South Pacific Studies Vol.36, No.1, 2015

Distributional Range Extension of Herklotsichthys
quadrimaculatus (Clupeiformes: Clupeidae) in Southern
Japan
Hata Harutaka1*, takayaMa Mayumi2 and MotoMura Hiroyuki2

1: Graduate School of Fisheries, Kagoshima University, 4-50-20 Shimoarata, Kagoshima
890-0056, Japan
2: The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan*
Corresponding author
E-mail: k2795502@kadai.jp

Abstract
Nine specimens (98.1-133.8 mm standard length) of Herklotsichthys quadrimaculatus (Rüppell,1837), previously recorded in Japanese waters only from the southern Ryukyu Islands (Okinawa
Prefecture and Yoron-jima island) and Ogasawara Islands, were collected from the northern Ryukyu
Islands (Tanega-shima and Amami-Oshima islands), thereby representing a ca. 450 km northward
distributional range extension of the species, with the Tanega-shima specimens being the northernmost
record. The northern Ryukyu Islands specimens are described in detail.
Key words: Amami Islands, distribution, new record, Osumi Islands, sardine, taxonomy

Introduction
The clupeoid fish genus Herklotsichthys Whitley, 1951 is characterized by two fleshy
outgrowths on the hind margin of the gill opening, 3-8 front parietal striae on the top of thehead, the lower part of the second supramaxilla larger than the upper part, the last dorsalfin ray not filamentous and the hypo-maxilla toothless (wHiteHead 1985, Munroe et al.
1999). It is currently represented in Japanese waters by a single species, H. quadrimaculatus(Rüppell, 1837) (aonuMa and yagisHita 2013), having been recorded from the Ogasawara
Islands, Yoron-jima island and the southern Ryukyu Islands (aonuMa and yagisHita 2013,
Hata 2014). However, nine new specimens recently collected from Tanega-shima island(Osumi Islands) and Amami-Oshima island (Amami Islands) represent the first records ofthe species from those islands, and are described herein.
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Received: 21 June, 2015
Accepted: 23 August, 2015

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South Pacific Studies Vol.36, No.1, 2015

Materials and Methods
Counts and proportional measurements, expressed as percentages of standard length(SL), followed Hata and MotoMura (2015), except in following cases: head length wasmeasured from the snout tip to the most posterior point of the opercle, and postorbital lengthfrom the most posterior point of the orbit to the most posterior point of the opercle. Counts
and proportional measurements are shown in Table 1. All measurements were made withdigital calipers to the nearest 0.1 mm. The Osumi Islands is treated as a part of the Ryukyu
Islands in this study. Institutional codes used in this study are as follows: Bishop Museum,
Honolulu (BPBM), Hiwa Museum for Natural History (HMNH), Kagoshima University
Museum, Kagoshima (KAUM), Kanagawa Prefectural Museum of Natural History (KPM),
National Museum of Nature and Science, Tsukuba (NSMT), and Okinawa Churashima
Foundation (URM).

Results and Discussion
Herklotsichthys quadrimaculatus (Rüppell, 1837)
Standard Japanese name: Mizun
Fig. 1; Table 1
Material examined. 9 specimens (98.1-133.8 mm SL) from the Ryukyu Islands,
Japan — OSUMI ISLANDS: KAUM–I. 62110, 133.8 mm SL, off Hirota Fishing Port,
Minamitane, Tanega-shima island, 30°25′29″N, 130°58′50″E, 20 m depth, set net, 13 June
2014, H. Hata and M. takayaMa; KAUM–I. 66994, 98.1 mm SL, off Hirota Fishing Port,
Minamitane, Tanega-shima island, 30°25′29″N, 130°58′50″E, 20 m depth, set net, 19 Nov.
2014, M. takayaMa; KAUM–I. 67210, 107.6 mm SL, off Hirota Fishing Port, Minamitane,
Tanega-shima island, 30°25′29″N, 130°58′50″E, 20 m depth, set net, 6 Dec. 2014, M.
takayaMa; KAUM–I. 67809, 102.6 mm SL, off Hirota Fishing Port, Minamitane, Tanegashima island, 30°25′29″N, 130°58′50″E, 20 m depth, set net, 7 Dec. 2014, M. takayaMa.
AMAMI ISLANDS: KAUM–I. 62615, 100.4 mm SL, KAUM–I. 63848, 100.6 mm SL,
KAUM–I. 63849, 106.0 mm SL, KAUM–I. 63850, 102.8 mm SL, KAUM–I. 63851, 104.4mm SL, Yui, Setouchi, Amami-Oshima island, 28°10′53″N, 129°17′35″E, 7 m depth, linefishing, 25 June 2014, B. Jeong.
Description. Body oblong, depth about equal to head length, deepest at origin of dorsal
fin. Dorsal profile of head and body convex from snout tip to caudal-fin base. Ventral profile
of head and body convex from lower-jaw tip to end of anal-fin base, straight at caudal
peduncle. Belly slightly rounded, covered by 30 to 32 scutes from isthmus to anus. Predorsal
scute absent. Head large, compressed. Snout rounded. Mouth terminal, small, posterior tipof maxilla reaching to level with anterior margin of iris. Second supramaxilla spoon-shaped,
lower part larger than upper part. Premaxilla and hypomaxilla without teeth. Lower edge ofHata et al.: Distributional Range Extension of Herklotsichthys quadrimaculatus

