Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 1–28, March 22, 2007

Four New Subspecies of Acheilognathus Bitterlings (Cyprinidae:
Acheilognathinae) from Japan
Ryoichi Arai1, Hiroshi Fujikawa2 and Yoshikazu Nagata3
1

Department of Zoology, University Museum, University of Tokyo,
7–3–1 Hongo, Bunkyo-ku, Tokyo 113–0033, Japan
E-mail: araryo@um.u-tokyo.ac.jp
2
Osaka Prefectural Board of Education, Chuo-ku, Osaka 540–8571, Japan3
Department of Biology, Osaka Kyoiku University, 4–698–1 Asahigaoka, Kashiwara, Osaka 582–8582, Japan

Abstract Four new bitterlings, Acheilognathus tabira erythropterus subsp. nov., Acheilognathus
tabira tohokuensis subsp. nov., Acheilognathus tabira jordani subsp. nov. and Acheilognathus tabira nakamurae subsp. nov., were described on the basis of more than 600 specimens from 26 localities in Japan. Acheilognathus tabira erythropterus, A. t. tohokuensis and A. t. jordani, formerly all
included in a single undescribed subspecies of Acheilognathus tabira, differ from other subspeciesof A. tabira in having a red-edged anal fin in nuptial males. Acheilognathus t. erythropterus, distributed on the Pacific Ocean side of eastern Honshu, is distinguished from A. t. tohokuensis and A.
t. jordani by having shorter ellipsoidal eggs (ratio of major axis to minor axis: 1.4–2.2 vs. 2.0–3.3
in A. t. tohokuensis and A. t. jordani). Acheilognathus t. tohokuensis, distributed on the Japan Sea
side of eastern Honshu, is distinguished from A. t. jordani in lacking a black blotch on the dorsal
fin in juveniles. Acheilognathus t. jordani, distributed on the Japan Sea side of western Honshu, isdistinguished from A. t. erythropterus and A. t. tohokuensis by having a black blotch on the dorsal
fin in juveniles. Acheilognathus t. nakamurae, distributed in the Kyushu area, differs from all othersubspecies by the following combination of characters: edge of the dorsal fin in males red, anal fin
in nuptial males edged with white, a conspicuous black blotch on the dorsal fin in both juveniles
and small adult females, and long ellipsoidal eggs (ratio of major axis to minor axis: 2.3–2.9).
Acheilognathus t. tabira Jordan and Thompson, 1914 differs from the 4 new subspecies by having
more branched dorsal fin rays (10 vs. 9), the edge of the dorsal fin in males (grayish vs. red) andshorter ellipsoidal eggs (ratio of major axis to minor axis: 1.4–1.7 vs. 1.4–3.3). A key is provided
for all Japanese species and subspecies of Acheilognathus.
Key words : Four new subspecies, Acheilognathus tabira erythropterus, Acheilognathus tabira
tohokuensis, Acheilognathus tabira jordani, Acheilognathus tabira nakamurae, Cyprinidae, Japan.

The genus Acheilognathus includes about 28
species and subspecies, all being restricted toEast Asia: 2 species are known from Russia(Bogutskaya and Naseka, 1996), 1 species from
the Mongolian People’s Republic (Sokolov et al.,1983), 4 species from Laos (Kottelat, 2001a),about 7 species from Vietnam (Mai, 1978; Kottelat, 2001b), about 18 species from China (Woo,1964; Lin, 1998), 6 species from South Korea(Kim and Park, 2002) and 9 species and subspecies from Japan (Nakamura, 1969; Hagiwara,
2002).
Regarding Acheilognathus from Japan, Hosoya(1993) described 8 species and subspecies including A. melanogaster Bleeker, 1860, A.
cyanostigma Jordan and Fowler, 1903, A.
rhombeus (Temminck and Schlegel, 1846), A.
longipinnis Regan, 1905, A. typus (Bleeker,
1863), A. tabira tabira Jordan and Thompson,
1914, A. tabira subsp. 1 and A. tabira subsp. 2.
Hagiwara (2002) later reported an introduced bitterling, A. macropterus (Bleeker, 1871), which
had become established in Lake Kasumigaura,
Ibaraki Prefecture, eastern Honshu.
Acheilognathus tabira Jordan and Thompson,1914 had earlier been divided into 3 subspecies

2

R. Arai, H. Fujikawa and Y. Nagata

by Nakamura (1969), with Japanese names for
each, i.e., Shirohire-tabira for A. tabira tabira,Akahire-tabira for A. tabira subsp. (a) and Seboshi-tabira for A. tabira subsp. (b). Although
the latter two subspecies have at no time formallydescribed, biological information on them has
been accumulated through various studies; interrelationships based on phenotypes and karyotypes (Arai, 1978, 1988), molecular phylogeny
based on a mitochondrial gene (Okazaki et al.,2001), allozymes (Fujikawa et al., 1984), karyotypes (Ojima et al., 1973), sensory canal system(Arai and Kato, 2003), pharyngeal teeth andmasticatory process of the basioccipital bone(Suzuki and Hibiya, 1985a), minute skin surface
tubercles on larvae (Fukuhara et al., 1982; Suzuki and Hibiya, 1985b), development (Suzuki,1985), and distribution (Nagata et al., 1981; Fujikawa, 1983; Suzuki, 1985; Saitoh et al., 1988;
Watanabe, 1998). Predictably, however, the lackof formal designation has resulted in some difficulties in comparing the above studies.
Accordingly, morphological reexaminationand formal taxonomic treatment of Nakamura’s(1969) unnamed subspecies is necessary. Nakamura (1969) defined Acheilognathus tabira subspp. (a) and (b) on egg shape immediately after
oviposition and presence or absence of a black
blotch on the dorsal fin in juveniles and small
adult females. According to Nakamura (1969),
the egg shape of A. t. subsp. (a) was short ellipsoidal (Japanese ‘keiran’ type), similar to that of
A. t. tabira, being distinguished from A. t. subsp.
(b) by a lower ratio of major to minor axes
(1.6–1.7 vs. 2.2–2.3 in A. t. subsp. (b)) (Nakamura, 1969). However, during the present study,
eggs of A. t. subsp. (a) from the Japan Sea side of
Honshu were found to be longer ellipsoid (ratioof major to minor axes, 2.0–3.3), similar to thoseof A. t. subsp. (b) (ratio of major to minor axes
2.3–2.9, present study). Furthermore, although
Nakamura (1969) considered that both A. t.
subsp. (a) and A. t. tabira lacked a black blotch
on the dorsal f in, whereas it occurred in both juveniles and small adult females of A. t. subsp.
(b), such were found in A. t. subsp. (a) juveniles

from the Japan Sea side of western Honshu.
Clearly, Nakamura’s (1969) concept of Acheilognathus tabira subspecies was in need of revision. The present study showed that A. t. subsp.
(b) and A. t. tabira (both Nakamura, 1969) differed from each other at the subspecific level,
being consistent in their characteristics, whereas
A. t. subsp. (a) comprised 3 hitherto unrecognized subspecies, identificable by a combinationof egg shape and presence or absence of a black
blotch on the dorsal fin in juveniles. One subspecies had a black dorsal fin blotch in juveniles,
previously considered an important diagnostic
character of A. t. subsp. (b) by Nakamura (1969).
The remaining 2 undescribed subspecies lacked ablack dorsal fin blotch in juveniles, but could be
distinguished from each other by ellipsoidal egg
shape, barbel length, and vertebral number. Thus,
a new classification of Acheilognathus tabira,which includes 4 new subspecies, is proposed
below.
Materials and Methods
More than 1000 specimens in total, from 38
localities, were examined (Table 1, Fig. 1). The
classification of genera of the subfamily
Acheilognathinae follows Arai and Akai (1988),
that of host mussels following Kondo (1982,
1998, 2006). Methods for counts and measurements follow Hubbs and Lagler (1958). Vertebrae and unpaired fin rays were counted from radiographs. Vertebral number includes the Weberian complex (as 4) and the terminal pleurostyle(1). The inserted position of the proximal segment of the first pterygiophore in the dorsal andanal fins, expressed as D-PTG-1 (Fig. 2) and APTG-1, respectively (see Arai et al., 1995), wasexamined from radiographs. D-PTG-1 was described in relation to the associated vertebra (vertebral number), suffix ‘a’ indicating insertion of
the second dorsal pterygiophore into the succeeding intervertebral space (Fig. 2-left), suffix‘b’ indicating no pterygiophore insertion into thesucceeding intervertebral space (Fig. 2-right).
Because proportional measurements of adults

Four New Acheilognathus Bitterlings from Japan
Table 1.

3

List of specimens examined in 5 subspecies of Acheilognathus tabira.

Loc.
No.*

Cat. No.

No. ofspecimens

Subspecies

Locality

Date ofcollection

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
21
22
22
22
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38

NSMT-P 74699
OKU-P 114
NSMT-P 74700
NSMT-P 72657, 74701
NSMT-P 27553
ZUMT 61513
NSMT-P 27043
ZUMT 61512
NSMT-P 72656, 74702
NSMT-P 74703
NSMT-P 74704
NSMT-P 74705
NSMT-P 74706
NSMT-P 72594
NSMT-P 74707
NSMT-P 74708
NSMT-P 72658, 74709
NSMT-P 74710
NSMT-P 72595
OKU-P 110ZUMT 61511
OKU-P 213
OKU-P 154
OKU-P 171
NSMT-P 74711
NSMT-P 11004
NSMT-P 72596
NSMT-P 74712
NSMT-P 72659, 74713
NSMT-P 74714
NSMT-P 72597
NSMT-P 74715
NSMT-P 74716
NSMT-P 74717
NSMT-P 74718
NSMT-P 74719
NSMT-P 74720
OKU-P 170
NSMT-P 74721
NSMT-P 74722
NSMT-P 74723
NSMT-P 74724

10
42
40
66
2
1
2
2
50
35
38
3
41
2
50
5
7
33
2
34
12
48
11
30
57
2
1
47
49
13
1
8
49
50
25
36
40
29
13
33
16
76

A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. erythropterus
A. t. erythropterus
A. t. erythropterus
A. t. erythropterus
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. nakamurae
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira
A. t. tabira

Lake Kaikon-numa
Lake Hachirogata
Lake Ogata
Lake Nishinuma
Yasuda-machi
near Teradomari
Natori City
Nasu-machi
Lake Kasumigaura
Seimeigawa River
Machinogawa River
Kawaradagawa River
Lake Akauragata
Himi City
Takaoka City
Lake Ouchigata
Lake Kibagata
Lake Shibayamagata
Ono City
Tanegaike Pond
Oharagawa River
Oharagawa River
Hatahokogawa River
Hatahokogawa River
Hatahokogawa River
Hatahokogawa River
Tataragawa River
Horikawayosui River
Futatsukawa River
Lake Shimoezuko
Kushidagawa River
Lake Yogo
Katayama, Lake Biwa
Sugaura, Lake Biwa
Setagawa River
Kanzakigawa River
Yodogawa River
Pond in Sanda City
Mukogawa River
Gion-yosui River
Nakajimaooike Pond
Ashimorigawa River

28 Aug. 1978
5 Aug. 1981
5 Aug. 1981
4 Aug. 1981
26 Oct. 1974
21 Oct. 2005
26 June 1975
28 May 2005
2 Apr. 1982
1–2 Apr. 1982
16 Oct. 1981
16 Oct. 1981
16 Oct. 1981
29 May 2005
7 Aug. 1981
17 Oct. 1981
18 Oct. 1981
18 Oct. 1981
12 June 2005
27 June 1981
4 July 2003
3 Aug. 1984
4 Apr. 1977
26 May 1982
28 July 1976
1 Sept. 1969
23 May 2005
16 Apr. 1982
15 Apr. 1982
14–15 Apr. 1982
10 June 2005
4 May 1982
1 May 1982
4 May 1982
21 Aug. 1981
27 Mar. 1982
10 Oct. 1980
18 May 1982
18 May 1982
21 Mar. 1982
21 Mar. 1982
18 Apr. 1980

* Locality numbers correspond to those in Fig. 1.

differed from those of juveniles, only specimens
longer than 40 mm in standard length (SL) wereused in this study. Ripe eggs were obtained bypressing the belly of a mature female (identified
by a characteristically long ovipositor) andplaced on a mesh slide glass or transferred into apetri dish filled with distilled water for subsequent observation. Ripe eggs of bitterlings are
non-spherical, the shape being indicated by theratio of the major to minor axes (Coleman, 1991),
following measurement of eggs fixed in 10%
formalin. When eggs were not available for direct
examination, measurements were taken fromphotographs. Eastern and western Honshu were
defined as the areas east and west of the Fossa
Magna (Shizuoka-Itoigawa Line), respectively.
Principal component analyses (PCA) were carried out on 6 selected log10-transformed morpho-

4

R. Arai, H. Fujikawa and Y. Nagata

Fig. 1. Map showing distribution of 5 subspecies of Acheilognathus tabira. Numbered symbols, corresponding
to locality numbers in Table 1, indicate collection localities of specimens examined; unnumbered symbols indicate collection locality of only eggs (Inabegawa River, new record) and literature records.

