See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/226875839

Revision of the cyprinid genus Placocheilus Wu, 1977 in China, with

description of a new species from Yunnan

Article  in  Hydrobiologia · November 2002

DOI: 10.1023/A:1022938132599

CITATIONS READS

21 74

3 authors, including:

Ee Zhang
Chinese Academy of Sciences
67 PUBLICATIONS   352 CITATIONS   

SEE PROFILE

All content following this page was uploaded by Ee Zhang on 09 July 2014.

The user has requested enhancement of the downloaded file.

 Hydrobiologia 487: 207–217, 2002.
© 2002 Kluwer Academic Publishers. Printed in the Netherlands.
207

Revision of the cyprinid genus Placocheilus Wu, 1977 in China, with

description of a new species from Yunnan

E. Zhang, Shun-Ping He & Yi-Yu Chen

Department of Ichthyology, Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan 430072,
Hubei Province, P. R. China
Tel: 86-27-87647725. Fax: 86-27-87875132. E-mail: zhange@ihb.ac.cn

Received 4 January 2001; in revised form 26 September 2002; accepted 21 October 2002

Key words: Placocheilus, taxonomy, new species, Yunnan, China

Abstract
Placocheilus is re-diagnosed on the basis of oromandibular structures compared to those of closely related groups.
It is represented in China by three species restricted to Yunnan, namely P. caudofasciatus from the Tengtiaohe River
(Nam Na), P. cryptonemus from the Nujiang Rive (upper Salween River basin), and P. robustus, herein described as
a new species from the Yuanjiang River (Red River basin). P. robustus n. sp. is differentiated from its congeners in
having a stout caudal peduncle (depth 72.4–82.5% of its length) and the medium-sized scaleless midventral region
of belly extended slightly beyond halfway from the pectoral- to pelvic-fin origin. It is further distinguished from
P. caudofasciatus in having a coarsely crenulated rostral fold with 18–23 indentations on its distal margin and a
smaller mental adhesive disc (length 66.7–71.4% of its width and 37.0–43.1% of the head length).

Introduction 1977) or a more thicker, flattened skin fold extended

Species of the cyprinid genus Placocheilus Wu,
1977 are elongate, small- to medium-sized, bottom-
dwelling fishes of swiftly flowing waters, known from
the upper Salween River basin in China, Red River
basin in both Vietnam and China, and Nam Ma basin
in Laos (Wu et al.,1977; Chu & Cui, 1989; Kottelat,
2001a, b). This genus has a lower lip modified into a
mental adhesive disc, by which it is distinguished from
all other currently known East and Southeast Asian
labeonine genera together with Discocheilus Zhang,
1997, Discogobio Fang, 1936 and Garra Hamilton,
1822. Placocheilus, as delimited by Wu et al. (1977),
is differentiated from Garra by having a narrower,
thicker skin fold formed by the anterior margin of the
mental adhesive disc and two rows of pharyngeal teeth
(Chu & Cui, 1989; Zhang et al., 2000). Placocheilus
shares with Discogobio and Discocheilus the presence
of two rows of pharyngeal teeth; however, its mental
adhesive disc is missing the horse-shoe shaped skin
fold diagnostic of Discogobio (Lin, 1931; Wu et al.,
posteriorly to overlap the anterior part of the central
fleshy pad characteristic ofDiscocheilus (Chen & Lan,
1992; Zhang, 1997).
Two species are currently recognized in Pla-
cocheilus. The first one, with a prominent semicircular
black mark on the middle of each caudal-fin lobe, was
originally described by Pellegrin & Chevey (1936) as
Discognathus caudofasciatus from Lai Chau (in the
Nam Na basin), Vietnam. Later, Wu et al. (1977)
reported D. caudofasciatus from the Yuanjiang River
(the main Red River before it enters into Vietnam) in
Yunnan, China and proposed Placocheilus as a new
genus for it. Chu & Cui (1989) identified the mater-
ial from the Yuanjiang River in Honghe and Jinping,
Yunnan as P. caudofasciatus. Mai (1978), however,
referred D. caudofasciatus to Garra, as was done by
Kottelat (2001a). P. caudofasciatus was recently re-
corded by Kottelat (2001b) as a Garra species from
the Nam Ma basin in Laos. The second species, P.
cryptonemus, was originally described by Cui & Li
(1984) and it is known only from the Nujiang River
208

Figure 1. Ventral views of structures in the oromandibular region in: (A)Placocheilus robustus, IHB 661132, 99.1 mm SL, holotype; (B) Garra
mirofrontis, IHB 78IV0389, 65.1 mm SL; (C) Discogobio tetrabarbus, IHB 81X0306, 64. 5 mm SL. (D) Discocheilus wui, IHB 89XII0011,
53.1 mm SL. af = anteromedian fold; al = anterolateral lobe; cp = central callous pad; lf = lateroposterior flap; mb = maxillary barbel; rf =
rostral fold; rb = rostral barbel.

