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Accepted by L. Page: 10 Mar. 2011; published: 17 May 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2881: 39–50 (2011)
www.mapress.com/zootaxa/Article
39
Cophecheilus bamen, a new genus and species of labeonine fishes
(Teleostei: Cyprinidae) from South China
YU ZHU1, E ZHANG2,5, MING ZHANG3 & YAO-QUAN HAN4
1Fishery and Livestock Farming School of Guangxi, Nanning 530021, P.R. China
2Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan 430072, Hubei Province, P.R. China
3Nanjing Institute of Environmental Sciences, Ministry of Environmental Protection, Nanjing 210042, P.R. China
4Guangxi Institute of Fisheries, Nanling 530021, P.R. China
5Correspondent author. E-mail: zhange@ihb.ac.cn
Abstract
A new garrain genus and species are described from a tributary of the Zuo-River of the Pearl River drainage in Guangxi Prov-
ince, South China. Cophecheilus, new genus, is separated from all other Asian garrains by its uniquely modified oromandib-
ular morphology; i.e., rostral cap with a shallow, arched, subdistal depression extending almost the full length of its ventral
margin; upper lip greatly reduced to a thin membranous fold medially adnate to the upper jaw, but reflected slightly from its
distal margin, and laterally produced into a frenum connecting the upper jaw with the lower lip. The new species, Cophechei-
lus bamen, has an inconspicuous longitudinal black stripe extending along each side of the body.
Key words: taxonomy, Garraina, Pearl River drainage, Guangxi Province
Introduction
The Labeonini (sensu Reid, 1982; Stiassny and Getahun, 2007) is composed of a large number of cyprinid genera
widely known from the freshwaters of tropical Africa and Asia. These taxa show a high degree of morphological
modification of their oromandibular structures not shared with other cyprinid fish groups (Zhang et al., 2000). Cur-
rently, this group is represented in Southeast Asia and South China by approximately 29 genera (Zhang et al.,
2008). Diversity of the Labeonini at the generic rank is concentrated in South China where 25 genera are found,
accounting for approximately 85.2% of the total number censused by Yang and Mayden (2010). Twenty genera of
the Labeoninae (which is essentially equal to the Labeonini sensu Reid, 1982, and Stiassny and Getahun, 2007)
were documented in the recent monograph of Chinese species of the Cyprinidae (Zhang et al., 2000). The generic
diversity of Chinese labeonins continues to rise as a result of ongoing taxonomic research; e.g., Zhang and Chen
(2004, 2006), Zhang and Kottetat (2006), Zhang et al. (2006), Zhang et al. (2008), and Yang et al. (2008). Five other
genera were added by these researchers to the tally of Chinese labeonins, of which three are new to science,
Akrokolioplax, Qianlabeo and Hongshuia, and two are newly recorded, Bangana and Mekongia. Undoubtedly,
there are still several taxa awaiting formal description.
Among specimens captured in July, 2008, from a tributary of the Zuo-Jiang of the Pearl River drainage in Jingxi
County, Guangxi Province, South China, nine share all of the characters diagnostic for the Garraina as defined by
Stiassny and Getahun (2007), but are unidentifiable to any currently recognized Asian garrains. These nine specimens
are therefore referred herein to a new genus. The addition of Cophecheilus brings the total number of Chinese
Labeonini genera to 26.
ZHU ET AL.
40 · Zootaxa 2881 © 2011 Magnolia Press
Material and methods
Measurements were taken point to point with digital calipers that were linked directly to a data recording computer,
and data were recorded to the nearest 0.1 mm. Measurements and counts, made on the left side of individuals
whenever possible, followed those of Kottelat (2001). Predorsal, prepectoral, prepelvic and preanal lengths were
taken, respectively, from the anteriormost tip of the snout to the dorsal-, pectoral-, pelvic- and anal-fin origins.
Interorbital width was measured between the upper margins of the eyes. Abdominal vertebrae and caudal vertebrae
were counted from radiographs following the method outlined by Roberts (1989). The Weberian and urostylar
complex were included in the counts of the abdominal vertebrae and caudal vertebrae, respectively. The pharyngeal
teeth were counted and presented in a formula utilizing Hubbs and Lagler’s (1947) method, and the formula 2, 3,
5–5, 3, 2 indicates that the pharyngeal bones of both left and right sides bear three rows, with five teeth in the inner,
three in the middle, and two in the outer row. The number of specimens with a given meristic count is indicated in
parentheses after the count. All values for the holotype are indicated by asterisks in the text. Measurements of parts
of the head are presented as proportions of the head length (HL). The head length and measurements of other parts
of the body are given as percentages of the standard length (SL).
