Wei
Zhou1,2,*
,
Xiao-Fu Pan1 and Maurice Kottelat3
1
Faculty of Conservation Biology,
2
Faculty of Life Science,
3
Route de
*To whom all
correspondence and reprint requests should be addressed:
E-mail: weizhou@public.km.yn.cn
Tel: 86-871-3862458.
Fax: 86-871-3863635.
Received: 1 June 2004 Accepted: 20 April 2005 Published online: 28 July
2005
Abstract Wei
Zhou, Xiao-Fu Pan, and Maurice Kottelat (2005)
Species of Garra and Discogobio (Teleostei: Cyprinidae) in
the Yuanjiang (Upper Red River) drainage of
Key words: Discogobio, Garra micropulvinus, Garra, Placocheilus,
----------------------------------------------------------------------------------------------------------------
Introduction
Three
nominal genera of the Labeoinae (family Cyprinidae) (Garra Hamilton, Placocheilus Wu,
Discogobio Lin) are characterized by
the lower lip which is modified into an adhesive disk. They are collectively called below the Garra group. Six species are known in the Yuanjiang
(also known as Hong He, the Red River in English, and the Song Hong in
Vietnamese) drainage in Yunnan Province, China: G. imberba Garman, G.
orientalis Nichols, G. alticorpora
Chu and Cui, P. caudofasciatus
(Pellegrin and Chevey), D. yunnanensis (Regan),
and D. brachyphysallidos Huang (see
Chu 1987, Chu and Cui 1987, Chu and Chen 1989, Chu et al. 1993).
Most
members of the Garra group are small-
to medium-sized fish with a rounded, cylindrical body, usually with a flat
belly. The paired fins extend
horizontally. The mouth is
inferior, transverse, and semicircular.
The rostral cap and lower lip are continuous at the mouth corner, but without
lateral lobes. The lower lip is
thick and fleshy and forms an oral sucking disc on the chin, which is of
variable shape but usually rounded.
This oral sucking disc allows these fishes to adhere to the surface of
rocks or stones and to maintain their position in rapidly flowing waters.
Since
1996, the authors have surveyed the ichthyofauna of the Yuanjiang drainage in Yunnan Province and have collected
various material of the Garra group
from the tributaries, including the Lixianjiang, Tentiaohe, Xinxianhe, Nanxihe,
Panlonghe, and Luzhijiang (the suffixes "-jiang" and "-he"
mean "river" in Chinese) (Fig. 1). Some specimens from the Panlonghe were
found to differ from all known members of the Garra group. They
represent a new species described below, in the context of a review of the
members of the Garra group in the
Yuanjiang drainage of

Fig. 1. Distribution of the Garra group in southern
The
examined material is are deposited in the collections of the Museum of the
Southwest Forestry College (SWFC), Kunming Institute of Zoology, Chinese
Academy of Sciences (KIZ), and of the 3rd author (CMK). The method for obtaining counts and
measurements followed Hubbs and Lagler (1947), except that the count of lateral
line scales includes the scales on the caudal-fin base. Abbreviations used in the text are: m
asl, meters above sea level; co., county; prov., province; r., river; and SL,
standard length.
Members
of the Garra group share the
following characters of the oral sucking disc. The rostral cap is curved ventrally and
connected with the lower lip at the corners of the mouth. The edge of the rostral cap is fringed
or crenulated, and covered with papillae.
The upper and lower jaws have a cutting horny sheath. The upper lip is missing. The lower lip extends backwards and
forms a round to elliptical oral sucking disc on the chin. The lateral and posterior edges are
free, around a median, fleshy pad whose shape is variable (and often diagnostic
for species).
In
the genus Discogobio (Fig.
In
the genus Garra (Fig.
In
G. micropulvinus (Fig. 2D), the
fleshy median pad is smaller, and its transverse diameter is almost equal to
half the mouth width. The median
pad is almost entirely surrounded by a skin fold, and there is only a narrow
posterior interruption in the groove around it. On both sides of the median pad there
are 2~7 small fleshy buds between the skin fold and the median pad. The part of the skin fold anterior to
the median pad is wider than the posterior part. The median pad is divided into two parts
by a shallow groove: the anterior part is semicircular and entirely covered by
flat and low papillae, while the posterior part is triangular and its surface
is smooth (without papillae). There
are densely set papillae on the skin fold.
There is no narrow skin flange between the lower jaw and the skin fold.

