinoerabu-jimaIsland; and 3 from Iriomote-jima Island. Aizawa
and Senou’s (1991: pl. 8, C) A. cyanosoma from
Oshima Island, Tokushima Prefecture is herein
identified as “A. cf properuptus, lined type” inhaving a narrow stripe on the cheek and flank.
Kuiter and Kozawa (1999) published underwaterphotographs of this species as Apogon sp. 12 from
Iriomote-jima Island. Although Ichikawa et al.
(1992) and Arai and Ida (1975) listed “Kinsenishimochi” as A. cyanosoma from Yaku-shimaIsland, and Kuniyasu (1999) listed it as A. properuptus from the island, taxonomic status of their“Kinsen-ishimochi” are unknown.
“Apogon cf properuptus, lined type” is similar
to A. properuptus in overall appearance. However,
although A. properuptus from the southwestern
Pacific (see e.g., Randall et al., 1997a: 145) has
a white line below the eye not extending beyond
the opercular margin, the Japanese population of“A. cf properuptus, lined type” has the white line
extending onto near the anal-fin base.
At Yaku-shima Island, three specimens of“A. cf properuptus, lined type” were collected
off Yudomari, but it has been observed at depthsof less than 25 m (vs. more than 25 m in “A. cf
properuptus, dotted type”) throughout the year,
usually sheltering among rocky reefs. At Yakushima Island, spawning of this species begins inearly summer.
Apogon selas Randall and Hayashi, 1990
[Japanese name: Nagareboshi]
Apogon selas Randall and Hayashi, 1990: 399,
figs. 1–2 (type locality: Tripod Reef, off Nagada Harbor, Madang, Papua New Guinea);
Kuniyasu, 1999: 12 (Kurio, Yaku-shima, Japan).
Remarks. Apogon selas was originally described by Randall and Hayashi (1990) on thebasis of 25 specimens, including 18 paratypes
44

A

B

Fig. 22. Apogon cf properuptus, lined type. A, KAUM–I.
20112, 32.3 mm SL, Yudomari; B, off Isso, 20 m, 13
Dec. 2004, S. Harazaki.

(19.4–25.7 mm SL) from Japan (Iriomote-jimaIsland in the Ryukyu Islands). Apogon selashas been known from Japan, Indonesia, Papua
New Guinea, the Solomon Islands, and New
Caledonia (Allen et al., 2005; Randall, 2005). In
Japanese waters, this species has been reported
only from the Ryukyu Islands [Amami-oshimaIsland (Hayashi, 2002), Miyako Group (Senou
et al., 2007) and Iriomote-jima Island (Hayashi
and Kishimoto, 1983, as Apogon sp.; Randall andHayashi, 1990; Hayashi, 2002)].
Identification of Kuniyasu’s (1999) A. selasis uncertain as the second author has dived toobserve fishes around Yaku-shima Island almost
every day for six years, but has never seen this
distinct species.
Apogon seminigracaudus Greenfield, 2007
[Japanese name: Oguro-tenjikudai](Fig. 23)
Apogon seminigracaudus Greenfield, 2007a: 362,
figs. 1–4 (type locality: north shore of Vanua
Levu, Fiji).
Material examined. KAUM–I. 21785, 26.8mm SL, Isso.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 14; pelvic-fin rays I,

Apogonid fishes in Yaku-shima Island

5; pored lateral-line scales 24 ; predorsal scales 6;
circumpeduncular scales 12; total gill rakers 17,
developed gill rakers 10; body reddish; and lower
caudal-fin lobe blackish distally.
Remarks. The Yaku-shima Island specimen
and other Japanese specimens, previously identified as Apogon fuscus Quoy and Gaimard, 1825 (orNectamia fusca), agree with A. seminigracaudus(see Greenfield, 2007a), with the exception of thenumber of pectoral-fin rays (14 in Yaku-shimaIsland specimen vs. 13 rays in type specimens;
Greenfield, 2007a). Apogon fuscus is currently
considered to be a valid species of the A. bandanensis group (T. Fraser, pers. comm. in Greenfield,
2007a).
Apogon seminigracaudus is widely distributedin the western Pacific where it ranges from theFiji and Tonga Islands north to Japan (Greenfield, 2007a). In Japanese waters, this species
has been reported as A. fuscus from Tateyama
Bay (Hagiwara and Kimura, 2005), the Sagami
Sea (Senou et al., 2006b), Shizuoka Prefecture(Hayashi, 2002), Shikoku [Ehime Prefecture(Hayashi, 2002) and Kashiwa-jima Island (Hirata
et al., 1996)], and the Ryukyu Islands [Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou and Nakamura, 2003; Yoshigou et al.,2005), and Iriomote-jima Island (Yoshigou et al.,2001)].
At Yaku-shima Island, a single specimen of
A. seminigracaudus was collected from Isso, representing the first reliable records of the species
from the island. This species lives solitarily (notforming schools) in the shade of rocky reefs at
a depth of 20 m off Isso, Yaku-shima Island; noother localities confirmed from the island.

Apogon semiornatus Peters, 1876
[Japanese name: Yami-tenjikudai](Fig. 24)
Apogon semiornatus Peters, 1876: 436 (type locality: Mauritius).
Remarks. The photographed individual (Fig.
24) is herein identified as Apogon semiornatus on
the basis of the following characters: transparent
pinkish body, with a broad reddish brown bandfrom the snout to the anal-fin base, and a broad
dark brown band at the middle of the body surface.
Apogon semiornatus is widely distributedin the Indo-West Pacific where it ranges fromthe Red Sea and Gulf of Oman east to Australia
and Japan (Allen et al., 2005; Randall, 2005). In
Japanese waters, this species has been reported
from the Izu Islands [Miyake-jima Island (Ida andMoyer, 1974; Hayashi, 2002) and Hachijo-jimaIsland (Senou et al., 2002)], Chiba Prefecture andsouthward (Hayashi, 2002), including Tateyama
Bay (Hagiwara and Kimura, 2005), the Sagami
Sea (Senou et al., 2006b), Kashiwa-jima Island(Hirata et al., 1996), and Ie-jima Island (Senou etal., 2006a).
Figure 24 represents the first confirmed recordof A. semiornatus from Yaku-shima Island, but nospecimens have been collected from the island.
This species has usually been found hiding in a
crack or hole on a large rock at depths of 10–20 m
at Yaku-shima Island.

Fig. 24. Apogon semiornatus. Off Isso, 15 m, 21 July 2006,
S. Harazaki.

Fig. 23. Apogon seminigracaudus. KAUM–I. 21785, 26.8mm SL, Isso.

