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A review of the genus Distoechodon (Teleostei: Cyprinidae),
and description of a new species
Yahui Zhao & Fang Kullander & Sven O Kullander &
Chunguang Zhang
Received: 18 September 2007 /Accepted: 22 September 2008 / Published online: 21 October 2008
#
Springer Science + Business Media B.V. 2008
Abstract Distoechodon species are widely distribut-
ed in rivers of southern China. Previous studies re-
cognized only two species in the genus, viz., D.
tum irostris and D. hupeinensis. A morphological
analysis of 164 specimens from different localities
permits recognition of four species. Distoechodon
tumirostris is mainly restricted to small coastal rivers
in the Zhejiang Province. Distoechodon multispinnis
is widely distributed in the Chang Jiang and Zhu
Jiang basins. Distoechodon compressus is known only
from the Fujian and Jiangxi Provinces, and Taiwan.
Distoechodon macrophthalmus, the new species, is
known only from the Chenghai Lake, Chang Jiang
basin in Yunnan Province.
Keywords China
.
Jiang river
.
Zhu Jiang river
.
Distoechodon
.
Systematics
Introduction
The endemic Chinese fish genu s Distoechodon
(Cypriniformes: Cyprinidae) was established by Peters
(1881) with the type species, D. tumirostris, type lo-
cality Ningpo (now known as Ningbo), located on the
Qiantang Jiang River, Zhejiang Province, China. It
can be distinguished from the most similar genus
Xenocypris by having a transverse instead of a curved
mouth and possessing only two instead of three rows
of pharyngeal teeth (Peters 1881). The status of
Distoechodon is ambiguous and confusing at both
the genus and species levels.
Nichols (1928) synonymized Distoechodon with
Xenocypris and treated it as a subgenus without pro-
viding any specific explanation. Yang (1964) revali-
dated Distoechodon on the basis of possessing only
two instead of three rows of pharyngeal teeth.
Bănărescu (1970) disagreed with Yang’s(1964) syno-
nymy because reduction of the number of rows of
pharyngeal teeth is common among cyprinid fishes. He
thus synonymized Distoechodon back as a subgenus
of Xenocypris. But almost all contemporary Chinese
systematic work on xenocyprinid genera follows Yang
(1964), recognizing Distoechodon as a valid genus
having two rather than three rows of pharyngeal teeth
(Lian 1984; Chen and Li 1989; Mao 1991; Ding
1994; Liu and He
1998).
Environ Biol Fish (2009) 86:31–44
DOI 10.1007/s10641-008-9421-z
Y. Zhao (*)
:
C. Zhang (*)
Institute of Zoology, Chinese Academy of Science,
Beijing 100101, China
e-mail: zhaoyh@ioz.ac.cn
F. Kullander
:
S. O. Kullander
Department of Vertebrate Zoology, Swedish Museum of
Natural History,
POB 50007, SE-104 05 Stockholm, Sweden
Nichols (1925) described Xenocypris compressus,
which has a deeper body and longer pectoral fins,
from Yenping (now known as Nanping), Min Jiang
River, Fujian Province. Later, he put the species into the
subgenus Distoechodon with X. tumirostris (Nichols
1928). Yang (1964) recognized both of them as species
of Distoechodon.Bănărescu (1970) synonymized X.
compressus with X. tumirostris, but retained it as a
subspecies. Lian (1984)followedYang(1964), recog-
nizing D. compressus as a valid species. Liu and He
(1998) synonymized D. compr essus with D. tumirostris,
based on the fact that the number of lateral-line scales
and gill rakers among populations of Distoechodon are
overlapping. Shan (1998) described differe nces between
D. compr essus and D. tumirostris in lateral-line scale
count, shapes of the dentary and basioccipital mastica-
tory plates, and revalidated D. compressus. Molecular
analysis of nucleotides of the mitochondrial D-loop
region and the complete cytochrome b gene supported
D. compr e ssus as a distinct species (Liu 2002).
Yih (1964) added another species to the genus,
Distoechodon hupeinensis, from the Liangzihu Lake,
Chang Jiang River basin, in Hubei Province. Com-
pared to all other Distoechodon species, D. hupei-
nensis has fewer lateral-line scales, fewer gill rakers,
and lacks the dark horizontal stripes along the sides of
the body. Liu and He (1998) accepted this description
in their review. In a phylogenetic study based on
morphological characters, Shan (1998) placed D.
hupeinensis under Xenocypris. A molecular study of
the cytochrome b gene from a number of xenocypri-
nid species showed that D. hupeiensis is indeed more
closely related to
X. argentea and X. davidi than to
other Distoechodon species (Xiao et al. 2001).
With synonymising Distoechodon to Xenocypris,
Bănărescu ( 1970) also recognized three subspecies of
X. (D.) tumirostris, based mainly on lateral-line scale
counts and number of gill rakers: X. (D.) t. tumirostris
(Zhejiang Province, 71–77 lateral-line scales, 80–85
gill rakers), X. (D.) t. compressus (Fujian Province;
74–75 lateral-line scales, 93–98 gill rakers), and X.
(D.) t. multispinnis, a new subspecies from the Chang
Jiang basin, Sichuan Province, with 77–84 lateral-line
scales and 91–108 gill rakers. Ding (1994) tentatively
synonymized X. tumirostris multispinnis with D.
tumirostris without providing any explanation.
In the course of reviewing the genus Distoechodon
as well as clarifying the species diversity of the genus,
we examined 164 specimens of Distoechodon from
different localities in China, allowing the recognition
of the genus, redescription of the type species, D.
tumirostris, revalidation of both D. comp ressus and
D. multispinnis, as well as a new species, which is
herein described.
Materials and methods
We examined 164 specimens of Distoechodon and 57
specimens of Xenocypris. Comparative material of
Xenocypris is listed separately. Measurements were
taken point to point with a digital caliper (Mitutoyo
CD-20C) to 0.1 mm. Morphometric and meristic char-
acters were selected according to methods described by
Chen (1998) and Fang (1997). The anterior-most mea-
suring point is at the tip of the snout instead of the
upper jaw symphysis. Post-predorsal length was mea-
sured from the posterior end of supraoccipital to the
dorsal-fin insertion. Measurements and counts were
taken on the left side of the body whenever possible.
Gill rakers were counted only in the lower part of the
first gill arch, on the ceratobranchial and hypobranchial
bones.
Specimens were radiographed with Philips MG 150
low voltage X-ray unit with Kodak X-Omat V films.
Dorsal, anal and caudal-fin rays and vertebrae were
counted on radiographs. Abdominal vertebrae include
the Weberian apparatus. Pr edorsal vertebrae were
counted from the first free vertebra to the vertebra
before the first dorsal-fin pterygiophore. Caudal verte-
brae were counted from the first vertebra with haemal
spine to the last half centrum.