Fig. 2. Oral sucking discs of (A) Garra imberba, SWFC0111233, Namoguo, Wenshan Co., Yunnan; (B) G. caudofasciata, SWFC9712992,
Baiheqiao, Pinbian Co., Yunnan; (C) G.
orientalis, SWFC97121423, Hekou, Hekou Co., Yunnan; (D) G. micropulvinus, paratype, SWFC0111008,
Gaji, Wenshan Co., Yunnan; (E) Discogobio
yunnanensis, SWFC0111317, Yaodian, Wenshan Co., Yunnan; (F) D. brachyphysallidos, SWFC0111031, Gaji,
Wenshan Co., Yunnan.
We
considered the possibility that G.
micropulvinus could represent a distinct genus. The reduced and divided central pad, the
presence of lateral buds, and the posterior notch are very distinctive, and
they constitute good diagnostic characters. But the general structure of the disc
remains the same as in other Garra. To treat G. micropulvinus as a distinct genus would require demonstrating
that the remaining species of Garra
form a monophyletic lineage and that G.
micropulvinus is the sister species of that lineage, or that several
lineages can be diagnosed and named within Garra. While it cannot be excluded that Garra is a polyphyletic genus, without a
proper phylogenetic analysis (which should also include Discogobio and similar-looking labeonine genera), such a decision
is not possible.
1b - Median pad not surrounded by a
horseshoe-shaped swelling; no dark submarginal stripe along each caudal fin
lobe (Garra). ……… 3
2b - Scales on belly extending
forward at most to midlength of pectoral fin, naked area sometimes with a few
isolated scales.……… D. brachyphysallidos
3b - No fleshy buds between median
pad and skin fold; posterior edge of oral sucking disc without a notch.……… 4
4b - No barbels; 46~52 lateral line
scales; anus close to pelvic-fin base.……… G.
imberba
5b - Snout without a notch; 37~39
lateral line scales; a dark semicircular mark on middle of each caudal-fin
lobe. ……… G. caudofasciata
Discogobio brachyphysallidos Huang 1989: 358 (Yiliang and
Luoping); Chu et al. 1993: 243 (Nanpanjiang, Yuanjiang, and Jinshajiang).
Material examined: All from China, Yunnan: SWFC
0111030~0111035, 0111105~0111108, 10, 93.0~
Diagnosis: See under D.
yunnanensis for characters distinguishing the two species.
Distribution: In
Remarks: Discogobio
brachyphysallidos was originally described on the basis of material from
Yiliang and Luoping (Huang 1989).
Chu et al. (1993) reexamined this material and showed that the specimens
from Yiliang in fact belong to D.
yunnanensis. The original
description (Huang 1989) does not mention the dark midlateral stripe which we
observe in our material. The caudal
fin is dark gray in specimens from Shundianhe (29~

Fig. 3. Discogobio
brachyphysallidos, SWFC0112087,
Discognathus yunnanensis Regan 1907: 63 (type locality:
Discogobio yunnanensis: Wu, in Wu et al. 1977: 386
(Yiliang, Yunnan Prov.); Chu and Cui 1989: 284 (Dianchi Lake and Nanpanjiang);
Chu et al. 1993: 243 (Jinshajiang, Pearl R., and Yuanjiang).
Material examined: All from China, Yunnan: SWFC
0111128~0111135, 0111188~0111195, 0111278~0111282, 0111286~0111288,
0111309~0111311, 27, 45.0~
Diagnosis: Distinguished from D.
brachyphysallidos, the only other species of the genus in the Yuanjiang
drainage in China, by scales on breast and belly extending forward at least to
posterior extremity of pectoral-fin base (vs. at most to midlength of pectoral
fin, except sometimes for a few isolated scales in naked area); an irregular
and faint stripe along flank from anal-fin origin backwards, darker
posteriorly, and with contrasting edges on caudal peduncle (vs. stripe
irregular, conspicuous on entire flank).
Distribution: In the Yuanjiang drainage, D. yunnanensis has been collected in the Namoguohe and Luzhijiang
(Fig. 1). It also occurs in the
Jinshajiang (upper Yangtze R.) and Nanpanjiang (upper Pearl R.) drainages.