45

T. Yoshida et al.

Apogon taeniophorus Regan, 1908
[Japanese name: Misuji-tenjikudai](Figs. 25A, B)
Apogon taeniophorus Regan, 1908: 226 (type locality: Maldives, Indian Ocean).
Apogon endekataenia (not of Bleeker): Arai and
Ida, 1975: 192 (Kusugawa, Yaku-shima Island, Japan).
Material examined. BSKU 96641, 37.8 mm
SL, Isso; FRLM 34677, 47.2 mm SL, Kurio;
KAUM–I. 11171, 59.3 mm SL, Kurio; KAUM–
I. 11358, 76.9 mm SL, Kurio; KAUM–I. 11359,
76.1 mm SL, Kurio; KAUM–I. 20208, 28.2 mm
SL, Yudomari; KAUM–I. 20240, 76.8 mm SL,
Kurio; KAUM–I. 20341, 90.4 mm SL, Isso;
KAUM–I. 20344, 88.8 mm SL, Isso; KAUM–I.
21718, 15.5 mm SL, Kurio; KAUM–I. 21719,
16.6 mm SL, Kurio; KAUM–I. 21720, 14.8 mm
SL, Kurio; KAUM–I. 25227, 87.4 mm SL, Nagata; KPM–NI 22577, 21.8 mm SL, Isso; KPM–
NI 22578, 25.8 mm SL, Isso; MUFS 25595, 57.7mm SL, Kurio; MUFS 25596, 56.6 mm SL, Kurio; MUFS 25597, 63.1 mm SL, Kurio; NSMT–P
17846, 4 specimens: 28.8–68.0 mm SL, Kusugawa; NSMT–P 91351, 45.9 mm SL, Yudomari;
NSMT–P 91682, 3 specimens: 58.4–69.3 mm SL,
Kurio; NSMT–P 95459, 2 specimens: 32.6–84.6mm SL, Haruo; NSMT–P 95463, 87.8 mm SL,
Hirauchi.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 24; predorsal scales
3–4; circumpeduncular scales 12; total gill rakers
18–20, developed gill rakers 11–13; body white,with 6 distinct dark brown stripes on lateral surface of body; posterior portion of third stripe notreaching to second stripe at below first dorsal fin;posterior end of fourth stripe (middle stripe) forming a poorly defined black spot, less than pupil
diameter, at caudal-fin base; and fourth stripe notextending onto caudal-fin membrane.
Remarks. Matsuura and Tachikawa (1994)
reported A. taeniophorus from Chichi-jima Island, the Ogasawara Islands, as the first record of
the species from Japan, on the basis of a single
specimen (NSMT–P 35302, 60.8 mm SL), and46

proposed a new Japanese name, Misuji-tenjikudai,
for the species. Subsequently, Mabuchi et al.
(2004) recognized two distinct species in specimens previously identified as A. taeniophorus
from Japan on the basis of analysis of nucleotide
sequence data of approximately 610 bp from theanterior half of the mitochondrial 16S rRNA gene.
The two species were named by Mabuchi et al.
(2004) as “Lined type of Misuji-tenjikudai, A.
taeniophorus” and “Spotted type of Misuji-tenjikudai, A. taeniophorus”. The two can be easily
distinguished by color patterns on the body andcaudal fin (Mabuchi et al., 2004; this study, see
Description of each species account).
Judging from numerous underwater photographs and published data for A. taeniophorus
from the Indo–Pacific, Mabuchi et al.’s (2004)“Spotted type of Misuji-tenjikudai” is herein identified as A. taeniophorus. However, the taxonomic
status of “Lined type of Misuji-tenjikudai” is still
unknown. Two syntypes of Amia fasciata stevensi
McCulloch, 1915 (AMS I. 4247, 48.3 mm SL,
Fiji; AMS I. 13461, 59.9 mm SL, Vanuatu) were
examined during this study and we confirmed that
A. fasciata stevensi is a junior synonym of A. taeniophorus (= “Spotted type of Misuji-tenjikudai”)
in having 14 pectoral-fin rays; white body with
six dark brown stripes on lateral surface of body;posterior portion of third stripe not reaching tosecond stripe below first dorsal fin; posterior endof fourth stripe forming a black spot, less than pupil diameter, at caudal-fin base; and fourth stripe
not extending onto caudal-fin membrane.
Because Matsuura and Tachikawa’s (1994)
specimen (NSMT–P 35302) from Chichi-jima Island is in very poor condition (lacking most bodyscales and stripes), it cannot be reliably identified
as either A. taeniophorus or “Lined type of Misuji-tenjikudai”. However, in their checklist of fishesof the Ogasawara Islands, Randall et al. (1997b)
regarded Kuwamura et al.’s (1983) A. robusta(Smith and Radcliffe, 1911) (currently regarded asa junior synonym of A. cookii) and Sato’s (1991)
A. cookii (both listed from the Ogasawara Islands)
as A. taeniophorus. Sato’s (1991: fig. 11) A. cookii
is also herein identified as A. taeniophorus. In
addition, Randall et al. (1997b) listed four newly
collected specimens of A. taeniophorus (BPBMApogonid fishes in Yaku-shima Island

35092; BPBM 35125, 2 specimens; BPBM
35297) suggesting that A. taeniophorus is common in the Ogasawara Islands. Accordingly, weregard here that Matsuura and Tachikawa’s (1994)
specimen is A. taeniophorus (not Mabuchi et al.’s“Lined type of Misuji-tenjikudai”) and the Japanese name proposed by Matsuura and Tachikawa(1994) is applied to A. taeniophorus.
At Yaku-shima Island, A. taeniophorus ismost similar to A. cookii, but differs in having 14
pectoral-fin rays (vs. 15 rays in A. cookii) and an
indistinct, smaller caudal-fin base spot (vs. a distinct, larger spot) (Gon, 1987; this study).
Apogon taeniophorus is widely distributed in
the Indo-West Pacific where it ranges from theeast coast of Africa east to the islands of Micronesia, the Line Islands, French Polynesia, and northto Japan (Allen et al., 2005; Randall, 2005). In
Japanese waters, this species has been reported
from the Ogasawara Islands (Matsuura and Tachikawa, 1994; Randall et al., 1997b; Mabuchi etal., 2004), Tateyama Bay (Hagiwara and Kimura,2005), the Sagami Sea (Senou et al., 2006b), andthe Ryukyu Islands [Kuchinoerabu-jima Island(Mabuchi et al., 2004), Okinoerabu-jima Island(Yoshigou et al., 2005), and Tokashiki-jima Island(Watai et al., 2009)].
At Yaku-shima Island, specimens of A. taeniophorus were collected from Hirauchi, Isso, Kurio,
Kusugawa, Nagata, and Yudomari, representing
the first records of the species from the island.
This solitary species lives in the shade of rocky
reefs at depths of less than 15 m at Yaku-shimaIsland; it can be observed in coastal areas of theisland throughout the year. This species spawns in
the early summer at the island.
Apogon cf taeniophorus Regan, 1908, lined type
[Japanese name: “misuji-tenjikudai, rain-gata”](Figs. 26A, B)
Apogon taeniophorus Regan, 1908: 226 (type locality: Maldives, Indian Ocean).
Material examined. KPM–NI 22579, 25.1mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin rays

A

B

Fig. 25. Apogon taeniophorus. A, KAUM–I. 11359, 76.1mm SL, Kurio; B, off Kurio, 2 m, 17 Dec. 2009, S.
Harazaki.