Morphometric and meristic information was analyzed
and organized using SYSTAT version 10. Most of the
coordinates provided in the material examined list were
researched by the authors. Institutional abbreviations
follow Leviton et al. (1985) and Leviton and Gibbs
(1988). The local Chinese toponymy is employed for all
localities. The Chinese names for the Yangtze and Pearl
Rivers are Chang Jiang and Zhu Jiang, r espectively.
Results
Key to Xenocyprinid genera
Previous keys were mainly based on the number of
pharyngeal tooth rows and thus impractical as they
32 Environ Biol Fish (2009) 86:31–44
require extensive dissection (Liu and He 1998). An
alternative key is proposed here, usin g external
characters only, which is more user-friendly and
practical.
Key to genera of the Xenoxyprininae
1. Abdominal keel longer than distance from pelvic-
fin base to vent …………… 2
1. Abdominal keel absent or rather short ……..…… 3
2. Lateral line scales fewer than 50; gill rakers on first
gill arch more than 100…………………………
Pseudobrama
3. Lateral line scales more than 70; gill rakers on
first gill arch fewer than 50 …………………
Plagiognathops
4. Mouth transverse; pectoral-fin axillary fold
present …………………Distoechodon
5. Mouth curved; pectoral-fin axillary fold absent
…………………………Xenocypris
Key to species of Distoechodon
Distoechodon Peters 1881
Distoechodon Peters, 1881, Mber. Akad. Wiss., Berl.
45: 912 (type species: Distoechodon tumirostris Peters)
Body elongated and compressed; abdomen round
without keel; mouth inferior, transverse (slightly
curved), with cuticular edge (Fig. 1A); snout round
(Fig. 1B); pectoral-fin axillary fold present (Fig. 1B);
pharyngeal teeth (Fig. 1C) in 2 rows, outer row well-
developed, deeply compressed; lateral line curved,
possessing scales more than 60. The genus includes
four species, all distributed in China (Chang Jiang
River basin, Zhu Jiang River basin and some coastal
rivers in south-east China; historical records in Taiwan
Island).
Key to species of Distoechodon
1. Eye relatively large among the same-sized indi-
viduals, 5.97–10.82% of standard length; distrib-
uted in Fujian Province and Chenghai Lake in
Yunnan Province ……………………………2
1. Eye relatively small among the same-sized indi-
viduals, 4.58– 9.08% of standard length; distrib-
uted Chang Jiang and Zhu Jiang River basins and
coastal rivers in Zhejiang Province ……….….
….3
2. Lateral line scales 67–74, predorsal scales 27–
33…………………… D. compressus
3. Lateral line scales 78–85, predorsal scales 34–
39…………..… D. macrophthalmus
4. Lateral line scales 68–77; dorsal and anal fins
relatively short, 16.49–21.73% and 11.47
–14.73%
of standard length respectively, distributed in costal
rivers in eastern China ……………………………
D. tumirostris
Pectoral-fin axillary fold
A
B
C
Fig. 1 Some typical characteristics of the Distoechodon species.
a Mouth shape. b Snout and pectoral-fin axillary. c Pharyngeal
teeth
Environ Biol Fish (2009) 86:31–44 33
Lateral lin e scales 72–82, dorsal and anal fins
relatively long, 18.97–24.13% and 12.22–16.58% of
standard length respectively, distributed in Chang
Jiang River and Zhu Jiang River basins………………
D. multispinnis sp.nov.
Distoechodon tumirostris Peters 1881 (Fig. 2A, Table 1)
Distoechodon tumirostris Peters 1881, Mber. Akad.
Wiss., Berl. 45: 925 (Ningpo, now known as Ningbo
in Zhejiang Province); Mao 1991, Fauna of Zhejiang,
Fresh Water Fishes: 76 (Anji, Tonglu, Tiantai and
Yunhe in Zhejiang Province)
Xenocypris (Distoechodon) tumirostris Nichols
1928, Bull. Am. Mus. Nat. Hist. 58(1): 24 (Ningpo);
Nichols 1943, Nat. Hist. Central Asia 9: 124 (Ningpo)
Xenocypris (Distoechodon) tum irostris tumirostris
Bănărescu 1970, Rev. Roum. Biol. (zool.) 15(6): 400
(Ningpo)
Specimens examined
All from China. Zhejiang Province: Ou Jiang River
basin: ASIZB 64104–64105, 2, 154.4–156.2 mm SL,
Lishui (28°27′ N, 119°54′ E), 27 Apr. 1972; ASIZB
61771–61773, 3, 111.5–126.2 mm SL, Qingtian (28°09′
N, 120°18′E), 28 Apr. 1972; IHB 0002420, 4, 159.2–
318.0 mm SL, Lishui and Jinyun (28°39′N, 120°04′E).
Ling Jiang River basin: ASIZB 73089–73092, 4, 99.1–
156.8 mm SL, Xianju (28°51′N, 120°44′E) and Linhai
(28°51′N, 121°07′E), 29 May 1972; ASIZB 62312–
62316, 5, 172.3–243 mm SL, Xianju and Linhai, 31
May 1972; IHB 0002422, 15, 91.3
–242.0 mm SL,
Linhai, Xianju and Tiantai (29°08′N, 121°02′E), Sep.
1974. Yong Jiang River basin: IHB 0002421, 5, 146.0–
245.0 mm SL, Ningbo (29 °52′N, 12 1°33′E) and
Fenghua (29°39′N, 121°24′E), Sep. 1974 and May
1976. Xin’an Jiang River basin: IHB 0002426, 1,
298.3 mm SL, Chun’an (29°37′N, 119°02′E), June
1977. Cao’e Jiang River basin: IHB 0002423, 2, 103.3–
149.5 mm SL, Shengzhou (29°36’N, 120°49’E), Sep.
1974. River unknown: ASIZB 62137, 1, 75.5 mm SL,
May 1972; ASIZB 62139, 1, 86.6 mm SL, May 1972;
IHB 0002425, 3, 146.1–203.1 mm SL.
Diagnosis Distoechodon tumirostris can be distin-
guished from all other congeners by having a
combination of the following characters: lateral-line
scales 68–77, modally 73; predorsal scales 30–36,
mostly 33 (Fig. 3); relatively smaller eye (5.07–9.08%
in SL). Similar to D. compressus in lateral-line scale
count, but is different from it by having more
predorsal scales (30–36 vs. 27–33, Fig. 3) and always
smaller eyes at the same body size (5.07–9.08% vs.
