Biogeography 9.
23–30. Aug. 20, 2007

In situ Ontogenetic Color Changes of Pentapodus aureofasciatus (Perciformes,
Nemipteridae) off Yakushima Island, Southern Japan and Comments on the
Biology of the Species
Hiroyuki Motomura1* and Shigeru Harazaki2
1The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima, 890-0065 Japan
2Mori-To-Umi, 1559-1 Miyanoura, Kamiyaku-cho, Kumage-gun, Kagoshima, 891-4205 Japan

Abstract. Ontogenetic color changes of Pentapodus aureofasciatus are documented on the basis of underwater observations off Yakushima Island, southern Japan from 2004 to 2006, and compared with those of aclosely-related species, P. caninus. A photograph of a mature breeding male P. aureofasciatus is provided forthe first time and sexual dichromatism in the species also discussed. Morphological and color variations
recorded during the study suggested that P. aureofasciatus includes more than one species. The distributional
range of P. aureofasciatus extends to Izu Peninsula and the Izu Islands, Yakushima Island being the most likely northernmost breeding ground, supplying juveniles and/or larvae to the Pacific coast of mainland Japan via
the Kuroshio Current.
Key words: Nemipteridae, Pentapodus aureofasciatus, Pentapodus caninus, coloration, comparisons, distribution, ecology.

Introduction
Recently described by Russell (2001), the western Pacific nemipterid, Pentapodus aureofasciatus,
had been previously reported and figured as an undescribed species by Russell (1990) and Randall et al.
(1990). Randall et al. (1990: unnumbered figs) provided underwater photographs of a juvenile andadult from northeastern Australia, and Russell (1990:
pl. 5h) and Randall (2005: unnumbered fig.) provided the same photograph of a fresh adult specimen(BPBM 27118) from New Caledonia. Randall(2005: unnumbered fig.) also included an underwaterphotograph of a juvenile from the Coral Sea. An
adult specimen (identified as P. nagasakiensis) from
Taiwan figured in Shen (1993: pl. 108, fig. 9) was reidentified as P. aureofasciatus by Russell (2001).
*Corresponding author: motomura@kaum.kagoshimau.ac.jp

Thus, color photographs of juveniles and adults of P.
aureofasciatus have been published on several occasions, the coloration of the species being relatively
well known. However, ontogenetic color changes inthe species, based on a series of life stages, have at
no time been documented, although distinct color
differences occur between juveniles and adults.
A year-long tracing survey of the P. aureofasciatus population off Yakushima Island revealed theextent of body color changes with growth in thespecies. In addition, numerous mature males inbreeding coloration were observed during the study,
such having not been previously reported. Underwater photographs of a series of life stages, including abreeding male, are herein provided for the first time.
An examination of published and unpublishedunderwater photographs indicated that the geographic distribution of P. aureofasciatus extends to Izu
Peninsula and the Izu Islands, Japan, approximately
1,400 km northeast of the previously recorded north-− 23 −

Coloration of Pentapodus aureofasciatus

ernmost record (Okinawa Island). Several otheraspects of the biology of P. aureofasciatus are also
reported herein.
As Russell (1990, 2001) pointed out, the coloration of P. aureofasciatus is similar to that of P.
caninus (Cuvier in Cuvier and Valenciennes, 1830),
the latter co-occurring with P. aureofasciatus off
Yakushima Island. A series of life stages of P. caninus from Yakushima Island is also provided andcomparisons with P. aureofasciatus made.

Materials and Methods
Terminology and counting and measurement
methods followed Russell and Gloerfelt-Tarp(1984), Russell and Golani (1993), and Russell(2001). Standard and total lengths are expressed as
SL and TL, respectively. A tracing survey of the P.
aureofasciatus population by underwater observations was conducted north of Yakushima Island from2004 to 2006, some of the observed individuals
being photographed. During the study, all stages ofP. aureofasciatus from juvenile (less than 5 mm TL)
to adult (over 25 cm TL) were observed, and two
specimens collected to confirm identification(deposited in the Kagoshima University Museum,Japan: KAUM).
Specimens of P. aureofasciatus examined:
KAUM–I. 35, female, 150.0 mm SL, off Shitogo,
Yakushima Island, Kagoshima, Japan, 30˚27´N,
130˚31´E, 25 m depth, hand line, Nov. 2005, collected by H. Chuuman (Fig. 1a). KAUM–I. 285, male,158.5 mm SL, 26 Jan. 2006, other data as for
KAUM–I. 35 (Fig. 1b). Data for underwater photographs included in this report: P. aureofasciatus—
Fig. 2a: 4 cm TL, 15 m depth, 30˚27.32´N, 130˚
29.19´E, 22 July 2004; Fig. 2b: 6 cm TL, 25 m, 30˚
27.32´N, 130˚29.19´E, 16 Jan. 2006; Fig. 2c: 10 cm
TL, 25 m, 30˚27.32´N, 130˚29.19´E, 15 Jan. 2006;
Fig. 2d: 15 cm TL, 25 m, 30˚27.56´N, 130˚30.11´E,