FMNH, Field Museum of Natural History,
Chicago; NSMT, National Museum of Natureand Science (formerly National Science Museum), Tokyo; OKU, Osaka Kyoiku University,
Osaka; ZUMT, Department of Zoology, University Museum, University of Tokyo, Tokyo.
Fig. 2. Insertion of first pterygiophore in dorsal
fin (D-PTG-1). Left, D-PTG-110tha; right,
D-PTG-110thb. 1st PTG, first pterygiophore;
D1–5, first 5 dorsal fin rays; numbers, vertebrae.

Acheilognathus tabira Jordan and Thompson,
1914(Japanese name: Tabira)(New English name: Tabira bitterling)

metric characters (see ‘Statistical analyses ofmorphometric and meristic characters’) and 5
meristic characters in order to clarify which character was different among subspecies.
Institutional abbreviations are as follows:
AKPM, Akita Prefectural Museum, Akita;(Figs. 3–12)

Proportional measurements of specimens
longer than 40 mm SL as % SL: body depth
29–38, head length (HL) 21–27, predorsal length
50–57, caudal peduncle depth 10–13; as % HL:

Four New Acheilognathus Bitterlings from Japan

orbit diameter (OD) 27–40; as % OD: snout
length 41–117 (relatively longer with growth)and barbel length 2–65. Dorsal rays, iii, 8–11;
anal rays, iii, 8–10; number of dorsal rays minus
number of anal rays, 1 to 2; first simple rays of
dorsal and anal fins very small, hidden underskin; longest simple rays of dorsal and anal fins,
soft; pectoral rays, i, 14–16; pelvic rays, i, 7; caudal rays, i9/8i; lateral line complete; lateralline scales, 33–40; scales on proximal part ofcaudal fin, 1 or 2; transverse scales, 10–11;
scales around caudal peduncle, 14; abdominal
vertebrae, 17–20; caudal vertebrae, 17–20; totalvertebrae, 35–38; first dorsal pterygiophore supporting first and second dorsal rays; insertion of
first pterygiophore in dorsal fin (D-PTG-1), 10th
to 13th; insertion of first pterygiophore in anal
fin (A-PTG-1), 17th to 20th; pharyngeal teeth,
0.0.5–5.0.0; chewing area on pharyngeal teeth
serrated; gill rakers on outer side of 1st gill arch,
9–11; a pair of maxillary barbels; pearl organs on
snout and around nostrils in adult males; a roundor elliptical black blotch immediately behind
upper corner of gill opening; a short longitudinal
stripe along mid-caudal peduncle to a point
under middle of dorsal fin, not reaching caudal
fin; edge of dorsal fin in males, red or grayish;
nuptial coloration in males conspicuous during
spawning season, distal anal fin margin being redor white (Figs. 3–4 and 7–11); female coloration
unremarkable, an ovipositor obvious during February to August (spawning season); a black
blotch absent or present on dorsal fin in juveniles
and/or small adult females (Fig. 5); eggs ellipsoid immediately after oviposition, ratio of major
axis to minor axis, 1.4–3.3 (Table 6, Fig. 6);
diploid chromosome number, 44.
Remarks. Acheilognathus tabira differsfrom other Japanese Acheilognathus species by
the following combination of characters:
branched dorsal rays 8–11, branched anal rays
9–10, number of dorsal rays minus number ofanal rays 1 to 2, and anal fin in nuptial males
edged with red or white. This species comprises
5 subspecies (Tables 1–11). As shown in Tables 4
and 11, the number of total vertebrae is almost

5equal to that of lateral-line scales.
According to many previous studies (see synonymies for subspecies of Acheilognathus tabira)and the present survey, this species is endemic toJapan. Katayama (1941), Nakamura (1977) and
the Research Group on Fishes and Shellfishes in
Nagano Prefecture (1980: 68) reported Acheilognathus tabira from the Maruyama River, Hyogo
Prefecture, from near Maizuru, Kyoto Prefecture,and from Lake Kizakiko, Nagano Prefecture,
Japan, respectively. Because the specimens usedby those authors were not available for the present study, they could not be identified to subspecific level.
This species has also been reported from
Korea and China. Mori (1935: 566–567, 1 table)
described Paracheilognathus tabira (Acheilognathus tabira) from a single male specimen
from Seoul, Korea. This Korean specimen, however, differed from A. tabira in Japan by having alonger longitudinal stripe on the body, i.e., reaching anteriorly beyond the dorsal fin origin (Mori,
1935), while it ends anteriorly below the middleof the dorsal fin in the latter (Jordan and Thompson, 1914), in that way being similar to A.
melanogaster, although sharing with A. tabira asimilar combination of dorsal and anal fin ray
numbers (D. iii, 9; A. iii, 9). Additional specimens of A. tabira from Korea have not yet beenreported. Woo (1964) reported Acanthorhodeus
tabiro (Acheilognathus tabira) from China, although this was later described as a new species,
Acheilognathus macromandibularis Doi, Arai,and Liu, 1999.
Subspecies of Acheilognathus tabira
Acheilognathus tabira tabira Jordan andThompson, 1914
(Japanese name: Shirohire-tabira)(New English name: White tabira bitterling)(Figs. 4C, 6 L–M, 7)
Acheilognathus tabira Jordan and Thompson, 1914: 220,
pl. 25, fig. 1 (Lake Biwa); Aoyagi, 1957: 100 (in part).
Acheilognathus tabira tabira: Nakamura, 1963: 149, fig.
81; Nakamura, 1969: 36 and 378, pls. 2 A–A, 14, 15,

6

R. Arai, H. Fujikawa and Y. Nagata

A

B

C

D

Fig. 3. Color in life of nuptial males of 2 subspecies of Acheilognathus tabira. A: A. t. tohokuensis subsp. nov.,
near Teradomari, collected by Nobuo Inoue, 21 Oct. 2005, photographed by Ryu Uchiyama, Oct. 2005 (after
artificial induction of nuptial coloration). B–D: A. t. erythropterus subsp. nov.; B, Natori, Miyagi Prefecture,collected by Kenji Saitoh, 11 June 2005, photographed by Ryu Uchiyama, June 2005; C, Inashiki, Lake Kasumigaura, collected by Tomiji Hagiwara, 21 May 2005, photographed by Ryu Uchiyama, June 2005; D,
Ichinosegawa River, Ibaraki Prefecture, collected and photographed by Yoshikazu Nagata, 2 Apr. 1986.

A

B

C

D

Fig. 4. Color in life of nuptial males of 3 subspecies of Acheilognathus tabira. A–B: A. t. jordani subsp. nov.; A,
Himi, collected by Osamu Inamura, 2 Feb. 1992, photographed by Osamu Inamura, 20 May 1992; B, Ono,collected by Jun-ya Kitazima, 12 June 2005, photographed by Ryu Uchiyama, June 2005. C: A. t. tabira,Inabegawa River, Mie Prefecture, collected by Jun-ya Kitazima, 23 June 2004, photographed by Ryu Uchiyama, May 2005. D: A. t. nakamurae subsp. nov., Tataragawa River, collected by Jun Nakajima, 23 May
2005, photographed by Ryu Uchiyama, May 2005.

Four New Acheilognathus Bitterlings from Japanand 92 A–B; Ojima et al., 1973: 172, fig. 7 (Osaka andOkayama); Fukuhara et al., 1982: 233, fig. 2B (Okayama); Hosoya, 1982: 29 (Okayama); Sawada, 1984: 54,
pl. 53 I–J; Fujikawa et al., 1984: 57 (Shiga, Osaka,
Hyogo, and Okayama); Suzuki and Hibiya, 1985a:
184, fig. 5A (Shiga and Okayama); Suzuki and Hibiya,
1985b: 339, fig. 3A (Shiga); Suzuki, 1985: 64, figs.
2–3 (Okayama); Arai and Akai, 1988: 201; Nagata,
1989: 373, 3 figs.; Doi, 1992: 36 (Wakayama); Hosoya,
1993: 216; Watanabe, 1998: 262; Okazaki et al., 2001:
92, figs. 3–4 (Shiga); Arai and Kato, 2003: 3, table 1,
fig. 6B (Shiga).
Paracheilognathus tabira tabira: Tsubokawa, 1988: 4(Okayama); Hosoya, 1988: 155.
Rhodeus (?) tabira tabira: Kawanabe, 1987: 26, 1 fig.
Acheilognathus lanceolata tabira: Niwa, 1967: 103 (Aichi
and Gifu).
Acheilognathus limbata: Jordan and Fowler, 1903: 818 (inpart: Shiga).
Material examined. FMNH 57071, holotype ofAcheilognathus tabira, male, 68.0 mm SL, Lake Biwa;

7

NSMT-P 74718 (ex OKU-P 101) (25 specimens: 5 males,
18 females, and 2 sex unknown), 26.3–75.8 mm SL, Setagawa River, Nango, Shiga Prefecture, 21 Aug. 1981;
NSMT-P 74715 (ex OKU-P 167) (8: 3M5F), 47.1–61.5mm SL, Lake Yogo, Shiga Prefecture, 4 May 1982;
NSMT-P 74716 (ex OKU-P 166) (49: 25M24F),
41.1–70.8 mm SL, Katayama, Lake Biwa, Shiga Prefecture, 1 May 1982; NSMT-P 74717 (ex OKU-P 168) (50:20M28F2 sex unknown), 31.8–79.3 mm SL, Sugaura,
Lake Biwa, Shiga Prefecture, 4 May 1982; NSMT-P
74719 (ex OKU-P 158) (36: 11M25F), 41.2–74.9 mm
SL, Kanzakigawa River, Yodogawa River System, Osaka
Prefecture, 27 Mar. 1982; NSMT-P 74720 (ex OKU-P
151) (40: 10M30F), 26.1–64.8 mm SL, floodplain pools,
Yodogawa River, Osaka Prefecture, 10 Oct. 1980; OKU-P
170 (29: 19M10F: voucher specimens lost after examination but radiographs still available), 39.1–72.8 mm SL,
pond in Sanda, Hyogo Prefecture, 18 May 1982; NSMT-P
74721 (ex OKU-P 169) (13: 2M11F), 44.8–75.8 mm
SL, Mukogawa River, Hyogo Prefecture, 18 May 1982;
NSMT-P 74722 (ex OKU-P 155) (33: 15M18F),
40.1–64.5 mm SL, Gion-yosui River, Okayama, Okayama

A

B

C

D

E

F

Fig. 5. Dorsal fin coloration in juveniles and adult females. A–B: Juvenile A. t. tohokuensis subsp. nov. (blotch
absent); A, AKPM-P 436-1, paratype, 13.0 mm SL, Gojome-machi, Akita Prefecture, collected by Hideki
Sugiyama, 27 June 2004; B, AKPM-P 438-2, paratype, 26.4 mm SL, Kisakata, Akita Prefecture, collected
and photographed by Hideki Sugiyama, 4 June 2006. C–E: Juvenile A. t. jordani subsp. nov. (black blotch
present); C, NSMT-PL 265-1, paratype, 19.5 mm SL, Lake Kibagata, collected by Kunihiko Yamamoto, 17June 2005; D, 22.1 mm SL, Oharagawa River, collected 3 Aug. 1984 (from Saitoh et al., 1988); E, 31.0 mm
SL, Oharagawa River, collected by Chika Oshiumi, Mar. 2003. F: Adult female of A. t. nakamurae subsp.
nov. (black blotch present), NSMT-P 75167-6, paratype, 50.2 mm SL, Futatsukawa River, collected by Hiroshi
Fujikawa, 3 Sep. 1980.