5.97–10.82% in SL, Table 1 and Fig. 4). Similar to D.
multispinnis in predorsal scale count and eye diame-
A
C
B
D
Fig. 2 Four species of Distoechodon. a Distoechodon tumirostris,
ASIZB 73089, 201.9 mm SL. b Distoechodon compressus,ASIZB
66087, 116.0 mm SL. c Distoechodon multispinnis, NRM T3486,
141.9 mm SL. d Distoechodon macrophthalmus sp. nov.; KIZ
797297, holotype, 155.4 mm SL
34 Environ Biol Fish (2009) 86:31–44
ter, but differs by having fewer lateral-line scales (68–
77 vs. 72–82, Fig. 3), shorter dorsal-fin and anal-fin
length (16.49–21.73% vs. 18.97–24.13% and 11.47–
14.73% vs. 12.22–16.58% in SL respective ly).
Description Body elongate and moderately com-
pressed; dorsal profile straight or slightly convex;
ventral profile concave; abdomen round; abdominal
keel absent. Head rather small and compressed; snout
round, anterior tip truncated; snout length longer than
eye diameter; mouth inferior, transverse, with cuticu-
lar edge; mouth wide, as wide as head width at that
point; no barbel; nostrils closer to eyes than to tip of
snout; eyes relatively small.
Scales moderately large. Lateral line complete, scales
68(5), 71(5), 72(7), 73(6), 74(7), 75(6), 76(6), 77(4).
Table 1 Morphometry of four species of the genus Distoechodon. Measurements are expressed as percents of SL
Distoechodon tumirostris Distoechodon compressus
N Min max mean SD N Min max mean SD
SL (mm) 46 75.5 318.0 171.2 56.3 60 81.2 247.7 145.8 37.6
Body depth 46 22.9 29.7 25.6 1.6 60 22.2 32.0 26.9 2.0
Predorsal length 46 50.0 56.3 53.3 1.4 60 50.6 59.6 53.8 1.7
Preanal length 46 73.6 83.1 77.3 2.2 60 73.1 82.9 77.9 1.9
Prepectoral length 46 20.8 26.1 23.8 1.3 60 21.2 28.2 25.2 1.5
Prepelvic length 46 50.3 56.2 53.0 1.5 60 51.0 57.6 53.7 1.5
Caudal-peduncle length 46 13.1 17.1 14.9 1.0 60 12.3 17.2 14.6 0.9
Caudal-peduncle depth 46 9.4 11.8 10.8 0.6 60 9.6 12.6 11.2 0.6
Pectoral-fin length 46 14.3 19.0 16.7 1.0 60 16.2 19.9 18.1 0.9
Pelvic-fin length 46 12.1 16.0 14.2 0.8 60 13.8 17.5 15.5 0.8
Dorsal-fin length 46 16.5 21.7 19.4 1.2 59 15.1 24.3 21.6 1.5
Anal-fin length 46 11.5 14.7 13.2 0.8 59 12.8 16.2 14.4 0.8
Head length 46 20.2 26.4 22.9 1.4 60 21.4 26.8 24.3 1.3
Head depth 46 15.4 19.7 17.7 0.8 60 15.9 20.7 18.0 1.0
Snout length 46 7.1 10.0 8.2 0.6 60 6.9 10.1 8.6 0.8
Eye diameter 46 5.1 9.1 7.0 1.1 60 6.0 10.8 8.2 1.0
Interorbital width 46 7.7 10.6 8.9 0.6 60 7.6 10.8 9.0 0.7
Mouth width 46 5.8 9.3 7.7 0.7 60 6.4 9.5 8.1 0.7
Distoechodon multispinnis Distoechodon macrophtalmus sp.nov.
N min max mean SD N min max mean SD
SL (mm) 52 111.6 326.8 191.7 43.8 15 104.2 200.1 153.0 31.1
Body depth 52 21.7 32.4 26.0 2.4 15 22.9 26.6 24.8 0.9
Predorsal length 52 49.7 56.4 52.5 1.5 15 51.6 55.8 53.5 1.2
Preanal length 52 73.7 81.7 76.6 1.7 15 74.7 78.8 76.7 1.3
Prepectoral length 52 21.6 26.4 23.5 1.1 15 21.0 25.6 23.7 1.5
Prepelvic length 52 49.9 55.4 52.2 1.3 15 49.2 56.6 52.4 1.8
Caudal-peduncle length 52 12.9 17.5 15.4 1.1 15 12.1 16.4 14.4 1.1
Caudal-peduncle depth 52 9.3 13.2 11.4 0.8 15 9.3 11.3 10.4 0.6
Pectoral-fin length 52 15.4 20.9 16.8 1.0 15 16.7 18.9 17.8 0.8
Pelvic-fin length 52 13.6 17.6 15.1 0.9 15 14.5 15.9 15.3 0.5
Dorsal-fin length 42 19.0 24.1 20.5 1.2 15 18.1 22.8 20.2 1.1
Anal-fin length 42 12.2 16.6 14.4 1.1 15 12.4 15.4 14.3 0.7
Head length 52 20.3 25.0 22.3 1.1 15 21.4 24.5 22.8 1.1
Head depth 52 16.0 19.7 17.4 0.9 15 16.2 17.8 17.1 0.5
Snout length 52 7.1 9.2 8.2 0.5 15 7.3 8.4 7.8 0.4
Eye diameter 52 4.6 8.3 6.4 0.8 15 6.5 9.4 8.0 1.0
Interorbital width 52 8.0 10.2 9.1 0.4 15 7.8 9.3 8.4 0.5
Mouth width 52 6.3 8.6 7.7 0.5 15 6.6 7.8 7.2 0.4
Environ Biol Fish (2009) 86:31–44 35
Lateral line descending from top of gill opening to
above pectoral-fin ray, then running parallel to horizon-
tal scale row and ascending to body midline above anal-
fin base and extending to end of caudal peduncle. Scales
above lateral line 12–13, rarely 14; scales below lateral
line 7–8, rarely 9; circumpeduncular scales 26; pre-
dorsal scales irregularly arranged, 30–36, mostly 33;
scales cover abdomen. Pectoral-fin axillary fold present;
pelvic-fin axillary scale well-developed. Gill rakers tiny,
numerous and triangular, 51(1), 52(3), 53(7), 54(3), 55
(1), 56(5), 57(1), 58(3), 59(6), 61(1), 62(3), 63(2), 64(1)
in lower part of first gill arch.
Pectoral-fin insertion slightly anterior to vertical
through posterior margin of opercle. Pectoral fin short,
reaching to midway between pectoral and pelvic-fin
insertions. Pelvic fin short, reaching to midway between
pelvic-fin insertion and vent. Dorsal-fin origin midway
between snout tip and caudal-fin base, and slightly
posterior to pelvic-fin insertion. Anal-fin insertion
midway between pelvic-fin origin and caudal-fin base.