I, 5; pored lateral-line scales 24; predorsal scales
4; circumpeduncular scales 12; total gill rakers
18, developed gill rakers 12; body white, with
6 dark brown stripes on lateral surface of body;posterior portion of third stripe curving upwardly
and reaching to second stripe at below first dorsal
fin; posterior end of fourth stripe (middle stripe)
reaching on caudal-fin margin.
Remarks. The Yaku-shima Island specimen (Fig. 26A) and a photographed individual(Fig. 26B) are herein identified as Mabuchi etal.’s (2004) “Lined type of Misuji-tenjikudai,
A. taeniophorus” on the basis of the following
combination of characters: a white body, with six
dark brown stripes on the lateral surface of thebody, the posterior portion of the third stripe curving upwardly and reaching to the second stripe
below the first dorsal fin, and the posterior end of
the fourth stripe (middle stripe) reaching to thecaudal-fin margin. Mabuchi et al. (2004) reported
four specimens of the species (2 specimens from
Misho, Ehime Prefecture, and 2 specimens from
Kuchinoerabu-jima Island, Kagoshima Prefecture)from Japan as A. taeniophorus (lined type).
The Yaku-shima Island specimen was collected at a depth of 4 m off Isso. Another individualof the species was observed by the second author
at a depth of 15 m off Isso.
47

T. Yoshida et al.

A

B

Fig. 26. Apogon cf taeniophorus, lined type. A, KPM–NI
22579, 25.1 mm SL, Isso; B, off Isso, 20 m, 14 Sept.
2009, S. Harazaki.

the east coast of Africa and the Red Sea east toAustralia and Japan (Allen et al., 2005; Randall
et al., 1997a). In Japanese waters, this species(mostly as Apogon timorensis) has been reported
from the Sagami Sea (Senou et al., 2006b),
Shizuoka Prefecture (Hayashi, 2002), Shikoku[Ehime Prefecture (Hayashi, 2002) and Kashiwajima Island (Hirata et al., 1996)], and the Ryukyu
Islands [Okinawa-jima Island (Yoshigou et al.,2005), Tokashiki-jima Island (Watai et al., 2009),
and Iriomote-jima Island (Hayashi and Kishimoto,
1983, as A. fraxineus)].
At Yaku-shima Island, three specimens of A.
timorensis were collected in less than 3 m depths
off Isso and Yudomari, representing the first reliable records of A. timorensis from the island.

Apogonichthyoides timorensis (Bleeker, 1854)[Japanese name: Kakure-ishimochi](Fig. 27)
Apogon timorensis Bleeker, 1854b: 207 (type locality: Kupang, Timor, Indonesia).
Material examined. KAUM–I. 20076, 59.9mm SL, Yudomari; KPM–NI 22542, 29.0 mm SL,
Yudomari; KPM–NI 22580, 21.9 mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; anal-fin
rays II, 8; pectoral-fin rays 16; pelvic-fin rays I, 5;
pored lateral-line scales 23–24; predorsal scales
3; circumpeduncular scales 12; total gill rakers
15, developed gill rakers 8–9; posterior tip of depressed pelvic fin extending beyond origin of anal
fin; body yellowish brown, with four broad, vertical, dark brown bands on lateral surface of body,first band from anterior portion of first dorsal fin
to pectoral fin, second band between bases ofsecond dorsal and anal fins, third band on middleof caudal peduncle, and fourth band at caudal-finbase; and head with a broad brownish band fromposterior margin of orbit to opercular margin, anda narrow brown stripe across cheek.
Remarks. Fraser and Allen (2010) allocatedthis species to the genus Apogonichthyoides.
Apogonichthyoides timorensis is widely distributed in the Indo-West Pacific where it ranges from48

Fig. 27. Apogonichthyoides timorensis. KAUM–I. 20076,
59.9 mm SL, Yudomari.

Archamia fucata (Cantor, 1849)[Japanese name: Sumitsuki-atohiki-tenjikudai](Figs. 28A, B)
Apogon fucata Cantor, 1849: 986 [type locality:
Sea of Pinang (Penang), Malaysia].
Material examined. BSKU 96549, 50.6 mm
SL, Yudomari; KAUM–I. 20044, 49.5 mm SL,
Yudomari; KAUM–I. 20074, 53.3 mm SL, Yudomari; KAUM–I. 20075, 73.5 mm SL, Yudomari.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 16; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 25; predorsal scales
6; circumpeduncular scales 12; total gill rakers
20–23, developed gill rakers 20; body pinkish,with numerous narrow, vertical or oblique reddish

Apogonid fishes in Yaku-shima Islandorange bars; an indistinct reddish black spot behind opercle; and no black band on cheek;.
Remarks. Archamia dispilus Lachner, 1951
was recently synonymized with A. fucata by Gon
and Randall (2003). Although the former (with anelongate spot behind head) is likely to be a valid
species, we tentatively follow Gon and Randall’s(2003) taxonomic decision.
Archamia fucata is widely distributed in the
Indo-West Pacific where it ranges from the Red
Sea and the east coast of Africa east to the Marshall Islands, the Samoa Islands, and Tonga, andnorth to Japan (Allen et al., 2005; Randall, 2005).
In Japanese waters, this species (as A. dispilus)
has been reported from Hachijo-jima Island (Senou et al., 2002), the Ogasawara Islands (Sato,
1991; Randall et al., 1997b), Miyazaki Prefecture(Motomura et al., 2001), Kagoshima Prefecture(Yamada, 2007), and the Ryukyu Islands [Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou et al., 2001; Yoshigou and Nakamura,
2002; 2003; Yoshigou et al., 2005), Miyako Group(Senou et al., 2007), and Iriomote-jima Island(Hayashi and Kishimoto, 1983)].
At Yaku-shima Island, specimens of A. fucata
were collected only from Yudomari, representing
the first reliable records of the species from theisland. This species forms schools in less than 5m on muddy substrates in river mouths and fishing ports of Yaku-shima Island; it can be observed
there in aggregations throughout the year.
Cheilodipterus artus Smith, 1961
[Japanese name: Kasumi-yarai-ishimochi](Fig. 29)
Cheilodipterus artus Smith, 1961: 409, pl. 50, fig.
F (type locality: Mahé, Seychelles)
Remarks. Hayashi (2002) reviewed the genus
Cheilodipterus in Japanese waters and regarded
four species as valid (Japanese names in parentheses), i.e., C. artus Smith, 1961 (Sudare-yarai-ishimochi), C. macrodon Lacepède, 1802 (Ryukyuyarai-ishimochi), C. quinquelineatus Cuvier, 1828
(Yarai-ishimochi), and C. subulatus Weber, 1909
(Kasumi-yarai-ishimochi). Incidentally, althoughGon (1993) regarded C. subulatus as a junior

A

B

Fig. 28. Archamia fucata. A, KAUM–I. 20075, 73.5 mm
SL, Yudomari; B, off Isso, 4 m, 26 Aug. 2006, S.
Harazaki.