8

R. Arai, H. Fujikawa and Y. Nagata

Prefecture, 21 Mar. 1982; NSMT-P 74723 (ex OKU-P
156) (16: 3M13F), 40.5–72.1 mm SL, Nakajimaooike
Pond, Okayama, Okayama Prefecture, 21 Mar. 1982;
NSMT-P 74724 (ex OKU-P 148) (76: 21M55F),
42.1–65.2 mm SL, Ashimorigawa River, Okayama Prefecture, 18 Apr. 1980; NSMT-P. 72597 (1M), 44.0 mm SL,
Kushidagawa River, Mie Prefecture, 10 June 2005.

data are shown in Tables 2–4. Proportional measurements as % SL: body depth 30–38, headlength 21–27, predorsal length 50–55, caudal peduncle depth 11–13; as % HL: orbit diameter
28–40; as % OD: snout length 41–106, barbel
length, 12–51. Dorsal rays, iii, 9–11; anal rays,iii, 8–10; number of dorsal rays minus number ofanal rays, 0–2; pectoral rays, i, 14–16; pelvic
rays, i, 7; caudal rays, i9/8i; abdominal vertebrae, 17–20; caudal vertebrae, 17–20; total vertebrae, 35–38; insertion of first pterygiophore in
dorsal fin (D-PTG-1), 10th to 13th (Table 5); insertion of first pterygiophore in anal fin (A-PTG-

Diagnosis. Dorsal rays, iii, 9–11; anal rays,iii, 9–10; dorsal fin in males grayish; anal fin in
nuptial males white; no black blotch on dorsal finin juveniles; eggs short ellipsoid, ratio of major
axis to minor axis, 1.4–1.7 (Table 11).
Description. Morphometric and meristic

Table 2. Averages of proportional measurements of body depth (BD), head length (HL), predorsal length(PredL), caudal peduncle depth (CPD), orbit diameter (OD), snout length (SnL), and barbel length (BarL) in
specimens 40 mm SL in 5 subspecies of Acheilognathus tabira.
Loc.
No.

Cat. No.

A. t. tohokuensis
1 NSMT-P 74699
2 OKU-P 114
3 NSMT-P 74700
4 NSMT-P 72657, 74701
A. t. erythropterus
9 NSMT-P 72656, 74702
10 NSMT-P 74703
A. t. jordani
11 NSMT-P 74704
12 NSMT-P 74705
13 NSMT-P 74706
15 NSMT-P 74707
16 NSMT-P 74708
17 NSMT-P 72658, 74709
18 NSMT-P 74710
20 OKU-P 110
21 OKU-P 213
A. t. nakamurae
22 OKU-P 154
22 OKU-P 171
24 NSMT-P 74712
25 NSMT-P 72659, 74713
26 NSMT-P 74714
A. t. tabira
28 NSMT-P 74715
29 NSMT-P 74716
30 NSMT-P 74717
31 NSMT-P 74718
32 NSMT-P 74719
33 NSMT-P 74720
34 OKU-P 170
35 NSMT-P 74721
36 NSMT-P 74722
37 NSMT-P 74723
38 NSMT-P 74724

Locality
No. of
BD/SL HL/SL PredL/SLCPD/SL OD/HL SnL/OD BarL/OD(Prefecture) specimens (%)
(%)
(%)
(%)
(%)
(%)
(%)
Akita
Akita
Akita
Akita

7
42
40
54

32
32
32
31

25
24
25
25

55
53
54
54

11
12
11
11

30
31
30
29

82
87
76
90

22
11
22
23

Ibaraki
Ibaraki

49
35

32
32

24
24

53
53

11
11

30
32

81
81

36
35

Ishikawa
Ishikawa
Ishikawa
ToyamaIshikawa
Ishikawa
Ishikawa
Tottori
Shimane

36
2
40
41
5
7
30
34
48

32
32
31
31
31
30
31
32
34

24
24
23
24
24
25
24
24
24

53
53
53
54
52
53
52
53
53

11
11
12
11
11
11
11
12
12

29
31
29
31
29
31
31
32
32

88
72
89
82
89
78
84
87
83

30
14
22
17
16
18
24
27
31

NagasakiNagasakiFukuoka
Fukuoka
Kumamoto

8
30
47
49
13

35
35
32
32
32

24
23
23
24
23

56
53
54
54
52

12
12
12
12
12

31
30
30
32
33

77
81
98
85
80

37
34
38
38
47

Shiga
Shiga
Shiga
Shiga
Osaka
Osaka
Hyogo
Hyogo
OkayamaOkayamaOkayama

7
49
44
21
33
21
27
13
30
12
75

33
34
33
35
33
33
34
35
34
35
36

24
24
24
23
24
24
24
22
25
24
25

51
52
51
52
51
54
54
53
53
53
53

11
11
11
12
12
11
12
12
12
12
12

32
33
33
35
32
33
31
32
32
37
31

81
85
83
75
78
75
84
80
76
57
76

35
35
37
26
30
33
22
34
35
32
32

Four New Acheilognathus Bitterlings from Japan

A

B

C

D

E

F

G

H

I

J

K

L

M

N

9

Fig. 6. Eggs of 5 subspecies of Acheilognathus tabira. A–D: A. t. tohokuensis subsp. nov.; A, Gojome-machi,
photographed by Hideki Sugiyama, 29 Apr. 2006; B, Akita, Akita Prefecture, photographed by Hideki
Sugiyama, 7 May 2006; C, Aizu area, Fukushima Prefecture, photographed by Makoto Kuraishi, 24 May
2006; D, near Teradomari, photographed by Ryu Uchiyama, 11 Dec. 2005 (after artificial induction of spawning). E–G: A. t. erythropterus subsp. nov.; E, Nasu-machi (A), photographed by Yoshikazu Nagata, 24 June
2006; F, Nasu-machi (B), photographed by Hitoshi Kubota, 15 June 2006; G, Lake Kasumigaura, photographed by Ryu Uchiyama, July 2005. H–K: A. t. jordani subsp. nov.; H, Himi, photographed by Ryu
Uchiyama, June 2005; I, Lake Kibagata, photographed by Kunihiko Yamamoto, 31 May 2001; J, Ono, photographed by Ryu Uchiyama, May 2005; K, Oharagawa River, photographed by Chika Oshiumi, 7 Apr. 2003.
L–M: A. t. tabira; L, Inabegawa River, photographed by Ryu Uchiyama, May 2005; M, Kushidagawa River,photographed by Ryu Uchiyama, June 2005; N, A. t. nakamurae subsp. nov., Tataragawa River, photographed
by Ryu Uchiyama, May 2005.

1), 17th to 20th. Eggs short ellipsoid, ratio ofmajor axis to minor axis, 1.4–1.7 (Table 6, Figs.
6 L–M).
Color in life. Dorsal fin in males marginedwith gray (red in other subspecies). Nuptial coloration strikingly presented in sexually mature
males during the spawning season: lateral surface
of body bright greenish-blue, belly black; pelvic
fin black proximally, margined with white along
anterior edge; anal fin proximally black, margin-

ed with white (hence derivation of English nameof subspecies). Females show no nuptial coloration, but develop an ovipositor to deposit eggs
into host mussels. A black blotch on the dorsal
fin is absent in juveniles.
Habitat. Widely distributed from the shore
to 30–40 m depth in Lake Biwa, and numerous
small nearby ponds continuous with the former.
Also along rocky shores of rivers and irrigation
creeks and in floodplain pools.

10

R. Arai, H. Fujikawa and Y. Nagata

A

B

Fig. 7. Acheilognathus tabira tabira. A: NSMT-P 74718-9, male, 75.8 mm SL, Nango, Setagawa River, collected
and photographed by Hiroshi Fujikawa, 21 Aug. 1981; B: NSMT-P 72597, male, 44.0 mm SL, Kushidagawa
River, collected by Jyun-ichi Kitamura, 10 June 2005. Photographed by Ryu Uchiyama, 14 June 2005.

Spawning season. From late April to earlyAugust, with peak spawning in May and June.
Host mussels. Six species and subspecies ofmussels have been reported as hosts for eggs of
A. tabira tabira. Obovalis omiensis (Fukuhara etal., 1982), Pronodularia japanensis and Anodonta woodiana (Nakamura, 1963, 1969) are usually
hosts for bitterling eggs. In Lake Biwa, Unio
douglasiae biwae, Inversiunio reinianus and A.
calipygos are hosts for bitterling eggs, there
being no particular host selectivity evident. Selectivity of O. omiensis as host is unknown in
Lake Biwa, however, because of the extremely
low density of the former (Hirai, 1964). In a
creek in Okayama City, Obovalis omiensis wasclearly selected as host over Unio douglasiae
nipponensis, in spite of the abundance of the latter (Kondo et al., 1984). No significant differences in mussel size preference by bitterlings
were apparent in either Lake Biwa or the above
creek (Hirai, 1964; Kondo et al., 1984).
Distribution. Western Honshu: Aichi, Gifu,
Mie, Shiga, Kyoto, Osaka, Hyogo and Okayama
Prefectures.
Remarks. Acheilognathus t. tabira is distinguished from the other 4 subspecies by the dorsal
fin color in males (grayish vs. red in the other 4
subspecies) and more branched dorsal rays (9–11
vs. usually 9). Nakamura (1969, pl. 92B) showedthat A. t. tabira lacked a black blotch on the dorsal fin in juveniles. The short ellipsoidal eggs of
A. t. tabira were described by Nakamura (1969:38) and Suzuki (1985), being the shortest among
all subspecies of Acheilognathus tabira (ratio ofmajor axis to minor axis: 1.4–1.7 vs. 1.4–3.3)(Table 6, Fig. 6).
Populations from Lake Biwa and adjacentareas (locality numbers 28–31, Fig. 1) were distinctive from those from the San-yo area (locality
numbers 36–38) by the shorter predorsal
length/SL on average (51–52 % vs. 53 % in Sanyo populations), and a greater average of totalvertebrae (36.4–36.9 vs. 35.9–36.1) (Tables 2
and 4). Twenty-five specimens (33% of 75 specimens) from the Ashimorigawa River had 11
branched dorsal rays. Twelve of 75 specimens
had 9 branched anal rays (Table 3). The combination of 11 and 9 branched dorsal and anal rays,
respectively, is similar to that in Paracheilognathus pseudorhombeus from Korea (Mori,
1935, table 1), which was synonymized with
Acheilognathus rhombeus by Uchida (1939:
159–160). However, A. t. tabira from the
Ashimorigawa River (75 specimens) differs fromP. pseudorhombeus (9 specimens) by the dorsal
fin (grayish vs. dark red in P. pseudorhombeus)
and anal fin (white vs. pale red) color patterns inmales, a shallower body (meanSD and rangeof body depth/SL: 35.71.1, 33.3–38.0% vs.
40.41.6, 38.5–43.5%), shorter snout (meanSD and range of snout length/HL: 23.52.1,
19.5–27.9% vs. 32.60.9, 31.3–34.5%) andlarger orbit (meanSD and range of OD/HL:
31.21.6, 27.5–35.0% vs. 26.71.2, 24.4–
28.6%).