Dorsal-fin rays iii, 7 (11), last unbranched dorsal-fin ray
ossified, smooth. Anal-fin rays iii, 9 (9), 8 (2). Pectoral-
fin rays i, 14 (6), 15 (3), 16 (2). Pelvic-fin rays i, 8 (11).
Caudal fin forked, lobes almost equal in length;
procurrent caudal-fin rays 9 (2), 10 (7), 11 (2) dorsally,
8 (2), 9 (9) ventrally.
Total vertebrae 25+20=45 (3), 26+19=45 (5), 26+
20=46 (2), 27+19=46 (1); predorsal vertebrae 9(3), 10
(7), 11 (1). Pharyngeal teeth 2 in rows, 6, 3; 3, 6. Outer
row of pharyngeal teeth strong or well developed,
narrow and compressed, with curved and pointed tip.
Inner row of pharyngeal teeth short, not compressed,
with very sharp tips.
Coloration in preservative Alcohol preserved speci-
mens generally grayish to brownish, without markings.
All fins light brownish.
Geographical distribution Coastal rivers (Ou Jiang,
Ling Jiang, Yong Jiang, Qiantang Jiang and Cao’eJiang
River basins) in Zhejiang Province (Fig. 5).
Remarks Characters agree with data from the original
description (Peters 1881), except for body depth, which
was stated as 4.75 in the original description, but comes
out as 3.4–4.3 in our study. The original description was
Lateral line scales
0
2
4
6
8
10
12
14
67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83
No.
Predorsal scales
0
2
4
6
8
10
12
14
16
27 28 29 30 31 32 33 34 35 36 37 38 39
Distoechodon tumirostris
Distoechodon compressus
Distoechodon multispinnis
Distoechodon macro
p
hthalmus
No.
a
b
Fig. 3 Differences in lateral-line scale (above, A) and predorsal
scale counts (below, B) among four species of Distoechodon
0 100 200 300 400
Standard len
g
th (mm)
5
6
7
8
9
10
11
Eye diameter (% of standard length
)
Fig. 4 Eye diameter (% SL) plotted against standard length
(mm) for Distoechodon tumirostris and D. compressus. ○
Distoechodon tumirostris. ● Distoechodon compressus
36 Environ Biol Fish (2009) 86:31–44
based on one specimen only. The difference in body
depth may reflect a different way of measuring standard
length. The holotype was not re-examined. Some of our
material is from Ningbo, the type locality of the species,
and serve as topotypes.
Distoechodon compressus (Nichols), 1925
(Fig. 2B, Table 1)
Xenocypris compressus Nichols 1925,Am.Mus.Novit.
(185):6 (Yenping, now known as Nanping in Fujian
Province);
Xenocypris (Distoechodon) compressus Nichols
1928, Bull. Am. Mus. Nat. Hist. 58(1): 24 (Fukien,
i.e. Fujian Province); Nichols 1943, Nat. Hist. Central
Asia 9: 124 (Kienning, now known as Jianning and
Yenping in Fujian Province).
Distoechodon compressus Yang 1964, The Cyprinid
fishes of China, 131 (Nanping); Lian 1984,Thefishes
of Fujian Province: 254 (Nanping, Jianyang, Jian’ou etc.
in Fujian Province).
Xenocypris (Distoechodon) tumirostris compressus
Bănărescu 1970, Rev. Roum. Biol. (zool.) 15(6): 400
(Yenping, Fukien)
Distoechodon tumirostris Lian 1984, The fishes of
Fujian Province: 253 (Nanping, Jianyang, Chong’an
etc. in Fujian Province); Shen and Tzeng 1993, Fishes
of Taiwan: 141 (Jilong River, Yilan in Taiwan Island).
Specimens examined All from China. Fujian Province:
Min Jiang River basin: ASIZB 67298–67305, 8, 134.2–
160.6 mm SL, Jianyang (27°20’ N, 118°07’E), Jianxi
River, Jun. 15, 1977; ASIZB 66084–66089, 6, 81.2–
115.0 mm SL, Jian’ou (27°03’N, 118° 19’E), Jianxi
River, Apr. and May, 1977; IHB 0002431, 3, 112.7–
153.6 mm SL, Nanping (26°38’N, 118°10’E), Apr.,
1957. Jiulong Jiang River basin: IHB 0002432, 4, 98.8–
134.2 mm SL, Zhangzhou (24°31’N, 117°39’E), May,
1974; SHFU A004217, 1, SHFU 6164–6169, 6, 130.8–
159.9 mm SL, Jimei (24°34’N, 118°07’E). Ting Jiang
River basin: IHB 0002423, 1, 96.9 mm SL, Shanghang
(25°03’N, 116°24’E), Jun., 1974; IHB 0002434, 7,
171.5–247.7 mm SL, Liancheng (25°42’N, 116°44’E),
Jun., 1974 and 1976; SHFU 6185, 1, SHFU 6212, 2,
SHFU uncat., 1, SHFU 6181, 1, SHFU 6184, 1, SHFU
6219, 1, SHFU 6199, 1, SHFU 6217, 1, SHFU 6204, 1,
SHFU 6201, 1, SHFU 6206, 1, 100.7–210.8 mm SL,
Liancheng, Jun., 1975. Ao Jiang River basin: IHB
Fig. 5 Distribution of four
Distoechodon species. ●
Distoechodon tumirostris. ★
Distoechodon compressus.
Distoechodon compressus. ✚
Distoechodon multispinnis.
▲ Distoechodon macro-
phthalmus sp. nov
Environ Biol Fish (2009) 86:31–44 37
0002437, 5, 115.9–139.8 mm SL, Lanjiang (26°12’N,
119°31’E), Jun. and Jul., 1974.
Diagnosis Distinguished from other Distoechodon
species by a combination of the foll owing characters:
fewer lateral – line scales (67–74, mostly 70, Fig. 3),
fewer predorsal scales (27–33, mostly 30, Fig. 3), and
relatively larger eye, 8.28% vs. 6.37–7.77% in SL.
The most similar species is Distoechodon tumirostris,
which can be distinguished by having more predorsal
scales (Fig. 3) and smal ler eyes (Fig. 4).
Description Body elongate and moderately com-
pressed; dorsal profile straight or slightly concave;
ventral profile convex; abdomen round; abdominal
keel absent. Head relatively large and compressed;
snout round, anterior tip truncated; snout length
longer than eye diameter; mouth inferior, transverse,
with cuticular edge; mouth wide, as wide as head
width at that point; no barbel; nostrils closer to eyes
than to tip of snout; eye diameter relatively greater
than that of D. tumiostris and D. multispinnis (Fig 4.
