Rhodeus shitaiensis, a new bitterling from China (Teleostei: Cyprinidae)

Abstract

Rhodeus shitaiensis, new species, is described from the Qiupu River, Anhui Province, China. It is distinguished from other Rhodeus species by the combination of branched dorsal-fin rays 9-10 (usually 9); branched anal-fin rays 9-10 (usually 9); longest simple rays of dorsal and anal fins strong and stiff, distally segmented; longitudinal scale series 35-36; pored scales 6-12; vertebrae 34-36; a transverse row of black spots on dorsal-fin membrane; colour pattern in nuptial males (iris, belly and central pat of caudal fin reddish-orange, lateral scales edged with deep blue).

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Copyright © Verlag Dr. Friedrich Pfeil
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© 2010 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
Rhodeus shitaiensis, a new bitterling from China
(Teleostei: Cyprinidae)
Fan Li* and Ryoichi Arai**
Rhodeus shitaiensis, new species, is described from the Qiupu River, Anhui Province, China. It is distinguished
from other Rhodeus species by the combination of branched dorsal-fin rays 9-10 (usually 9); branched anal-fin rays
9-10 (usually 9); longest simple rays of dorsal and anal fins strong and stiff, distally segmented; longitudinal scale
series 35-36; pored scales 6-12; vertebrae 34-36; a transverse row of black spots on dorsal-fin membrane; colour
pattern in nuptial males (iris, belly and central pat of caudal fin reddish-orange, lateral scales edged with deep
blue).
* Institute of Biodiversity Science, Ministry of Education Key Laboratory for Biodiversity Science and Eco-
logical Engineering, Fudan University, Shanghai 200433, China. E-mail: lfaqua@gmail.com
** 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
Introduction
The cyprinid genus Rhodeus belongs to the sub-
family Acheilognathinae and is separated from
the other 2 genera of the subfamily, Acheilo gnathus
and Tanakia, by having an incomplete lateral line,
no barbels, and wing-like yolk sac projections in
larvae (Arai & Akai, 1988). Rhodeus is distributed
in Eurasia and includes about 17 species/subspe-
cies (Arai et al., 2001; Bogutskaya & Komlev, 2001;
Bohlen et al., 2006; Kottelat & Freyhof, 2007).
Seven species of Rhodeus have been reported
in China: R. sinensis (Günther, 1868; Akai & Arai,
1998, Chen et al., 2005), R. sericeus (Woo, 1964;
Lin, 1998), R. ocellatus (Kner, 1867; Lin, 1935; Woo,
1964; Lin, 1998; Chen & Chang, 2005), R. spinalis
(Arai et al., 1990), R. haradai (Arai et al., 1990),
R. fangi (Miao, 1934; Woo, 1964; Lin, 1998 ; Chen
et al., 2005) and R. notatus (Nichols, 1929; Arai et
al., 2001). We describe here a new species from
the Qiupu river, Anhui Province.
Materials and methods
The classification of the genera of the subfamily
Acheilognathinae follows Arai & Akai (1988).
Methods for counts and measurements follow
Hubbs & Lagler (2004). Last two rays of dorsal
and anal fins are counted as one ray. Vertebrae
and unpaired fin rays of type specimens were
counted from radiographs. Some non-type spec-
imens were dissected for observation of the
pharyngeal teeth, gill rakers and vertebrae. Ver-
tebrae number includes the Weberian complex
(as 4) and the terminal pleurostyle (as 1). As os-
teological approach for the original point of
dorsal and anal fins, the inserted position of the
proximal segment of the first pterygiophore in
the dorsal and anal fins is expressed according
to Arai et al. (1995, 2007). The positions of the first
dorsal- and anal-fin ray pterygiophore (D-PTG-1
and A-PTG-1, respectively) are examined from
radiographs. When proximal radial of D-PTG-1

304 Copyright © Verlag Dr. Friedrich Pfeil
Li & Arai: A new Rhodeus from China
is inserted between neural spines of the vertebral
centra n and (n+1), the position of D-PTG-1 is
expressed as D-PTG-1 = n. When the proximal
radial of A-PTG-1 is inserted between haemal
spines of the vertebral centra m and (m+1), or in
front of the first haemal spine supported by ver-
tebral centrum (m+1), the position of A-PTG-1 is
expressed as A-PTG-1 = m. Measurements of
unfertilized eggs were taken from photographs.