synonym of C. singapurensis Bleeker, 1859–60,
Hayashi (2002) overlooked Gon’s (1993) taxonomic decision.
Hayashi and Kishimoto’s (1983: pl. 3, figs.
10a–b; IORD 77-748, 77-20) Cheilodipterussp., type A, and Hayashi’s (2002) ‘Sudare-yaraiishimochi, C. artus’ are herein identified as C. intermedius Gon, 1993 (see Gon, 1993: pl. 1, fig. B).
In addition, Hayashi and Kishimoto’s (1983: pl. 3,
fig. 9; IORD 76-435) C. subulatus, and Hayashi’s(2002) ‘Kasumi-yarai-ishimochi, C. subulatus’ areherein identified as C. artus (see Gon, 1993: pl. 4,
fig. C). Thus, the Japanese names ‘Kasumi-yaraiishimochi’ and ‘Sudare-yarai-ishimochi’ correspond to C. artus and C. intermedius respectively(Senou et al., 2006a, 2007; this study).
Although C. artus is very similar to C. intermedius in juvenile coloration, C. artus differsfrom the latter in ontogenetic color changes of thecaudal peduncle. Black pigments around a black
spot at the caudal-fin base of C. artus increase
with growth and the posterior portion of the caudal peduncle finally becomes black, whereas noblack pigments usually occur around the caudalfin base spot of C. intermedius and the posterior
portion of the caudal peduncle usually becomeswhitish (Gon, 1993; H. Senou, pers. comm.). The
49

T. Yoshida et al.

figured individual (Fig. 29) may be an adult of
C. intermedius (rather than C. artus); at this life
stage, specimens are required for accurate identification.
Cheilodipterus artus is widely distributed in
the Indo-West Pacific where it ranges from theeast coast of Africa east to the Marshall Islands
and Tuamotu Archipelago, and north to Japan(Allen et al., 2005; Randall, 2005). In Japanesewaters, reliable records of the species are from
Yaku-shima Island (this study) and the Ryukyu
Islands [Ie-jima Island (Senou et al., 2006a), Miyako Group (Senou et al., 2007), and Iriomotejima Island (Hayashi and Kishimoto, 1983, as C.
subulatus)].
This species inhabits rocky and coral reefs
at depths of less than 20 m at Yaku-shima Island; common in the island and can be observed
throughout the year.

spot at caudal fin base in adults (Fig. 30B) (seeRemarks of C. artus).
Cheilodipterus intermedius is widely distributed in the western Pacific where it ranges fromthe Solomon Islands to the South China Sea andJapan (Allen et al., 2005). In Japanese waters, thisspecies has been reported from the Sagami Sea(Senou et al., 2006b, as C. artus) and the Ryukyu
Islands [Sesoko-jima Island (Gon, 1993), Ie-jimaIsland (Senou et al., 2006a), and Miyako Group(Senou et al., 2007)].
This species is solitary on rocky and coral
reefs at depths of less than 20 m at Yaku-shimaIsland; common in the island throughout the year.

A

B

Fig. 29. Cheilodipterus cf artus. Off Isso, 8 m, 11 June
2005, S. Harazaki.

Cheilodipterus intermedius Gon, 1993
[Japanese name: Sudare-yarai-ishimochi](Figs. 30A, B)
Cheilodipterus intermedius Gon, 1993: 19, pl. 1,
figs. 3B, 8 (type locality: east side of Marinelab., Sesoko-jima Island, Okinawa, Ryukyu
Islands, Japan).
Remarks. Photographed individuals (Figs.
30A, B) are identified as C. intermedius on thebasis of the following combination of characters:nine blackish brown broad and narrow stripes onlateral surface of body; a black spot, surroundedby yellow, at midbase of caudal fin in young (Fig.
30A); and no black vertical bar or distinct black
50

Fig. 30. Cheilodipterus intermedius. A, off Isso, 10 m,
5 Nov. 2005, S. Harazaki; B, off Isso, 10 m, 5 Nov.
2005, S. Harazaki.

Cheilodipterus macrodon (Lacepède, 1802)[Japanese name: Ryukyu-yarai-ishimochi](Figs. 31A, B)
Centropomus macrodon Lacepède, 1802: 252, 273(type locality: Mauritius or Réunion Island).
Cheilodipterus lineatus; Arai and Ida, 1975: 184,
pl. 17, fig. 1 (Kusugawa, Yaku-shima Island,
Japan).
Cheilodipterus macrodon; Ichikawa et al., 1992: 9(Yaku-shima Island, Japan); Kuniyasu, 1999:
12 (Kurio, Yaku-shima Island, Japan).

Apogonid fishes in Yaku-shima Island

Material examined. BSKU 96548, 58.7 mm
SL, Yudomari; KAUM–I. 11453, 70.5 mm SL,
Isso.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 12; pelvic-fin rays I,
5; pored lateral-line scales 25; predorsal scales 6;
circumpeduncular scales 12; total gill rakers 20,
developed gill rakers 9; large canine-like teeth onjaws; preopercular margin serrated; body white,with nine poorly defined dark brown stripes onlateral surface of body; a broad, black band on
caudal-fin base in young.
Remarks. Cheilodipterus macrodon is widelydistributed in the Indo-West Pacific where it ranges from the Red Sea and east coast of Africa east
to French Polynesia (expect for the Marquesas Islands), and the Pitcairn Islands, and north to Japan(Allen et al., 2005; Randall, 2005). In Japanesewaters, this species has been reported from the Izu
Islands [Hachijo-jima Island (Senou et al., 2002)],
the Ogasawara Islands (Kuwamura et al., 1983;
Sato, 1991; Randall et al., 1997b), Tateyama Bay(Hagiwara and Kimura, 2005), the Sagami Sea(Senou et al., 2006b), Shikoku [Kochi (Kamohara,
1964, as Jadamga quinquelineatus) and Kashiwajima Island (Hirata et al., 1996)], Yaku-shimaIsland (Ichikawa et al., 1992; Kuniyasu, 1999),
and the Ryukyu Islands [Amami-oshima Island(Kamohara and Yamakawa, 1968), Ie-jima Island(Senou et al., 2006a), and Tokashiki-jima Island(Watai et al., 2009)].
At Yaku-shima Island, two specimens of C.
macrodon were collected from Isso and Yudomari.
This species occurs in less than 20 m at rocky orcoral reefs at Yaku-shima Island throughout theyear.
Cheilodipterus quinquelineatus Cuvier, 1828
[Japanese name: Yarai-ishimochi](Figs. 32A, B)
Cheilodipterus quinquelineatus Cuvier in Cuvier
and Valenciennes, 1828: 167 (type locality:
Bora Bora, Society Islands); Ichikawa et al.,
1992: 9 (Yaku-shima Island, Japan); Kuniyasu, 1999: 12 (Kurio, Yaku-shima Island,
Japan).

A

B

Fig. 31. Cheilodipterus macrodon. A, BSKU 96548, 58.7mm SL, Yudomari; B, off Isso, 8 m, 22 June 2004, S.
Harazaki.