41

maxilla toothed. Lower jaw with several conical teeth anteriorly. Posterior ramus of lower
jaw elevated. Orbit elliptical, eye and pupil round. Eye large, covered with adipose eyelid;
visible in dorsal and ventral views. Interorbital space flat. Nostrils elliptical, close to eachother, positioned anterior to orbit. Upper point of pectoral-fin insertion between verticallines through posteriormost points of preopercle and opercle. Lower point of pectoral-fin
insertion posterior to a vertical line through posteriormost point of opercle. Posterior tipof pectoral fin pointed. Pectoral-fin axillary scale present. Dorsal-fin origin posterior to avertical line through posterior tip of pectoral fin. Posterior end of dorsal-fin base posterior toa vertical line through posterior end of pelvic fin. Origin of pelvic fin located below originof 9-10th dorsal-fin ray. Posterior end of pelvic-fin base below 11th dorsal-fin ray. Posterior
end of depressed pelvic fin behind that of dorsal-fin base. Pelvic-fin axillary scale present.
Anal-fin origin positioned just below 24-25th lateral scale. Posterior end of anal-fin basebelow 30-31st lateral scale. Bases of dorsal and anal fins sheathed in a row of deciduous
scales. Caudal fin forked. Anus situated just anterior to anal-fin base, posterior to midpointof body. Scales cycloid, thin and deciduous except ventral scutes; body scales with 7
continuous striae across center and a slightly serrated posterior edge without perforations;Fig. 1. Fresh specimens of Herklotsichthys quadrimaculatus. A, KAUM–I. 62110,
133.8 mm SL, Tanega-shima island, Kagoshima Prefecture, Japan; B, KAUM–I.
62615, 100.4 mm SL, Amami-Oshima island, Kagoshima Prefecture, Japan.