Four New Acheilognathus Bitterlings from Japan

11

Acheilognathus tabira erythropterus subsp. nov.
(Japanese name: Akahire-tabira)(New English name: Red tabira bitterling)(Figs. 3 B–D, 6 E–G, 8)
Acheilognathus tabira: Okada and Ikeda, 1938: 102 (inpart: Miyagi); Kuronuma, 1940: 235 (Chiba); Hubbs
and Kuronuma, 1943: 184, figs.1–2 (Chiba).
Acheilognathus tabira subsp.: Nakamura, 1963: 149, fig.
82 (in part); Ojima et al., 1973: 172 (Ibaraki); Saito,
1979: 164, 1 fig. (Tochigi).
Acheilognathus tabira subsp. (a): Nakamura, 1969: 42
and 380, pl. 92C (in part); Fujikawa et al., 1984: 54 (inpart: Ibaraki); Suzuki and Hibiya, 1985a: 184, fig. 5B(Ibaraki); Suzuki, 1985: 64, figs. 4–5 (Ibaraki); Arai

Fig. 8. Acheilognathus
tabiraerythropterus
subsp. nov., ZUMT 61512, paratype, male,64.6 mm SL, Nasu-machi, collected by
Takayoshi Ueda, 28 May 2005. Photographed
by Ryu Uchiyama, June 2005.

Table 3. Frequency distributions of branched dorsal fin rays/branched anal fin rays in 5 subspecies of Acheilognathus tabira.
Loc. No.

Cat. No.

A. t. tohokuensis
1 NSMT-P 74699
2 OKU-P 114
3 NSMT-P 74700
4 NSMT-P 72657, 74701
5 NSMT-P 27553
A. t. erythropterus
7 NSMT-P 27043
9 NSMT-P 72656, 74702
10 NSMT-P 74703
A. t. jordani
11 NSMT-P 74704
12 NSMT-P 74705
13 NSMT-P 74706
15 NSMT-P 74707
16 NSMT-P 74708
17 NSMT-P 72658, 74709
18 NSMT-P 74710
20 OKU-P 110
21 OKU-P 213
A. t. nakamurae
22 OKU-P 154
22 OKU-P 171
22 NSMT-P 11004
24 NSMT-P 74712
25 NSMT-P 72659, 74713
26 NSMT-P 74714
A. t. tabira
28 NSMT-P 74715
29 NSMT-P 74716
30 NSMT-P 74717
31 NSMT-P 74718
32 NSMT-P 74719
33 NSMT-P 74720
34 OKU-P 170
35 NSMT-P 74721
36 NSMT-P 74722
37 NSMT-P 74723
38 NSMT-P 74724

Locality
No. of
8/8(Prefecture) specimens
Akita
Akita
Akita
Akita
Niigata

10
42
40
52
2

Miyagi
Ibaraki
Ibaraki249
35

Ishikawa
Ishikawa
Ishikawa
ToyamaIshikawa
Ishikawa
Ishikawa
Tottori
Shimane

38
3
39
49
5
7
32
33
48

NagasakiNagasakiNagasakiFukuoka
Fukuoka
Kumamoto

11
30
2
48
49
13

Shiga
Shiga
Shiga
Shiga
Osaka
Osaka
Hyogo
Hyogo
OkayamaOkayamaOkayama

8
49
50
25
36
40
29
13
34
17
75

8/9

9/8

2
2
3

1

2
1

2
1

1
1

1
3

1

5
2

1

1

1

2

9/9 9/10 10/8 10/9 10/10 11/9 11/10

10
37
34
47
2
1
44
32
21
3
37
35
3
4
22
28
45
10
23
2
43
43
12

2
3

1
1

2

10

5

1

1
13

1

1

8

1

1

4

1

1

1
1
1

2
4
1

3
3
3
2
1

2
3
2

1
1

1

2

8
39
40
20
28
36
24
10
26
15
42

5
5
2
4
2
1

1
1
1

1
1

1

2

1

6

12

13

1

2
1

12

R. Arai, H. Fujikawa and Y. Nagata

and Akai, 1988: 201 (in part); Okazaki et al., 2001: 92,
figs. 3–4 (Ibaraki); Arai and Kato, 2003: 3, table 1, fig.
6C (Ibaraki).
Acheilognathus tabira subsp. 1: Sawada, 1984: 55, pl. 53
K–L (in part); Hosoya, 1993: 216 (in part); Watanabe,
1998: 262 (in part); Nagata and Fujikawa, 2000: 120, 1
fig. (in part); Ogawa, 2001: 35 (Tochigi).
Acheilognathus tabira subsp. R: Nagata, 1989: 374, 2 figs.
(in part).
Paracheilognathus tabira subsp. 1: Hosoya, 1988: 155 (inpart).

male, 49.5 mm SL, Lake Kasumigaura, Hama, Tamatsukuri, Ibaraki Prefecture, collected on 2 April 1982.
Paratypes. NSMT-P 74702 (ex OKU-P 160) (49: 29
males and 20 females), 41.4–62.8 mm SL, Lake Kasumigaura, Hama, Tamatsukuri, Ibaraki Prefecture, 2 Apr.
1982; NSMT-P 74703 (ex OKU-P 159) (35: 17M18F),
40.5–56.8 mm SL, Seimeigawa River, Ami, Ibaraki Prefecture, 1–2 Apr. 1982; NSMT-P 27043 (2), 30.1 and 34.0mm SL, Natori, Masudagawa River, Miyagi Prefecture, 26
June 1975; ZUMT 61512 (2: 1M1F), 64.6 and 58.2 mm
SL, Nasu-machi, Nakagawa River system, Tochigi Prefecture, 28 May, 2005.

Holotype. NSMT-P 72656 (ex OKU-P 160-41),
Table 4. Frequency distributions and averages of numbers of vertebrae and avarages of lateral-line scales (LlSc)
in 5 subspecies of Acheilognathus tabira.
Loc.
No.

Cat. No.

A. t. tohokuensis
1 NSMT-P 74699
2 OKU-P 114
3 NSMT-P 74700
4 NSMT-P 72657, 74701
5 NSMT-P 27553
A. t. erythropterus
7 NSMT-P 27043
9 NSMT-P 72656, 74702
10 NSMT-P 74703
A. t. jordani
11 NSMT-P 74704
12 NSMT-P 74705
13 NSMT-P 74706
15 NSMT-P 74707
16 NSMT-P 74708
17 NSMT-P 72658, 74709
18 NSMT-P 74710
20 OKU-P 110
21 OKU-P 213
A. t. nakamurae
22 OKU-P 154
22 OKU-P 171
22 NSMT-P 11004
24 NSMT-P 74712
25 NSMT-P 72659, 74713
26 NSMT-P 74714
A. t. tabira
28 NSMT-P 74715
29 NSMT-P 74716
30 NSMT-P 74717
31 NSMT-P 74718
32 NSMT-P 74719
33 NSMT-P 74720
34 OKU-P 170
35 NSMT-P 74721
36 NSMT-P 74722
37 NSMT-P 74723
38 NSMT-P 74724

Locality(Prefecture)

No. ofspecimens

Akita
Akita
Akita
Akita
Niigata

10
42
40
52
2

Miyagi
Ibaraki
Ibaraki249
35

Ishikawa
Ishikawa
Ishikawa
ToyamaIshikawa
Ishikawa
Ishikawa
Tottori
Shimane

38
3
41
48
5
7
33
33
48

NagasakiNagasakiNagasakiFukuoka
Fukuoka
Kumamoto

10
27
2
46
49
13

Shiga
Shiga
Shiga
Shiga
Osaka
Osaka
Hyogo
Hyogo
OkayamaOkayamaOkayama

8
49
50
24
36
38
29
13
33
16
74

Number of vertebrae
35

1
5
8

36

37

38

Average Average

5
16
2
7

5
25
22
38
2

1
16
7

36.5
36.6
37.4
37.0
37.0

1
37
22
5
6
7
1

1
1
5
12
2

1
3
9
3
16

LlSc

8
24
37

7
5
31
3
35
32
4
6
18
9
9

2
9
1
6
1

5
15
2
8
5

38
36
8

5
13
14
6
25
23
17
4
27
14
49

3
30
32
14
7
6
11
9
3
2
9

6
5

5
4
4
1
1

35.9
36.1
37.2
36.4

35.5
36.0
35.9

35.5
35.8

36.9
37.0
36.9
37.0
36.8
37.1
36.9
36.3
36.2

36.4
36.5
36.4
36.3
36.8
36.3
36.5
36.1
35.4

35.5
35.6
35.0
37.1
36.9
36.6

35.0
35.3

36.4
36.8
36.8
36.9
36.2
35.9
36.4
36.7
36.0
36.1
35.9

36.3
36.8
37.0
36.7
35.8
35.5
36.1
36.1
35.9
35.9
35.6

37.6
36.8
36.5

Four New Acheilognathus Bitterlings from Japan

Diagnosis. Dorsal rays, usually iii 9; analrays, usually iii 9; fewer total vertebrae than in allother subspecies, 36.00.6 (meanSD); dorsal
fin in males, red; anal fin in nuptial males, red;no black blotch on dorsal fin in juveniles; eggs
ellipsoid, ratio of major axis to minor axis,
1.4–2.2 (Table 11).
Description. Morphometric and meristic
data are shown in Tables 2–4. Paratype data are
given in parentheses. Proportional measurements
as % SL: body depth 32 (30–34), head length 24(23–26), predorsal length 53 (50–55), caudal peduncle depth 11 (10–13); as %HL: orbit diameter
30 (28–35); as % OD: snout length 103 (51–116),barbel length 31 (24–53). Dorsal rays, iii, 9 (iii,
8–9); anal rays, iii, 9 (iii, 8–9); number of dorsal
rays minus number of anal rays, 0 (1 to 1); pectoral rays, i, 14 (i, 14–15); pelvic rays, i, 7 (i, 7);
caudal rays, i9/8i (i9/8i); abdominal vertebrae, 18 (17–19); caudal vertebrae, 18 (17–19);
total vertebrae, 36 (35–37); insertion of firstpterygiophore in dorsal fin (D-PTG-1), 11th
(10th to 12th) (Table 5); insertion of first pterygiophore in anal fin (A-PTG-1), 18th (17th or18th). Eggs ellipsoid, ratio of major axis to minor
axis, 1.4–2.2 (Table 6, Figs. 6 E–G).
Color in life. Dorsal fin in males marginedwith red. Lateral body surface in males greenish
in spawning season (similar to that of A. t. jordani and A. t. tohokuensis); pelvic fin black proximally, margined with white along anterior edge;
edge of anal fin varying from red to pale red, but
usually margined with red (hence deviation ofEnglish name of subspecies). Black blotch absent
from dorsal fin in juveniles.
Habitat. Rivers, lakes and ponds in plains;
creeks and rivers near coasts, and mouths ofrivers draining into lagoons.
Spawning season. From April to June.
Host mussels. No information available.
Etymology. The subspecific name, erythropterus, is a Latin adjective referring to the nuptial
color of males (erythrored, pterusfin).
Distribution. Pacific Ocean side of eastern
Honshu: Miyagi, Tochigi, Ibaraki, Chiba Prefectures and Tokyo Metropolis.