Fig. 4. Discogobio yunnanensis, SWFC0111316,
Discognathus caudofasciatus Pellegrin and Chevey 1936: 223 (
Placocheilus caudofasciatus: Wu, in Wu et al. 1977: 382
(Yuanjiang);
Garra caudofasciatus: Kottelat
Material examined: All from China, Yunnan: SWFC
9712992, 1,
Diagnosis: Garra caudofasciata
is distinguished from all other species of Garra
and Discogobio by the presence of two
dark, semicircular marks on the middle of each lobe of the caudal fin (vs.
absence). The following characters
(none of which is unique to the species) further help to distinguish it from
the other species of Garra in the
Yuanjiang drainage in Yunnan: two pairs of barbels (vs. none in G. imberba); 37~39 lateral line scales
(vs. 46~
Remarks: Wu (in Wu et al. 1977) established the genus Placocheilus, with G. caudofasciata as the type species. It was diagnosed as having two (vs.
three) rows of pharyngeal teeth and a different structure of the oral sucking
disc. We have not observed significant
differences in the oral sucking disc morphology (see Fig. 2). All species of Garra in China have three rows of pharyngeal teeth (Wu et al. 1977,
Chu 1987, Chu and Cui 1987, Chu and Chen 1989), but we collected material of a
still unidentified Garra species from
the Nu Jiang (the upper Salween River) in Yunnan which has two rows of
pharyngeal teeth, suggesting that variation in the number of rows of pharyngeal
teeth occurs in the genus. Kottelat
(

Fig. 5. Garra
caudofasciata, SWFC9712992,
Garra (Ageneiogarra) imberba
Garman 1912: 114 (Kiating [Loshan], Minjiang, Sichuan Prov.).
Discognathus pingi Tchang 1929: 241 (
Garra pingi pingi: Wu et al. 1977: 373 (Jinshajiang,
Yuanjiang, and Lancangjiang);
Garra alticorpora
Chu and Cui 1987: 96 (Pinbian).
Material examined: All from China, Yunnan: SWFC
97121150~97121157, 8, 91~
Diagnosis: Garra imberba is
distinguished from the species of Garra
in the Yuanjiang drainage in Yunnan Province by the combination of the
following characters: no barbels (vs. two pairs); 46~52 lateral line scales
(vs. 32~
Remarks: The descriptions of both G.
imberba and G. pingi are based on
material from the Yangtze R. drainage in
Wu
et al. (1977), Chu and Cui (1987), and Chu and Cui (in
Garra alticorpora was described by