42South Pacific Studies Vol.36, No.1, 2015

predorsal scales paired; elongate, wing-like scales present beneath normal paired predorsal
scales; no lateral line; no scales on head; no scales on fins, except a broad triangular sheathof scales on caudal fin. 3-5 frontparietal striae on top of head. Gill rakers long, slender,rough. Pseudobranchial filaments present. Two fleshy outgrowths on posterior border of gill
opening and a large papilla on lower edge of gill opening. Posterior margins of preopercle
and opercle smooth.
Color when fresh (Fig. 1): Head and body dark blue dorsally, whitish-silver ventrally.
A single dark blue stripe running from upper end of gill-opening to caudal-fin base. Twoyellow blotches on posterior margin of gill-opening. Tip of lower-jaw, lower edge of upperjaw and pupil black. Dorsal and caudal fin rays yellowish-black. Pectoral, pelvic and anal
fin rays white. All fin membranes translucent. Iris silver.
Color in alcohol: Head and body dark purple dorsally, elsewhere pale brown.
Remarks. The nine specimens were identified as H. quadrimaculatus on the basis of
the following combination of characters, which closely match the diagnostic features of H.
quadrimaculatus given by wongratana (1980), wHiteHead (1985) and Munroe et al. (1999):
body depth 25.9-29.2% of SL; elongate, wing-like scales present beneath normal paired
predorsal scales; no black spots on flank; lower gill rakers 32-34; no dusky tips on dorsal
and caudal fins; two yellow blotches on posterior margin of gill opening; and a single dark
blue stripe running from upper end of gill opening to caudal-fin base.
The genus Herklotsichthys contains 10 valid species (wHiteHead 1985, wongratana
1987). Although H. quadrimaculatus is most similar to H. collettei Wongratana, 1987 insharing elongate wing-like scales beneath normal paired predorsal scales and lacking blackspots on the flank (wongratana 1987, Munroe et al. 1999), the former is distinguished fromthe latter by having a lower count of lower gill rakers on the first gill arch (30-36 vs. 38-40
in H. collettei), yellow blotches on the posterior margin of the gill-opening (vs. no blotches),and dorsal fin without prominent markings (vs. tip of dorsal fin jet black; wongratana 1987,
Munroe et al. 1999).
Meristic and morphometric data for the present specimens were within the rangeof those for comparative specimens from Yoron-jima and Okinawa-jima islands, Japan,
Phuket, Thailand, and the Vava’u Islands, Tonga. However, the former (from Tanega-shima
and Amami-Oshima islands) had higher ranges of lower gill rakers on the second (33-38 vs.
27-37) and third gill arches (27-30 vs. 21-29), in addition to a slightly shorter head (25.027.8% of SL vs. 26.5-30.5%), smaller orbit diameter (11.8-12.9% of SL vs. 12.1-15.8%) andshorter mandible (12.1-13.8% of SL vs. 12.7-16.2%) (Table 1). These minor differences are
regarded herein as intraspecific variations of H. quadrimaculatus.
kisHinouye (1907) described Clupea mizun as a new species from Okinawa-jimaisland and proposed a Japanese name “Mizun” for the species. Clupea mizun is currently
recognized as a junior synonym of H. quadrimaculatus (wongratana 1980, wHiteHead
1985). The kisHinouye’s (1907) record of C. mizun representeds the first voucher specimenbased record of H. quadrimaculatus from Japanese waters. Subsequently, kaMoHara (1965)
and kurata et al. (1971) reported H. quadrimaculatus (as Harengula ovalis) from Ishigakijima and Okinawa-jima islands, and the Ogasawara Islands, respectively. Although yosHino

Hata et al.: Distributional Range Extension of Herklotsichthys quadrimaculatus

Table 1.

43

Counts and measurements, expressed as percentages of standard length, of
Herklotsichthys quadrimaculatus. Modes for counts and means of measurements are inparentheses.
Tanega-shima island,
Kagoshima, Japan

Amami-Oshimaisland, Kagoshima,
Japan

Yoron-jima island
and Okinawa, Japan,
Thailand, and Tonga

n=4

n=5

n = 2498.1-133.8

100.4-106.0

24.9-121.4

Dorsal-fin rays (unbranched)44-5 (4)3-5 (4)

Dorsal-fin rays (branched)

15-16 (15, 16)

14-15 (15)13-16 (15)Anal-fin rays (unbranched)332-3 (3)Anal-fin rays (branched)

14-16 (14)14-15 (14)14-17 (16)Pectoral-fin rays (unbranched)

1

1

1

Pectoral-fin rays (branched)

14-16 (15)15-16 (15)14-16 (15)Pelvic-fin rays (unbranched)

1

1

1

Pelvic-fin rays (branched)

7

7

7

Caudal-fin rays (upper + lower)

10 + 910 +910 + 9Gill rakers on 1st gill arch (upper)

15-16 (15, 16)

15-16 (15)

11-16 (15)Gill rakers on 1st gill arch (lower)

32-34 (33)33-34 (34)28-34 (32)Gill rakers on 1st gill arch (total)

48-49 (48, 49)

48-49 (49)

40-50 (46)Gill rakers on 2st gill arch (upper)

14-15 (14)14-15 (15)12-16 (15)Gill rakers on 2st gill arch (lower)

33-35 (33, 35)

37-38 (38)27-37 (36)Gill rakers on 2st gill arch (total)

47-49 (49)51-53 (53)39-53 (51)Gill rakers on 3rd gill arch (upper)

14-15 (14)14-17 (17)10-15 (14, 15)

Gill rakers on 3rd gill arch (lower)

27-30 (28)27-29 (27)21-29 (29)Gill rakers on 3rd gill arch (total)

42-44 (42)42-44 (43, 44)