(the upper Salween River in China) in Yunlong and
Lushui, Yunnan (Chu & Cui, 1989; Chen, 1998). This
species was included by Yue & Chen (1998) in the
red data book of China, but is so far found to be
an abundant species in the Nujiang River (Cui, pers.
comm.).
Placocheilus caudofasciatus, as presently recog-
nized by Chinese authors, is known from the Yuanji-
ang River and Tengtiaohe River (or Nam Na, a trib-
utary of the Red River basin) in Yunnan (Wu et al.,
1977; Chu & Cui, 1989). Our comparison of the
material from the Yuanjiang and Tengtiaohe Rivers in-
dicates that two distinct species are involved. The type
locality of P. caudofasciatus is Lai Chau, Vietnam, in
the Nam Na basin. The material from the Yuanjiang
River formerly identified by both Wu et al. (1977)
and Chu & Cui (1989) as P. caudofasciatus actually
represents an unnamed species, herein described as P.
robustus.
Materials and methods
Measurements were taken point to point with digital
calipers and data recorded to 0.1 mm. Counts and
measurements were made on the left side of speci-
mens whenever possible. Methods of taking counts
and measurements follow Kottelat (2001b) with the
addition of measurements of disc width and length.
Disc width is the distance between the bases of the
two maxillary barbels; in the case of the non-barbelled
species P. cryptonemus it is measured between the
lateralmost ends of each side of the mental adhesive
disc. Disc length is the distance between the anterior-
mid end of the anteromedian fold (explained below)
and the posteriormost end of the mental adhesive disc.
Subunits of the head are given as percentages of head
length (HL). Head length itself and measurements of
body parts are expressed as proportions of the standard
length (SL) unless otherwise stated. The holotype of
D. caudofasciatus is not well preserved and therefore
excluded in the statistics of morphometric data.
The oromandibular structures are useful for the
identification of most labeonine genera. The termin-

Figure 2. Dorsal view of: (A) P. caudofasciatus, IHB 92VI0001,
65.0 mm SL; (B)P. cryptonemus, IHB 81X 4303, 110.2 mm SL.
ology of the oromandibular structures and their defin-
itions follow Reid (1985) and Siebert (1998). In Pla-
cocheilus, major oromandibular structures involve: a
rostral fold (or rostral cap), a lower lip (or lower super-
ficial labial fold), an upper and lower jaw and barbels
(maxillary or rostral). Placocheilus possesses a lower
lip modified to form a mental adhesive disc, as found
in Garra, Discocheilus and Discogobio. This structure
is actually a soft-tissue complex consisting of many
components. To provide an adequate description of the
mental adhesive disc, four new terms are employed
here and defined as follows: The anteromedian fold
is a fleshy flap of skin which is formed by the an-
terior border of the mental adhesive disc, posterior to
the horny sheath on the lower jaw and anterior to the
central callous pad. The anterolateral lobe is a fleshy
lobe of skin with discrete margins, lateral to the end of
the anteromedian fold and bridging the rostral fold and
lateroposterior flap of the mental adhesive disc around
the corner of the mouth. The central callous pad is a
fleshy pad of thickened skin on the central portion of
209
the mental adhesive disc, anteriorly separated from the
anteromedian fold by a transverse groove. The latero-
posterior flap is a free, loose, membranous, peripheral
flap of skin surrounding lateral and posterior portion
of the central callous pad, superficially covered with
numerous papillae.
The material examined for the present study
is deposited in the following collections: Institute
of Hydrobiology, Chinese Academy of Sciences,
Wuhan (IHB); Kunming Institute of Zoology, Chinese
Academy of Sciences (KIZ); Department of Biology,
Yunnan University, Kunming (YU); Muséum national
d’Histoire Naturelle, Paris (MNHN).
Placocheilus Wu, 1977
Type species: Discognathus caudofasciatus Pellegrin
& Chevey, 1936: Wu et al., 1977: 382)
Diagnosis
Placocheilus belong to Labeonins (sensu Reid, 1982,
1985) defined by having four unique characters: (1)
vomer-palatine organ present; (2) neural complex of
Weberian apparatus in direct contact with supraoc-
cipital region; (3) terete process of basioccipital; (4)
superficial labial fold developed posterior to lower
jaw. Placocheilus is clearly distinguished from other
labeonine genera in East and Southeast Asia by having
a mental adhesive disc modified from the lower lip,
a character also shared with Discocheilus, Discogo-
bio and Garra. Among these genera, Placocheilus is
differentiated by possessing a combination of the fol-
lowing characters: two rows of pharyngeal teeth; a
crenulated rostral fold; a mental adhesive disc with a
crescent, narrower, thicker anteromedian fold anteri-
orly not continuous with horny sheath on lower jaw;
a large central callous pad (width greater than half
of disc width); an elliptical anterolateral lobe; a wide
mouth opening (width greater than two-thirds of head
width).
Remarks
Discognathus caudofasciatus Pellegrin & Chevey,
1936 has usually been accepted as the type species of
Placocheilus (Wu et al., 1977; Eschmeyer, 1998). The
type locality is Lai Chau (in the Nam Na basin), Viet-
nam. However, the erection of Placocheilus by Wu et
al.(1977) was based on the material from the Yuanji-
ang River in Yunnan identified as D. caudofasciatus.
The Yuanjiang River basin material under the name is
in fact a distinct species from P. caudofasciatus dis-
tributed in the Nam Na basin in Vietnam and China,
210