The mitochondrial 16S ribosomal RNA (rRNA) gene sequences for species of the subfamily Cyprininae
including the Labeonini are available from Genbank. This gene was thus selected for the molecular analysis. Mito-
chondrial DNA was extracted from one specimens of Cophecheilus bamen (IHBCY0857780). All tissues used for
DNA extraction were stored in 95% ethanol and deposited in the Freshwater Organism Museum of Institute of
Hydrobiology of Chinese Academy of Sciences (IHB). Total DNA extraction from muscles followed general pro-
tocols (Sambrook et al., 1989). The mitochondrial 16S rRNA gene was amplified using standard PCR techniques.
Thermal cycle amplifications were performed in 60 µL reaction volumes containing 6 µL 10 × buffer, 0.75 mM of
each dNTP, 1.5 U of Taq polymerase (Biostar), 3 µL of each primer, and roughly 100 ng of genomic DNA. Ampli-
fication proceeded with a primary denaturation step at 95ºC for 3 minutes, then 30 cycles of denaturation at 94°C
for 30 seconds, annealing at 58-62° C for 30 seconds and extension at 72°C for 60 seconds, with a final extension
of 5 minutes at 72° C. After amplification, a 4 µL sample of each PCR product was detected by 0.8% low-melting
agarose gels; the remaining product was purified using the BioStar Glassmilk DNA Purification Kit following the
manufacturer’s protocols and sequenced directly by United Gene Corporation. Two pairs of primers (16Sp1F: 50-
CTTACACCGAGAARACATC-30 and 16 Sp1R: 50-CTTAAGCTCCAAAGGGTC-30; 16Sp2F: 50-GACCTG-
TATGAATGGCTAA-30, and 16Sp2R: 50-TRGGAAGAGGATTTGAACC-30) were used to amplify and sequence
the mitochondrial 16S rRNA gene.
DNA sequences retrieved from GenBank and the specimens used for molecular analysis are listed in Table 1.
Although the 16S rRNA gene was sequenced by Zheng et al. (2010) for most of the Chinese labeonin species, these
sequences have not been released online. Thereby, most DNA sequences utilized here are from Li et al. (2005);
however, some of their species were incorrectly identified. The voucher specimen recognized by Li et al. (2005) as
Sinilabeo laticeps is Bangana lippa (see Zhang and Chen, 2006); the species identified as Crossocheilus latius is C.
burmanicus (see Kottelat, 2003); and Epalzeorhynchos bicornis is referred to Akrokolioplax (Zhang and Kottelat,
2006). DNA sequences were aligned by CLUSTALX v1.83 (Thompson et al., 1997) with the default parameters,
and the alignments were revised by eye in an effort to maximize the positional homology.
Phylogenetic analyses were performed using Bayesian inference (BI). BI was run with MrBayes 3.0 (Ronquist
and Huelsenbeck, 2003) to calculate posterior probabilities of recovered clades, with the optimal model of
sequence evolution determined from the likelihood-ratio tests (LRTs). In all Bayesian analyses, starting trees were
random, and one cold and three heated Markov chains were run simultaneously for 1x106 generations. Trees were
saved every 100 generations for a total size of 10,000 in the initial sample. Graphical inspection of tree log-likeli-
hood in this sample revealed that stationarity was reached within 100,000 generations. Thus, we discarded the first
100,000 generations (1000 sampled trees) as burn-in and used the remaining 900,000 generations (9000 sampled
trees) in all subsequent analyses. A majority rule consensus tree calculated from the 9000 remaining trees was used
to determine the posterior probabilities of clades.
The Chinese toponymy is employed for the distribution data, and the international English toponymy, if avail-
able, is also given in parentheses following the Chinese river name when it first appears in the present paper. The
data for the latitude and longitude coordinates of each locality were not provided in the original collection data, and
are thus inferred by the authors based on the best information available. The examined specimens are stored in the
Zootaxa 2881 © 2011 Magnolia Press · 41
COPHECHEILUS BAMEN
collection of the Freshwater Fish Museum at the Institute of Hydrobiology (IHB), Chinese Academy of Sciences,
Wuhan City, Hubei Province, China.