31-44 (43)Gill rakers on 4th gill arch (upper)

12-14 (13)12-13 (13)8-14 (12, 13)

Gill rakers on 4th gill arch (lower)

19-21 (19)20-22 (22)16-23 (21)Gill rakers on 4th gill arch (total)

31-35 (31, 32, 33, 35)

32-35 (35)24-35 (34)Gill rakers on posterior face of 3rd gill arch

10-11 (10)8-9 (9)

7-11 (9)Prepelvic scutes

17-18 (17)18

17-19 (18)

Postpelvic scutes

13

12-14 (13, 14)

12-14 (13)30-31 (31)31-32 (32)30-32 (31)Branchiostegal rays

broken6-7 (7)5-7 (6)Lateral-scale series

40-41 (41)38-40 (38)38-41 (40)

Pseudobranchial filaments

19-20 (20)20-21 (20)

14-21 (20)Total length

124.6-124.9 (124.8)

123.2-125.8 (124.4)

123.1-130.9 (126.4)

Head Length25.0-27.2 (26.5)

26.7-27.8 (27.2)

26.5-30.5 (28.1)

Body depth

25.9-28.6 (27.2)

26.0-29.2 (27.5)

18.5-29.5 (25.6)

Pre-dorsal-fin length44.7-47.1 (45.8)

45.4-47.3 (46.4)

44.2-50.1 (46.6)

Standard length
CountsTotal scutes

Measurements (%SL)

44South Pacific Studies Vol.36, No.1, 2015

Snout tip to pectoral insertion

24.9-27.7 (26.6)

27.6-28.9 (28.0)

26.1-32.8 (28.3)

Snout tip to pelvic insertion

52.9-54.1 (53.4)

55.5-57.7 (56.9)

52.4-60.9 (55.5)

Pre-anal-fin length77.7-79.8 (79.2)

80.1-81.3 (80.9)

77.7-82.6 (80.0)

Dorsal-fin base length15.6-15.9 (15.7)

15.4-17.0 (16.0)

15.3-18.3 (16.4)

Anal-fin base length12.9-15.4 (13.8)

12.9-13.5 (13.1)

12.9-16.1 (14.2)

Caudal-peduncle length

8.7-9.4 (9.0)

8.3-10.0 (9.5)

6.3-9.8 (8.3)

Caudal-peduncle depth

8.2-9.8 (9.1)

8.4-9.3 (8.8)

7.7-10.8 (8.9)

Orbit diameter

11.8-12.8 (12.3)

12.5-12.9 (12.7)

12.1-15.8 (13.1)

Eye diameter

7.5-7.9 (7.7)

8.0-8.7 (8.3)

7.4-10.5 (9.1)

Snout length

3.7-4.3 (4.0)

3.6-4.6 (4.2)

3.4-4.7 (4.1)

D-P1

29.8-32.8 (31.3)

30.1-32.1 (31.1)

26.1-33.0 (30.4)

D-P2

25.2-27.3 (26.2)

25.7-28.5 (27.2)

17.3-30.1 (25.1)

D-A

39.9-41.1 (40.4)

39.4-40.6 (39.9)

26.5-41.6 (38.7)

P1-P2

28.3-29.9 (29.1)

29.8-31.1 (30.4)

19.6-31.3 (28.5)

P2-A

26.7-27.7 (27.2)

25.1-26.6 (25.9)

23.7-28.2 (26.3)

Pectoral-fin length

17.6

18.9-19.9 (19.4)

16.1-21.4 (19.6)

Pelvic-fin length12.4-13.0 (12.7)

12.3-13.1 (12.8)

11.1-13.6 (12.5)

Interorbital width

3.9-4.4 (4.1)

3.6-4.2 (3.9)

3.1-4.5 (4.1)

Postorbital length10.2-11.1 (10.7)

10.1-11.1 (10.6)

10.1-12.5 (10.9)

Upper-jaw length12.0-13.0 (12.4)

12.7-13.4 (13.0)

11.9-14.7 (13.3)

Mandible length12.1-13.4 (13.0)

12.9-13.8 (13.2)

12.7-16.2 (13.6)

1st unbranched dorsal-fin ray length

1.4-1.7 (1.6)

0.8-2.4 (1.6)

0.8-4.6 (1.8)

2nd unbranched dorsal-fin ray length

3.9-4.9 (4.3)