Abbreviations used: AMNH, American Mu-
seum of Natural History; New York; BMNH, The
Natural History Museum, London; NMW, Natur-
historisches Museum, Wien; NSMT-P, National
Museum of Nature and Science, Tokyo (= Na-
tional Science Museum, Tokyo); NTUM, Na-
tional Taiwan University, Taipei; SMU, Sang
Myung University, Seoul; SOU, Shanghai Ocean
University, Shanghai; ZUMT, Department of
Zoology, University Museum, University of To-
kyo, Tokyo.
Fig. 1. Rhodeus shitaiensis, China: Anhui Province: Shitai County: Qiupu River. a, SOU 0811001, holotype, 59.9 mm
SL, male; b, SOU 0905001, paratype, 50.1 mm SL, female.
a
b

305
Ichthyol. Explor. Freshwaters, Vol. 21, No. 4
Copyright © Verlag Dr. Friedrich Pfeil
Rhodeus shitaiensis, new species
(Figs. 1, 6a-b)
Holotype. SOU 0811001, 59.9 mm SL, male; Chi-
na: Anhui Province: Shitai county: Qiupu River,
Changjiang River system, 30°11' N 117°30' E; Y.
Liu, Nov. 2008.
Paratypes. SOU 0811002-0811005, 4 males, 48.4-
53.9 mm SL; same data as holotype. – SOU
0905001-0905006, 6 females, 41.7-54.8 mm SL; SOU
0905007-0905010, 4 males, 42.1-61.5 mm SL; NS-
MT-P 98063, 1 male, 57.5 mm SL; NSMT-P 98064,
1 female, 44.4 mm SL; ZUMT 61947, 4 males,
40.4-56.7 mm SL; same locality as holotype; F. Li,
3 May 2009.
Non-type specimens. SOU 0811006-0811012, 7 males,
45.7-56.2 mm SL; same data as holotype. – SOU 0905011-
0905013, 3 males, 40.5-44.4 mm SL; SOU 0905014-
0905015, 2 females, 47.2-52.7 mm SL; same locality as
holotype, F. Li, 3 May 2009.
Diagnosis. Rhodeus shitaiensis is distinguished
from all other Rhodeus species by the following
combination of characters: branched dorsal fin
rays 9-10 (usually 9); branched anal fin rays 9-10
(usually 9); longitudinal scale series 35-36; pored
scales 6-12; vertebrae 34-36 (usually 34); a trans-
verse row of black spots on dorsal fin membrane;
in nuptial males, iris, belly and central part of
caudal fin reddish-orange, dorsal and anal fins
margined with black, lined proximally by a
Fig. 2. Rhodeus shitaiensis, SOU 0811001, holotype, male.
Simple and first branched rays of dorsal (a) and anal
(b) fins.
a
b
Fig. 3. Rhodeus shitaiensis, SOU 0811001, holotype, 59.9 mm SL, male. Radiograph.

306 Copyright © Verlag Dr. Friedrich Pfeil
0.5 mm
2 mm
Fig. 4. Rhodeus shitaiensis, SOU 0811107. Pharyngeal
teeth.
Fig. 5. Rhodeus shitaiensis, eggs.
Table 1. Morphometric and meristic measurements of holotype and selected paratypes in Rhodeus shitaiensis.
Numbers in parentheses are number of specimens with a given count.
holotype paratypes
male males
n = 13 females
n = 7
Morphometrics
Standard length (mm) 59.9 42.1-61.5 41.7-54.8
In percent of standard length
Head length 22.3 22.1-23.8 21.9-23.1
Body depth 35.7 36.3-41.3 35.9-37.9
Snout length 5.8 5.7-6.4 5.4-5.8
Orbit diameter 7.8 7.6-8.1 7.9-8.8
Predorsal length 47.4 46.3-49.5 47.4-51.0
Caudal peduncle length 22.7 21.3-22.8 19.9-22.7
Caudal peduncle depth 12.8 12.1-13.9 11.9-13.3
Meristics
Branched dorsal-fin rays 9 9 (13) 9 (6), 10 (1)
Branched anal-fin rays 9 9 (13) 9 (6), 10 (1)
Number of anal-fin rays minus number of dorsal-fin rays 0 0 -1 to 1
Pectoral-fin rays i 13 i 12-13 i 12-13
Pelvic-fin rays i 7 i 7 i 7
Vertebrae 35 34 (10), 35 (2), 36 (1) 34 (6), 35 (1)
Insertion of 1st dorsal-fin pterygiophore above vertebral centrum 10 10 (6), 11 (7) 10 (5), 11 (2)
Insertion of 1st anal-fin pterygiophore below vertebral centrum 16 16 (5), 17 (6), 18 (2) 16 (4), 17 (3)
Scales in lateral series 35 35-36 35-36
Pored lateral line scales 9 6-12 8-11
Scales around caudal peduncle 14 14 (19) 14 (7)
Predorsal scales 14 13-15 13-15
reddish-orange band, lateral scales edged with
deep blue. Eggs spindle-shaped, ratio of major
axis to minor axis, 2.1-2.3.