Fig. 6. Garra imberba, SWFC0111237,
|
|
G.
micropulvinus |
G. imberba |
G.
alticorpora |
||||||||||
|
n |
29 |
8 |
8+4 |
23 |
2 |
||||||||
|
Locality |
Gaji |
Xinxian |
Baizhai and Baiheqiao |
Namoguo |
Baiheqiao |
||||||||
|
Drainage |
|
|
|
|
|
||||||||
|
|
|
|
|
|
|
||||||||
|
Dorsal fin rays |
iii,7~8½ |
iii,9½ |
iii,9½ |
iii,9 |
iii,8~9½ |
||||||||
|
Pectoral fin rays |
i,14 |
i,15~16 |
i,15~16 |
i,15-16 |
i,16 |
||||||||
|
Lateral line scales |
40~42 |
49~50 |
48~50 |
48~50 |
48~49 |
||||||||
|
Predorsal scales |
14 |
15 |
15 |
15 |
15 |
||||||||
|
Circumpeduncular scales |
12 |
16 |
16 |
16 |
16 |
||||||||
|
In percent
of standard length |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
range |
|
Predorsal length |
46.5 |
41.0~51.2 |
2.16 |
45.9 |
42.3~48.4 |
1.65 |
46.3 |
41.8~50.0 |
2.10 |
48.0 |
45.5~51.6 |
1.55 |
47.3~48.9 |
|
Body depth |
22.7 |
19.0~26.2 |
1.54 |
18.1 |
15.9~20.1 |
1.28 |
18.2 |
15.2~25.1 |
2.21 |
19.5 |
17.6~22.1 |
1.04 |
23.6~24.2 |
|
Head length |
24.0 |
19.0~27.3 |
1.63 |
22.9 |
21.3~24.2 |
0.82 |
23.7 |
18.3~25.6 |
1.36 |
24.3 |
22.6~25.5 |
0.76 |
22.5~23.0 |
|
Caudal peduncle length |
23.8 |
22.0~25.5 |
0.97 |
16.7 |
15.7~17.7 |
0.69 |
16.7 |
13.6~18.6 |
1.48 |
17.2 |
15.3~18.6 |
0.91 |
17.8~17.9 |
|
Caudal peduncle depth |
12.4 |
10.0~14.0 |
0.75 |
10.5 |
9.8~11.3 |
0.57 |
10.8 |
9.6~12.7 |
0.94 |
11.2 |
10.3~12.6 |
0.54 |
11.5~11.8 |
|
In percent
of head length |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
range |
|
Snout length |
53.4 |
46.0~66.7 |
3.79 |
57.0 |
53.2~61.1 |
2.25 |
59.6 |
54.9~80.8 |
2.23 |
58.4 |
55.6~61.7 |
1.91 |
55.0~56.4 |
|
Eye diameter |
12.4 |
9.1~16.3 |
1.63 |
22.2 |
19.6~23.4 |
1.16 |
22.8 |
20.6~36.9 |
1.69 |
23.9 |
21.5~25.9 |
1.34 |
18.8~20.5 |
|
Interorbital width |
48.4 |
45.0~57.1 |
2.36 |
45.4 |
42.7~49.6 |
2.04 |
46.0 |
42.2~53.4 |
2.93 |
48.0 |
44.4~52.9 |
2.28 |
50.0~51.3 |
|
Body depth |
95.1 |
83.0~117.0 |
7.71 |
79.4 |
68.2~87.2 |
6.19 |
77.0 |
65.6~90.5 |
7.16 |
80.6 |
71.7~96.7 |
5.34 |
102.5~107.7 |
|
In percent of body depth in front anal
fin |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
range |
|
Distance between snout and postorbital |
- |
- |
- |
124.8 |
112.3~139.7 |
8.56 |
129.8 |
114.8~148.1 |
10.08 |
121.8 |
112.7~129.7 |
4.94 |
107.1~110.7 |
|
In percent of length of caudal
peduncle |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
mean |
range |
S.D. |
range |
|
Depth of caudal peduncle |
52.2 |
45.0~59.4 |
3.45 |
63.1 |
55.1~68.8 |
4.52 |
65.2 |
54.5~80.9 |
8.75 |
65.3 |
56.7~76.9 |
4.56 |
64.5~66.1 |
Holotype: SWFC 0111011,
Paratypes: SWFC
0111001~0111010, 0111012~0111029, 26, 63.5~
Diagnosis: Garra micropulvinus
is distinguished from all species of Garra
and Discogobio by having a median
notch in the posterior margin of the posterior free fold of the oral sucking
disc (vs. no notch), and 2~7 small fleshy buds between the skin fold and sides
of the median pad (vs. none).