Material examined. KAUM–I. 11289, 50.0mm SL, Yudomari; KAUM–I. 20049, 80.3 mm
SL, Yudomari; KAUM–I. 20068, 101.9 mm SL,
Yudomari; KAUM–I. 20070, 85.8 mm SL, Yudomari; KAUM–I. 20073, 42.0 mm SL, Yudomari;
KAUM–I. 20535, 65.5 mm SL, Yudomari; KPM–
NI 22511, 40.0 mm SL, Yudomari.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 12; pelvic-fin rays I, 5;
pored lateral-line scales 24–25 ; predorsal scales
6; circumpeduncular scales 12; total gill rakers
18, developed gill rakers 12; large canine-liketeeth on jaws; preopercular margin serrated; bodywhite, with five black stripes on lateral surface ofbody, posterior end of third stripe (middle stripe)
reaching a black spot on caudal-fin base; and acaudal-fin base spot, less than pupil diameter, surrounded by poorly defined yellow blotch.
Remarks. Cheilodipterus quinquelineatus is
widely distributed in the Indo–Pacific, expect for
the Persian Gulf and Hawaiian Islands (Allen etal., 2005; Randall, 2005). In Japanese waters, thisspecies has been reported from the Izu Islands[Miyake-jima Island (Ida and Moyer, 1974) andHachijo-jima Island (Senou et al., 2002)], theOgasawara Islands (Kuwamura et al., 1983; Sato,
1991; Matsuura and Tachikawa, 1994; Randall et51

T. Yoshida et al.

al., 1997b), the Sagami Sea (Senou et al., 2006b),
Shikoku [Kochi (Kamohara, 1964) and Kashiwajima Island (Hirata et al., 1996)], Yaku-shimaIsland (Ichikawa et al., 1992; Kuniyasu, 1999),
and the Ryukyu Islands [Yagaji-shima Island(Yoshigou et al., 2005), Ie-jima Island (Senou etal., 2006a), Okinawa-jima Island (Yoshigou et al.,
2001; Yoshigou and Nakamura, 2002; Yoshigou
et al., 2005), Tokashiki-jima Island (Watai et al.,2009), Miyako Group (Senou et al., 2007), andIriomote-jima Island (Hayashi and Kishimoto,
1983; Yoshigou et al., 2001)].
At Yaku-shima Island, specimens of C. quinquelineatus were collected from Kurio and Yudomari. This species occurs in less than 20 m on
rocky or coral reefs at Yaku-shima Island throughout the year. The population of C. quinquelineatus
at Yaku-shima Island is the largest among the four
species of Cheilodipterus occurring around theisland.

A

B

Foa brachygramma (Jenkins, 1903)[Japanese name: Taiwan-mato-ishimochi](Fig. 33)
Fowleria brachygrammus Jenkins, 1903: 447, fig.
20 (type locality: Honolulu, Oahu Island, Hawaiian Islands).
Remarks. The photographed individual (Fig.
33) is herein identified as Foa brachygramma on
the basis of the following combination of characters: lateral line incomplete; no large black blotch
on opercle; body blackish brown, with numerous
small white blotches on head and caudal-fin base;and no vertical bands on lateral surface of body.
Foa brachygramma is widely distributed in
the Indo-West Pacific where it ranges from theeast coast of Africa east to the Hawaiian Islands,
and north to Japan (Hayashi, 2002; Randall,
1997a). In Japanese waters, this species has beenreported from Shimoda Bay, Shizuoka (Masuda
et al., 1984) to the Ryukyu Islands [Ie-jima Island (Senou et al., 2006a), Okinawa-jima Island(Yoshigou et al., 2005), Tokashiki-jima Island(Watai et al., 2009), Miyako Group (Senou et al.,2007), and Iriomote-jima Island (Hayashi andKishimoto, 1983; Yoshigou et al., 2001)].
Figure 33 represents the first confirmed
record of F. brachygramma from Yaku-shimaIsland. This species can be observed in aggregations, each with about 20 individuals, on sandy
substrates in less than 20 m off Isso, Yaku-shimaIsland, throughout the year; no other localities
confirmed from the island.

Fig. 32. Cheilodipterus quinquelineatus. A, KAUM–I.
11289, 50.0 mm SL, Yudomari; B, off Isso, 8 m, 26
Jan. 2005, S. Harazaki.

Fig. 33. Foa brachygramma. Off Isso, 20 m, 15 Sept. 2009,
S. Harazaki.

52

Apogonid fishes in Yaku-shima Island

Fowleria isostigma (Jordan and Seale, 1906)[Japanese name: Naha-mato-ishimochi](Fig. 34)
Apogonichthys isostigma Jordan and Seale, 1906:
251, fig. 45 (type locality: Apia, Upolu Island,Western Samoa).
Material examined. BSKU 96573, 17.0 mm
SL, Yudomari.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 11; predorsal scales
6; circumpeduncular scales 12; total gill rakers
4, developed gill rakers 4; palatine without teeth;
lateral line incomplete; body whitish brown, without small white blotches; a large ocellated black
blotch on opercle; no vertical bands on lateral surface of body; and no spots or lines on fins.
Remarks. Two color variations of F. isostigma are known in Japanese waters: one has a roundblack spot basally on each scale of the lateral surface of the body, the spots forming several longitudinal stripes; the other lacks the spots (Hayashi,
2002). The Yaku-shima Island specimen is identified as Hayashi’s (2002) “unmarked form”.
Fowleria isostigma is widely distributed in
the western Pacific where it ranges from Australia
east to Mangareva and the islands of Micronesia,
and north to Japan (Randall, 2005). In Japanesewaters, this species has been reported from Wakayama Prefecture and southward (Hayashi,
2002), including Kashiwa-jima Island (Hirata etal., 1996) and the Ryukyu Islands [Amami-oshimaIsland (Kamohara and Yamakawa, 1968), Iejima Island (Senou et al., 2006a), Okinawa-jimaIsland (Yoshigou and Nakamura, 2003; Yoshigou
et al., 2005), Ishigaki-jima Island (Yoshigou etal., 2001), and Iriomote-jima Island (Hayashi andKishimoto, 1983)].
At Yaku-shima Island, a single specimen of
F. isostigma was collected from Yudomari, representing the first reliable record of F. isostigma
from the island. Ecological information on thespecies at Yaku-shima Island is unknown.

Fig. 34. Fowleria isostigma. BSKU 96573, 17.0 mm SL,
Yudomari.

Fowleria cf marmorata (Alleyne and Macleay, 1877)[Japanese name: Obi-shibori](Fig. 35)
Apogonichthys marmorata Alleyne and Macleay,
1877: 268, pl. 5, fig. 2 (type locality: Cape
Grenville, Queensland, Australia).
Material examined. KAUM–I. 21784, 39.8mm SL, Isso.
Description. Dorsal-fin rays VII-I, 9; analfin rays II, 8; pectoral-fin rays 14; pelvic-fin raysI, 5; pored lateral-line scales 6 (7 on right sideof body); predorsal scales 6; circumpeduncularscales 12; total gill rakers 13, developed gill rakers 4; palatine without teeth; lateral line incomplete; body red, with several vertical dark red bars
on lateral surface of body; 3 black bands radiatingfrom posterior margin of eye; and a large ocellated black spot on opercle.
Remarks. The Yaku-shima Island specimen
is herein identified as F. marmorata in having an
incomplete lateral line; red body, with several vertical dark red bars on lateral surface of body anda large ocellated black spot on the opercle. However, the number of pored lateral-line scales in the
Yaku-shima Island specimen is six (seven on rightside of body), whereas those described by Hayashi(2002) and Randall (2005) are 12 and 10–13 respectively. In addition, the number of developed
gill rakers in the Yaku-shima Island specimen isfour, whereas those described by Hayashi (2002)
is seven. More Yaku-shima Island specimens are
required to determine whether these characters
represent intraspecific variations or distinct species.
53