Table 1. Genbank accession numbers for 16S rRNA sequences used in this paper.
Cophecheilus, new genus
Type species: Cophecheilus bamen, new species
Diagnosis. Cophecheilus can be distinguished from all other genera of the Garraina (sensu Stiassny and Getahun,
2007) by the presence of a shallow, arched, subdistal depression extending almost the entire length of the ventral
margin of the rostral cap and partitioning this margin into a soft, slightly papillated anterior portion and a flexible,
cornified, medially slightly furrowed but non-fimbriate posterior portion, and an upper lip reduced to a thin membra-
nous fold which is medially adnate to the upper jaw but reflected slightly from its distal edge, and laterally produced
into a frenum connecting the upper jaw with the lower lip (Fig. 1). The genus is further separated by having a unique
combination of the following characters: lower lip partitioned into one median lobe and two lateral lobes by long
Taxon Voucher specimens Accession number
Myxocyprinus asiaticus IHBCY0305001 DQ845896
Procypris rabaudi IHBCY0308496 DQ845846
Sinocyclocheilus grahami IHBCY0410014 DQ845924
Sinocyclocheilus yangzongensis IHBCY0410008 DQ845926
Cyprinus carpio NC_001606
Spinibarbus sinensis IHBCY0403005 DQ845864
Spinibarbus hollandi IHBCY0205001 DQ845865
Schizothorax molesworthi IHBCY0305085 DQ845848
Schizothorax labiatus IHBCY0380470 DQ845858
Barbus barbus DQ845879
Onychostoma simum IHBCY0306001 DQ845861
Schizopygosis pylzovi IHBCY0308716 DQ845856
Platypharodon extremus IHBCY0308702 DQ845855
Oxygymnocypris stewartii IHBCY0510084 DQ845918
Labeo yunnanensis IHBCY0301133 DQ845881
Garra kempi IHBCY0309091 DQ845885
Garra orientalis IHBCY0403441 DQ845884
Epalzeorhynchus frenatus IHBCY0408007 DQ845905
Labiobarbus lineatus IHBCY0407001 DQ845914
Osteochilus salsburyi IHBCY0308001 DQ845892
Sinilabeo lippa IHBCY0210030 DQ845904
Placocheilus cryptonemus IHBCY0504726 DQ845915
Discogobio bismargaritus hap1 IHBCY0308821 DQ845900
Discogobio bismargaritus hap2 IHBCY0308734 DQ845890
Discogobio tetrabarbatus IHBCY0308003 DQ845888
Discogobio laticeps IHBCY0308942 DQ845889
Rectoris posehensis IHBCY0204016 DQ845891
Pseudogyrinocheilus prochilus IHBCY0405017 DQ845894
Parasinilabeo assimilis IHBCY0308002 DQ845887
Ptychidio jordani IHBCY0308004 DQ845893
Pseudocrossocheilus bamaensis IHBCY0509003 DQ845895
Crossocheilus burmanicus IHBCY0308005 DQ845882
Akrokolioplax bicornis IHBCY0505291 DQ845919
Discogobio sp. IHBCY0308882 DQ845901
Lobocheilus melanotaenia IHBCY0405266 DQ845902
Garra gravelyi DQ845907
Cirrhinus molitorella DQ845883
Semilabeo notabilis DQ845886
Cophecheilus bamen IHBCY 0857780 DQ1380940
ZHU ET AL.
42 · Zootaxa 2881 © 2011 Magnolia Press
postlabial grooves extending anteromedially close to the anterolateral margin of the median lobe, with a gap between
their anterior margins wider than one-third of the mouth gape; many lobate papillae scattered over anterior part of
median lobe of lower lip; lower jaw separated from lower lip by a deep groove along its entire length, with a thick,
flexible, cornified sheath on its anterior margin; rostral cap completely overlying upper jaw and laterally connected
with lower lip around corners of mouth; mouth opening arched, width greater than three-fourths of corresponding
head width; three rows of pharyngeal teeth; two pairs of well-developed barbels, antero-rostral position of rostral
barbels; and eight branched dorsal-fin rays.