Fig. 1. Fresh specimens of Pentapodus aureofasciatus from Yakushima Island, Kagoshima, Japan.
a: KAUM–I. 35, female, 150.0 mm SL (frozen for about 12 months after collection). b:
KAUM–I. 285, male, 158.5 mm SL (frozen for about 6 months). Photos by H. Motomura.

− 24 −

Hiroyuki Motomura and Shigeru Harazaki

16 Jan. 2006; Fig. 2e: 20 cm TL, 20 m, 30˚27.32´N,
130˚29.19´E, 9 June 2005; Fig. 2f: female, 25 cm
TL, 20 m, 30˚27.32´N, 130˚29.19´E, 24 May 2005;
Fig. 2g: male, 25 cm TL, 25 m, 30˚27.32´N, 130˚
29.19´E, 5 Oct. 2004. P. caninus—Fig. 3a: 4 cm TL,
15 m, 30˚27.32´N, 130˚29.19´E, 12 Aug. 2004; Fig.
3b: 15 cm TL, 30 m, 30˚27.32´N, 130˚29.19´E, 11
Nov. 2005; Fig. 3c: 20 cm TL, 20 m, 30˚27.32´N,
130˚29.19´E, 14 Apr. 2005; Fig. 3d: 25 cm TL, 20 m,
30˚27.32´N, 130˚29.19´E, 24 May 2005; Fig. 3e: 40
cm TL, 20 m, 30˚27.32´N, 130˚29.19´E, 16 Feb.
2005. All underwater photographs were taken off
Issou, Yakushima Island, Japan, by H. Harazaki.

Results and Discussion
Identification—Both specimens collected(KAUM–I. 35, I. 285) were identified as P. aureofasciatus, confirming to the species diagnosis provided
by Russell (2001): head scales not reaching forward
dorsally to a line between posterior nostrils; lower
limb of preopercle scaly; posterior tip of depressed
pelvic fin just short of level of anus; caudal fin
forked, without filamentous extensions, upper andlower lobes subequal in length; and no dark bar at
pectoral-fin base. The following counts and proportional measurements of the specimens also agreed
with those of the type specimens given by Russell(2001): dorsal-fin rays X–XI, 8–9; anal-fin rays III,7; pectoral-fin rays 16–17; pelvic-fin rays I, 5; pored
lateral-line scales 46; transverse scale rows above
lateral line 3, below 11; transverse scale rows on preopercle 3–5+3; body depth 3.6–3.7 in SL; headlength 3.5–3.6 in SL; snout 3.2–3.4 in head length;eye diameter 3.3–3.4 in head length, 1.0–1.1 in snout
length; interorbital width 2.8–3.0 in head length;
suborbital depth 8.1–8.9 in head length; caudalpeduncle depth 2.2–2.5 in caudal-peduncle length;
dorsal-fin base length 1.9 in SL; fourth dorsal-fin
spine longest, 2.0–2.1 times length of first dorsal-fin
spine; sixth dorsal-fin soft ray longest, 0.8–0.9 times
length of longest dorsal-fin spine; anal-fin baselength 6.1 in SL; first anal-fin spine 1.9–2.1 in lengthof second anal-fin spine; second anal-fin spine