herein described as P. robustus. Be that as it may, the mental adhesive disc. The central callous pad of
according to the article 70.3.1 of ICZN (International the mental adhesive disc is anteriorly overlapped by
Code of Zoological Nomenclature), D. caudofasciatus the anteromedian fold in Discocheilus. But this over-
Pellegrin & Chevey, 1936 is here selected as the type lap is actually absent in Discogobio, Discocheilus and
species of Placocheilus. Placocheilus, all of them having a central callous pad
To date, there remains a controversy on the generic anteriorly or anteriorly and antero-laterally bordered
status of Placocheilus in the known literatures. Since with the anteromedian fold. The mouth opening of
its erection, Placocheilus has been generally treated by Placocheilus is wider than two-thirds of the head
Chinese authors as a distinct genus from Garra. This width, a state shared with Discogobio and Garra; but
point, however, was recently challenged by Kottelat a narrower mouth opening is present in Discocheilus,
(2001), who maintained that Placocheilus is unable to its width being less than or equal to half of the head
be accepted as different from Garra. The reason is that width.
the loss of one row of pharyngeal teeth is a reductive Additionally, the rows of pharyngeal teeth are also
character that is uninformative to diagnose the evol- useful for distinguishing Garra from Placocheilus,
utionary lineage (genera) and also that the difference Discogobio and Discocheilus. Three rows of pharyn-
in the anteromedian fold on the mental adhesive disc geal teeth are present in Garra but two rows in
is quite subtle between Placocheilus and Garra. In Placocheilus, Discogobio and Discocheilus.
my opinion, it is the best way to evaluate the validity
of Placocheilus in the phylogenetic framework. The
Species account
decisive answer to the generic status of Placocheilus
can’t be provided, unless the phylogenetic analysis is
Placocheilus caudofasciatus (Pellegrin & Chevey,
conducted for this genus and its relatives (Discogo-
1936) (Figs 2A, 3A, and 4A)
bio, Discocheilus and Garra). While awaiting this
analysis, we tentatively accept Placocheilus as a valid
Discognathus caudofasciatus Pellegrin & Chevey,
genus.
1936, 61:223 (Vietnam: Lai Chau)
There are morphological differences in the oro-
Garra caudofasciatus, Chevey & Lemasson, 1937, 33:
mandibular structures within Discocheilus, Discogo-
26.
bio, Garra and Placocheilus (Fig. 1). Like Garra and
Garra caudofasciatus, Mai, 1978: 58
Discogobio, Placocheilus has a crenulated rostral fold
Garra angulostoma Mai, 1978: 59
instead of a non-crenulated one in Discocheilus. In
Placocheilus caudofasciatus, Chu & Cui, 1989: 278
Placocheilus the mental adhesive disc has a antero-
(Yunnan: Jinping in Nam Na basin).
median fold apart from the lower jaw and anteriorly
Garra caudofasciatus, Kottelat, 2001a (northern Viet-
not continuous with the horny jaw sheath, with a
nam)
transverse deep groove between the horny jaw sheath
Garra caudofasciatus, Kottelat, 2001b (Laos: Nam
and anteromedian fold as present in both Garra and
Ma)
Discogobio; this kind of groove is absolutely absent
in Discocheilus, in which the anteromedian fold of
Material examined: MNHN 85-93-13-1, holotype,
the mental adhesive disc is separated from the lower
78.3 mm SL, Lai Chau, Vietnam. IHB 92VI001-04, 4
jaw but continuous with the horny jaw sheath. Dis-
specimens, 64.0–76.4 mm SL and YU, not numbered,
cocheilus has a large, triangular anterolateral lobe on
6 specimens, 23.0–65.0 mm SL; Tengtiaohe River in
the mental adhesive disc; whereas, this lobe is small,
Jinping County, Yunnan, China; Ji-Cang, He et al.,
elliptical in Placocheilus and Garra, even absent in
Jul., 1992. KIZ 20005352-56 20005981 20005985-
Discogobio. Placocheilus and Garra develop a lar-
86 20005972 20005358-60, 12 specimens, 49.1–87.
ger central callous pad of the mental adhesive disc
3 mm SL; Jinshuihe River, a left bank tributary of
than both Discogobio and Discocheilus, its width be-
Tengtiaohe River in Jinping County, Yunnan; China;
ing greater than half of the disc width. Placocheilus
Xiao-Yong, Chen, Jun., 2000.
shares with Garra a mental adhesive disc that has a
crescent anteromedian fold, with an arched, thicker Diagnosis
one occurring in Discocheilus or a horse-shoe shaped
one in Discogobio. Placocheilus differs from Garra Placocheilus caudofasciatus is distinguished from its
in having a narrower, thicker anteromedian fold on congeners by possessing a smaller scaleless midvent-
 211