Table 2 Morphometric data for Cophecheilus bamen.
Etymology. The generic name is a Latinized combination of the Greek words kpheia (depression) and cheilos
(lips), the latter a commonly used suffix for Labeonin genera, alluding to the presence of a shallow, arched, subdis-
tal depression along the ventral margin of the rostral cap. Gender: masculine.
FIGURE 1. Diagrammatic illustrations of oromanibular structures of Cophecheilus bamen. Ventral view. The rostral cap is
removed in the right side. drc = depression on ventral subdistal margin of rostral cap; fr = frenum connecting upper and lower
lips; lj = lower jaw; lll = lateral lobe of lower lip; mb = maxillary barbel; mll = median lobe of lower lip; pg = postlabial groove;
rb = rostral barbel; rc = rostral cap; uj = upper jaw; ul = upper lip.
Characters Holotype Paratypes (8)
Range Mean SD
Standard length (mm) 105.7 65.3–84.5 78.8 11.49
Percentage of standard length
Body depth 26.7 25.7–31.0 27.4 1.45
Head length 22.3 22.6–24.8 23.4 0.76
Caudal-peduncle length 16.2 15.8–19.0 17.3 0.98
Caudal-peduncle depth 12.8 12.5–13.9 13.0 0.45
Dorsal length 24.7 21.4–23.5 22.8 1.01
Pectoral length 22.0 10.3–23.0 21.5 1.17
Ventral length 19.4 17.3–20.9 18.7 1.18
Anal length 17.3 16.0–19.1 17.7 0.99
Predorsal length 49.8 48.3-51.8 49.5 1.14
Prepectoral length 22.7 21.8–24.1 22.8 0.69
Preventral length 55.3 50.1–54.6 52.6 1.50
Preanal length 77.4 71.0–77.4 74.7 1.97
Percentage of head length
Head depth 67.2 63.2–70.0 65.9 2.14
Head width 67.6 62.1–68.3 64.8 2.13
Snout length 36.9 32.1–37.5 36.1 2.21
Eye diameter 20.4 20.2–25.0 22.5 1.59
Postorbial length 51.3 44.2–50.6 48.1 2.32
Zootaxa 2881 © 2011 Magnolia Press · 43
COPHECHEILUS BAMEN
Cophecheilus bamen, new species
(Figs. 2a, 3; Table 2)
Holotype: IHB 2000800009, 105.7 mm SL, a tributary flowing into Zuo-Jiang of Zhu-Jiang (Pearl River) drainage
at Jingxi County, Guangxi, South China; coll. J.H. Lan July, 2008.
Paratypes: IHB 2000800010–7, 8, 65.3–84.5 mm SL, same data as holotype.
Diagnosis. See the generic diagnosis.
Description. Morphometric data for nine type specimens are given in Table 2. Body elongate, compressed lat-
erally, with greatest depth at dorsal-fin origin and least depth of caudal peduncle slightly closer to caudal-fin base
than to posterior end of anal-fin base. Dorsal profile of body gradually ascending from snout tip to dorsal-fin ori-
gin; from there to origin of dorsal procurrent caudal-fin rays profile slightly concave. Ventral profile from tip of
snout to pectoral-fin insertion nearly straight; from there to anal-fin origin convex; anal-fin base straight; from pos-
terior end of anal-fin base to origin of ventral procurrent caudal-fin rays slightly concave.
FIGURE 2. Ventral view of oromandibular structures in: (a) Cophecheilus bamen, IHB 2000800009, holotype, 105.7 mm SL;
(b) Semilabeo notabilis, IHB 86082394, 167.1 mm SL, Hongshui-He of Pearl River drainage at Bama County, Guangxi Prov-
ince; (c) Pseudogyrinocheilus prochilus, IHB 79III0068, 185.6 mm SL, upper Yangtze River drainage at Yibing City, Sichuan
Province; (d) Parasinilabeo assimilis, IHB 75IV2445, 93.3 mm SL, Pearl River drainage at Xiuyun County, Guangxi Province.
ZHU ET AL.