1.3–1.4 in length of third anal-fin spine. However,
although Russell (2001: 55) stated “Maxilla reachingalmost to or just behind anterior margin of eye”, bothof the Yakushima specimens had the posterior margin of the maxilla well short of a vertical through theanterior margin of the eye (Fig. 4).
Sexual dichromatism in fresh adult specimens(Fig. 1)—Figure 1 shows fresh adult female andmale specimens of P. aureofasciatus collected off
Yakushima Island, although their being maintained
frozen for a long period before being photographed
may have resulted in slightly changed colorationfrom that occurring naturally. Nevertheless, thefemale specimen (Fig. 1a: 150.0 mm SL) was wellconsistent with the specimen (BPBM 27118, 156
mm SL) figured by Russell (1990, 2001) and Randall(2005), both having reddish caudal fins. The caudal
fin of the male specimen (Fig. 1b: 158.5 mm SL),tinged with reddish on the upper lobe and yellowishgreen on the lower lobe, with the posterior margin of
the fin translucent was consistent with an adult specimen photograph of P. aureofasciatus (143 mm SL,labeled as P. nagasakiensis) given by Shen (1993).
Bruce Carlson (pers. comm. in Russell, 2001) reported that some individuals in Fiji had the upper lobe of
the caudal fin blackish. Such a color pattern, reddish(or blackish in the Fijian population) upper caudalfin lobe and yellowish or yellowish-green lower
lobe, is most likely to be that normally characteristicof adult male P. aureofasciatus, although colorationof the caudal-fin posterior margin seems to be variable. The Yakushima male and that figured in Shen
(1993) both had the posterior margin of the caudal
fin translucent (Fig. 1b), whereas Russell (2001) stated that an adult specimen (BPBM 29186, 155.4 mmSL) from Tonga had a blackish margin on the upper
caudal-fin lobe. In addition, the caudal fin of theYakushima female was entirely reddish, including
the posterior margin (Fig. 1a), whereas that of thepresumed female specimen figured by Russell (1990,
2001) and Randall (2005) was reddish with a whitish
posterior margin.
The Yakushima male had a greenish body (Fig.
1b). However, Shen’s male specimen had a similar− 25 −

Coloration of Pentapodus aureofasciatus

Fig. 2. Life stages of Pentapodus aureofasciatus off Yakushima Island, Japan. a: 4 cm TL. b: 6 cm
TL. c: 10 cm TL. d: 15 cm TL. e: 20 cm TL. f: 25 cm TL. g: 25 cm (male). Photos by H.
Harazaki (photo data in text).

body color to the Yakushima female (Fig. 1a); i. e.
pinkish dorsal-fin membranes and pectoral fin,whitish pelvic and anal fins, brownish dorsal and silvery ventral body, and a distinct yellow stripe on theside of the body from behind the eye to the anterior
portion of the caudal peduncle. The Yakushima
male specimen was probably exhibiting nuptial col-ors when it was collected, judging from the coloration evident in an underwater photograph of abreeding male (Fig. 2g).
All published photographs of fresh specimens of
P. aureofasciatus have shown the upper lobe (atleast) of the caudal fin to be reddish. However,underwater photographs showing a reddish caudal− 26 −

Hiroyuki Motomura and Shigeru Harazaki

Fig. 3. Life stages of Pentapodus caninus off Yakushima Island, Japan. a: 4 cm TL. b: 15 cm TL.
c: 20 cm TL. d: 25 cm TL. e: 40 cm TL. Photos by H. Harazaki (photo data in text).

fin have never been reported (Randall et al., 1990:
unnumbered figs.; Randall, 2005: unnumbered fig.).
In addition, all adults, except breeding males,
observed during this study had a translucent caudal
fin (see Fig. 2). Apparently, the reddish caudal-fin
coloration in fresh female specimens is a postmortem development, paralleling a similar condition
known to occur in the mullid fish, Mulloidichthys
vanicolensis (Valenciennes in Cuvier and Valenciennes, 1831).
In situ color changes with growth (Figs 2–3) —Four cm TL juveniles of P. aureofasciatus (Fig. 2a)
had a deep purplish blue head and body with two
brown-edged bright yellow stripes, the uppermost of
the latter extending on each side from the upper partof the caudal peduncle over the eye to above theanterior eye margin, thereafter joining in the dorsal

midline on the head. The width of the stripe wasnarrower than the pupil diameter, that part of the
stripe above the eye being the narrowest. The lower
bright yellow stripe ran from the tip of the upper lip
to the anteroventral margin of the eye, and from theposteroventral margin of the eye through above thepectoral-fin base to the middle of the posterior margin of the caudal-fin base. The posterior part (posterior to eye) of the lower strip was wider than both theanterior part (anterior to eye) and the upper stripe.
The widest part of the lower stripe, between the pectoral- and anal-fin bases, was subequal to the pupil
diameter. The eye, except for the black pupil, wasusually bright yellow. All fins were transparent.
Although four cm TL juveniles of P. caninus(Fig. 3a) shared almost identical coloration with P.
aureofasciatus (Fig. 2a), differences existed in the− 27 −

Coloration of Pentapodus aureofasciatus

Fig. 4. Head of Pentapodus aureofasciatus. KAUM–I.
285, male, 158.5 mm SL. Left and right verticalbars indicate locations of the posterior margin of
the maxilla and anterior margin of the eye,respectively.