3.6-7.2 (5.1)

3.0-9.6 (5.0)

3rd unbranched dorsal-fin ray length

7.1-9.2 (8.2)

8.1-15.8 (10.5)

5.6-16.7 (9.6)

1st unbranched anal-fin ray length

1.2-1.3 (1.3)

0.7-1.9 (1.2)

0.6-2.6 (1.3)

2nd unbranched anal-fin ray length

2.7-2.7 (2.7)

2.8-5.2 (4.0)

2.0-5.8 (3.5)

3rd unbranched anal-fin ray length

6.0-6.8 (6.5)

5.7-6.7 (6.2)

5.5-7.0 (5.9)

1st pectoral-fin ray length

broken18.5-19.7 (19.1)

18.4-21.4 (19.2)

1st pelvic-fin ray length

11.9

12.5-13.1 (12.9)

11.1-13.5 (12.3)

et al. (1975) also reported the species (as Harengula ovalis) from the Ryukyu Islands,
detailed locality and voucher specimens were not indicated. senou and suzuki (1980)
reported Herklotsichthys quadrimaculatus (as Harengula ovalis) from the mouth of Nakama
River, Iriomote-jima island on the basis of six specimens (61.3-141.3 mm total length).
Subsequently, wHiteHead (1985) showed that applicable scientific name for this species is
Herklotsichthys quadrimaculatus.
Herklotsichthys quadrimaculatus was recorded from the southern Ryukyu Islands(Okinawa Prefecture) by the following authors on the basis of voucher specimens and/or
photographs: kon et al. (1998): Nakagusuku Bay, Okinawa-jima island, URM-P 38640,
60.8 mm SL; tacHiHara et al. (2001): mangrove estuary of Gesashi River, Okinawa-jimaisland, 4 specimens; Maeda and tacHiHara (2006): mouth of Teima River, Okinawa-jimaisland, juvenile specimens; senou et al. (2006): Ie-jima island, underwater photograph,

Hata et al.: Distributional Range Extension of Herklotsichthys quadrimaculatus

45

KPM-NR 36550; senou et al. (2007): Miyako Islands, underwater photographs, KPMNR 68271, 69489, 69807, 69969; watai et al. (2009): Tokashiki-jima island, 1 specimen,
KPM-NI 21601, underwater photograph, KPM-NR 44033; sakai et al. (2001): Iriomotejima island, NSMT-P 28501; oHta (2007): Yaeyama Islands, 17 kg catches in 2006; nanjo
et al. (2008): Urauchi River, Iriomote-jima island, 7 specimens, 30-40 mm SL; kanda etal. (2009): Hirakubo River, Ishigaki-jima island, 1 specimen; Miura (2012): Chinen Fish
Market, Nakagusuku Bay, Okinawa-jima island, photograph; and oka and MiyaMoto (2014):
Okinawa-jima island, 2838 juvenile specimens, 11.5-29.7 mm SL.
r andall et al. (1997) reported four specimens (BPBM 35347, 107-127 mm
SL; NSMT-P 46670, 84 mm SL; NSMT-P 46673, 99 mm SL) of Herklotsichthys
quadrimaculatus from the Ogasawara Islands and yosHigou and nakaMura (2002) noted
six specimens (HMNH-P 2816-2819, 2952, 2953) from Haha-jima and Chichi-jima islands,
Ogasawara Islands. Hata (2014) reported two specimens (KAUM–I. 52218, 52219) of H.
quadrimaculatus from Yoron-jima island, in the mid latitude of the Ryukyu Islands.
Although Hata and MotoMura (2011) reported four specimens (KAUM–I. 1014, 79.8mm SL; KAUM–I. 1015, 83.7 mm; KAUM–I. 7390, 38.7 mm; KAUM–I. 7391, 39.8 mm)from the mainland of Kagoshima, southern Kyushu, re-examination of those specimens
revealed them to be Sardinella lemuru Bleeker, 1853. However, aonuMa and yagisHita(2013) cited Hata and MotoMura’s (2011) records, including the Kagoshima mainland, inthe distributional range of H. quadrimaculatus. Although kawano et al. (2011) listed H.
quadrimaculatus from the Sea of Japan (coast of Yamaguchi Prefecture), no specimens wereretained.
Accordingly, H. quadrimaculatus has been recorded in Japanese waters only from theOgasawara Islands, Yoron-jima island, and Okinawa Prefecture, on the basis of voucher
specimens. Thus, the present (voucher) specimens from Tanega-shima and Amami-Oshimaislands represent a northward distributional range extension for the species, the Tanegashima specimens representing the northernmost record of the species.
Comparative material examined. Herklotsichthys quadrimaculatus (24 specimens,
24.9-121.4 mm SL): KAUM–I. 28792, 83.6 mm SL, reservoir tank of Okinawa Yambaru
Seawater Pumped Storage Power Station, Kunigami, Okinawa Prefecture, Japan(26°40′25″N, 128°15′56″E), 0.5 m depth, 11 July 2008, hand net, I dea C onsultant
I ncorporated ; KAUM–I. 52218, 104.2 mm SL, KAUM–I. 52219, 110.1 mm SL, off
Hakibina beach, south coast of Yoron-jima island, Kagoshima Prefecture, Japan (27°01′45″N,
128°25′07″E), 1.5 m depth, seine net, 23 Oct. 2012, FrlM FisH TeaM; KAUM–I. 55086, 37.5mm SL, KAUM–I. 55118, 35.8 mm SL, Chabana Port, west coast of Yoron-jima island,
Kagoshima Prefecture, Japan (27°02′57″N, 128°24′19″E), 0.5 m depth, fishing light, 30
June 2013, H. Hata et al.; KAUM–I. 60969, 104.3 mm SL, off Phuket Island, Thailand;
KAUM–I. 71459, 56.8 mm SL, Itoman Fishing Port, Itoman, Okinawa-jima island, Okinawa
Prefecture, Japan (26°07′N, 127°39′E), 3 Oct. 1988, fishing light, M. KiMura; KAUM–I.
71460, 24.9 mm SL, KAUM–I. 71461, 30.4 mm SL, Itoman Fishing Port, Itoman, Okinawajima island, Okinawa Prefecture, Japan (26°07′N, 127°39′E), 17 Sept. 1992, fishing light, M.
kiMura; URM-P 33372, 121.4 mm SL, URM-P 33373, 115.5 mm SL, URM-P33374, 113.3