44 · Zootaxa 2881 © 2011 Magnolia Press
Head relatively small, longer than wide. Eye medium-sized with a wide, slightly convex interorbital space, dor-
solaterally placed in anterior half of head. Snout blunt in lateral view and pointed in dorsal view, with a shallow
groove running along anteroventral border of lachrymal from rostral-barbel base to lateral extremity of rostral cap;
no tubercles on its tip and anterior portion of lachrymal. Two pairs of well-developed barbels; rostral barbel posi-
tioned at anterior end of shallow sublachrymal groove on each side of snout and extending to posterior margin of
eye or not; maxillary barbel rooted in corner of mouth, originating from exterior surface of confluence between lat-
eral lobe of lower lip and lateral extremity of rostral cap, and extending beyond posterior margin of eye to opercle.
Mouth inferior and arched.
FIGURE 3. Lateral view of Cophecheilus bamen, IHB 2000800009, holotype, 105.7 mm SL.
FIGURE 4. (a) Left side of pharyngeal tooth plate in dorsomedial view, and (b) gas bladder in ventral view, of Cophecheilus
bamen, IHB 2000800009, holotype, 105.7 mm SL.
Zootaxa 2881 © 2011 Magnolia Press · 45
COPHECHEILUS BAMEN
Dorsal-fin with four unbranched and 8* (9) branched rays, last one split to base; origin slightly nearer to tip of
snout than to caudal-fin base; last unbranched ray longer than HL; distal margin slightly concave. Pectoral fin fal-
cate, with one unbranched and 14 (1), 15* (6) or 16 (2) branched rays, inserted posterior to vertical through poste-
rior-most point of opercle, reaching three-fourths of distance to pelvic-fin insertion and as far as third or fourth
scale anterior to pelvic-fin insertion. Pelvic fin with one unbranched and 7* (9) branched rays, inserted vertically
posterior to second branched dorsal-fin ray base, and extending to vent, but not to anal-fin origin. Anal fin with
three unbranched and 5* (9) branched rays, last one split to base; distal margin slightly concave; origin placed equi-
distant between pelvic-fin insertion and caudal-fin base. Caudal deeply forked; upper and lower lobes equal in
length and shape, with tapering rounded tips.
Body scales moderately sized; chest and belly scaled, scales slightly smaller than ones on flank. Lateral line
complete, horizontal 43* (6), 44 (1) or 45 (1) plus three scales on caudal-fin base; scale rows above lateral line 7
(3) or 8* (6) and below 6 (3) or 7* (6). Predorsal scales 16 (4) or 17* (5), smaller than ones on flank, and anteriorly
embedded under skin. Circumpeduncular scale rows 22* (9). Axillary scale present at pelvic-fin base, extending
beyond base of last ray. Two scales between vent and anal-fin origin.
Vertebrae 41–42 (6) [26+15 = 41* (1), 25+16 = 41 (2), 26+16 = 42 (2), or 25+17 = 42 (1)]. Pharyngeal teeth
triserial; tooth pattern 2, 3, 4–4, 3, 2 (1), with compressed and pointed tips (Fig. 4a). Gas bladder bipartite; anterior
chamber oval, wider than posterior one; posterior chamber stick-like or elongate with an enlarged, rounded distal
end, about twice as long as anterior one (Fig. 4b). Intestine long, forming many coils. Gill rakers on outer side of
first gill arch 15–17 (4), short and small.
FIGURE 5. Map showing the distribution of Cophecheilus bamen in Guangxi Province of South China.
ZHU ET AL.
46 · Zootaxa 2881 © 2011 Magnolia Press
Color pattern in formalin. Top of head grayish brown, snout and lip grayish; cheek with sparse brownish pig-
ments; opercle and subopercle brownish. Ground color of body grayish brown dorsally and laterally, yellowish
white ventrally. An indistinct longitudinal black stripe, about one and half scales in width, originating near upper
extremity of gill opening and terminating at caudal-fin base in a vertical bar extending along first scale row directly
above lateral line anterior to vertical through anal-fin origin, thereafter along lateral line to caudal-fin base. Each
scale on flank with dark chromatophores along exposed part of its posterior margin constituting a dark, crescentic,
vertically-elongate mark; marks on scales above lateral line and directly below more prominent. Dorsal and caudal
fins with dark chromatophores on fin rays, giving them a dusky appearance, and pectoral and pelvic fins with dark
chromatophores on fin rays, more densely on anterior fin rays, giving them a dusky appearance. Anal fins hyaline.