Description. Morphometric and meristic data of
holotype and paratypes are shown in Table 1.
Body compressed. Mouth sub-inferior. Barbels
absent. Dorsal fin with 3 simple and 9-10 branched
rays, usually 9. Anal fin with 3 simple and 9-10
branched rays, usually 9. Pectoral fin with 1 sim-
ple ray and 12 (12-13) branched rays. Pelvic fin
with 1 simple ray and 7 branched rays. Caudal
fin with 1 simple, 9 + 8 branched, 1 simple rays.
First simple ray in dorsal and anal fins very small,
hidden under skin. Longest simple ray of dorsal
Li & Arai: A new Rhodeus from China

307
Ichthyol. Explor. Freshwaters, Vol. 21, No. 4
Copyright © Verlag Dr. Friedrich Pfeil
Fig. 6. Four Rhodeus species from Changjiang River system. a-b, R. shitaiensis, male (a) and (b) female, Anhui
Province: Shitai County; c-d, R. sinensis, male (c) and (d) female, Shanghai City; e-f, R. ocellatus, male (e) and (f)
female, Jiangsu Province: Zhenjiang City; g-h, R. fangi, male (g) and (h) female, Jiangsu Province: Chinkiang
(= Zhenjiang) City.
Fig. 7. Rhodeus shitaiensis, juvenile, 21 mm SL; Anhui
Province: Shitai County.
fin strong and stiff, distally segmented: seg-
mented from a point corresponding to third
branching points of first branched ray (Fig. 2).
Width of basal part of longest simple ray of dor-
sal fin much wider than that of first branched ray.
Lateral line incomplete: pored scales 6-12. Lon-
gitudinal scale series 35-36 (33-34 on body + 2 on
caudal fin). Transverse scales 10. Predorsal scales
a
e
g
b
h
c
cd
d
f
f
gh

308 Copyright © Verlag Dr. Friedrich Pfeil
13-15. Scale rows around caudal peduncle 14.
Abdominal vertebrae 16-18; caudal vertebrae
17-19; total vertebrae 34-36. Inserted position of
first pterygiophore of dorsal fin 10th to 11th.
Inserted position of first pterygiophore of anal
fin 16th to 18th (Fig. 3). Pharyngeal teeth in one
row, 0.0.5-5.0.0 (Fig. 4), occlusal grooves rela-
tively reduced. Gill rakers on external side of first
gill arch 11-12. Pearl organs on snout and area
between nostril and eye in adult males, absent in
females (Fig. 6a-b). Eggs spindle-shaped, ratio of
major axis to minor axis, 2.1-2.3 (Fig. 5).
Coloration in life. Body gray. Blue longitudinal
stripe on each side of body running from below
dorsal fin to end of caudal peduncle, broader in
males than in females. Dorsal and anal fins gray
with two pale stripes; a transverse row of black
spots on dorsal-fin membrane. Nuptial coloration
in adult males: iris, belly, end of caudal peduncle,
and central part of caudal fin reddish-orange;
reddish-orange band on caudal fin extending to
tip of caudal-fin rays or not. Dorsal and anal fins
margined with black, lined proximally by a
reddish-orange band; two additional whitish
bands between a reddish-orange band and base
of fin. Posterior portion of scales on flank deep
blue, which may reflect color of scale pocket cor-
responding to posterior portion of scale (Fig. 6a).
Adult females grayish, dorsal and anal fins lack-
ing colored band. No black blotch on anterior part
of dorsal fin. Scales on flank grayish (Fig. 6b).
Pelvic fin hyaline in females, in males usually
turning to pale reddish-orange and margined
with black in spawning season. Caudal fin in
females hyaline. A large black blotch on anteri-
ormost part of dorsal fin in juveniles (Fig. 7).
Colour in preservative. Ground colour of body
yellowish. A dark longitudinal stripe on flank,
broader in males than in females. Posterior part
and margin of scales on flank grayish, darker in
males than in females. Dorsal and anal fins gray-
ish with two pale stripes, margined with black in
males, indistinctly margined with black in fe-
males.
Distribution and ecology. Rhodeus shitaiensis is
known only from Qiupu River, a tributary of
Changjiang River (Yangtze) in Shitai County,
Table 2. Comparison of Rhodeus species from China. * from Arai et al. (2001), ** from T. Ishinabe (pers.
comm.).