Table 1.)
Scales moderately large. Lateral-line complete,
scales 67(6), 68(5), 69(8), 70(10), 71(13), 72(11), 73
(4), 74(2). Lateral line descending from top of gill
opening to above pectoral-fin ray, then running
parallel to the horizontal scale row and ascending to
body midline above anal-fin base, and running until
end of caudal peduncle. Scales above lateral line 12–
13; scales below lateral line 7–8; circumpeduncular
scales 26; predorsal scales irregularly arranged, 27–
33, mostly 30; scales cover abdomen. Pectoral-fin
axillary fold present; pelvic-fi n axillary scales well-
developed. Gill rakers tiny, numerous and triangular,
51(1), 52(1), 53(3), 54(2), 55(5), 56(6), 57(4), 58(6),
59(4), 60(3), 61(4), 62(2), 63(9), 64(2), 65(1), 66(1)
in lower part of first gill arch.
Pectoral-fin insertion slightly anterior to vertical
through posterior margin of opercle. Pectoral fin short,
reaching to midway between pectoral and pelvic-fin
insertions. Pelvic fin short, reaching to midway between
pelvic-fin insertion and vent. Dorsal-fin origin midway
between snout tip and caudal-fin base, and slightly
posterior to pelvic-fin insertion. Anal-fin insertion
midway between pelvic-fin origin and caudal-fin base.
Dorsal-fin rays iii, 7 (8), last unbranched dorsal-fin ray
ossified but smooth. Anal-fin rays iii, 9 (8). Pectoral-fin
rays i, 14 (3), 15 (2), 16 (3). Pelvic-fin rays i, 8 (8).
Caudal fin forked, lobes almost equal in length;
procurrent caudal-fin rays 9 (1), 10 (6), 11 (1) dorsally,
8 (3), 9 (5) ventrally.
Total vertebrae 25+19=44 (1), 26+18=44 (1), 26+
19=45 (5), 26+20=46 (1); predorsal vertebrae 9(2), 10
(5), 11 (1). Pharyngeal teeth in 2 rows, 3, 6; 6, 3. Outer
row of pharyngeal teeth strong, deeply compressed,
with curved and pointed tips; inner row of pharyngeal
teeth not well-developed, with very sharp tips.
Coloration in pr eservative Alcohol preserved specimens
generally grayish to brownish, all fins light yellowish.
Geographical distribution Coastal rivers (Min Jiang,
Jiulong Jiang, Ting Jiang and Ao Jiang River basins) in
Fujian Province, and northern Taiwan Island (Yilan,
Jilong River, Shen and Tzeng 1993, Chen and Chang
2005)(Fig.5)
Remarks The characters observed for this study gener-
ally agree with the original description (Nichols 1925
).
The holotype was not re-examined but topotypes are
included in the material. It was difficult to check the
specimens from Taiwan. Based on the data from the
Fish Database of Taiwan (http://fishdb.sinica.edu.tw),
nine specimens have 66–74 lateral line scales, 30–32
predorsal scales. The rest of the characters are also
consistent with our description. Therefore, we put them
under Distoechodon compressus. It still needs to be
further studied in the future.
Distoechodon multispinnis (Bănărescu), 1970
(Fig. 2C, Table 1)
Distoechodon tumironstris Tchang 1933, Zool. Sinica
(B) 2 (1): 110 (Kaiting, should be Kiating); He 1989,
Fishes of Zhujiang River: 115 (Longsheng and
Rong’an counties, Guangxi); Ding 1994, The fishes
of Sichuan, China: 163 (Huanglongxi, Leshan, Yibin,
Luzhou and Shengzhong in Sichuan Prvince)
Xenocypris compressus Tcha ng 1933,Zool.Sinica
(B) 2 (1): 117 (Szechuan i.e. Sichuan Province)
Xenocypris (Distoechodon) tumirostris multispinnis
Bănărescu 1970, Rev. Roum. Biol. (zool) 15 (6): 401
(Suifu, Szechwan, now known as Yibin City in Sichuan
Province).
Specimens examined All from China. Chang Jiang
River basin: Sichuan Province: NRM T3483, 1; NRM
38 Environ Biol Fish (2009) 86:31–44
T3486, 1; NRM T3489–3495, 7, 141.9–187.6 mm SL,
Leshan (29°34’00”N, 103°43’59”E), 7 Aug 2002, S. O.
Kullander et al.; ASIZB 39779, 1, 183.0 mm SL; Min
Jiang River, Jun 1957; ASIZB 55822, 1, 214.0 mm SL,
Tuo Jiang River; IHB 0002417, 6, 111.6–224.1 mm SL,
Tuo Jiang River, 1958. IHB 0002578, 3, 120.3–
200.8 mm SL, Leshan, 1978, 1982 and 1987. Shaanxi
Province: ASIZB 61555, 2, 135.0–146.7 mm SL,
Hanzhong (33°04’00”N, 107°02’00”E), Hanshui River,
Apr 1990, C. G. Zhang and W. L. Jia; ASIZB 73551–
73554, 4, 146.6–199.6 mm SL, Yangxian (33°13’00”N,
107°33’00”E), Hanshui River, 12 Jul 2003 T. Q. Xu;
IHB 0002439, 2, 210.9–326.8 mm SL, Hanzhong,
Hanshui River, 1966 and May 1980. Hubei Province:
IHB 0002444, 7, 198.4–260.2 mm SL, Danjiangkou
(32°33’00N, 108°30’00E), Hanshui River, Jun 1976.
Guizhou Province: IHB 0002418, 1, 201.35 mm SL,
Tongren (27°43’N, 109°12’E), Yuan Jiang River, Apr
1988. Hunan Province: IHB 0002436, 1, 145.2 mm SL,
Luxi (28°17’N, 110°09’E), Yuan Jiang River, Jul 1987.
Zhu Jiang River basin: Guangxi Zhuang Autonomous
Region: ASIZB 63456–63457, 2, 188.4–237 mm SL,
Sanjiang (25°47’N, 109°36’E), Rong Jiang River, 14
May 1975. IHB 0002441, 2, 193.1–214.5 mm SL,
Longsheng (25°47’N, 110°02’E), Rong Jiang River,
Apr 1975; IHB 0002440, 3, 173.1–319.7 mm SL,
Guilin (25°17’N, 110°17’E) and Yangshuo (24°46’N,
110°28’
E), Li Jiang River, 1958, 1975 and 1981; IHB
0002442, 5, 125.9–179.9 mm SL, Rong’an (25°13’N,
109°22’), Siweihe reservoir, Jul 1973 and Nov 1974.