Distribution. Known only from a tributary flowing into the Zuo-Jiang of the Pearl River drainage at Jingxi
County, Guangxi Province, South China (Fig. 5).
Etymology. The specific epithet, used as a noun in apposition, is made from Ba Men, the local name of this and
similar species.
Discussion
Stiassny and Getahun (2007) provided morphological evidence for the monophyletic nature of the Garraina as well as
the Labeonini. Synapomorphic characters for the Garraina include: mediation of contact of neural complex with
neurocranium; fimbriate or invecked ventral margin of rostral cap; antero-rostral position of rostral barbels; papil-
lae densely scattered over ventral margin of rostral cap and lower lip; upper lip fully regressed and absent medially,
and upper lip frenum attaching to lower lip and /or rostral fold cap usually retained at lateral margin of lower jaw.
These characters are shared with Cophecheilus, which is referred to the Garraina.
The Garraina, as delimited by Stiassny and Getahun (2007), includes the following Chinese genera: Crossochei-
lus, Hongshuia, Mekongina, Akrokolioplax, Rectoris, Parasinilabeo, Ptychidio, Pseudocrossocheilus, Pseudogyri-
nocheilus, Sinocrossocheilus, Garra, Semilabeo, Discogobio, Discocheilus, and Placocheilus. Qianlabeo and
Sinilabeo should be placed in the Garraina as well. Zhang and Chen (2004) held that Qianlabeo likely represents an
intermediate form between the Labeonini (= Labeonina) and the Garrini (= Garraina). Zhang et al. (2006) and
Zhang et al. (2008) advocated that Sinilabeo and Qianlabeo formed a third subgroup of the Labeonini according to
the morphology of oromandibular structures. Stiassny and Getahun (2007) allocated these two genera to the
Labeoina in terms of literature information as materials were unavailable for their study; their data for Sinilabeo
did not come from Zhang et al. (2006), who indicated that the generic name was previously misapplied to Chinese
species referred to Bangana. Our external and radiographic examinations of the type materials the type species for
Qianlabeo and Sinilabeo revealed that both have all aforementioned characters diagnostic for the Garraina, thus
suggesting that they should be placed in the subtribe. This placement is partially confirmed by Zheng et al.’s (2010)
molecular phylogenetic analysis of the Chinese Labeoninae in which Qianlabeo was shown to be closely allied to
Chinese garrains such as Pseudocrossocheilus, Semilabeo, Ptychidio, Pseudogyrinocheilus, Rectoris and Parasini-
labeo.
Within the Garraina, Cophecheilus has unusual oromandibular structures (Fig.1). A combination of the following
five main characters clearly separates the genus from all other East Asian genera of this subtribe: (1) rostral cap with
a shallow, arched, subdistal depression extending nearly the full length of its ventral edge; (2) upper lip greatly
reduced to a thin membranous fold medially adnate to upper jaw but reflected slightly from its distal edge and later-
ally produced into a frenum directly connecting the upper jaw and lower lip; (3) lower lip partitioned into one median
and two lateral lobes by prolonged post-labial grooves extending anteromedially close to the anterolateral margin of
the median lobe, with a gap between their anterior margins wider than one-third of the mouth gape; (4) two pairs of
barbels well-developed; and (5) a rostral fold with a non-fimbriate ventral margin. The first two characters are unique
among the Garraina or Labeonini and, thus, diagnostic for Cophecheilus. An analogous depression is seen only in
Linichthys, an endemic Chinese barbine cyprinid genus so far known only from the upper and middle Pearl River
drainage, and the upper Yangtze River drainage in Guiyang City, Guizhou Province, South China (Zhang and Fang,
2005). In this taxon, the depression is present on the upper lip instead of the rostral cap. Given the distant relationship
of this barbin, we interpret the acquisition of this depression in these two genera to represent homoplasy under shared
selection pressure. The upper lip exhibited by Cophecheilus is distinct from Kullander and Fang’s (2004) observa-
Zootaxa 2881 © 2011 Magnolia Press · 47
COPHECHEILUS BAMEN
tions for Myanmarese Garra species where the upper lip forms a narrow band of papillated folds along the upper jaw,
or irregular transverse ridges or patches away from the mouth, and from Stiassny and Getahun’s (2007) observations
for garrains where the upper lip is reduced to a very thin membranous fold that may either be completely adnate to
the proximal margin of the keratinized upper jaw in adults, or overhangs ventrally as a short membranous fringe
fully overlain by the rostral cap.