Fig. 7. Garra
micropulvinus, holotype, SWFC0111011,
Description: Dorsal fin iii,7~8½; anal fin iii,5½; pectoral fin i,11;
pelvic fin i,8; branched caudal rays 9+8.
Lateral line scales 40~42 (including scales on caudal-fin base), 5
scales from lateral line to dorsal-fin origin, 3 to pelvic-fin origin, 12
circumpeduncular scales. Gill
rakers on outer side of 1st gill arch 9 or 10, on inner side 11 or 12. Pharyngeal teeth in 2 rows, 3.5-5.3,
2.4-4.2, or 2.3-3.2 (Fig.
Body
relatively elongate; cylindrical anteriorly, gradually compressed behind
dorsal-fin base, abdomen smooth before pelvic-fin origin. Head moderately compressed, longer than
wide. Interorbital area convex,
snout rounded without depression in front of nostril or secondary rostrum. Tip of snout with numerous, small,
irregularly distributed tubercles.
Gill opening extending to ventral surface of head. Nostrils closer to anterior margin of
eye than to tip of snout. Two pairs
of barbels, rostral pair longer than or equal to half of eye diameter,
maxillary pair shorter than rostral pair.
Eye moderately large, in posterior half of head, lateral, not visible in
ventral view. Postorbital length of
head shorter than snout length.
Mouth
inferior, transverse, semicircular.
Lower jaw covered with sharp horny edge. Rostral cap curved ventrally, covering
upper jaw and continuous with lower lip at corner of mouth. Edge of rostral cap with small
crenulations. Oral sucking disk
with a notch in posterior margin, width equal to head width at level of mouth,
posterior edge reaching to or beyond vertical through middle of eye. Gill rakers short, small, and widely
set. Pharyngeal teeth small,
surface smooth with sharp point at tip; pharyngeal bone thin and narrow (Fig.
Dorsal-fin
origin near middle of body, slightly closer to snout tip than to caudal-fin
base, in front of vertical through pelvic-fin origin, posterodorsal margin
slightly concave. Distance between
anal-fin origin and pelvic-fin base about half of distance between anal-fin
origin and caudal-fin base, distal margin slightly concave. Tip of pectoral fin rounded, not
reaching pelvic-fin origin, separated from it by 4 or 5 scales. Pelvic fin reaching to or beyond
anal-fin origin. Caudal fin deeply
forked, tip slightly pointed.

Fig. 8. Garra
micropulvinus, paratype, SWFC0111021,
Lateral
line complete and almost straight, with 40~42 scales. Scales on isthmus small, embedded in
skin, gradually increasing in size until pelvic-fin base. Scales on breast and belly smaller than
those on flank. Pelvic-fin base
with axillary scales. Anus
immediately in front of anal-fin origin.
Color pattern: Body brown and yellow, suffused with blue and
black. Dorsum relatively dark and
belly grayish. Scales on flank with
bronze spots. Dorsal and caudal
fins grayish black; pectoral and pelvic fins grayish black with light yellow
hue; anal fin gray.
Distribution: Garra micropulvinus
is presently only known from Panlonghe (a tributary of the upper Yuanjiang),
Habitat and ecology: Garra
micropulvinus prefers stream stretches with a rapid current and stony
substrate. They feed mainly on
algae, occasionally on aquatic plants and larvae of aquatic insects. According to villagers, adults swim
upstream in September~December and spawn in pools of clear water.
Etymology: Micropulvinus is
derived from the latinized Greek prefix micro
(small) and pulvinus (cushion or small
pillow), alluding to the small central pad of the oral sucking disc. It is a noun in apposition.
Garra orientalis: Nichols 1925: 4 (
Material examined: All from China, Yunnan: SWFC
9812024, 1,
Diagnosis: Garra orientalis
is distinguished from the species of Garra
in the Yuanjiang drainage in Yunnan by the following combination of characters:
two pairs of barbels (vs. none in G.
imberba); 32~35 lateral line scales (vs. 46~

Fig. 9. Garra orientalis, SWFC97121423,
Remarks: In his synopsis of the freshwater fishes of northern
Vietnam, Kottelat (
Acknowledgments: This
study was supported by a grant of the National Basic Research Program of China
(No. 2003CB415100).
----------------------------------------------------------------------------------------------------------------
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