Guizhou Province: ASIZB 65694–65697, 4, 146.6–
224.0 mm SL, Duliujiang River, 1973.
Diagnosis Distinguished from other Dis toe chodo n
species by a combination of characters: lateral-line
scales 72–82, mostly 77; predorsal scales 29–37,
mostly 34 (Fig. 3); relatively small eye (6.37% vs.
6.98–8.28% in SL). Distoechodon multispinnis can be
distinguished from another similar species, D. tumir-
ostris, by having more lateral-line scales (Fig. 3), and
longer dorsal and anal fins. (Table 1).
Description Body elongate and moderately com-
pressed; dorsal profile straight or slightly convex;
ventral profile concave; abdomen round; abdominal
keel absent. Head rather small and compressed; snout
round, anterior tip truncated; snout length longer than
eye diameter; mouth inferior, transverse, with cuticular
edge; mouth wide, as wide as head width at that point;
no barbel; nostrils closer to eyes than to tip of snout;
eyes relatively small.
Sca les moderate ly small. Lateral-line complete,
scales 72(2), 73(8), 74(6), 75(3), 76(9), 77(8), 78(3),
79(2), 80(4), 81(6), 82(2). Lateral line descending from
top of gill opening to above pectoral-fin ray, then
running parallel to horizontal scale row and ascending to
body midline above anal-fin base and extending to end
of caudal peduncle. Scales above lateral line 12–13;
scales below lateral line 6–8; circumpeduncular scales
26; predorsal scales irregularly arranged, 29–37, mostly
34; scales cover abdomen. Pectoral-fin axillary fold
present; pelvic-fin axillary scales well-developed. Gill
rakers tiny, numerous and triangular, 52(2), 53(1), 54(5),
55(5), 56(3), 57(8), 58(4), 59(8), 60(2), 61(1), 62(6), 63
(2), 64(2), 65(2), 66(1) in lower part of first gill arch.
Pectoral-fin insertion slightly anterior to vertical
through posterior margin of opercle. Pectoral fin short,
reaching to midway between pectoral and pelvic-fin
insertions. Pelvic fin sho rt, reaching t o midway
between pelvic-fin insertion and vent. Dorsal-fin origin
midway between snout tip and caudal-fin base, and
slightly posterior to pelvic-fin insertion. Anal-fin
insertion midway between pelvic-fin origin and cau-
dal-fin base. Dorsal-fin rays iii, 7 (17), last unbranched
dorsal-fin ray ossified but smooth. Anal-fin rays iii, 8,
(1), 9 (14), 10 (2). Pectoral-fin rays i, 14 (3), 15 (10), 16
(1), 17 (3). Pelvic-fin rays i, 8 (17). Caudal fin forked,
lobes almost equal in length; procurrent caudal-fin rays
8 (1), 9 (2), 10 (14) dorsally, 7 (1), 8 (7), 9 (8), 10 (1)
ventrally.
Total vertebrae 26+19=45 (3), 26+20=46 (7), 27+
19=46 (2); 26+21=47 (2), 27+20=47 (3); predorsal
vertebrae 9 (1), 10 (13), 11 (3). Pharyngeal teeth in 2
rows, 3, 7; 7, 3. Outer row of pharyngeal teeth strong,
compressed, with curved and pointed tips; inner row of
pharyngeal teeth not well-developed, with very sharp
tips.
Coloration in preservative Alcohol preserved speci-
mens generally grayish to brownish; a dark spot usually
present on each side of gill cover; Horizontal scale-rows
above the lateral line are marked by dark spots along the
posterior margin of each scale, forming 10 to 12 dark
stripes. In poorly preserved specimens, this character is
not distinct.
Geographical distribution Upper Chang Jiang basin
(Min Jiang, Tuo Jiang in Sichuan Province, Hanshui
Environ Biol Fish (2009) 86:31–44 39
Rivers in Shanxi and Hubei Provinces, Yuan Jiang
River in Guizhou and Hunan Provinces) and Zhu
Jiang basin (Rong Jiang and Li Jiang Rivers in
Guangxi Zhuang Autonomous Region and Duliu
Jiang River in Guizhou Province) (Fig. 5).
Remarks Characters observed generally agree with
the original description (Băn ă rescu 1970), except for
caudal peduncle length and preanal length (19.5–
21.9% and 69.8–72.5%, respectively in the original
description). Differences in these two characters may
be explained by different measuring methods. Eight
specimens were included in the original description
but were not re-examined by us. Several specimens
from near the type locality are included in our
material.
Nucleotide sequences of the mitochondrial D-loop
region and complete cytochrome b gene show that the
p-distances between Distoechodon specimens from
the upper Chang Jiang basin in Sichua n and the Zhu
Jiang basin in Guangxi Zhuang Autonomous Region,
and D. compressus from costal rivers in Fujian
Province are higher than the common intraspecific
difference (Liu 2002), which supports the current
recognition of D. multispinnis (including the speci-
mens both from upper Chang Jiang and Zhu Jiang
basins) as distinct from D. compressus. D. multi-
spinnis is most similar to D. tumirostris in body shape
and scales count. Principal Component Analysis
clearly separated the two species on PCII, character-
ized by high loadings of pectoral fin-base length,
dorsal-fin length as well as anal-fin length (Table 2,
Fig. 6).
Distoechodon macrophthalmus sp. nov. (Fig. 2D,
Table 1)
Distoechodon tumirostris Chen and Li, 1989, The
fishes of Yunnan, China, Part I Cyprinidae: 96
(Changhai Lake, Yunnan Province).
Holotype KIZ 797297, 155.4 mm SL. Chenghai Lake,
Chang Jiang basin in Yongsheng County (26°42’N,
100°44’E), Yunnan Province, P. R. China, 1979.
Paratypes KIZ 797298, 1; KIZ 799738, 1; KIZ
797295, 1; KIZ 818413, 1; 104.2− 155.4 mm SL,
same data as holotype; IHB 0002446, 10, 115.2
−200.2 mm SL, same locality as holotype.
Diagnosis Distinguished from other species of Distoe-
chodon in possessing the highest number of lateral line
scales (78–85, mostly 82), more pre-dorsal scales (34–
39, mostly 36) (Fig. 3) and relatively bigger eye, being
7.97% vs. 6.37
–6.98% in SL. Distoechodon macro-
phthalmus is similar to D. multispinnis in having
higher lateral line and predorsal scale counts, and both
occur in the upper Chang Jiang basin. However, D.
macrophthalmus is easily distinguished by having
larger eyes at the same body size (Fig. 7, Table 1).