Prolonged postlabial grooves are found in Qianlabeo, extending anteromedially close to the anteriormost end of
the midline of the lower lip, but not meeting with each other (Zhang and Chen, 2004: 30, fig. 1C). In contrast, post-
labial grooves of Cophecheilus terminate immediately posterior to the anterior margin of the lower lip in a gap
between their anterior margins wider than one-third of the mouth gap (Fig. 1). Other differences between these two
genera are: eight (vs. seven in Qianlabeo) branched dorsal-fin rays, and many lobate papillae anteriorly scattered
over the median lobe of the lower lip (vs. absent).
Two pairs of well-developed barbels are shared with four other Chinese garrain genera: Pseudocrossocheilus,
Sinocrossocheilus, Hongshuia and Qianlabeo, and with Parasinilabeo longibarbus Zhu, Lan and Zhang, 2006.
Pseudocrossocheilus (Zhang and Chen: 2004: 30, fig. 1F) has a fimbriate, greatly papillated (vs. non-fimbriate,
slightly papillated) rostral cap, no upper lip (vs. present but greatly regressed) and the postlabial groove restricted
only to the side of the lower jaw (vs. prolonged and extending anteromedially close to the anterolateral margin of the
median lobe of the lower lip), thereby differing from Cophecheilus. Both Sinocrossocheilus and Hongshuia contrast
sharply with Cophecheilus in lower lip morphology. The median lobe of the lower lip is produced into a densely
papillated, greatly protruded, crescentic anterior fold and a papillated, somewhat protruded, triangular fleshy poste-
rior pad posteriorly continuous with the mental region in Sinocrossocheilus (Yuan et al., 2008: 38, fig. 1A). This
lobe is modified into a round, fleshy plate peripherally greatly protruded to form a ring-like fold, and centrally
sunken to form a round, flat fleshy pad which is posteromedially continuous with the mental region in Hongshuia
(Zhang et al., 2008: 35, fig. 1A). These two genera further differ from Cophecheilus in having a rostral cap laterally
disconnected from (vs. connected with) the lower lip at the corners of the mouth, two (vs. three) rows of pharyngeal
teeth, and seven (vs. eight) branched dorsal-fin rays. Here, it is worthwhile to point out that Sinocrossocheilus sensu
Su, Yang and Cui, 2003, is a catch-all genus including some species of other garrain genera, e.g. Pseudocrossocheilus
and Hongshuia (Yuan et al., 2008). Inconsistent information about the state of the rostral cap of Sinocrossocheilus
in the literature, as pointed out by Stiassny and Getahun (2007), has been clarified (Yuan et al., 2008). This genus,
as currently conceived, consists of two species, S. guizhouensis and S. labiatus, known only from the Wu River of
the upper Yangtze River drainage in Guizhou Province, South China. The shared presences of long rostral barbels
extending beyond the midpoint of the eye can easily separate Cophecheilus bamen and Parasinilabeo longibarbus
from all other garrain species. These two genera are distinguished below.
A non-fimbriate ventral margin of the rostral cap is present in four garrain genera: Semilabeo, Pseudogyrinochei-
lus, Parasinilabeo (Fig.2b-d) and Discocheilus (Zheng et al., 2010:11, fig.6B). They have a densely papillated ventral
margin on the rostral cap, a densely papillated lower lip, and a thin, flexible horny sheath on the anterior margin of
the lower jaw posteriorly continuous with the anterior edge of the lower lip (Zhang, 2005: 137); all three of these
characters distinguish them from Cophecheilus. Stiassny and Getahun (2007) stated: “Even in those taxa in which
the upper lip is fully regressed and absent medially, an upper lip frenum attaching to the lower lip and/or rostral cap
is usually retained at the lateral margin of the lower jaw.” An upper lip frenum occurs in Cophecheilus and in other
four genera, Pseudocrossocheilus, Sinilabeo, Qianlabeo and Rectoris. Contrary to Stiassny and Getahun’s (2007)
observations, however, the frenum is missing in Pseudogyrinochilus, Discocheilus and Semilabeo. Discocheilus pos-
sesses biserial (vs. triserial) pharyngeal teeth and a mental adhesive disc modified from the lower lip (see Zhang,
2005: 137, fig. 11C) (vs. absent), thereby differing from Cophecheilus. The mouth gape is less than half of the corre-
sponding head width in Semilabeo and Pseudogyrinocheilus, and the dorsal fin has seven branched rays in Parasini-
labeo. In contrast, Cophecheilus has a wide mouth, with its width more than three-fourths of the corresponding
head width, and eight branched rays in the dorsal fin.