T. Yoshida et al.

Fowleria marmorata is widely distributedin the Indo-West Pacific where it ranges fromthe Red Sea east to the Line, Society, and Marquesas Islands, and north to Japan (Allen et al.,2005; Randall, 2005). In Japanese waters, thisspecies has been reported from Kashiwa-jima Island (Hirata et al., 1996) and the Ryukyu Islands[Okinawa-jima Island (Yoshigou et al., 2005) andSesoko-jima Island (Hayashi, 2002)]. At Yakushima Island, a single specimen was collectedfrom Isso at a depth of 30 m.

to Japan (Allen et al., 2005; Randall, 2005). In
Japanese waters, this species has been reported
from the Ogasawara Islands (Sato, 1991; Matsuura and Tachikawa, 1994; Randall et al., 1997b)
and the Ryukyu Islands [Amami-oshima Island(Yamakawa, 1971; Yoshigou et al., 2001), Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou et al., 2001; Yoshigou and Nakamura,
2003; Yoshigou et al., 2005), Miyako Group(Senou et al., 2007), and Iriomote-jima Island(Hayashi and Kishimoto, 1983; Yoshigou et al.,
2001; Yoshigou and Nakamura, 2002)].
At Yaku-shima Island, two specimens of F.
variegata were collected from Yudomari, representing the first reliable records of F. variegata
from the island. Ecological information on thespecies at Yaku-shima Island is unknown.

Fig. 35. Fowleria cf marmorata. KAUM–I. 21784, 39.8mm SL, Isso.

Fowleria variegata (Valenciennes, 1832)[Japanese name: Shibori](Fig. 36)

Fig. 36. Fowleria variegata. KPM–NI 22505, 28.8 mm SL,
Yudomari.

Apogon variegata Valenciennes, 1832: 55 (typelocality: Mauritius).
Material examined. BSKU 96547, 33.8 mm
SL, Yudomari; KPM–NI 22505, 28.8 mm SL, Yudomari.
Description. Dorsal-fin rays VI-I, 9; anal-fin
rays II, 8; pectoral-fin rays 13; pelvic-fin rays I,
5; pored lateral-line scales 10 ; predorsal scales
4–5; circumpeduncular scales 12; total gill rakers
14–15, developed gill rakers 6–8; palatine without teeth; lateral line incomplete; body blackish
brown, with numerous black spots and blotcheson lateral surface of body; a large dark ocellated
spot on opercle; fins brown, with numerous whitespots.
Remarks. Fowleria variegata is widely distributed in the Indo-West Pacific where it ranges
from the Red Sea and the east coast of Africa,east to the Mariana and Samoa Islands, and north54

Neamia octospina Smith and Radcliffe, 1912
[Japanese name: Yatsutoge-tenjikudai](Figs. 37A, B)
Neamia octospina Smith and Radcliffe in Radcliffe, 1912: 441, pl. 36, fig. 2 (type locality:
Rasa Island, off Mantaguin Bay, Palawan,
Philippines).
Remarks. Photographed individuals (Figs.
37A, B) are herein identified as N. octospina on
the basis of the following combination of characters: a semitranslucent white body, with numerous
small reddish spots on the body and fins, and threeindistinct reddish bands radiating from the posterior margin of the eye. Specimens have not beencollected from Yaku-shima Island.
Neamia octospina is widely distributed in

Apogonid fishes in Yaku-shima Island

the Indo-West Pacific where it ranges from the
Red Sea east to Fiji, and north to Japan (Allenet al., 2005; Randall, 2005). In Japanese waters,
this species has been reported from the Ryukyu
Islands [Okinawa-jima Island (Yoshigou et al.,2005) and Ishigaki-jima Island (Ida and Moyer,1974)].
Figures 37A, B represent the first confirmed
records of N. octospina from Yaku-shima Islandand the northernmost records for the species in
the western Pacific. The photographed individuals
were found among branches of the Alcyonacea,
Cladiella digitulata, at a depth of 2 m off Motoura
Beach, Isso, Yaku-shima Island on 23 Oct. 2006.
These individual disappeared after C. digitulata’
death confirmed by the second author; no otherindividuals have been observed at the island.

A

B

Fig. 37. Neamia octospina. A and B, off Isso, 5 m, 23 Oct.
2006, S. Harazaki.

Nectamia bandanensis (Bleeker, 1854)[Japanese name: Banda-ishimochi](Figs. 38A, B)
Apogon bandanensis Bleeker, 1854a: 95 (typelocality: Banda Neira, Banda Islands, Indonesia).

Material examined. BSKU 96538, 66.0 mm
SL, Yudomari; KAUM–I. 20069, 67.8 mm SL,
Yudomari.
Description. Dorsal-fin rays VII-I, 9 (onespecimen with VII-II, 9); anal-fin rays II, 8;
pectoral-fin rays 13; pelvic-fin rays I, 5; pored
lateral-line scales 24 ; predorsal scales 3–4; circumpeduncular scales 12; total gill rakers 25–26,
developed gill rakers 23; body depth 43.2–43.7% of SL; body silvery brown, with 3 dark brownbands, widths subequal to pupil diameter, on
lateral surface of body; and an oblique, straight,
brownish bar from posteroventral margin of eye.
Remarks. Fraser (2008) recently allocatedthis species to Nectamia from Apogon. Nectamia
bandanensis has been known from the western
Pacific from Japan to American Samoa (Randall,2005). In Japanese waters, this species has beenreported from the Ogasawara Islands (Matsuura
and Tachikawa, 1994, as A. bandaensis; Randall etal., 1997b, as A. bandaensis) and the Ryukyu Islands [Amami-oshima Island (Kamohara and Yamakawa, 1968, as A. bandaensis), Ie-jima Island(Senou et al., 2006a, as A. bandaensis), Okinawajima Island (Hayashi, 2002, as A. bandaensis),
Sesoko-jima Island (Fraser, 2008), Tokashiki-jimaIsland (Watai et al., 2009), Miyako Group (Senou
et al., 2007, as A. bandaensis), and Iriomote-jimaIsland (Hayashi and Kishimoto, 1983; Hayashi,
2002, as A. bandaensis)].
At Yaku-shima Island, two specimens of N.
bandanensis were collected from Isso and Yudomari, representing the first reliable records ofthe species from the island and the northernmost
records for the species. This species is observed in
the shade of rocky reefs in the daytime and aroundthe reefs at night; it can be observed in less than
10 m at Yaku-shima Island throughout the year.
Pseudamia gelatinosa Smith, 1956
[Japanese name: Numeri-tenjikudai](Figs. 39A, B)
Pseudamia gelatinosa Smith, 1956: 690, pl. 18,
fig. A (type locality: Aldabra Islands).
Material examined. BSKU 96540, 42.9mm SL, Yudomari; KAUM–I. 20120, 31.2 mm
55

T. Yoshida et al.

A

B

Fig. 38. Nectamia bandanensis. A, KAUM–I. 20069, 67.8mm SL, Yudomari; B, off Isso, 10 m, 22 Oct. 2009, S.
Harazaki.