Description Body elongate and moderately com-
pressed; dorsal profile straight or slightly convex;
Table 2 Character Loadings on Principal Components I–III for
Measurement from specimens of Distoechodon tumirostris (46
specimens) and D. multispinnis (36 specimens)
PC I PC II PC III
Standard length 0.3172 0.0157 0.0135
Body depth 0.3353 −0.0315 0.0212
Predorsal length 0.3042 0.0141 0.0124
Post-predorsal length 0.3226 0.0119 0.0252
Prepectoral length 0.2787 0.0149 −0.0054
Prepelvic length 0.3005 0.0161 0.0074
Preanal length 0.3102 0.0151 0.0131
Caudal peduncle length 0.3347 0.0015 0.0297
Caudal peduncle depth 0.3461 −0.0328 0.0123
Dorsal-fin base length 0.3210 0.0036 0.0027
Anal-fin base length 0.3122 −0.0061 −0.0003
Pectoral-fin base length 0.3240 0.0384 −0.0103
Pelvic-fin base length 0.3458 −0.0123 0.0095
Dorsal-fin length 0.3181 −0.0375 −0.0113
Anal-fin length 0.3542 −0.0452 −0.0140
Pectoral-fin length 0.3032 −0.0087 −0.0151
Pelvic-fin length 0.3193 −0.0327 −0.0086
Head length 0.2703 0.0134 −0.0063
Head dorsum length 0.2774 0.0241 −0.0142
Head depth 0.3042 0.0061 0.0108
Head width 0.3308 0.0183 0.0158
Snout length 0.3316 0.0069 −0.0368
Eye diameter 0.1735 0.0176 −0.0069
Interorbital width 0.3546 −0.0026 0.0144
Upper jaw length 0.2963 −0.0165 0.0050
Mouth width 0.3554 0.0286 −0.0087
Prenostril length 0.3323 0.0008 −0.0605
Variance explained by
components
2.6935 0.0122 0.0095
Percent of total variance
explained
97.4509 0.4421 0.3446
Three highest loadings on PC II indicated by boldface
40 Environ Biol Fish (2009) 86:31–44
ventral profile concave; abdomen round; abdominal
keel absent. Head rather small and compressed; snout
round, anterior tip truncated; snout length longer than
eye diameter; mouth inferior, transverse, with cuticular
edge; mouth wide, as wide as head width at that point;
no barbel; nostrils closer to eyes than to tip of snout, first
pair of nostrils small and round, second pair of nostrils
transverse, after first pair immediately with membranes;
eye relatively bigger than that of D. tumirostris and D.
multispinnis (Table 1,Fig.7).
Scales moderately large. Lateral-line complete,
scales 78(2), 79(1), 81(1), 82(4), 84(6), 85(1). Lateral
line descending from top of gill opening to above
pectoral-fin ray, then running parallel to horizontal
scale row and ascending to body midline above anal-
fin base and extending to end of caudal peduncle.
Scales above lateral line 12–13; scales below lateral
line 7–8; circumpeduncular scale s 26 (14), rarely 24
(1); predorsal scales irregularly arr anged, 34–39,
mostly 36; scales cover abdomen. Pectoral-fin axillary
fold present; pelvic-fin axillary scales well-developed.
Gill rakers tiny, numerous and triangular, 53(3), 55
(2), 56(6), 57(2), 59(1) in lower part of first gill arch.
Pectoral-fin insertion slightly anterior to vertical
through posterior margin of opercle. Pectoral fin short,
reaching to midway between pectoral and pelvic-fin
insertions. Pelvic fin short, reaching to midway
between pelvic-fin insertion and vent. Dorsal-fin origin
midway between snout tip and caudal-fin base, and
slightly posterior to pelvic-fin insertion. Anal-fin
insertion midway between pelvic-fin origin and cau-
dal-fin base. Dorsal-fin rays iii, 7 (15), last unbranched
dorsal-fin ray ossified but smooth. Anal-fin rays iii, 9
(15). Pectoral-fin rays i, 14 (6), 15 (9). Pelvic-fin rays i,
8 (18). Caudal fin forked, lobes almost equal in lengt h.
Pharyngeal teeth in 2 rows, 2, 7; 7, 2. Outer row of
pharyngeal teeth strong, deeply compressed, with
curved and pointed tips; inner row of pharyngeal
teeth not well-developed, with very sharp tips.
Color in life. Dorsum dark grayish; abdomen silvery
whitish; side with 10 to 12 horizontal stripes composed of
black dots; dorsal and caudal fin light yellowish; pectoral,
pelvic and anal fin light reddish (Chen and Li 1989).
Coloration in preservative Alcohol-preserved speci-
mens generally brownish, dorsum blackish, no spe-
cific color pattern observed.
Geographical distribution Distoechodon macro-
phthalmus is known only from Chenghai Lake, upper
Chang Jiang basin in Yunnan Province (Fig. 5).
Chenghai Lake is situated about 1,503 m above sea
level, with an average depth at 25.7 m (Tao et al.,
1999). The lake was isolated from the Jinsha Jiang
-5 -4 -3 -2 -1 0 1 2 3 4
Principal Component I
-0.4
-0.3
-0.2
-0.1
0.0
0.1
0.2
0.3
Principal Component II
Fig. 6 Distoechodon tumirostris and D. multipinnis. Plot of
scores of principal component II on principal component I. ○
Distoechodon multispinnis. ● Distoechodon tumirostris
100 150 200 250
Standard len
g
th (mm)
0.05
0.06
0.07
0.08
0.09
0.10
Eye diameter (% of standard length)
Fig. 7 Eye diameter (% SL) plotted against standard length
(mm) for Distoechodon multispinnis and D. macrophthalmus
sp. nov. ○ Distoechodon multispinnis. ● Distoechodon macro-
phthalmus sp. nov.
Environ Biol Fish (2009) 86:31–44 41
River (belonging to Chang Jiang drainage) at least
400 years ago (Wang and Dou, 1998).
Etymology The specific name macrophthalmus is a
noun composed of the Greek words macros, meaning
big or long, and ophthalmus meaning eye, referring to
its large eyes.
Local name Hongchiyu (Red-wing fish) in Yongsheng
County, obviously referring to the light reddish pec-
toral, pelvic and anal fins in live specimens.
Remarks The population is shrinking quickly. It was a
very important component of the local fishery before the
1990s. But during our field survey in 2004, we could
not find any adult specimen. The primary threat might
be the introduction of icefishes (family Salangidae),
based on our survey.
Discussion
Compared to Xenocypris, Distoechodon has a trans-
verse instead of concave mouth, a shorter dentary
(Fig. 8), a well-developed (vs. weak or not developed)
cutting edge on the lower jaw, distinct pectoral-fin
axillary folds, more gill rakers (85–122 vs. 39–61),
and more predorsal vertebrae (Fig. 9). Distoechodon
and Xenocypris are both diagnosible and obviously
valid genera.