R. shitaiensis R. sericeus*R. sinensis
n = 21 n = 43 n = 28
Branched dorsal-fin rays 9-10 (usually 9) 8-10 (usually 9) 9-11
Branched anal-fin rays 9-10 (usually 9) 8-10 (usually 9) 9-12
Number of anal-fin rays minus number of dorsal-fin rays -1 to 1 -1 to 1 0 to 2
Longitudinal scale series 35-36 36-40 32-35
Predorsal scales 13-15 13-15 13-16
Pored scales 6-12 4-7 0-6
Gill rakers 11-12 10-14 7-8
Vertebrae 34-36 36-37 32-34
Insertion of 1st dorsal-fin pterygiophore above vertebral centrum 10-11 10-12 9-11
Insertion of 1st anal-fin pterygiophore below vertebral centrum 16-18 17-19 15-16
Body depth (% SL) 35-41 32-40 38-50
Longest simple dorsal fin ray intermediate soft soft
Male nuptial color
Iris reddish-orange red reddish-orange
Lateral band of caudal fin reddish-orange absent reddish-orange
Belly reddish-orange red yellow
Color in formalin
Dark spot behind gill opening absent absent present
Vertical blotches on 4th-5th scales in lateral series vague present present
Dorsal fin membrane blackish blackish clear
Diploid chromosome number unknown 48 48
Li & Arai: A new Rhodeus from China

309
Ichthyol. Explor. Freshwaters, Vol. 21, No. 4
Copyright © Verlag Dr. Friedrich Pfeil
Anhui Province (Figs. 8-9). It was mainly col-
lected in riffle areas, while R. o. ocellatus and
R. sinensis were collected on calm water areas in
Qiupu River. Spawning season is from April to
June. Rhodeus shitaiensis usually spawns several
times, about 10 eggs, as inferred from number of
ripe eggs obtained by pressing the belly of mature
females in one spawning. Host mussels are not
known.
Etymology. The species is named after Shitai, its
type locality. An adjective.
Discussion
Rhodeus shitaiensis closely resembles the R. sericeus
complex from northeastern Asia and Europe in
sharing the number of branched dorsal and anal
fin rays (D. 9-10, A. 9-10 vs. D. 8-10, A. 8-10) and
a transverse row of black spots on the membrane
of the dorsal fin. The R. sericeus complex includes
4 species, i. e., R. sericeus from northeastern Asia,
R. colchicus from west Transcaucasia (Bogutskaya
& Komlev, 2001), R. meridionalis from the southern
Balkan Peninsula in Europe, and R. amarus from
the rest of Europe (Bohlen et al., 2006). Rhodeus
colchicus was described on the basis of morpho-
logical evidence (Bogutskaya & Komlev, 2001),
while R. meridionalis was separated from R. ama-
rus mainly on molecular characters (Kottelat &
Freyhof, 2007). Rhodeus shitaiensis is distinguished
from R. colchicus in having fewer vertebrae (34-36,
mode 34 vs. 33-36, mode 35) and more pored
lateral line scales (6-12 vs. 3-7), and from R. sericeus,
R. amarus and R. meridionalis by the presence of a
red longitudinal band on caudal fin (vs. absent),
fewer longitudinal scale series (35-36 vs. 36-40),
fewer vertebrae (34-36 vs. 36-37) (Arai et al., 2001;
Bogutskaya & Komlev, 2001; Kottelat & Freyhof,
2007).
Rhodeus shitaiensis is distinguished from oth-
er Rhodeus species as follows: from R. pseudo-
sericeus in having fewer branched anal fin rays
(9-10, mode 9 vs. 9-11, mode 10) and more pored
lateral line scales (6-12 vs. 2-7), a red longitudinal
band on caudal fin (vs. absent) (Arai et al., 2001);