SL, Yudomari; KAUM–I. 20163, 16.4 mm SL,
Yudomari; KAUM–I. 20164, 21.0 mm SL, Yudomari; KAUM–I. 20165, 14.9 mm SL, Yudomari;
KAUM–I. 20166, 16.5 mm SL, Yudomari; KPM–
NI 22529, 18.7 mm SL, Yudomari.
Description. Dorsal-fin rays VI-I, 9; analfin rays II, 8; pectoral-fin rays 15; pelvic-fin raysI, 5; pored lateral-line scales 35; predorsal scales
5–6; circumpeduncular scales 12; total gill rakers
11, developed gill rakers 8; caudal fin lanceolate;anterior nostril with a long flap; semitranslucent
slender body, with numerous small blackish spots
forming several longitudinal stripes; caudal fin
with a large black blotch dorsally; and a black
blotch on caudal-fin base.
Remarks. Pseudamia gelatinosa differs fromits close relative, P. amblyuroptera, in having 8
developed gill rakers and 8 anal-fin rays (vs. 11
rakers and usually 9 rays; Yoshigou and Yoshino,2004).
Pseudamia gelatinosa is widely distributedin the Indo-West Pacific where it ranges from the
Red Sea and east coast of Africa, east to the Line
Islands, Society Islands and Rapa, and north toJapan (Allen et al., 2005; Randall, 2005). In Japanese waters, this species has been reported from56

the Izu Islands [Hachijo-jima Island (Senou et al.,2002)], the Sagami Sea (Hayashi, 2002; Senou
et al., 2006b), and the Ryukyu Islands [Ie-jimaIsland (Senou et al., 2006a), Okinawa-jima Island(Yoshigou et al., 2005), Miyako Group (Senou etal., 2007), Ishigaki-jima Island (Ida and Moyer,1974), and Iriomote-jima Island (Yoshigou et al.,2001)].
At Yaku-shima Island, specimens of P. gelatinosa were collected from Yudomari, representing the first reliable records of the species fromthe island. This species is solitary in the shade of
rocky reefs (not forming schools) at depths of lessthan of 10 m off Isso and Yudomari, Yaku-shimaIsland; no other localities confirmed from the island.

A

B

Fig. 39. Pseudamia gelatinosa. A, BSKU 96540, 42.9mm SL, Yudomari; B, off Isso, 10 m, 16 Oct. 2009, I.
Takaku.

Rhabdamia gracilis (Bleeker, 1856)[Japanese name: Sukashi-tenjikudai](Figs. 40A, B)
Apogonichthys gracilis Bleeker, 1856b: 371 (typelocality: Ternate, Moluccas, Indonesia).
Remarks. Photographed individuals (Figs.
40A, B) are identified as R. gracilis by having atranslucent silvery body, with a bluish longitudinal
stripe on the middle of the body. No specimens
have been collected from Yaku-shima Island.
Mabuchi (2001) regarded that R. gracilis contained two distinct species: one has a black spotApogonid fishes in Yaku-shima Islandon the posteroventral caudal peduncle, and theother lacks the spot. However, because these twohave been observed as members of a school (seeFig. 40A) and the second author confirmed thatthey have often paired with each other at Yakushima Island, we tentatively treat them as conspecific in this study.
Rhabdamia gracilis is widely distributed in
the Indo-West Pacific where it ranges from theeast coast of Africa east to the Marshall Islands,
Caroline Islands and Fiji, and north to Japan (Allenet al., 2005; Randall, 2005). In Japanese waters,
this species has been reported from the Sagami
Sea (Senou et al., 2006b), Wakayama Prefectureand southward (Hayashi, 2002), including the
Ryukyu Islands [Ie-jima Island (Senou et al.,2006a), Okinawa-jima Island (Ida and Moyer,
1974; Yoshigou et al., 2005), Tokashiki-jima Island (Watai et al., 2009), Miyako Group (Senou
et al., 2007), and Iriomote-jima Island (Ida andMoyer, 1974; Hayashi and Kishimoto, 1983;
Yoshigou and Nakamura, 2002)]. Figures 40
represent the first confirmed record of R. gracilis
from Yaku-shima Island.
Rhabdamia gracilis, forming a school of ca.
50 individuals, lives sympatrically with A. notatus
in depths of 20–25 m at two diving spots off Isso,
Yaku-shima Island; no other localities confirmed
from the island. At Yaku-shima Island, the spawning period of the species is in July and August.
Siphamia majimai Matsubara and Iwai, 1958
[Japanese name: Majimakuro-ishimochi](Figs. 41A–C)
Siphamia majimai Matsubara and Iwai, 1958:
603, fig.1 (type locality: Urasokari, Amamioshima Island, Japan, ca. 28°07’N, 129°20’E).
Material examined. BSKU 96557, 16.9 mm
SL, Yudomari; KAUM–I. 20158, 20.3 mm SL,
Yudomari; KAUM–I. 20211, 19.3 mm SL, Yudomari; KPM–NI 22582, 16.9 mm SL, Isso.
Description. Dorsal-fin rays VI-I, 9; analfin rays II, 8; pectoral-fin rays 15; pelvic-fin raysI, 5; pored lateral-line scales 23; predorsal scales
6; circumpeduncular scales 12; total gill rakers
13–14, developed gill rakers 7–8; luminous organ

A

B

Fig. 40. Rhabdamia gracilis. A, off Isso, 20 m, 15 Sept.
2009, S. Harazaki; B, off Isso, 20 m, 15 Sept. 2009, S.
Harazaki.

inside translucent muscles of ventral body fromthroat to caudal peduncle; and blackish silver
body, with numerous small red spots on body andfins when fresh.
Remarks. Siphamia majimai is distributed in
the western Pacific from Australia to Japan (Randall et al., 1997a). In Japanese waters, this species
has been reported from the Ogasawara Islands(Randall et al., 1997b), Kashiwa-jima Island (Hirata et al., 1996), and the Ryukyu Islands [Amami-oshima Island (Matsubara and Iwai, 1958;
Hayashi, 2002), Tokashiki-jima Island (Watai etal., 2009), Miyako Group (Senou et al., 2007),and Iriomote-jima Island (Hayashi and Kishimoto,
1983; Hayashi, 2002)].
At Yaku-shima Island, specimens of S. majimai were collected from Isso and Yudomari, representing the first reliable records of the species
from the island. Siphamia majimai usually forms
a school with five or six individuals and lives
among spines of Diadema setosum at depths ofless than 15 m at Yaku-shima Island. This species
spawns in July and juveniles are observed in the
middle of summer at the island.

57

T. Yoshida et al.

AB

has been reported from Kashiwa-jima Island (Hirata et al., 1996; Hayashi, 2002).
Figure 42 represents the first confirmed recordof S. tubulata from Yaku-shima Island. Siphamia
tubulata is usually solitary in the shade of rocky
reefs at depths of 25–30 m at Yaku-shima Island.
A school with five or six young individuals wasobserved around Padina arborescens at depths of
25–30 m only once over the last six years of underwater observations.

C

Fig. 42. Siphamia tubulata. Off Isso, 25 m, 28 Oct. 2005, S.
Harazaki.