Figure 3. Lateral view of: (A)P. caudofasciatus, IHB 92VI0001, 65.0 mm SL. (B) P. robustus IHB 601132, 99.1 mm SL, holotype; (C) P.
cryptonemus, IHB 81X 4303, 110.2 mm SL.

ral region only restricted to chest (vs. larger, extended
slightly beyond halfway from pectoral- to pelvic-fin
origin in P. robustus or as far as to reach anus in P.
cryptonemus). It is further differentiated from P. ro-
bustus in having a slightly crenulated rostral fold with
12–15 (vs. 18–23) indentations on its distal margin, a
shallower caudal peduncle (depth 63.7–71.3% of its
length vs. 72.4–82.5) and a larger mental adhesive
disc (length 72.7–80.7% of its width vs. 66.7–71.4 and
44.5–49.2% of HL vs. 37.0–43.1).
Color pattern in formalin
Body black above and gray and silvery underside.
Dorsal fin across with a black band; caudal fin with a
prominent semicircular black mark on middle of each
lobe; other fins gray and silvery.
Distribution
Placocheilus caudofasciatus is known from the Nam
Na basins in China and Vietnam (Fig. 5). This spe-
cies was first described by Pellegrin & Chevey (1936)
from Lai Chau (in the Nam Na basin), Vietnam and
it was later recorded by Mai (1978) from northern
Vietnam, without precise locality data. In China, this
species was documented by Chu & Cui (1989) from
the Tengtiaohe River in Jinping, Yunnan. The ex-
amined specimens come from the Tengtiaohe River of
the Nam Na basin in Jinping, Yunnan. Recently, this
212

Figure 4. Ventral view of: (A)P. caudofasciatus, IHB 92VI0001, 65.0 mm SL; (B) P. robustus, IHB 661132, 99.1 mm SL, holotype; (C) P.
cryptonemus, IHB 81X 4303, 110.2 mm SL.

Figure 5. Map showing distribution of three Placocheilus species in China. Hollow triangle indicates the type locality.