13

Remarks. Acheilognathus t. erythropterus
subsp. nov. differs from the other 4 subspecies by
the following combination of characters: anal fin
in nuptial males edged with red, no black blotch
on the dorsal fin in juveniles, fewer total vertebrae, and eggs intermediate in shape between
short and long ellipsoid. Acheilognathus t. erythropterus (85 specimens examined) differs from
A. t. tohokuensis (127 specimens examined) by
having fewer vertebrae (meanSD and range:
36.00.6, 35–37 vs. 37.00.6, 36–38 in A. t. tohokuensis) and a longer barbel (meanSD andrange of BarL/OD: 35.86.5, 22–56% vs.
18.77.8, 2–39%) (Tables 2, 4, 8, and 11). Furthermore, eggs of the former are shorter thanthose of the latter (ratio of major axis to minor
axis, 1.4–2.2 vs. 2.0–3.3, respectively). Eggs oftabira bitterlings from Natori City, Miyagi Prefecture were similar in shape to those from Nasumachi, Tochigi Prefecture (Table 6, Fig. 6; Saitoh
et al., 2007). Kuronuma (1940) and Hubbs andKuronuma (1943) reported the shape and size ofeggs of A. t. erythropterus from Chiba Prefecture, but those eggs were still within the ovipositor. Therefore, such data is not compatible here.
Komakine (1976) reported that anal fin edge inmales in populations from Lake Kasumigaura
was red during the spawning season, becoming
pale red at the spawning peak. Ogawa (2001)
suggested Margaritifera laevis as a host mussel
for eggs of A. t. erythropterus from Nasu-machi,
Tochigi Prefecture. Saitoh et al. (2007) reported
that juveniles of A. t. erythropterus from Natori
basin, Miyagi Prefecture lacked a dark blotch on
the dorsal fin.
Acheilognathus tabira tohokuensis subsp. nov.
(New Japanese name: Kitanoakahire-tabira)(New English name: Northern red tabira bitterling)(Figs. 3A, 5 A–B, 6 A–D, 9)
Acheilognathus tabira: Ikeda and Ide, 1937: 27 (Akita);
Okada and Ikeda, 1938: 102 (in part: Akita and Yamagata); Aoyagi, 1957: 100 (in part); Fujita, 1990: 213,
fig. 32 (Akita); Matsuura and Arai, 1993: 62 (in part);
Sugiyama, 2000: 382, 1 fig. (Akita).
Acheilognathus tabira subsp.: Nakamura, 1963: 149 (in14

R. Arai, H. Fujikawa and Y. Nagata

A

B

C

D

Fig. 9. Acheilognathus tabira tohokuensis subsp. nov. A: AKPM-P 435-2, paratype, male, 64.0 mm SL, Gojomemachi, collected and photographed by Hideki Sugiyama, 1 May 2004; B: AKPM-P 437-2, paratype, male,48.2 mm SL, Kisakata, collected and photographed by Hideki Sugiyama, 4 June 2006; C: male, Aizu area,collected and photographed by Osamu Inaba, 13 May 2003; D: ZUMT 61513, paratype, male, 57.2 mm SL,near Teradomari, collected by Nobuo Inoue, 21 Oct. 2005, photographed by Ryu Uchiyama, 8 Nov. 2005
(after artificial induction of nuptial coloration).
part); Sugiyama, 1981: 20 (Akita); Sugiyama, 1985:
76, 2 figs. (Akita); Matsumoto et al., 1988: 73 (Niigata); Shimizu and Matsumoto, 1991: 287, fig. 6-9 (Niigata); Inaba, 2003: 51, fig. 3 (Fukushima).
Acheilognathus tabira subsp. (a): Nakamura, 1969: 42, pl.
92 D–E (in part: Akita, Niigata, and Fukushima).
Acheilognathus tabira subsp. 1: Nagata and Fujikawa,
2000: 120, 1 fig. (in part).
Rhodeus tabira: Sugihara, 1944: 12 (Yamagata).
Acheilognathus moriokae: Ikeda and Ide, 1937: 27(Akita); Honma, 1952: 142 (Niigata); Kataoka, 1959: 8(Akita); Matsumoto et al., 1988: 73 (Niigata).
Acheilognathus cyanostigma: Kataoka, 1959: 8 (Akita).
Holotype. NSMT-P 72657 (ex OKU-P 103-11),
male, 59.0 mm SL, Lake Nishinuma, Yokote, Akita Prefecture, collected on 4 Aug. 1981.
Paratypes. NSMT-P 74701 (ex OKU-P 103) (65: 41
males and 24 females), 29.9–69.7 mm SL, Lake Nishinuma, Yokote, Akita Prefecture, 4 Aug. 1981; NSMT-P
74699 (ex OKU-P 109) (10: 5M5F), 30.0–67.5 mm SL,
Lake Kaikon-numa, Yamamoto-machi, Akita Prefecture,
28 Aug. 1978; NSMT-P 74700 (ex OKU-P 113) (40:25M15F), 42.9–62.9 mm SL, Lake Ogata, Kanaashi,
Akita Prefecture, 5 Aug. 1981; AKPM-P 435 (5:3M2F), 61.8–72.6 mm SL, Gojome-machi, Akita Prefecture, 1 May, 2004; AKPM-P 436 (3 juveniles),11.0–13.0 mm SL, Gojome-machi, Akita Prefecture, 27
June, 2004; AKPM-P 437 (5: 3M2F ), 39.8–53.3 mm

SL, Kisakata, Nikaho, Akita Prefecture, 4 June, 2006;
AKPM-P 438 (2 juveniles), 23.4–26.4 mm SL, Kisakata,
Nikaho, Akita Prefecture, 4 June, 2006; NSMT-P 27553(2), 57.2 and 61.0 mm SL, Yasuda-machi, Aganogawa
River, Niigata Prefecture, 26 Oct. 1974; ZUMT 61513(1M), 57.2 mm SL, near Teradomari, Niigata Prefecture,
21 Oct. 2005.
Non-type specimens. OKU-P 114 (42: 9M33F:
voucher specimens lost after examination but radiographs
still available), 45.8–71.2 mm SL, Lake Hachirogata,
Akita Prefecture, 5 Aug. 1981.

Diagnosis. Dorsal rays, usually iii 9; analrays, usually iii 9; more total vertebrae than in allother subspecies, 37.00.6 (meanSD); dorsal
fin in males, red; anal fin in nuptial males, red;no black blotch on dorsal fin in juveniles; eggs
long ellipsoid, ratio of major axis to minor axis,
2.0–3.3 (Table 11).
Description. Morphometric and meristic
data are shown in Tables 2–4. Paratype and nontype data are given in parentheses. Proportional
measurements as % SL: body depth 32 (29–34),
head length 24 (23–26), predorsal length 53(51–58), caudal peduncle depth 10 (10–13); as% HL: orbit diameter 31 (27–33); as % OD:

Four New Acheilognathus Bitterlings from Japan

15

Table 5. Frequency distributions of insertion of first pterygiophore in dorsal fin (D-PTG-1) in 5 subspecies ofAcheilognathus tabira.
Loc.
No.

Cat. No.

Locality
No. of(Prefecture) specimens

A. t. tohokuensis
1
NSMT-P 74699
Akita
2
OKU-P 114
Akita
3
NSMT-P 74700
Akita
4
NSMT-P 72657, 74701 Akita
5
NSMT-P 27553
Niigata
A. t. erythropterus
7
NSMT-P 27043
Miyagi
9
NSMT-P 72656, 74702 Ibaraki
10
NSMT-P 74703
IbarakiA. t. jordani
11
NSMT-P 74704
Ishikawa
12
NSMT-P 74705
Ishikawa
13
NSMT-P 74706
Ishikawa
15
NSMT-P 74707
Toyama16
NSMT-P 74708
Ishikawa
17
NSMT-P 72658, 74709 Ishikawa
18
NSMT-P 74710
Ishikawa
20
OKU-P 110
TottoriA. t. nakamurae
22
OKU-P 154
Nagasaki
22
OKU-P 171
Nagasaki
22
NSMT-P 11004
Nagasaki
24
NSMT-P 74712
Fukuoka25
NSMT-P 72659, 74713 Fukuoka
26
NSMT-P 74714
Kumamoto
A. t. tabira
28
NSMT-P 74715
Shiga
29
NSMT-P 74716
Shiga
30
NSMT-P 74717
Shiga
31
NSMT-P 74718
Shiga
32
NSMT-P 74719
Osaka
33
NSMT-P 74720
Osaka
34
OKU-P 170
Hyogo
35
NSMT-P 74721
Hyogo
36
NSMT-P 74722
Okayama37
NSMT-P 74723
Okayama38
NSMT-P 74724
Okayama

10tha

10thb

10
42
40
52
2

7
1

2
49
35

1

38
3
41
50
5
7
33
33

1

10
27
2
46
49
13
8
49
50
24
36
38
29
13
33
16
76

snout length 86 (57–106), barbel length 20(2–33). Dorsal rays, iii, 9 (iii, 8–9); anal rays, iii,
9 (iii, 8–9); number of dorsal rays minus numberof anal rays, 0 (1 to 1); pectoral rays, i, 14 (i,14–15); pelvic rays, i, 7 (i, 7); caudal rays,i9/8i (i9/8i); abdominal vertebrae, 19(18–19); caudal vertebrae, 18 (17–19); total vertebrae, 37 (36–38); insertion of first pterygiophore in dorsal fin (D-PTG-1), 12th (10th to12th) (Table 5); insertion of first pterygiophore inanal fin (A-PTG-1), 19th (18th or 19th). Eggs
long ellipsoid, ratio of major axis to minor axis,

11tha

2
15
5
2
2
2
11
5
2
11
9
5
14

1
1
4
3
1
1
5

1
1
5
4
12
5
1
3
12

11thb

12tha

12thb

6
25
9
15
1

3
7
19
26
1

1
3
11
10

21
21

11
9

1

6

3

15
2

27
1
21
34

9
17

5
11
7

2

2
12
2
4
3
5

6
37
33
12
19
22
18
8
27
9
56

1
8
7
7
1
5
8
4
3
3
2

3
1

13tha

1

2

1

3
1
29
35
5

12
11
1

1

2
4
1
3
2
1
1
1
1

2.0–3.3 (Table 6, Figs. 6 A–D).
Color in life. Dorsal fin in males marginedwith red. Lateral body surface in males greenish
in spawning season, similar to A. t. erythropterus
and A. t. jordani; pelvic fin black proximally,
margined with white along anterior edge; anal fin
margined with white, lined proximally by a red
band. Black blotch absent from dorsal fin in juveniles (Figs. 5 A–B).
Habitat. Rivers, lakes and ponds in plains;
creeks and rivers near coasts, and mouths ofrivers draining into lagoons. This subspecies co-

16

R. Arai, H. Fujikawa and Y. Nagata
Table 6.
Subspecies
A. t. tabira
A. t. tabira
A. t. tabira
A. t. erythropterus
A. t. erythropterus
A. t. erythropterus
A. t. erythropterus
A. t. erythropterus
A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. tohokuensis
A. t. jordani
A. t. jordani
A. t. jordani
A. t. jordani
A. t. nakamurae
A. t. nakamurae

Size and shape of eggs in 5 subspecies of Acheilognathus tabira.