condition of the anterior portion of the upper stripe(extending anterior to the eye and not joining in the
middle) and the presence of faint narrow yellow
stripe extending from the lip along the dorsal midlineof the head (Fig. 3a).
In P. aureofasciatus, the deep purplish-blue bodyfound in four cm TL juveniles gradually becamebrownish dorsally and bluish-white ventrally withgrowth (Figs. 2a–f). The anterior portion of theupper yellow stripe (above anterior margin of eye)disappeared at ca 15 cm TL, the anterior portion of
the lower yellow stripe (anterior to eye) also being
lost at ca 25 cm TL (Fig. 2f). The body color change
pattern with growth of P. caninus was similar to thatof P. aureofasciatus, although the upper body stripeof P. caninus between the eye and caudal peduncle
remained throughout life, despite fading somewhat(Figs 3a–e). The mid-dorsal head stripe in P. caninus juveniles subsequently disappeared (Figs 3a, b).
With the exception of the nuptial colored males,
the caudal-fin coloration of live P. aureofasciatus
was always transparent or translucent throughout life
(Figs 2a–f). In contrast, P. caninus over ca 15–20
cm TL had reddish anterior margins on the upperand lower caudal-fin lobes (Figs 3b–e).
During this study, numerous breeding males of P.
aureofasciatus were observed off Yakushima. A
photograph of a nuptial colored male (Fig. 2g), taken

Fig. 5. Map of southern and central Japan. Enlargement
indicates positions of Yakushima and Okinawa
Islands, location of underwater observations andspecimens collected during the study (star) anddirection of the Kuroshio Current (arrow).

at 25 m on 5 October 2004, is the first publishedrecord of such. That individual (25 cm TL) was thesame size as the female (or normal colored male?)illustrated in Fig. 2f. Nuptial colored males wereclearly distinct from normal colored adults,
subadults and juveniles, being characterized by thefollowing color patterns: head and body yellowishgreen dorsally and yellowish-white ventrally, withthree blue stripes; the first stripe from posterodorsal
margin of eye to a vertical through anal-fin origin,
the second from above upper lip to anterior marginof eye and subsequently from posterior margin ofeye to caudal peduncle, the third from below lower
lip passing below eye to opercular margin; caudal fin
reddish on upper lobe and upper part of lower lobe,
remaining lower lobe and posterior margin of fintranslucent; yellow stripe absent (replaced by second− 28 −

Hiroyuki Motomura and Shigeru Harazaki

blue stripe). The nuptial coloration of a male P. caninus is shown in Fig. 3e.
Russell (2001: 55) described “pearly blue band on
ventral surface from behind pelvic fins to caudal
peduncle present in larger specimens” as the life
color of P. aureofasciatus. Although such a blue
band is not found in the P. aureofasciatus population
off Yakushima Island (Fig. 2), it is found in nuptial
colored P. caninus males (Fig. 3e). As mentioned
above, several color pattern and morphological (Fig.
4) differences occur among P. aureofasciatus populations from different localities, indicating that ‘P.
aureofasciatus’ includes more than one species orgeographic subspecies or, at least, genetically distinct populations. Molecular analysis may berequired in future to assess these possibilities.
Distribution and ecological notes—P. aureofasciatus is widely distributed in the western Pacific,
where it ranges from Okinawa Island south to northern New South Wales, and around New Caledonia, Fiji, Tonga and Samoa (Russell, 2001).
Yakushima Island, where the present specimens
were collected or photographed, is located approximately 500 km north-northeast from the previouslyknown northernmost record (Okinawa Island) of thespecies (Fig. 5).
North of Yakushima Island P. aureofasciatus isone of the most abundant fish species. Numerous
epibenthic juveniles occur on the rocky reef slopes,
they gradually moving into mid-water with growth.
While epibenthic juveniles have a bright blue body(Figs 2a–b), this changes to brownish with habitatchange (Figs 2c–f). Subadult and adult fish form
mid-water schools, usually comprising 200–300
individuals (but often over 500) off YakushimaIsland. Although large numbers of P. aureofasciatus
are found north of Yakushima Island throughout theyear, few occur south of the island and at otherJapanese localities, including the Ryukyu Islands.
Only juveniles of P. aureofasciatus have beenreported from the Pacific coast of mainland Japan,
the northernmost records, based on underwaterobservations, being from Izu Peninsula (several
underwater photographs on web sites) and the Izu