species was recorded by Kottelat (2001b) (as Garra
caudofasciatus) from the Nam Ma basin in Laos.
Remarks
In the original description of Discognathus caudo-
fasciatus by Pellegrin & Chevey (1936), there is a
disagreement between the text and figures in the count
of the lateral line scales. It was reported in the text to
have 36 lateral line scales, but 40–41 were illustrated
by its accompanying figures (1, 2). We observed the
holotype of Discognathus caudofasciatus and found
that it has 40 lateral line scales; this number falls
within the range (40–41) found in our examined spe-
cimens. Apparently, the count of the lateral line scales
by Pellegrin & Chevey (1936) is not accurate, or by a
different method, with the exclusion of scales on the
caudal peduncle base.
Rows of pharyngeal teeth are an important char-
acter diagnostic of Placocheilus. This character was
not mentioned in the original description by Pel-
legrin & Chevey (1936) and some subsequent authors
(Menon, 1964; Mai, 1978). Our observation of the
Nam Na basin specimens identified as P. caudofasci-
atus revealed that this species possesses two rows of
pharyngeal teeth, as speculated by Wu et al. (1977).
Garra angulostoma was first described by Mai
(1978) based on a single 80.0 mm SL specimen from
northern Vietnam and the original description is very
vague. However, this species was clearly illustrated
in his Figure 24 to share with P. caudofasciatus the
same caudal-fin color pattern including a semicircular
black mark on the middle of each lobe, by which it
is easily identified as a Placocheilus species. In addi-
tion, G. angulostoma has a scaled belly according to
the translation of Mai (1978) from Kottelat (2001a),
as found in P. caudofasciatus. Due to the vague in-
formation of the original description provided by Mai
(1978) for G. angulostoma and no accessibility to the
holotype, we have no reason not to follow Kottelat
(20001a,b) in tentatively considering it as a synonym
of P. caudofasciatus.
Placocheilus cryptonemus Cui & Li, 1984 (Figs 2B,
3C and 4C)
Placocheilus cryptonemus Cui & Li, 1984, 9:110
(Yunnan: Yunlong and Lushui in upper Salween River
basin)
Placocheilus cryptonemus, Chu & Cui, 1989:280
(Yunnan: Yunlong and Lushui in upper Salween River
basin)
213
Placocheilus cryptonemus, Chen, 1998: 180 (Yunnan:
Lushui and Yunlong in upper Salween River basin)
Material examined: KIZ 742035, holotype, 75.0 mm
SL, Nujiang River in Yunlong county, Yunnan, China;
KIZ 20007154 20007146 20007171, 3 ex., 84.2–100.3
mm SL, Nujiang River in Gongshan county, Yunnan,
China; KIZ 20007372–73, 2 ex., 69.3–80.5 mm SL,
Nujiang River in Liuku county, Yunnan, China; IHB
84VIII0010 84VIII0027, 2 ex. 84.4–86.8 mm SL,
Nujiang River in Fugong county, Yunnan, China. IHB
81X4311–12 81X4329–30, 81X4303–04 81X4308, 7
ex., 58.0–115.5 mm SL, Nujiang River in Lushui
county, Yunnan, China.
Diagnosis
Placocheilus cryptonemus is distinguished from its
congeners by having no barbel (vs. two pairs), 45–47
lateral line scales (vs. 39–42), no semicircular black
mark on middle of each caudal lobe (vs. present), no
blackish band across dorsal fin (vs. present), a shallow
(vs. deep) groove between anterior edge and central
callous pad of mental adhesive disc, a mental adhesive
disc reaching beyond vertical of posterior margin of
eye (vs. not reaching), predorsal scales smaller than
flank scales (vs. of same size) and embedded in skin
(vs. not embedded), a larger scaleless midventral re-
gion extended as far as to reach anus (vs. restricted
to chest P. caudofasciatus or reaching slightly bey-
ond halfway from pectoral- to pelvic-fin origin in P.
robustus), and a more slender caudal peduncle (depth
46.4–56.1% of its length vs. 63.7–82.5).
Color pattern in formalin
Body gray and black dorsally and laterally, yellowish
ventrally. Dorsal, anal and caudal fins gray. Pectoral
and pelvic fins gray dorsally and yellowish ventrally.
Distribution
Placocheilus cryptonemus is known only from the
Nujiang River in Yunnan, China (Fig. 5). This species
was first described by Cui & Li (1984) from the Nuji-
ang River in Yunlong and Lushui county, Yunnan; it
was subsequently reported in the same localities (Chu
& Cui, 1989; Chen, 1998). The examined material
includes three specimens from Gongshan county and
two specimens from Fugong county, Yunnan, in the
Nujiang River basin.
214
Remarks
In the original description by Cui & Li (1984), P.
cryptonemus is diagnosed as having no barbels, 45–
47 lateral line scales, and a shallow groove between