Siphamia versicolor (Smith and Radcliffe, 1911)[Japanese name: Hikari-ishimochi]
Fig. 41. Siphamia majimai. A, KAUM–I. 20158, 20.3mm SL, Yudomari; B, off Isso, 5 m, 6 June 2005, S.
Harazaki; C, off Isso, 5 m, 3 June 2006, S. Harazaki.

Siphamia tubulata (Weber, 1909)[Japanese name: Inazuma-hikari-ishimochi](Fig. 42)
Apogon tubulatus Weber 1909: 160 (type locality:
Sapeh Strait, Sumbawa Island, Lesser Sunda
Islands, Indonesia).
Remarks. The photographed individual (Fig.
42) is herein identified as Siphamia tubulata on
the basis of the unique coloration, viz., body silvery, mottled with reddish brown blotches. Specimens have not been collected from Yaku-shimaIsland.
Siphamia tubulata is distributed in the western Pacific, from Indonesia and Australia to Japan(Hayashi, 2002). In Japanese waters, this species
58

Amia versicolor Smith and Radcliffe in Radcliffe,
1911: 257, fig. 3 (type locality: Cataingan
Bay, Masbate, Philippines).
Siphamia versicolor: Ichikawa et al., 1992: 9(Yaku-shima Island, Japan).
Remarks. Identification of Ichikawa et al.’s(1992) S. versicolor is uncertain. The second
author has dived for fish watching around Yakushima Island almost everyday for six years, but
has never found the species.
Siphamia versicolor is widely distributed in
the Indo-West Pacific where it ranges from theGulf of Oman and Maldive Islands east to the
Caroline Islands, and north to Japan (Allen et al.,2005; Randall, 2005). In Japanese waters, thisspecies has been reported from the Izu Islands[Hachijo-jima Island (Senou et al., 2002)], theOgasawara Islands (Kuwamura et al., 1983; Randall et al., 1997b; Yoshigou and Nakamura, 2002),

Apogonid fishes in Yaku-shima Islandand the Ryukyu Islands [Yagaji-shima Island(Yoshigou and Nakamura, 2002; Yoshigou et al.,2005), Ie-jima Island (Senou et al., 2006a), Okinawa-jima Island (Yoshigou et al., 2005), Tokashikijima Island (Watai et al., 2009), and Iriomote-jimaIsland (Hayashi and Kishimoto, 1983)].
Conclusion
Fifteen genera with 94 species of the family
Apogonidae have been recorded from Japanesewaters (see Introduction). During this study, 45
species of the family were confirmed to occur
around Yaku-shima Island, representing nearly
half of all known Japanese species. Three species,
Apogon parvulus, A. selas and Siphamia versicolor, previously reported by Ichikawa et al. (1992)
and Kuniyasu (1999) from the island on the basisof underwater observations, were not observed orcollected during this study (2004–2010). Thirteenof the 45 species, Apogon apogonides, A. chrysotaenia, A. fasciatus, A. fraenatus, A. kallopterus,
A. notatus, A. semiornatus, Cheilodipterus artus,
C. intermedius, Foa brachygramma, Neamia
octospina, Rhabdamia gracilis, and Siphamia
tubulata, were herein recorded only on the basisof underwater photographs, and records of theremaining species were based on collected specimens. Twenty-seven species represent the firstrecords from Yaku-shima Island.
Recently, Yoshida and Motomura (2009)
considered the northernmost recorded range of
Apogon amboinensis to be Yaku-shima Island. In
addition, our survey revealed that the northward
distributional range of the following six species
extended to Yaku-shima Island: Apogon caudicinctus, A. chrysotaenia, A. exostigma, A. moluccensis, Neamia octospina, and Nectamia bandanensis. Of these, A. chrysotaenia was recorded
from Japanese waters for the first time. Althoughno specimens of three species, A. chrysotaenia, A.
exostigma, and N. octospina, were collected during this study, underwater photographs facilitatedtheir accurate identification. Siphamia tubulata,
reported only from Kashiwa-jima Island (Hirata etal., 1996; Hayashi, 2002) in Japanese waters, wasfound at Yaku-shima Island. Incidentally, none of
the 45 species found at Yaku-shima Island represents the southernmost record of the species.

Most of the 45 species found at Yaku-shimaIsland have been observed throughout the year atvarious life stages by scuba diving, with the exception of the following four: Apogon fasciatus, A.
cf taeniophorus, Neamia octospina, and Siphamia
tubulata. Only a single, small specimen of A. cf
taeniophorus was collected and no adults of thespecies have been observed or collected at theisland. The remaining three species have neverbeen collected and have been observed by scuba
diving only once for each species during the last
six years. This suggests that the four species weremost likely transported from around Taiwan or
China by the Kuroshio Current, which normallyflows well west of the Ryukyu Islands. It is highly
unlikely that these species reproduce aroundYaku-shima Island.
Although the second author has confirmed
by underwater observations for the last six years
that the following three species occur at Yakushima Island, no specimens or photographs ofthese species were taken from the island: Apogon
fukuii Hayashi, 1990 (Futasuji-ishimochi), A. rubrimacula Randall and Kulbicki, 1998 (Akahoshikinsen-ishimochi), and Pseudamia hayashii
Randall, Lachner, and Fraser, 1985 (Sabikudaribouzugisumodoki). These species may be collected from the island in future.
Acknowledgments
We would like to thank M. Yamada (Kagoshima Aquarium, Japan), K. Hidaka, K. Miyamoto,
S. Yoshinaga, and K. Kudo (formerly MUFS),
H. Endo, E. Katayama, M. Nakayama, and M.
Yamamura (BSKU), K. Matsuura, G. Shinohara,
K. Kuriiwa, T. Sato, Y. Takata, and M. Nakae(NSMT), S. Kimura (FRLM), H. Senou (KPM), Y.
Iwatsuki and H. Izumi (MUFS), and G. Ogrihara,
M. Matsunuma, M. Meguro and M. Yamashita(KAUM) for their assistance with collecting
apogonid fishes from Yaku-shima Island during
this study. We are grateful to H. Senou (KPM)
for providing valuable information on taxonomy
of apogonids, M. McGrouther (AMS) for opportunity to examine two syntypes of Amia fasciata
stevensi, A. Y. Suzumoto (BPBM) for a specimenloan, T. Yoshino (Department of Chemistry, Biology and Marine Science, Faculty of Science, Uni59

T. Yoshida et al.

versity of the Ryukyus, Japan) and A. Yamane and
M. Kawakami (MUFS) for providing literature, I.
Takaku (Yakushima-Diving-Life, Japan) for providing an underwater photograph of Figure 39B,
T. Hashimoto (KAUM) for X-ray photographs,
G. Yearsley (Hobart, Australia) for checking ourmanuscript, and Y. Haraguchi (KAUM) and H.
Iwatsubo and T. Haraguchi (Kagoshima University, Japan) for their curatorial assistance. This
study was supported in part by a Grant-in-Aid for
Scientific Research (A) (19208019) from the Japan Society for the Promotion of Science, Tokyo,
Japan, and a Grant-in-Aid for Young Scientists (B)(19770067) from the Ministry of Education, Science, Sports and Culture, Japan.
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