Islands (S. Kato, pers. comm., 14 Oct. 2004). Suchjuveniles are most likely to have been accidentallytransported from Yakushima Island by the Kuroshio
Current, normally flowing from west of the Ryukyu
Islands through Yakushima and TanegashimaIslands (Fig. 5, gray arrow) to the Pacific coast ofJapan. Northern reefs off Yakushima Island appear
to be the northernmost (and largest in Japanesewaters) breeding ground for P. aureofasciatus.
Because of their abundance north of YakushimaIsland, P. aureofasciatus is often collected by local
fishermen for use as bait for the Bigfin Reef Squid,
Sepioteuthis lessoniana (Lesson, 1830).
Masuda and Kobayashi (1994: 166) provided
underwater photographs of Pentapodus sp. 1 (10 cm
TL) and Pentapodus sp. 2 (2 cm TL) from the
Yaeyama Islands (south of Okinawa Island), bothfish identified here as P. caninus. Okamura (1997:364) described P. paradiseus (7 cm TL) from Kochi
and Pentapodus sp. (8 cm TL) from Iriomote Island,
providing underwater photographs. These species
are identified here as P. aureofasciatus and P. caninus respectively.
Names—Russell (1990), Randall et al. (1990)
and Randall (2005) proposed different English
names, Yellow-striped Whiptail, Blue Butterfish andGoldstripe Bream, respectively, for P. aureofasciatus. Recently, Yellowstripe Threadfin Bream has
been officially registered in the “list of standardised
Australian fish names” provided by CSIRO Marine
and Atmospheric Research, Fisheries Research andDevelopment Corporation, and Seafood Services
Australia (Yearsley et al., 2006). Yakushimakitsuneuo is herein proposed as the new standard
Japanese name for the present specimens of P. aureofasciatus, although local fishermen at YakushimaIsland refer to the species (and also P. caninus) as
Maddo-Ibo.

Acknowledgments
We are grateful to Graham S. Hardy, who readthe initial manuscript and offered helpful comments,
and also thank Yuko Motomura, Kazue Harazaki,− 29 −

Coloration of Pentapodus aureofasciatus

Mayumi Takayama, Yuriko Haraguchi, Masahide
Ito and Mizuki Matsunuma for their assistance, andSyouichi Kato for providing information on P. aureofasciatus from the Izu Islands. This study was supported in part by Grant-in-Aids for Scientific Research (A) by the Japan Society for the Promotion of
Science, Tokyo, Japan (19208019) and for Young
Scientists (B) by the Ministry of Education, Science,Sports and Culture, Japan (19770067).

Literature Cited
Masuda, H. & Kobayashi, Y., 1994. Grand Atlas ofFish Life Modes. 465 pp. Tokai Univ. Press,
Tokyo.
Okamura, O., 1997. Pentapodus paradiseus, Pentapodus sp. In Okamura, O. & Amaoka, K. (Eds),
Sea Fishes of Japan: 364. Yama-kei, Tokyo.
Randall, J. E., 2005. Reef and shore Fishes of theSouth Pacific. New Caledonia to Tahiti and the
Pitcairn Islands. xii, 707 pp. Univ. Hawaii
Press, Honolulu.
Randall, J. E., Allen, G. R. & Steene, R. C., 1990.
Fishes of the Great Barrier Reef and Coral Sea.
xx, 557 pp. Univ. Hawaii Press, Honolulu.
Russell, B. C., 1990. Nemipterid Fishes of the World(threadfin breams, whiptail breams, monocle

breams, dwarf monocle breams and coralbreams). An annotated and illustrated Catalogueof nemipterid Species known to Date. FAO
Species Catalogue, 12. FAO Fish. Synopsis, No.
125. v, 149 pp., FAO, Rome.
—, 2001. A new species of Pentapodus(Teleostei: Nemipteridae) from the western
Pacific. Beagle, Rec. Mus. Art Galleries North.
Territory, 17: 53–56.
Russell, B. C. & Gloerfelt-Tarp, T., 1984. A new
species of Parascolopsis (Pisces: Nemipteridae)
from Indonesia. Beagle, Occas. Pap. North. Territory Mus. Arts Sci., 1: 111–114.
Russell, B. C. & Golani, D., 1993. A review of the
fish genus Parascolopsis (Nemipteridae) of thewestern Indian Ocean, with description of a new
species from the northern Red Sea. Israel J.
Zool., 39: 337–347.
Shen, S.-C., 1993. Fishes of Taiwan. xx, 960 pp.
Taiwan Univ., Taipei.
Yearsley, G. K., Last, P. R. & Hoese, D. F. (Eds),
2006. Standard Names of Australian Fishes. 65
pp. CSIRO Marine and Atmospheric Research,
Hobart.

− 30 −(Accepted January 31, 2007)