the anteromedian fold and the central callous pad of
the mental adhesive disc. Besides, the mental adhesive
disc of P. cryptonemus has a posterior margin reach-
ing beyond the vertical of the posterior margin of the
eye, no black band across the dorsal fin, no semicircu-
lar black mark on the middle of each caudal-fin lobe,
and predorsal scales smaller than the flank scales and
embedded in the skin, all these characters that clearly
distinguish it from P. caudofasciatus and P. robustus.
Also, in P. cryptonemus the caudal peduncle is more
slender, with its depth 46.4–56.1% of its length instead
of 63.7–71.3% in P. caudofasciatus and 72.4–82.5% in
P. robustus.
Placocheilus robustus n. sp. (Figs 1A, 3B and 4B)
Placocheilus caudofasciatus, Wu et al., 1977: 382
(Yunnan: Yuanjiang in Red River basin).
Placocheilus caudofasciatus, Chu & Cui, 1989: 278
(Yunnan: Honghe in Red River basin).
Placocheilus caudofasciatus, Zhang et al., 2000: 266
(Yunnan: Yuanjiang in Red River basin).
Holotype: IHB 601132, 99.1 mm SL, Yuanjiang River
in Yuanjiang county, Yunnan, China: in 1960.
Paratypes: IHB 601128–31, 60 1133–34, 6 ex., 48.6–
109.1 mm SL, same date as holotype.
Nontypes: YU 81122080, 1 ex., 77.1 mm SL, Yuanji-
ang River in Xinping county, Yunnan, China. KIZ
60201 60205–6 60219 60223–4 60227 60253 60233
60209 8340525, 12 ex., 53.1–117.4 mm SL, Yuanjiang
River in Honghe county, Yunnan, China.
Etymology The name is from the Latin robustus,
meaning stout, alluding to the shape of the caudal
peduncle. An adjective.
Diagnosis
Placocheilus robustus n. sp is differentiated from its
congeners by having a stouter caudal peduncle (depth
72.4–82.5% of its length vs. 63.7–71.3 in P. caudo-
fasciatus or 46.4–56.1% in P. cryptonemus) and the
medium-sized scaleless midventral region extended
slightly beyond halfway from pectoral- to pelvic-fin
origin (vs. extended as far as to reach anus in P.
cryptonemus or restricted to chest in P. caudofasci-
atus). It is further distinguished from P. caudofasci-
atus in having a coarsely crenulated rostral fold with
18–23 (vs.12–15) indentations on distal margin and a
smaller mental adhesive disc (length 66.7–71.4% of
its width vs. 72.7–80.7 and 37.0–43.1% of HL vs.
44.5–49.2).
Description
All morphometric and meristic data are summarized
in Table 1. Body elongate, depth 17.2–24.2% of SL;
cylindrical anteriorly and slightly compressed posteri-
orly; dorsal profile slightly convex and ventral surface
flattened from head to pelvic-fin origin. Head relat-
ively small and depressed; width greater than height;
length 17.9–22.2% of SL; interorbital space, slightly
convex, width 47.8–60.0% of HL. Snout obtuse and
round, length 46.4–59.3% of HL, bearing a few large
tubercles on its tip and anterior portion of lachrymal,
with a shallow furrow extending obliquely from base
of rostral barbels to lateral extremity of rostral fold.
Mental adhesive disc oval, small, not extending back-
ward beyond vertical of posterior margin of eye, with
a deep, arched groove separating anteromedian fold
from central callous pad, length 66.7–71.4% of its
width and 37.0–43.1% of HL, width 54.3–60.5% HL.
Rostral fold coarsely crenulated, with 18–23 indent-
ations on its distal margin, covered by numerous
papillae. Anteromedian fold, anterolateral lobe and
lateroposterior flap of mental adhesive disc papillose.
Two pairs of barbels, minute; rostral ones originating
at anterior end of lateral groove of snout, equal to
or longer than eye diameter; maxillary ones rooted at
corner of mouth, shorter than rostral ones. Eye small,
diameter 15.2–22.0% of HL, laterodorsally located,
closer to posterior margin of gill cover than to tip of
snout. Caudal peduncle stouter, length 13.2–16.1%
of SL and depth 11.0–13.4% of SL; caudal peduncle
depth 72.4–82.5% of its length.
Dorsal fin with 3 simple and 8 branched rays, last
one split to base, origin closer to snout tip than to
caudal base, distal margin slightly concave. Pectoral
fin falcate, with 1 simple and 12 – 14 branched rays,
longer than HL, reaching beyond halfway to pelvic-fin
origin. Pelvic fin round, with 1 simple and 13 or 14
branched rays, equal to or longer than HL, reaching
greatly beyond midway to anal-fin origin to exceed
anus, origin vertically well behind dorsal-fin origin;
axillary scales present and long, reaching beyond basis
of last ray. Anal-fin with 3 simple and 5 branched rays,
last one split to base, origin in midway between pelvic-
 215
Table 1. Comparisons of the morphometric and meristic data within three Chinese species of Placocheilus