from R. sinensis by the absence of a spot behind
gill opening (vs. presence) and more gill rakers
(11-12 vs. 7-8) (Akai & Arai, 1998); from both
R. o. ocellatus and R. o. kurumeus by the iris color
of males (reddish-orange vs. red), more longitu-
dinal scale series (35-36, mode 36 vs. 32-34, mode
about 32) and fewer branched anal fin rays (9-10
vs. 10-12) (Arai et al., 2001; Hosoya, 2002; Chen
& Chang, 2005); from the R. smithii complex
(R. fangi, R. notatus, R. smithii, R. atremius and
R. suigensis) in having more gill rakers (11-12 vs.
4-8) and a shorter longitudinal stripe on flank
(not extending beyond the origin of the dorsal fin
vs. extending beyond) (Arai et al., 2001); from
R. haradai, R. spinalis, R. laoensis, R. rheinardti and
R. kyphus in having fewer branched dorsal-fin
rays (9-10, usually 9 vs. 10-14) and branched anal
fin rays (9-10, usually 9 vs. 11-17); and from
R. elongatus in having more branched dorsal fin
rays (9-10 vs. 8) (Mai, 1978, Arai et al., 1990; Kotte-
lat, 1998; Kottelat, 2001). Four Chinese species of
Rhodeus (R. maculatus, R. hwanghoensis, R. wang-
kinfui, R. pingi) are currently synonymized with
R. ocellatus. Rhodeus shitaiensis differs from R. ma-
culatus, R. wangkinfui and R. pingi by having
fewer branched dorsal fin rays (9-10, usually 9
vs. 10-11) (Fowler, 1910; Wu, 1930; Miao, 1934).
Rhodeus hwanghoensis shares numbers of branched
dorsal and anal fin rays with R. shitaiensis. How-
ever, it differs from R. shitaiensis by having fewer
longitudinal scale series (32 vs 35-36), a small dark
spot above the gill opening, and a yellowish color
in formalin. Rhodeus hwanghoensis seems to be
more similar to a northern type of R. sinensis
R. ocellatus R. haradai*R. spinalis*R. fangi R. notatus
n = 37 n = 11 n = 39 n = 19 n = 4
10-12 12-13 10-13 9-12 9
10-12 12-14 13-17 10-11 9
-1 to 1 0 to 2 2 to 4 -1 to 1 0
33-35 33-34 34-36 33-34 33-34
14-17 13-14 14-15 13-15 14-16
3-7 5-7 5-8 4-6 4-5
10-13 14-15 8-12 7-8 7
33-34 33-34 33-35 32-33 32
10 9-10 10-11 10 9-10
17 15-16 16-17 15-16 16
39-49 48-56 41-51 35-40 32-38
soft hard intermediate hard hard
red unknown red** red blackish
red unknown red** blackish
or absent blackish
red unknown red** pale pale
absent absent absent present absent
present unknown unknown present absent
clear clear clear clear clear
48 unknown unknown 46 46

310 Copyright © Verlag Dr. Friedrich Pfeil
(= R. lighti) than to R. shitaiensis (Mori, 1928, Akai
& Arai, 1998).
Lin (1998) reviewed the classification of the
Chinese bitterlings and considered R. lighti as a
valid species and synonymized R. sinensis and
R. notatus with R. ocellatus. In fact, R. sinensis is
not a junior synonym of R. ocellatus, but a valid
species and senior synonym of R. lighti (Akai &
Arai, 1998). Rhodeus notatus is not a junior syno-
nym of R. ocellatus but a valid species distin-
guished by the length of the longitudinal stripe
on flank (extending beyond the origin of dorsal
fin vs. not extending beyond) and the diploid
chromosome number (2n = 46 in notatus vs. 2n = 48
in ocellatus) (Hong et al., 1983; Lee et al., 1982).
Rhodeus fangi, R. notatus, and R. atremius belong
to the R. smithii complex (Nichols, 1929; Miao,
1934; Kimura & Nagata, 1992; Okazaki et al. 2001).
This contradicts Lin (1998) who considered
R. fangi as a valid species and synonymized
R. notatus and R. atremius with R. ocellatus. Rho-
deus notatus might be a valid species or a subspe-
cies of R. fangi or R. smithii. The characters distin-
guished the eight Rhodeus species recorded in
China are summarized in Table 2.
Comparative materials. Rhodeus fangi: SOU 200905501,
9 males and 4 females, 31.4-34.5 mm SL; China: Jiangsu
Province: Chinkiang (Zhenjiang) City; May 2009. – SOU
200706501, 1 male, 38.5 mm SL; China: Zhejiang Prov-
ince: Shengzhou City; 9 Jun. 2007. – SOU 201005504, 3
males and 2 females, 30.6-45.0 mm SL; China: Anhui
Province: Xiuning County; 4 May 2010.
120°E
120°E
112°E 114°E 116°E 118°E 120°E 122°E
Changjiang
basin
Qiantangjiang
basin
China
34°N –
32°N –
30°N –
28°N –
100 km
@
Fig. 8. Sampling locality of Rhodeus shitaiensis (@).
Rhodeus haradai: NTUM 7600, holotype, male,
56.0 mm SL; China: Hainan: Longtang; June 1942.
Rhodeus notatus: AMNH 9654, holotype, male,
33.0 mm SL (x-ray); AMNH 10812, paratypes, 5 males
and 1 female, 24.3-29.2 mm SL; China: Shandong Prov-
ince: Tsinan (Jinan); April-July 1924. – SOU 200702501,
2 males and 2 females, 42.6-46.0 mm SL; China: Beijing
City; Feb. 2007.