Locality

No. ofeggs

Lake Biwa
4
Inabegawa River
3
Kushidagawa River
38
Natori City
28
Nasu-machi (A)79
Nasu-machi (B)6Lake Kasumigaura unknown
Lake Kasumigaura
2
Gojome-machi
21
Akita City
18
Aizu area
5
near Teradomari
7
Himi City
8
Lake Kibagata
19
Ono City
9
Oharagawa River
16
Tataragawa River
19
Yanagawa City
20

Major
axis
(mm)

Minor
Ratio ofaxis major axis to(mm) minor axis

2.4–2.6
1.8–2.1
1.8–2.2
2.3–2.8
2.2–2.9

1.5
1.2–1.3
1.1–1.4
1.3–1.7
1.3–1.7

ca. 2.3
2.1–2.2
3.1–3.5
3.2–3.8

ca. 1.35
1.1–1.3
1.3–1.6
1.1–1.4

2.8–3.0
2.4–2.9

1.0–1.1
1.0–1.3

2.7–2.9
3.3–4.0
2.5–2.7
2.5–2.8

1.1–1.3
1.5–1.6
1.0–1.1
1.1–1.3

1.6–1.7
1.4–1.7
1.5–1.7
1.4–2.0
1.6–2.1
1.8–2.2*
ca. 1.7
1.6–1.9
2.0–2.6
2.5–3.3
2.5–2.8*
2.6–3.0
2.2–2.5
2.4–2.8*
2.2–2.6
2.2–2.6
2.3–2.9
2.2–2.3**

SourcesNakamura 1969
present study
present study
H. Sugiyama, pers. comm.
present study
present study
Nakamura 1969
present study
present study
present study
present study
present study
present study
present study
present study
C. Oshiumi, pers. comm.
present study
Nakamura 1969

* Measurement based on photographs.
** Range of averages.

exists with a bitterling, Tanakia lanceolata,
in large rivers such as the Omonogawa River, andwith a cyprinid, Pseudorasbora pumila pumila,in small ponds or reservoirs, often man-made(so-called “tameike”) (Hideki Sugiyama, pers.
comm.).
Spawning season. From May to July in
Akita Prefecture (Sugiyama, 2000: 382). April
and May in Fukushima Prefecture (Osamu Inaba,
pers. comm.).
Host mussels. Two mussels, Unio douglasiae nipponensis and Anodonta woodiana, havebeen found to function as hosts for eggs of A. t.
tohokuensis in Akita Prefecture (Hideki Sugiyama, pers. comm.). This bitterling subspecies also
spawns eggs into Anemina arcaeformis, Pronodularia japanensis and Inversiunio jokohamensis in the Aizu area, Fukushima Prefecture(Osamu Inaba, pers. comm.).
Etymology. The subspecific name, tohokuensis, refers to the Tohoku area, where this subspecies is distributed.
Distribution. Japan Sea side of eastern Honshu: Akita, Yamagata, Fukushima (western area)
and Niigata Prefectures.

Remarks. Acheilognathus t. tohokuensis
subsp. nov. differs from the other 4 subspecies by
the following combination of characters: anal fin
in nuptial males edged with red, eggs long andellipsoid, and a black blotch absent from the dorsal fin in juveniles. Nakamura (1969, pl. 92 D–E)
reported 2 juveniles (21.5 mm and 36.5 mm intotal length) from Lake Goshikinuma lacking a
black blotch on the dorsal fin. Osamu Inaba(pers. comm.) also noted the absence of a black
blotch on the dorsal fin in juveniles from the
Aizu area, Aganogawa River and Lake Goshikinuma, western Fukushima Prefecture. Acheilognathus t. tohokuensis (127 specimens examined) differed from A. t. erythropterus (85 specimens examined) by the shorter barbel(meanSD and range of barbel length/OD:
18.77.8, 2–39% vs. 35.86.5, 22–56 % in A. t.
erythropterus) and more vertebrae (meanSD
and range: 37.00.6, 36–38 vs. 36.00.6,
35–37) (Tables 2, 4, 8, and 11). Furthermore, theeggs of the former were longer than those of thelatter (Table 6, Fig. 6).

Four New Acheilognathus Bitterlings from Japan

A

B

C

D

17

Fig. 10. Acheilognathus tabira jordani subsp. nov. A: NSMT-P 72658, holotype, male, 45.1 mm SL, Lake Kibagata, collected by Hiroshi Fujikawa, 18 Oct. 1981; B: NSMT-P 72594, paratype, male, 73.9 mm SL, Himi,
collected by Osamu Inamura and Masaki Nishio, 29 May 2005, photographed by Ryu Uchiyama, May 2005;
C: NSMT-P 72595, paratype, male, 53.1 mm SL, Ono, collected by Jun-ya Kitazima, May 2005, photographed by Ryu Uchiyama, 22 May 2005; D: OKU-P 212-1, male, 70.4 mm SL, Oharagawa River, collected and photographed by Yoshikazu Nagata, Mar. 1984.

Acheilognathus tabira jordani subsp. nov.
(New Japanese name: Minamiakahire-tabira)(New English name: Southern red tabira bitterling)(Figs. 4 A–B, 5 C–E, 6 H–K, 10)
Acheilognathus tabira: Aoyagi, 1957: 100 (in part); Matsuura and Arai, 1993: 72 (in part).
Acheilognathus tabira subsp.: Nakamura, 1963: 148 (inpart); Nagata et al., 1981: 48 (Tottori); Yamaguchi,
1999: 12 (Shimane); Kawamura, 2003: 195 (Tottoriand Shimane).
Acheilognathus tabira subsp. (a): Nakamura, 1969: 42 (inpart); Fukuhara et al., 1982: 233, fig. 2C (Tottori); Fujikawa et al., 1984: 54 (in part: Toyama, Ishikawa, andTottori); Saitoh et al., 1988: 59, fig. 2 (Shimane).
Acheilognathus tabira subsp. 1: Tanaka, 1989: 32, fig. 9(Toyama); Watanabe, 1998: 262 (in part); Nagata andFujikawa, 2000: 120, 1 fig. (in part).
Acheilognathus tabira subsp. R: Nagata, 1989: 374 (inpart); Oshiumi, 2003 (Shimane).
Rhodeus tabira: Yanai, 1950: 18 (Shimane); Hirai, 1980:
22 (Ishikawa).
Paracheilognathus tabira: Mori, 1956: 8 (Shimane); Kato,
1985: 81, fig. 33, pl. III-13 (Fukui).
Holotype. NSMT-P 72658 (ex OKU-P 121-5), male,45.1 mm SL, Lake Kibagata, Komatsu, Ishikawa Prefecture, collected on 18 Oct. 1981.

Paratypes. NSMT-P 74707 (ex OKU-P 116) (49: 33
males and 16 females), 39.4–63.3 mm SL, Takaoka, Toyama Prefecture, 7 Aug. 1981; NSMT-P 74705 (ex OKU-P
117) (3: 2M1F), 66.0–70.3 mm SL, Suegawa River,
Kawaradagawa River system, Mii, Wajima, Ishikawa Prefecture, 16 Oct. 1981; NSMT-P 74704 (ex OKU-P 118)
(38: 26M12F), 48.5–63.4 mm SL, Machinogawa River,
Machino, Ishikawa Prefecture, 16 Oct. 1981; NSMT-P
74706 (ex OKU-P 119) (39: 19M20F), 41.5–64.1 mm
SL, Lake Akauragata, Nanao, Ishikawa Prefecture, 16
Oct. 1981; NSMT-P 74708 (ex OKU-P 120) (5: 4M1F),
51.4–58.4 mm SL, Lake Ouchigata, Hakui, Ishikawa Prefecture, 17 Oct. 1981; NSMT-P 74709 (ex OKU-P 121)
(6: 4M2F), 40.8–49.9 mm SL, Lake Kibagata, Komatsu,
Ishikawa Prefecture, 18 Oct. 1981; NSMT-PL 265 (exOKU-P 254) (7 juveniles), 17.5–19.5 mm SL, Lake Kibagata, Komatsu, Ishikawa Prefecture, 17 June, 2005;
NSMT-P 74710 (ex OKU-P 123) (33: 16M17F),
36.7–79.0 mm SL, Lake Shibayamagata, Ishikawa Prefecture, 18 Oct. 1981; NSMT-P 72594 (2: 1M1F), 73.9 and58.5 mm SL, Himi, Toyama Prefecture, 29 May, 2005;
NSMT-P 72595 (2: 1M1F), 53.1 and 50.0 mm SL, Ono,
Fukui Prefecture, 12 June, 2005; ZUMT 61511 (12:7M5F), 32.8–61.1 mm SL, Ohda, Oharagawa River,Shimane Prefecture, 4 July, 2003.
Non-type specimens. OKU-P 110 (34: 28M6F:
voucher specimens lost after examination but radiographs
still available), 42.0–59.5 mm SL, Tanegaike Pond, Tottori, Tottori Prefecture, 27 June 1981; OKU-P 213 (48:

18

R. Arai, H. Fujikawa and Y. Nagata
Table 7. Number of vertebrae (NV) and frequency distributions of insertion of first pterygiophore in dorsal fin in
specimens characterised by 10/9 branched dorsal rays/branched anal rays in 2 subspecies of Acheilognathus
tabira.

Loc. No.

Cat. No.

A. t. jordani
11
NSMT-P 74704
11
NSMT-P 74704
11
NSMT-P 74704
15
NSMT-P 74707
15
NSMT-P 74707
15
NSMT-P 74707
18
NSMT-P 74710
18
NSMT-P 74710
18
NSMT-P 74710
A. t. tabira
28
NSMT-P 74715
28
NSMT-P 74715
29
NSMT-P 74716
29
NSMT-P 74716
29
NSMT-P 74716
31
NSMT-P 74718
31
NSMT-P 74718
31
NSMT-P 74718

NV

No. of
10tha
specimens

38
37
36
38
37
36
38
37
36

1
8
1
3
9
1
2
3
3

37
36
38
37
36
38
37
36

3
5
3
25
11
4
10
6

10thb

11tha

11thb

12tha

12thb

13tha

1
8
1
3
2
1
1
1

21M27F: voucher specimens lost after examination),
42.4–66.7 mm SL, Ohda, Oharagawa River, Shimane Prefecture, 3 Aug. 1984.

Diagnosis. Dorsal rays, iii, 9–10; anal rays,iii, 9; dorsal fin in males, red; anal fin in nuptial
males, red; a black blotch on dorsal fin in juveniles; eggs long ellipsoid, ratio of major axis tominor axis, 2.2–2.8 (Table 11).
Description. Morphometric and meristic
data are shown in Tables 2–4. Paratype and nontype data are given in parentheses. Proportional
measurements as % SL: body depth 30 (29–37),
head length 24 (23–26), predorsal length 51(51–56), caudal peduncle depth 11 (11–13); as% HL: orbit diameter 33 (27–36); as % OD:
snout length 72 (58–108), barbel length 14(9–42). Dorsal rays iii, 9 (iii, 8–10); anal rays iii,
9 (iii, 8–10); number of dorsal rays minus number of anal rays, 0 (1 to 1); pectoral rays, i, 14(i, 14–15); pelvic rays, i, 7 (i, 7); caudal rays,i9/8i (i9/8i); abdominal vertebrae, 18(17–20); caudal vertebrae, 19 (17–20); total vertebrae, 37 (35–38); insertion of first pterygiophore in dorsal fin (D-PTG-1), 11th (10th to