Specimens examined P. caudofasciatus P. robustus P. cryptonemus

22 20 15

SL (mm) 48.0–76.5 48.6–117.2 58.0–115.5

HL (mm) 10.2–18.0 10.0– 25.4 9.0– 21.0

Proportion of SL (%)
Body depth 21.5–26.8 17.2– 24.2 16.7– 19.9
Head length 19.6–22.9 17.9– 22.2 17.8– 20.5
Length of caudal peduncle 16.7–18.9 13.2– 16.1 14.7– 17.0
Depth of caudal peduncle 12.0–13.3 11.0– 13.4 9.1– 10.3

Proportion of HL (%)
Snout length 45.2–50.0 46.4– 59.3 43.9– 54.0
Eye diameter 18.1–21.7 15.2– 22.0 14.6– 18.2
Interorbital width 58.1–65.4 47.8– 60.0 51.3– 54.1
Disc width 54.8–62.2 54.3– 60.5 58.9– 65.2
Disc length 44.5–49.2 37.0– 43.1 41.2– 48.8

Proportion of caudal peduncle length (%)

Caudal peduncle depth 63.7–71.3 72.4– 82.5 46.4– 56.1

Proportion of disc width (%)

Disc length 72.7–80.7 66.7– 71.4 66.5– 77.2

Dorsal-fin rays 3, 8 3. 8 3, 7–8

Anal-fin rays 3, 5 3.5 3.5
Pectoral-fin rays 1, 13–14 1, 13–14 1, 13–14
Pelvic-fin rays 1, 7 1, 8 1, 7
Lateral line scales 40–41 39–42 45–47
Scales above lateral line 5 5 or 5 12 5
Scales below lateral line 2 or 2 12 2 or 2 12 3 or 4
Predorsal scales 12 12–14 embedded
in the skin
Circumpeduncular scales 12 12 14
Pharyngeal teeth 4·5–5·4 (5) 3·5–5·3 (5) 3·5–5·3 (4)

fin origin and caudal-fin base, with a truncated distal
edge. Anus closer to anal-fin origin than to pelvic
origin. Caudal deeply forked.
Scales moderately large; lateral line complete, per-
forating 39–42 scales, 5 or 5 12 scales between dorsal-
fin origin and lateral line; 2 or 2 12 scales between
pelvic-fin origin and lateral line. Predorsal scales 12–
14, of same size as flank scales, not embedded in skin.
Circumpeduncular scales 12. Chest scaleless; belly
midventrally with a scaleless region extended slightly
beyond halfway from pectoral- to pelvic-fin origin.
Air bladder bipartite, anterior chamber round and pos-
terior chamber slender or stick-like and twice as long
as anterior one. Gill rakers sparse, short and small.
Two rows of pharyngeal teeth, teeth pattern 3·5–5·3
(5), with a compressed and pointed tip.
Color pattern in formalin
Body brown dorsally and laterally, gray and white
ventrally. Dorsal surface of pectoral and ventral fin
blackish. A blackish band across dorsal fin and one
prominent semicircular black mark on middle of each
caudal-fin lobe.
216
Distribution
Placocheilus robustus is known only from the Yuanji-
ang River basin in Yunnan, China (Fig. 5). This
species (misidentified as P. caudofasciatus) was repor-
ted by Wu et al. (1977) from the Yuanjiang River in
Yuanjiang, Yunnan and by Chu & Cui (1989) from the
same river in Honghe, Yunnan. It is also found in the
Yuanjiang River in Xinping, Yunnan. The Red River
flows from China to Vietnam and thus this species is
expected to occur in the Red River basin in Vietnam.
Remarks
Placocheilus robustus n. sp. is most similar to P.
caudofasciatus. Both has two pairs of barbels, lateral
line scales 37–42, one prominent semicircular black
mark on the middle of each caudal-fin lobe, a blackish
band across the dorsal fin, a deep groove between the
central callous pad and the anteromedian fold on the
mental adhesive disc, predorsal scales of same size as
the flank scales and not embedded in the skin, and a
mental adhesive disc not reaching beyond the vertical
of the posterior margin of the eye. All these characters
distinguish it from both P. cryptonemus. But compared
to P. caudofasciatus whose scaleless midventral region
is only confined to the chest, P. robustus n. sp. has
a larger scaleless midventral region extended slightly
beyond halfway from the pectoral- to pelvic-fin origin.
In P. robustus n. sp, the rostral fold is coarsely crenu-
lated with 18–23 indentations on its distal margin, but
slightly with 12–15 in P. caudofasciatus. P. robustus n.
sp. has a stouter caudal peduncle (depth 2.4–82.5% of
its length) than P. caudofasciatus, the ratio of which is
63.7–71.3%. A small mental adhesive disc is present
in P. robustus n. sp., its length being 66.7–71.4% of
its width and 37.0–43.1% of HL; but P. caudofasciatus
has a larger one with its length 72.7–80.7% of its width
and 44.5–49.2% of HL.
Key to the Chinese species of Placocheilus
1a No barbel; lateral line scales 45-47; no semicir-
cular black mark on middle of each caudal-fin
lobe; no blackish band across dorsal fin; a shallow
groove between central callous pad and antero-
median fold on mental adhesive disc; scaleless
midventral region extended as far as to reach
anus; predorsal scales smaller than flank scales
and embedded in skin; mental adhesive disc reach-
ing beyond vertical of posterior margin of eye;
caudal peduncle depth 46.4–56.1% of its length
.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cryptonemus
1b Two pairs of barbels; lateral line scales 37–
42; one prominent semicircular black mark on
middle of each caudal-fin lobe; a blackish band
across dorsal fin; a deep groove between cent-
ral callous pad and anteromedian fold on men-
tal adhesive disc; scaleless midventral region
restricted to chest or extended slightly beyond
midway from pectoral- to pelvic-fin origin; pre-
dorsal scales of same size as flank scales and
not embedded in skin; mental adhesive disc not
reaching beyond vertical of posterior margin of
eye; caudal peduncle depth 63.7–82.5% of its
length... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.
2a Scaleless midventral region merely restricted to
chest; 12–15 indentations on distal margin of
rostral fold; mental adhesive disc length 72.7–
80.7% of its width and 44.5–49.2% of HL;
caudal peduncle depth 63.7–71.3% of its length
... . . . . . . . . . . . . . . . . . . . . . . . . . . . P. caudofasciatus
2b Scaleless midventral region extended slightly bey-
ond halfway from pectoral- to pelvic-fin origin;
18–23 indentation on distal margin of rostral fold;
mental adhesive disc length 66.6–70.7% of its
width and 37.0–42.1% of HL; caudal peduncle
depth 72.4–82.5% of its length ............P. robustus
Acknowledgements
We are grateful to two reviewers, M. Kottelat and Fang
Fang, for their critical comments and making valuable
suggestions on this manuscript. We thank J. C. He
(YU) and G. H. Cui (KIZ) for permitting us to examine
the material under their care. Our thanks are also given
to J. X. Yang (KIZ), X. Y. Chen (KIZ) and Mai Dinh
Yen for their useful advices on this manuscript. Tech-
nical assistance by M. J. Cai is greatly appreciated
for drawing some of the illustrations. The project is
supported by Chinese National Natural Sciences Fund
No. 49832010 and No. 39870124 and also partially
supported by a grant for systematic and evolutionary
biology, Chinese Academy of Sciences.
References
Chen, J. X. & J. H. Lan, 1992. Description of a new genus and
three new species of fishes from Guangxi, China (Cypriniformes:
Cyprinidae, Cobitidae). Acta. Zootaxon. Sin. 17: 104–109 (in
Chinese).
Chen, Y. R., 1998. Labeoninae. In Chen, Y. Y. & Y. R. Chen (eds),
The Fishes of the Hengduan Mountains Region. Science Press,
Beijing: 168–183 (in Chinese).