Rhodeus ocellatus: NMW 10837, holotype, male,
51.6 mm SL (x-ray); China: Shanghai City. – SOU
200705501, 10 males and 7 females, 41.8-57.9 mm SL;
China: Shanghai City; May 2007. – SOU 200706501, 3
males and 5 females , 36.8-56.3 mm SL; China: Zhejiang
Province: Shengzhou City; 9 Jun. 2007. – SOU 200811501,
5 males and 4 females, 44.7-55.6 mm SL; China: Anhui
Province: Shitai County; Nov. 2008. – SOU 201005504,
1 male and 2 females, 35.8-48.8 mm SL; China: Anhui
Province: Xiuning County; 4 May 2010.
Rhodeus pseudosericeus: SMU 211, holotype, male,
45.5 mm SL; Korea: Gangwon-do: Hoengsong-gun:
Gonggun-myon: Hakdam-ri: Gumgye River, tributary
of Som River, Namhan River system; S. R. Jeon, 16 Oct.
1999. – ZUMT 61149-61151, 3 paratypes, 41.0-41.9 mm
SL, 1 male and 2 females; same data as holotype.
Rhodeus sericeus: ZUMT 61153, 7 males and 5 fe-
males, 46.1-60.7 mm SL; Russia: Ussuri: Jablonovka; 17
June 1997.
Rhodeus sinensis: BMNH 1868.10.19-150, lectotype,
male, 51.6 mm SL; China: Chikiang (Zhejiang); 19 Oct.
1868. – SOU 201005504, 4 males and 1 female, 35.2-
58.2 mm SL; China: Anhui Province: Xiuning County;
4 May 2010. – SOU 200705501, 10 males and 4 females,
43.6-58.5 mm SL; China: Shanghai City; May 2007 – SOU
200905501, 6 males and 3 females, 32.1-44.7 mm SL;
China: Jiangsu Province: Zhenjiang City; May 2009.
Rhodeus spinalis: NSMT-P 31906, neotype, male,
49.7 mm SL; Hainan: Dingan; 27 March 1966.
Li & Arai: A new Rhodeus from China

311
Ichthyol. Explor. Freshwaters, Vol. 21, No. 4
Copyright © Verlag Dr. Friedrich Pfeil
Fig. 9. Habitat of Rhodeus shitaiensis, China: Anhui Province: Shitai County.
Acknowledgements
We thank James Chambers (BMNH), Norma Feinberg
(AMNH), B. Herzig (NMW) and Shih Chieh Shen
(NTUM) for the loan of type specimens or photographs
of types of Rhodeus species; Y. Liu (Zhenjiang), H. L.
Wu (SOU) and J. S. Zhong (SOU) for collecting speci-
mens; and T. Ishinabe (Kannonzasi Nature Museum,
Japan) for information on the color pattern of Rhodeus
spinalis. The first author is also grateful to J. K. Chen,
C. Z. Fu, and J. G. Xiangyu (Fudan University, Shang-
hai) for providing valuable comments on the manu-
script, T. J. Chu (Fudan University) and M. Q. Wu (SOU)
for assistance in photographing, and Y. Q. Chang (Tai-
wan) and H. Zhou (Shenzhen) for sending literatures.
Literature cited
Akai, Y. & R. Arai. 1998. Rhodeus sinensis, a senior
synonym of R. lighti and R. uyekii (Acheilognathinae,
Cyprinidae). Ichthyological Research, 45: 105-110.
Arai, R. & Y. Akai. 1988. Acheilognathus melanogaster, a
senior synonym of A. moriokae, with a revision of
the genera of the subfamily Acheilognathinae (Cy-
priniformes, Cyprinidae). Bulletin of the National
Science Museum, Tokyo, Series A, 14: 199-213.
Arai, R., H. Fujikawa & Y. Nagata. 2007. Four new
subspecies of Acheilognathus bitterling (Cyprinidae:
Acheilognathinae) from Japan. Bulletin of the Na-
tional Museum of Nature and Science, Tokyo, Series
A. Supplement, 1: 1-28.
Arai, R., S. R. Jeon & T. Ueda. 2001. Rhodeus pseudosericeus
sp. nov., a new bitterling fom South Korea (Cyprini-
dae, Acheilognathinae). Ichthyological Research,
48: 275-282.
Arai, R., N. Suzuki & S. C. Shen. 1990. Rhodeus haradai,
a new bitterling from Hainan Island, China, with
notes on the synonymy of Rhodeus spinalis (Pisces,
Cyprinidae). Bulletin of the National Science Mu-
seum, Tokyo, Series A, 16: 141-154.