1
1
3

2
4
3
20
8
2
5
3

1
1
1

2
6
1
1

1

1

1
4
1
2
3

1
1

13th) (Table 5); insertion of first pterygiophore inanal fin (A-PTG-1), 18th (18th to 20th). Eggs
long ellipsoid in populations from Himi (Toyama), Lake Kibagata (Ishikawa), Ono (Fukui) andthe Oharagawa River (Shimane), ratio of major
axis to minor axis, 2.2–2.8 (Table 6, Figs. 6
H–K).
Color in life. Dorsal fin in males marginedwith red. Lateral body surface in males more
greenish in spawning season than in A. tabiratabira; pelvic fin black proximally, marginedwith white along anterior edge; anal fin generally
margined with red, varying from red to pale red
in nuptial males. Black blotch on the dorsal fin injuveniles from Lake Kibagata, Ishikawa Prefecture, and the Oharagawa River, Shimane Prefecture (Figs. 5 C–E), but not in adult females.
Habitat. Rivers, lakes and ponds in plains:
creeks near coasts, and mouths of rivers draining
into lagoons.
Spawning season. From April to June.
Host mussels. Unio douglasiae nipponensis
was found to be a host mussel for eggs of thisbitterling subspecies (Nagata et al., 1981).
Etymology. The subspecific name, jordani,Four New Acheilognathus Bitterlings from Japanis named after David Starr Jordan, who made agreat contribution to Japanese ichthyology andwas the first author of the original description of
Acheilognathus tabira.
Distribution. Japan Sea side of western
Honshu: Toyama, Ishikawa, Fukui, Tottori and
Shimane Prefectures.
Remarks. Acheilognathus t. jordani subsp.
nov. differs from the other 4 subspecies by thefollowing combination of characters: anal fin in
nuptial males edged with red, eggs long ellipsoid,a black blotch on the dorsal fin in juveniles, butlacking in adults. Although the whitish-edged
anal fin of a nuptial male from Himi, Toyama
Prefecture, is figured here (Fig. 4A), red-edged
anal fins in males from Himi have also been reported (Tanaka, 1997: 240, fig. 1). Osamu Inamura (pers. comm.) noted that the edge of theanal fin in nuptial males from Himi changed
from red to pale red. This subspecies is clearly
separated from A. t. erythropterus and A. t. tohokuensis by the black blotch presentation on thedorsal fin in juveniles. Saitoh et al. (1988) reported short ellipsoidal eggs (Japanese ‘keiran’ type)of A. t. jordani from the Oharagawa River, but
the eggs examined by them were still ovarian (notfully ripened). Oshiumi (2003) first reported thatripened eggs of bitterling from this area were, infact, long ellipsoid. The number of total vertebrae of populations from the San-in area (locality
numbers 20–21) was fewer than that of populations north of the Hokuriku area (Table 4). As
shown in Table 3, 31 specimens (26% of 118
specimens examined) of A. t. jordani from 3 pop-

A

19

ulations (locality numbers 11, 15 and 18) in the
Hokuriku area had a combination of 10 brancheddorsal rays and 9 branched anal rays, such being
characteristic of A. t. tabira. The combination ofcharacters such as total vertebral number, the DPTG-1 (Table 7) and the whitish-edged anal fin
in nuptial males from Toyama Prefecture, which
is very similar to that of A. t. tabira (Nakamura,
1969: 43), suggests genetic introgression from A.
t. tabira to A. t. jordani in the Hokuriku area (locality numbers 15 and 18) or vice versa. A
boundary between the distributions of A. t. jordani and A. t. tohokuensis corresponds to the
Fossa Magna, considered important for speciation of Japanese freshwater fishes (Watanabe,
1998; Watanabe et al., 2006). Molecular analysesof lactate and malate dehydrogenase isozymes in
Acheilognathus tabira by Fujikawa et al. (1984)
indicated that among the presently-recognizedsubspecies, A. t. jordani is more closely related
to A. t. erythropterus than to A. t. nakamurae.
Acheilognathus tabira nakamurae subsp. nov.
(Japanese name: Seboshi-tabira)(New English name: Blotched tabira bitterling)(Figs. 4D, 5F, 6N, 11)
Acheilognathus tabira: Aoyagi, 1957: 100 (in part).
Acheilognathus tabira subsp.: Nakamura, 1963: 149, fig.
83; Ojima et al., 1973: 172 (Fukuoka); Kimura, 2003:
150.
Acheilognathus tabira subsp. (b): Nakamura, 1969: 45
and 380, pls. 2 C–C, 16, 17, and 93 (Fukuoka); Tomoda, 1970: 203 (Nagasaki); Fukuhara et al., 1982: 233,
fig. 2D (Fukuoka); Fujikawa et al., 1984: 54 (Fukuoka

B

Fig. 11. Acheilognathus tabira nakamurae subsp. nov. A: male, 94 mm total length, Futatsukawa River, collected
and photographed by Ryu Uchiyama, June 1995; B: OKU-P 171-21, male, 59.8 mm SL, Hatahokogawa
River, collected by Hiroshi Fujikawa, 26 May 1982, photographed by Hiroshi Fujikawa, 27 May 1982.

20

R. Arai, H. Fujikawa and Y. Nagata

and Nagasaki); Suzuki and Hibiya, 1985a: 184, fig. 5C(Fukuoka); Suzuki and Hibiya, 1985b: 339, fig. 3B(Fukuoka); Suzuki, 1985: 64, figs. 6–7 (Fukuoka); Araiand Akai, 1988: 201; Okazaki et al., 2001: 92, figs.
3–4 (Fukuoka); Arai and Kato, 2003: 3, table 1(Fukuoka).
Acheilognathus tabira subsp. 2: Sawada, 1984: 55, pl. 53
M–N; Hosoya, 1993: 216; Watanabe, 1998: 262;
Tashima, 1999: 9 (Fukuoka); Imasaka, 1999: 23(Fukuoka); Nagata and Yokoyama, 2000: 122.
Acheilognathus tabira subsp. S: Nagata, 1989: 376, 4 figs.
Paracheilognathus tabira subsp. 2: Hosoya, 1988: 155.
Rhodeus (?) tabira: Azuma et al., 1977: 317 (Nagasaki).
Rhodeus (?) tabira subsp. 2: Kawanabe, 1987: 27, 3 figs.
Holotype. NSMT-P 72659 (ex OKU-P 163-36),
male, 54.0 mm SL, Futatsukawa River, Yanagawa, Fukuoka Prefecture, collected on 15 Apr. 1982.
Paratypes. NSMT-P 74713 (ex OKU-P 163) (48: 30
males and 18 females), 40.3–64.9 mm SL, FutatsukawaRiver, Yanagawa, Fukuoka Prefecture, 16 Apr. 1982;
NSMT-P 75167 (ex OKU-P 138) (11: 3M8F),
47.2–59.7 mm SL, Futatsukawa River, Yanagawa, Fukuoka Prefecture, 3 Sep. 1980; NSMT-P 74712 (ex OKU-P
165) (48: 10M38F), 45.9–74.0 mm SL, Horikawayosui
River, Asakura, Fukuoka Prefecture, 16 Apr. 1982;
NSMT-P 74714 (ex OKU-P 162) (13: 5M8F),
47.8–64.6 mm SL, Lake Shimoezuko, Kumamoto Prefecture, 14–15, Apr. 1982; NSMT-P 74711 (ex OKU-P 253)(57), 50.8–68.8 mm SL, Hatahokogawa River, Iki Islands,
Nagasaki Prefecture, 28 July 1976; NSMT-P 11004 (2),68.8 and 73.5 mm SL, Hatahokogawa River, Iki Islands,
Nagasaki Prefecture, 1 Sep. 1969; NSMT-P 72596 (1M),
76.4 mm SL, Tataragawa River, Fukuoka Prefecture, 23
May 2005.
Non-type specimens. OKU-P 154 (11: 5M5F1
sex unknown: voucher specimens lost after examination
but radiographs still available), 16.1–72.4 mm SL, Hatahokogawa River, Iki Islands, Nagasaki Prefecture, 4 Aug.
1977; OKU-P 171 (30: 8M22F: voucher specimens lost
after examination but radiographs still available),
42.2–73.5 mm SL, Hatahokogawa River, Iki Islands, Nagasaki Prefecture, 26 May 1982.

Diagnosis. Dorsal rays, iii 9; anal rays, iii 9;
dorsal fin in males, red; anal fin in nuptial males,
white; a black blotch on dorsal fin in juveniles
and small adult females; eggs long ellipsoid, ratioof major axis to minor axis, 2.3–2.9 (Table 11).
Description. Morphometric and meristic
data are shown in Tables 2–4. Paratype and nontype data are given in parentheses. Proportional
measurements as % SL: body depth 32 (30–37),

head length 24 (22–25), predorsal length 55(51–57), caudal peduncle depth 12 (11–13); as% HL: orbit diameter 34 (27–36); as % OD:
snout length 74 (63–117), barbel length 37(18–59). Dorsal rays, iii, 9 (iii, 9–10); anal rays,iii, 9 (iii, 8–10); number of dorsal rays minus
number of anal rays, 0 (1 to 1); pectoral rays, i,
14 (i, 14–15); pelvic rays, i, 7 (i, 7); caudal rays,i9/8i (i9/8i); abdominal vertebrae, 19(18–20); caudal vertebrae, 18 (17–19); total vertebrae, 37 (35–38); insertion of first pterygiophore in dorsal fin (D-PTG-1), 12th (11th to13th) (Table 5); insertion of first pterygiophore inanal fin (A-PTG-1), 19th (18th or 19th). Eggs
long ellipsoid, ratio of major axis to minor axis,
2.3–2.9 (Table 6, Fig. 6N).
Color in life. Dorsal fin in males marginedwith red. In spawning males, lateral body surface
brightly greenish-blue; anal fin margined withwhite; pelvic fin black proximally, margined withwhite along anterior edge. A black blotch on thedorsal fin (from which the English name was derived) was observed in an adult female (50.2 mmSL) from Fukuoka Prefecture (Fig. 5F), and in ajuvenile (16.1 mm SL) and 2 small adult females(37.2 and 39.9 mm SL) from the Iki Islands, Nagasaki Prefecture.
Habitat. Rivers and creeks in plains.
Spawning season. From February to August, peaking from April to June in the Futatsukawa River, Yabegawa River system, Yanagawa, Kyushu (Nagata and Nakata, 1988). This
subspecies starts spawning earlier than all othersubspecies of A. tabira.
Host mussels. Four mussel species, Obovalis
omiensis, Pronodularia japanensis, Anodonta
woodiana and Corbicula leana have been reported as hosts for eggs of this subspecies, O. omiensis being the primary host, in spite of its low density (4% of all mussels), in the FutatsukawaRiver, where coexisting C. leana was rarely utilized (Matsushima, 1980). Obovalis omiensis has
also been reported as a primary host in othercreeks. It is interesting that Acheilognathus t.
nakamurae has not been collected at stations
where O. omiensis was not found (Fukuhara et40.1–79.3

28.2–38.0
20.5–27.3
48.8–57.2
9.6–13.3

28–40

40–109
9–60

9–11
8–10
17–20
17–20
34–39

34.31.5
24.11.1
52.51.4
11.80.6

32.22.2

78.911.7
32.18.4

10.10.4
9.10.4
18.20.6
18.20.6
36.21.0

Range

54.08.7

MeanSDtabira323

9.00.2
9.00.3
17.90.4
18.10.535.70.8

88.614.6
35.86.5

31.12.0

31.91.1
24.10.752.81.2
11.40.6

48.13.9

MeanSD

8–10
7–10
17–19
17–19
33–38

38–127
22–56

27–35

29.6–35.8
22.6–26.3
50.2–55.8
10.2–13.0

40.5–62.8

Range

erythropterus
85

84.64184.641
0.429
0.431
0.431
0.418
0.388
0.345

Proportion
Cumulative proportion

Loadings
Body depth
Head length
Predorsal length
Caudal peduncle depth
Orbit diameter
Snout length

PC 1

0.1580.011
0.013
0.123
0.4580.866

8.365
93.007PC 2

0.27899
0.114620.14441
0.55569
0.71983
0.247443.770696.7771PC 3

9.10.4
9.00.3
18.50.5
18.20.6
36.21.085.09.8
24.38.6

30.61.9

31.91.6
23.90.8
53.11.2
11.70.7

51.66.2

MeanSD

Range

8–10
8–10
17–20
17–20
33–39

56–113
8–65

27–36

28.0–37.3
21.4–26.9
49.7–59.1
9.9–14.2

40.1–71.2

jordani249

9.00.2
9.00.3
18.60.5
18.00.6
36.61.2

87.815.5
38.18.4

31.12.1

33.11.5
23.41.0
53.81.4
12.00.6

55.07.9

Loadings
Branched dorsal rays
Branched anal rays
Abdominal vertebrae
Caudal vertebrae
Lateral-line scales

0.392
0.2500.674
0.1290.560

30.80330.803

PC 1

0.525
0.606
0.190
0.302
0.480

25.67756.480

PC 2

8–10
8–10
18–20
17–20
33–40

49–139
18–58

25–37

29.6–37.2
20.8–26.2
50.6–57.3
10.0–13.6

40.3–74.0

Rangenakamurae
144
MeanSD

Table 10. Loadings of the meristics on the first 2
principal components of Acheilognathus tabiratabira (323 specimens), A. t. erythropterus (85),
A. t. tohokuensis (127), A. t. jordani (249) and
A. t. nakamurae (144).