Chu, X. L. & G. H. Cui, 1989. Labeoninae. In Chu, X. L. & Y.
R. Chen (eds), The Fishes of Yunnan, China. Part I. Cyprinidae.
Science Press, Beijing: 229–286 (in Chinese).
Cui, G.H. & Z.Y. Li, 1984. Description of a new cyprinid fish of the
subfamily Barbinae from China. Acta Zootaxon. Sin. 9: 110–112
(in Chinese).
Eschmeyer, W. N., 1998. Catalog of fishes. California Academy of
Sciences, San Francisco: 2905 pp.
Kottelat. M., 2001a. Freshwater Fishes of Northern Vietnam. Report
prepared for the world Bank, as part of the project Fresh-
water Biodiversity Overlay to Support the Vietnam national
Hydropower Study. World Bank, Washington: 122 pp.
Kottelat, M., 2001b. Fishes of Laos. Wildlife Heritage Trust Public-
ations, Colombo: 196 pp.
Lin, S. Y., 1931. Carps and carp-like fishes of Kwangtung and
adjacent inlands: 167 pp (in Chinese).
Mai, D. Y., 1978. Identification of Freshwater Fishes of North
Provinces of Vietnam. Scientific & Technology Publisher, Hanoi:
339 pp (in Vietnamese).
Menon, A. G. K., 1964. Monograph of the cyprinid fishes of the
genus Garra, Hamilton. Mem. Indian Mus. 14: 173–260.
Pellegrin, J. & P. Chevey, 1936. Poissons nouveaux ou rares du
Tonkin et de l’Annam. Bull. Soc. Zool. Fr. 61: 219–231.
Reid, G. M., 1982. The form, function and phylogenetic significance
View publication stats
217
of the vomero-palatine organ in cyprinid fishes. J. Nat. Hist. 16:
497–510.
Reid, G. M., 1985. A revision of African species Labeo (Pisces:
Cyprinidae) and a redefinition of the genus. Theses. Zoologicae,
Cramer, Braunschweig, 6: 1–322.
Roberts, T. R., 1982. Unculi (horny projection arising from single
cells), an adaptive feature of the epidemis of Ostariophysan
fishes. Zool. Scr. 11: 55–76.
Siebert, D. J., 1998. Revision of Schismatorhynchos Bleeker, 1855
(Teleostei, Cyprinidae), with the description of two new species
from Borneo. Bull. Hist. Mus. Lond. (Zool.) 64: 97–109.
Wu, H. W., R. D. Lin, J. X. Chen, X. L. Chen & M. Q. He, 1977.
Barbinae. In Wu, H. W. (ed.), The Cyprinid Fishes of China,
Volume 2. People’s Press, Shanghai: 229–394 (in Chinese).
Yue, P. Q. & Y. Y. Chen, 1998. Pisces. In Wang, S. (ed.), China Red
Data Book of Endangered Animals. Sciences Press, Beijing: 246
pp (in Chinese and English).
Zhang, E., 1997. A new replacement name for the genus Dis-
cocheilus Chen & Lan, 1992. Acta. Zootaxon. Sin. 22: 224 (in
Chinese).
Zhang, E., P. Q. Yue & J. X. Chen, 2000. Labeoninae. In Yue, P. Q.
(ed.), Fauna Sinica (Osteichthyes: Cypriniformes III). Science
Press, Beijing: 171–272 (in Chinese).