Arai, R., Y. Xie & Y. Akai. 1995. Rediscovery of the bit-
terling, Tanakia lanceolata, in China (Pisces, Cyprini-
dae). Japanese Journal of Ichthyology, 42: 196-199.
Bogutskaya, N. G. & A. M. Komlev. 2001. Some new
data to morphology of Rhodeus sericeus (Cyprinidae:
Acheilognathinae) and a description of a new spe-
cies, Rhodeus colchicus, from West Transcaucasia.
Proceedings of the Zoological Institute, St. Peters-
burg, 287: 81-97.
Bohlen, J., V. Šlechtová, N. Bogutskaya & J. Freyhof.
2006. Across Siberia and over Europe: phyloge-
netic relationships of the freshwater fish genus
Rhodeus in Europe and the phylogenetic position of
R. sericeus from the River Amur. Molecular Phylo-
genetics and Evolution, 40: 856-865.
Chen, I. S. & Y. C. Chang. 2005. A photographic guide
to the inland-water fishes of Taiwan. Vol. 1. Cyprini-
formes. Sueichan Press, Taiwan, xx + 280 pp.

312 Copyright © Verlag Dr. Friedrich Pfeil
Chen, X. H., Y. Ni & H. L. Wu. 2005. Fishes of genus
Rhodeus Agassiz in Jiangsu Province, China. Marine
Fisheries, China, 27: 89-97 [in Chinese, English
abstract].
Fowler, H. W. 1910. Description of four new cyprinoids
(Rhodeinae). Proceedings of the Academy of Natu-
ral Sciences of Philadelphia, 62: 476-486.
Günther A. 1868. Catalogue of the fishes in the British
Museum. VII. British Museum, London. xx + 512 pp.
Hong, Y. H., M. Zhou & T. Zhou. 1983. Studies on the
karyotypes of Chinese cyprinid fishes. III. Com-
parative analysis of the chromosomes of seven
species of acheilognathid fishes. Journal of Wuhan
University (Natural Science), 1983 (2): 96-102, 116
[in Chinese, English abstract].
Hosoya, K. 2002. Acheilognathinae. Pp. 255-260 in:
T. Nakabo (ed.), Fishes of Japan with pictorial keys
to the species, English edition. Tokai University
Press, Tokyo.
Hubbs, C. L. & K. F. Lagler. 2004. Fishes of the Great
Lakes region. Revised edition. University of Mich-
igan Press, Bloomfield Hills, xxxii + 276 pp.
Kimura, S. & Y. Nagata. 1992. Scientific name of Nippon-
baratanago, a Japanese bitterling of the genus
Rhodeus. Japanese Journal of Ichthyology, 38: 425-
429.
Kner, R. 1867. Fische. Reise der Österreichischen Fre-
gatte “Novara” um die Erde in den Jahren 1857,
1858, 1859, unter den Befehlen des Commodore B.
von Wüllerstorf-Urbair. Zoologischer Theil, Abtei-
lung 3. Staatsdruckerei, Wien, pp. 275-433, pls. 12-
16.
Kottelat, M. 1998. Fishes of the Nam Theun and Xe
Bangfai basins, Laos, with diagnoses of twenty-two
new species (Teleostei: Cyprinidae, Balitoridae,
Cobitidae, Coiidae and Odontobutidae). Ichthyo-
logical Exploration of Freshwaters, 9: 1-128.
— 2001. Fishes of Laos. WHT Publications, Colombo,
Sri Lanka, 198 pp.
Kottelat, M. & J. Freyhof. 2007. Handbook of European
freshwater fishes. Kottelat, Cornol & Freyhof, Ber-
lin, xiii + 646 pp.
Lee, G. Y., J. N. So & S. Z. Kim. 1982. Chromosomes
and arm number of the acheilognathine fishes
(Cyprinidae) in Korea. Basic Science Review, Insti-
tute of Basic Science, Jeonbug National University,
5: 61-69 [in Korean, English abstract].
Lin, R. D. 1998. Acheilognathinae. Pp. 413-454, 504-506
in: Y. Y. Chen (ed.), Fauna Sinica, Osteichthyes,
Cypriniformes II. Science Press, Beijing [in Chinese,
English abstract].
Lin, S. Y. 1935. Contribution to a study of Cyprinidae
of Kwangtung and adjacent provinces. Lingnan
Science Journal, 14: 249-260.
Mai, D. Y. 1978. Identification of the freshwater fishes
of the North of Vietnam. Science & Technics Pub-
lishing House, Ha Noi, 339 pp. [in Vietnamese].