9–10
8–10
18–20
17–19
33–39

53–111
2–39

27–34

28.9–35.6
22.9–26.9
51.2–57.9
8.9–13.0

42.9–71.2

Range

Proportion
Cumulative proportion

9.00.2
9.00.3
18.80.5
18.20.6
36.60.984.011.4
18.77.8

30.21.7

31.91.2
24.60.8
54.01.3
11.20.7

53.76.4

MeanSD

tohokuensis
127

Morphometrics and meristics of specimens 40 mm SL in 5 subspecies of Acheilognathus tabira.

Table 9. Loadings of the log-transformed measurements on the first 3 principal components of
Acheilognathus tabira tabira (323 specimens),
A. t. erythropterus (85), A. t. tohokuensis (127),A. t. jordani (249) and A. t. nakamurae (144) .

SL (mm)Morphometrics% SL
Body depth
Head length
Predorsal length
Caudal peduncle depth% Head length
Orbit diameter% Orbit diameter
Snout length
Barbel length
Meristics
Branched dorsal rays
Branched anal rays
Abdominal vertebrae
Caudal vertebrae
Lateral-line scales

Subspecies
No. of specimens

Table 8.

Four New Acheilognathus Bitterlings from Japan21

22

R. Arai, H. Fujikawa and Y. Nagataal., 1998).
Etymology. The subspecific name, nakamurae, is named after Morizumi Nakamura, who
contributed greatly to the systematics of Japanese
bitterlings.
Distribution. Kyushu area: Fukuoka, Sagaand Kumamoto Prefectures, and Iki Islands in
Nagasaki Prefecture.
Remarks. Acheilognathus t. nakamurae
subsp. nov. differs from the other 4 subspecies by
the following combination of characters: edge ofthe dorsal fin in males red, anal fin in nuptial
males edged with white, and a black blotch on
the dorsal fin in juveniles and small adult females(Nakamura, 1969, pl. 93 B and D). Nakamura(1969: 37) reported that the anal fin in nuptial

males was rarely reddish-white. Nakamura(1969: 46), Suzuki (1985) and Hosoya (1988,
1993) all described eggs of this subspecies as
being usually long ellipsoid, but short ellipsoidal
eggs have been infrequently observed. Nobuhiro
Suzuki (pers. comm.) suggested that such shortellipsoidal eggs may not have been fully ripe.
Populations from the Iki Islands (locality number22) differed from those on the Kyushu mainland(locality numbers 24–26) by having deeper body,fewer total vertebrae and different insertion of
the first pterygiophore in the dorsal fin (D-PTG1) (Tables 2, 4, and 5).

Fig. 12. Statistical analyses of morphometric and meristic characters in 5 subspecies of Acheilognathus tabira.
A: Relationship between log10 standard length (SL) and log10 barbel length (BarL); B: average and standard
deviation of scores on PC2 and PC3 of 6 morphometric characters; C: average and standard deviation ofscores on PC1 and PC2 of 5 meristic characters.

Four New Acheilognathus Bitterlings from Japan

Statistical Analyses of Morphometric andMeristic Characters
Statistical analyses of 7 morphometric and 5
meristic characters were carried out (Table 8).
Although the barbel length tended to be shorterin large specimens of A. t. tohokuensis than in theother 4 subspecies (Fig. 12A), the large variancein barbel length in A. t. tohokuensis and large
overlap among the subspecies (Fig. 12A) precluded the treatment of that character as diagnostic for A. t. tohokuensis.
In a principal component analysis (PCA) on
the 6 remaining log10-transformed morphometriccharacters, the loadings of the variables on thefirst principal component (PC1) were all positive
and of similar magnitude (0.345–0.431) (Table9), indicating that the axis can be interpreted as aproxy for general size. The PC2 and PC3 scores
largely overlapped among the subspecies (Fig.
12B), indicating the absence of diagnostic characters.
Table 11.

23

PCA on the 5 meristic characters showed A. t.
erythropterus to overlap less with A. t. tohokuensis on the PC1 (Fig. 12C). The loadings of abdominal vertebral and lateral-line scale numbers
were high on PC1 (Table 10), indicating thatthose characters were useful for separating A. t.
erythropterus from A. t. tohokuensis. In fact, raw
data of the number of lateral-line scales separated A. t. erythropterus from A. t. tohokuensis(Table 4). The PC2 of meristic characters showed
A. t. tabira to be largely separated from A. t. erythropterus (Fig. 12C). Loadings of branched anal
ray and branched dorsal ray numbers were highon this principal axis (Table 10). However, raw
data for the number of branched dorsal rays separated A. t. tabira from all of the other 4 subspecies (Table 3).
The diagnostic characters and distribution of 5
subspecies of Acheilignathus tabira are detailedin Table 11.

Diagnostic characters (*) and distribution of 5 subspecies of Acheilognathus tabira.

Subspecies
Egg shape (Ratio of major
axis to minor axis)
Black blotch on dorsal finin juveniles
Black blotch on dorsal finin small adult females
Color of dorsal fin in males
Color of anal fin in nuptial
males
Spawning season

tabiranakamurae

erythropterus

tohokuensis

jordanishort ellipsoid(1.4 to 1.7)
absentlong ellipsoid(2.3 to 2.9)
present

middle ellipsoid long ellipsoid(1.4 to 2.2)
(2.0 to 3.3)
absentabsentlong ellipsoid(2.2 to 2.8)
present

absentpresent

absentabsentgrayish

red

red

red

red

white

white

red

red

red

April to June

May to July

April to June

April to August February toAugustBarbel length (BarL/OD)
long
long
(32.18.4%)(38.18.4%)
Branched dorsal rays
10.10.4
9.00.2
Number of dorsal rays minus 0.960.43
0.010.29
number of anal rays
Insertion of 1st pterygiophore 10th to 12th
11th to 13thin dorsal fin
Total vertebrae
more
more
(36.40.7)(36.60.8)
Lateral-line scales
36.21.0
36.61.2
Distribution
Western Honshu Kyushu

absent

long
short
long
(35.86.5%)(18.77.8%)(24.38.6%)
9.00.2
9.00.2
9.10.4
0.010.36
0.010.29
0.160.41
10th to 12th

11th to 13th

fewer
more
(36.00.6)(37.00.6)
35.70.8
36.60.9
Pacific Ocean
Japan Sea side
side of eastern of eastern
HonshuHonshu10th to 13th
more
(36.70.6)
36.21.0
Japan Sea sideof western
Honshu*Proportional measurements of barbel length and meristic characters taken from specimens 40 mm SL.

24

R. Arai, H. Fujikawa and Y. Nagata

Key to 11 Species and Subspecies of Acheilognathus from Japan
1a. Scales in lateral series more than 55 .......................................................A. typus (eastern Honshu)
1b. Scales in lateral series less than 41 .................................................................................................22a. Branched dorsal rays 12–13 .............................................................A. rhombeus (western Japan*)
2b. Branched dorsal rays more than 13.................................................................................................32c. Branched dorsal rays less than 12...................................................................................................43a. Number of dorsal rays minus number of anal rays 3–6 ...................................................................
................................................................................................A. macropterus (Lake Kasumigaura)
3b. Number of dorsal rays minus number of anal rays 0–2 ......A. longipinnis (Kinki and Chubu area)
4a. Branched dorsal rays usually 10; edge of dorsal fin in males grayish; edge of anal fin in males
white during spawning season ....................................................A. tabira tabira (western Honshu)
4b. Branched dorsal rays less than 10...................................................................................................55a. Branched dorsal rays 8 ...................................................................A. cyanostigma (western Japan)
5b. Branched dorsal rays usually 9 .......................................................................................................66a. Branched anal rays 8; edge of dorsal fin in males black ...........A. melanogaster (eastern Honshu)
6b. Branched anal rays usually 9, edge of dorsal fin in males red........................................................7
7a. Edge of anal fin in males white during spawning season; a black blotch on dorsal fin in small
adult females ................................................................A. tabira nakamurae subsp. nov. (Kyushu)
7b. Edge of anal fin in males red during spawning season; no black blotch on dorsal fin in small
adult females.................................................................................................................................. 88a. A black blotch on dorsal fin in juveniles ..........................................................................................
..................................................A. tabira jordani subsp. nov. (Japan Sea side of western Honshu)
8b. No black blotch on dorsal fin in juveniles. .................................................................................... 99a. Ellipsoidal eggs short, 1.4–2.2 in ratio of major axis to minor axis; total vertebrae fewer in number, 36.00.6 (meanSD) ...............................................................................................................
...................................A. tabira erythropterus subsp. nov. (Pacific Ocean side of eastern Honshu)
9b. Ellipsoidal eggs long, 2.0–3.3 in ratio of major axis to minor axis; total vertebrae more numerous, 37.00.6 (meanSD)...............................................................................................................
...........................................A. tabira tohokuensis subsp. nov. (Japan Sea side of eastern Honshu)*Distribution expanded to eastern Honshu by artificial transplantation.

Acknowledgments
We would like to thank Ryu Uchiyama (Shirahama-machi, Wakayama Prefecture), Hideki
Sugiyama (Akita Prefectural Institute of Fisheries, Oga), Osamu Inamura (Uozu City Board of
Education, Uozu), Jyun-ichi Kitamura (Department of Zoology, Kyoto University, Kyoto),
Osamu Inaba (Minamisoma City Museum, Minamisoma), Makoto Kuraishi (Marine Science

Museum, Fukushima Prefecture, Iwaki), Hitoshi
Kubota (Tochigi Prefectural Fisheries Experimental Station, Ohtawara), Kunihiko Yamamoto(Ishikawa Zoo, Komatsu) and Chika Oshiumi(Japan Corbicula Research Institute, Matsue) for
specimens and photographs of adults, juveniles
and/or eggs of tabira bitterlings, and Mikio
Azuma (Shizukuishi-machi, Iwate Prefecture),
Tomiji Hagiwara (Global Environmental Forum,
Tsukuba), Nobuo Inoue (Biodiversity Network

Four New Acheilognathus Bitterlings from Japan

Niigata, Niigata), Jun-ya Kitazima (Gifu University, Gifu), Jun Nakajima (Fisheries ResearchLaboratory, Kyushu University, Fukutsu), Masaki
Nishio (Himi City Board of Education, Himi),
Kenji Saitoh (Tohoku National Fisheries Research Institute, Shiogama), Takayoshi Ueda
(Utsunomiya University, Utsunomiya) and Katsuhide Yamaguchi (Hoshizaki Green Foundation, Hirata) for gifts of tabira bitterlings. We arealso much obliged to M. A. Rogers (FMNH,
Chicago) and Gento Shinohara (NSMT, Tokyo)
for the loan of specimens, Kazumi Hosoya(Kinki University, Nara) for taking radiographs,
Toshihiro Ishinabe (Kannonzaki Nature Museum,
Yokosuka), Nobuhiro Suzuki (Tokai University,
Shimizu), Shiro Matsumoto (Niigata Seiryo Senior High School, Niigata), Osamu Inaba andKenji Saitoh for valuable information, and Toshihiko Tsujino (Wakayama) for his help in preparation of the figures. The principal component
analyses were carried out by Tetsumi Takahashi(Department of Zoology, Kyoto University,
Kyoto), to whom we wish to express our sincere
gratitude.
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Manuscript received 27 January 2006; revised 23 January2007; accepted 29 January 2007.
Associate editor: K. Matsuura.