46

South Pacific Studies Vol.36, No.1, 2015mm SL, URM-P 33419, 94.8 mm SL, URM-P 33420, 86.1 mm SL, URM-P 33421, 90.8
mm SL, URM-P 33422, 74.4 mm SL, URM-P 33490, 115.6 mm SL, URM-P 33491, 114.8mm SL, Vava’u Islands, Kingdom of Tonga; URM-P 43968, 118.6 mm SL, URM-P 43969,
101.2 mm SL, URM-P 43970, 104.0 mm SL, URM-P 43971, 99.2 mm SL, URM-P 43972,
99.1 mm SL, Ou-jima island, Tamagusuku, Nanjo, Okinawa Prefecture, Japan, 19 June
2007; URM-P 44695, 83.7 mm SL, Chinen-umino, Nanjo, Okinawa-jima island, Okinawa
Prefecture, Japan, 18 Nov. 2008.

Acknowledgements
We are especially grateful to B. jeong (KAUM) for collecting specimens and K.
MiyaMoto (URM) for specimen loans. We also thank Y. HaragucHi and other volunteers,and students of KAUM for their curatorial assistance. We greatly appreciated comments on
the manuscript by K. koeda (KAUM) and G. Hardy (Ngunguru, New Zealand), who readthe manuscript and provided help with English.
This study was supported in part by Grants-in-Aid for Scientific Research (A:26241027, B: 24370041 and C: 23580259 and 26450265) from the Japan Society for the
Promotion of Science, Tokyo, Japan (JSPS); the JSPS Asian Core Program, “Establishmentof Research and Education Network on Coastal Marine Science in Southeast Asia”, the JSPS
International Training Program “Protect Diversity of Bioresources in the Tropical Area”; the“Coastal Area Capability Enhancement in Southeast Asia Project” of the Research Institutefor Humanity and Nature, Kyoto, Japan; the “Biological Properties of Biodiversity Hotspots
in Japan” project of the National Museum of Nature and Science, Tsukuba, Japan; and“Establishment of Research and Education Network on Biodiversity and Its Conservationin the Satsunan Islands” project of Kagoshima University adopted by the Ministry of
Education, Culture, Sports, Science and Technology, Japan.

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