Distoechodon, and the other three genera, Xeno-
cypris, Pseudobrama, and Xenocyprioides, was clas-
sified into the subfamily Xenocypridinae (Liu and He,
1998). The number of rows of pharyngeal teeth has
been considered to be a main diagnostic character in
the subfamily (sens u Liu and He 1998): species from
Xenocyprioides and Xenocypris possess three rows of
pharyngeal teeth; Distoechodon,tworows;and
Pseudobrama one row. Recent phylogenetic studies
based on both morphological and molecular charac-
ters have shown that the number of pharyngeal teeth
rows is not a phylogenetically informative character
(Shan 1998; Xiao et al. 2001).
The genus Xenocyprioides was established and
designated into the subfamil y Xenocypridinae by
Chen in 1982, mainly because of the similar pharyn-
geal teeth pattern to other members of the subfamily.
Xenocyprioides includes 2 min species (21–30 mm
Fig. 8 Radiograph of Xenocypris argentea (ASIZB 63153,
above) and Distoechodon compressus (ASIZB 67303, below),
showing the difference in the dentary
18
24
4
9
31
5
0
5
10
15
20
25
30
35
61011
No. of Predorsal Vertebrae
No. of individuals
Xenoc
y
pris
Distoechodon
789
Fig. 9 Differences in the number of predorsal vertebrae
between Xenocypris argentea and Distoechodon species (in-
cluding D. tumirostris, D. compressus, and D. multispinnis)
42 Environ Biol Fish (2009) 86:31–44
SL) from southern China. They possess a flexible last
unbranched dorsal-fin ray, and fewer vertebrae (33–35
vs. 40–48) and scales in a lateral row (31–35 vs. 57–
84,) and lack lateral line, in contrast wi th all other
xenocypridines. Shan’s phylogenetic work based on
morphology demonstrated that this genus falls outside
the true Xenocypridinae, although she thought it
might be a primary clade (Shan, 1998). The phyloge-
netic tree of Xenocypridinae based on combined Cyt
b, ND4L, ND4, tRNA
Ser
, and tRNA
His
data sets also
had a similar result (Xiao et al, 2001). The position of
the genus Xenocyprioides in the family Cyprinidae
still need to be further discussed. It might be closer to
Danio or Rasbora. Therefore, four genera were
included in the subfamily Xenocypridinae by this study.
Except for Distoechodon and Xenocypris, Pseudobrama
is diagnosed by an abdominal keel from the pelvic-fin
base to the vent. It also has the lowest number of
lateral-line scales (41–47) and the highest number of
gill rakers (128–139) among the xenocypridin genera.
In addition, based on the development degree of the
abdominal keel and cutting edge on the lower jaw, we
are apt to admit Plagiognathops as a valid genus.
In a recent review (Liu and He 1998), the genus
Distoechodon included two species, D. hupeinensis
and D. tumirostris. D. hupeinensis has a curved
mouth and fewer lateral line scales and gill rakers,
distinguished from D. tumirostris, but close to the
species of Xenocypris. D. hupeinensis was arranged
under the genus Di stoechodon by Yang (1964) mainly
because of two rows of pharyngeal teeth, the same as
the species from Distoechodon. In fact, it has many
characters in common with the species of Xenocypris
including mouth shape, premaxilla and dentary
morphology, and the number of lateral-line scales.
Therefore, Shan (1998) thought two rows of pharyn-
geal teeth might be the result of parallel evolution and
suggested placing D. hupeinensis within Xenocypris.
We accepted her opinion in the presen t study. The
evidence from molecular systematics work shows that
D. hupeinensis is closely related to X. argentea and X.
davidi, but is distant from the species of Distoe cho-
don (Xiao et al 2001). Even the study of the related
parasite fauna also supports this viewpoint (Xiao &
Wang 2000).
After removing Distoechodon hupeinensis, four
species can now be recognized in the genus Distoecho-
don. Bănărescu (1970) used the number of gill rakers
and lateral-line scales to distinguish them. Our study,
however, shows no particular variation in the gill-raker
count among the different species. The number of
lateral-line scales, on the other hand, does show great
variation among the different species. In addition, the
four species occupy different areas. D. multispinnis has
the widest distribution among them. The new species is
the westernmost record of the genus (Fig. 5).
Comparative material examined All from China.
Xenocypris argentea: Hunan Province: Yuan Jiang
River: NRM-EC2002-2 (4), 12.9–24.5 mm SL,
Taoyuan (28°53′59″N,′ 111°2 8 ′59″E), 18–19 Mar,
2002, F. Fang et al. NRM-EC2002-3 (13), 63.0–
123.0 mm SL, Taoyuan (28°53′59″N, 111°28′59″E),
18–19 Mar, 2002, F. Fang. Hebei Province: IZB
15758–15763 (6), 74.5– 131.5 mm SL, Tangshan.
Xenocypris davidi: Hunan Province: Lishui River:
NRM-EC2002-18 (2), 111–112 mm SL, Shimen (29°
35′21″N, 111°17′43″E), 25 Mar, 2002, F. Fang et al.
Guangxi Zhuang Autonomous Region: IZB 63153–
63156 (4), 123–185 mm SL. Xenocypris sp.: Hunan
Province: Yuan Jiang River: NRM-EC2002-5 (1),
50.0 mm SL, Chehuyuan (28°59′46″N, 111°31′35″E),
Taoyuan, 18 Mar, 2002, A. Malzahn et al. NRM-
EC2002-9 (25), 50–78 mm SL, Xinlongjie (28°47′54″
N, 111°7′25″E), Taoyuan, 20 Mar, 2002, C. G. Zhang.
NRM-EC2002-41 (2), 134.0–146.0 mm SL, Hengdong
(27°4′59″
N, 112°56′59″E), 9 Apr, 2002, F. Fang et al.
Acknowledgments The work was carried out within the
ECOCARP project (New native fish species for Asian aquacul-
ture: conserving natural genetic reserves and increasing options
for sustainable use of aquatic resources), European Commission
INCO-DEV contract number ICA4-CT-2001-10024, and also
supported by Swedish Biodiversity Centre, Swedish University of
Agricultural Sciences and Chinese National Science Fund for
Fostering Talents in Basic Research (NSFC-J0030092). Many
thanks go to Liu H. Z. for constructive discussions and
suggestions, to Tang W. Q., Cai M. J., Zhang E., Xu T. Q., Cui
G. H. and Yang J. X. for allowing to access specimens under their
care, to M. Höggren and V. Areskoug for their guidance, and kind
concern and help. A. Silfvergrip helped to take fish photos, and E.
Åhlander helped to take all the radiographs.
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