Miao, C. P. 1934. Notes on the fresh-water fishes of the
southern part of Kiangsu I., Chinkiang. Contribu-
tions from the Biological Laboratory of the Science
Society of China, Zoological Series, 10: 111-244.
Mori, T. 1928. Fresh water fishes from Tsi-nan, China,
with descriptions of five new species. Japanese
Journal of Zoology, 2: 61-72, pl. 2.
Nichols, J. T. 1929. Some Chinese fresh-water fishes.
American Museum Novitates, 377: 1-11.
Okazaki, M., K. Naruse, A. Shima & R. Arai. 2001.
Phylogenetic relationships of bitterlings based on
mitochondrial 12S ribosomal DNA sequences.
Journal of Fish Biology, 58: 89-106.
Woo, Q. J. 1964. Acheilognathinae. Pp. 199-211, 64-67
in: H. W. Wu (ed.), Cyprinid fishes of China. Volume
1. Shanghai Science Technology Press, Shanghai,
[in Chinese].
Wu, H. W. 1930. On some fishes collected from the
upper Yangtse valley. Sinensia, 1: 65-86.
— 1931. Notes on the fishes from the coast of Foochow
region and Ming River. Contributions from the
Biological Laboratory of the Science Society of
China, Zoological Series, 7: 1-64.
Received 14 July 2010
Revised 30 November 2010
Accepted 7 January 2011
Li & Arai: A new Rhodeus from China

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INSTRUCTIONS TO CONTRIBUTORS
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology

Articles appearing in this journal are indexed in:
AQUATIC SCIENCES and FISHERIES ABSTRACTS
BIOLIS - BIOLOGISCHE LITERATUR INFORMATION SENCKENBERG
CAMBRIDGE SCIENTIFIC ABSTRACTS
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C O N T E N T S
Costa, Wilson J. E. M. and Andre C. De Luca: Rivulus schuncki, a new species of the killi -
fish subgenus Melanorivulus, from eastern Brazilian Amazon (Cyprinodontiformes:
Rivulidae) ..................................................................................................................................... 289
Bohlen, Jörg and Radovan Harant: Microcobitis, a new genus name for Cobitis misgurnoides
(Teleostei: Cobitidae) .................................................................................................................. 295
Costa, Wilson J. E. M.: Ophthalmolebias ilheusensis, new combination, with additional notes
on colour pattern of live males (Cyprinodontiformes: Rivulidae) ..................................... 301
Li, Fan and Ryoichi Arai: Rhodeus shitaiensis, a new bitterling from China (Teleostei:
Cyprinidae) .................................................................................................................................. 303
Kottelat, Maurice and Helmut Steiner: Bangana musaei, a new cave fish from central Laos
(Teleostei: Cyprinidae) ............................................................................................................... 313
Anganthoibi, Nongmaithem and Waikhom Vishwanath: Two new species of Glyptothorax
from the Koladyne basin, Mizoram, India (Teleostei: Sisoridae) ........................................ 323
Delling, Bo: Diversity of western and southern Balkan trouts, with the description of a
new species from the Louros River, Greece (Teleostei: Salmonidae) ................................. 331
Park, Jong-Young and Su-Hwan Kim: Liobagrus somjinensis, a new species of torrent catfish
(Siluriformes: Amblycipitidae) from Korea ............................................................................ 345
Costa, Wilson J. E. M. and Andre C. De Luca: Rivulus cajariensis, a new killifish from the
Guiana Shield of Brazil, eastern Amazon (Cyprinodontiformes: Rivulidae).................... 353
Vreven, Emmanuel and Armel Ibala Zamba: Synodontis carineae, a new species of mochokid
catfish from the Kouilou-Niari River basin, Africa (Siluriformes: Mochokidae) ............. 359
Bragança, Pedro H. N. and Wilson J. E. M. Costa: Poecilia sarrafae, a new poeciliid from
the Parnaíba and Mearim river basins, northeastern Brazil (Cyprinodontiformes:
Cyprinodontoidei) ...................................................................................................................... 369
Radhakrishnan, K. V., S. Sureshkumar and Heok Hee Ng: Pseudolaguvia austrina, a new
species of sisorid catfish (Osteichthyes: Siluriformes) from Peninsular India .................. 377
Kottelat, Maurice: Claea, a new replacement name for Oreias Sauvage, 1874 (Teleostei:
Nemacheilidae) ........................................................................................................................... 384
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 21 • Number 4 •December 2010
Cover photograph:
Microcobitis misgurnoides (photograph by Jörg Bohlen)
Jörg Bohlen and Radovan Harant
(this volume pp. 295-300)