Despite morphological evidence provided by Stiassny and Getahun (2007) in support for the monophyletic
nature of the Garraina, little is known about phylogenetic relationships within the subtribe at the generic level. In
this context, it is not possible to reach a decision about the phylogenetic position of Cophecheilus among Chinese
or East Asian garrain genera. Likely, this genus, along with Qianlabeo and Sinilabeo, assumes the basal position in
the Garraina. All three genera possess a slightly crenulated or non-crenulated, slightly papillated rostral cap and a
greatly reduced upper lip, laterally produced into a frenum directly connecting the upper jaw and the lower lip. Most
ZHU ET AL.
48 · Zootaxa 2881 © 2011 Magnolia Press
garrain genera have a densely papillated, crenulated rostral cap and no upper lip frenum. A phylogenetic analysis of
all garrain genera based on morphological and/or molecular data is essential to test this proposed relationship.
FIGURE 6. Bayesian inference tree derived from 16S rRNA for all species. The nodal numbers are posterior probability val-
ues. Only values above 50% are given.
Recently, molecular phylogenetic analyses of the Cyprinidae and one of its subgroups, the Cyprininae, from
East Asia, provided strong support in favor of the monophyly of the Labeonini (Li et al., 2005; Wang et al., 2007;
Li et al., 2008). The same conclusion was reached by Yang and Mayden (2010) and Zheng et al. (2010) in their
molecular phylogenetic analyses of the Labeoninae. However, no clade recovered in these analyses matches Stia-
ssny and Getahun’s (2007) concept of the Garraina. Our molecular phylogenetic analysis with the addition of
Cophecheilus bamen to the dataset utilized by Li et al. (2008) placed the species with the monophylectic group
including Rectoris, Parasinilabeo, Semilabeo, Pseudogyrinocheilus, Ptychidio and Pseudocrossocheilus, in which
Cophecheilus is sister to Semilabeo in the molecular tree (Fig.5). The close relationships of Cophecheilus with
some garrain genera such as Pseudogyrinocheilus, Parasinilabeo, Semilabeo and Pseudocrossocheilus are also
indicated by their similarities of oromandibular structures (see above for details). However, this is not the final
decision about the phylogenetic position of Cophecheilus, given that the Labeonini (sensu Yang and Mayden,
Zootaxa 2881 © 2011 Magnolia Press · 49
COPHECHEILUS BAMEN
2010) includes about 400 species in 34 genera widely distributed in the fresh waters of tropical Africa and Asia,
and few species (representing no more than 17 genera) were sampled by Li et al. (2005) and Li et al. (2008). The
phylogenetic position of Cophecheilus among other garrain or labeonin genera needs to be further investigated
based on sampling more included species and genera.
Comparative material
Discocheilus wui: IHB 89XII0023, 89XII0008, 89XII0010, 3, 60.5–66.8 mm SL, Hongshui-He of Pearl River
drainage at Donglan County, Guangxi Province; IHB 85VIII068–70, 3, 36.5–44.5 mm SL, Hongshui-He of Pearl
River drainage at Tian’e County, Guangxi Province. Hongshuia paoli: IHB 200210911 (holotype), 54.3 mm SL;
Hongshui-He of Pearl River drainage at Fengshan County, Guangxi Province. Parasinilabeo assimilis: IHB
75IV229–32, 75IV626–7, 6, 78.0–93.0 mm SL, Pearl River drainage at Xiuyun County, Guangxi Province. Paras-
inilabeo longibarbus: IHB 20024051387 (holotype), 2002051405–7, 2002051413, 2002051410, 2002051424–5, 8,
43.9–53.3 mm SL, Gui-Jiang of Pearl River drainage at Fuchuan County, Guangxi Province. Pseudocrossocheilus
bamaensis: IHB 743119, 743136, 73422, 76092, 4, 100.0–108.0 mm SL, Pearl River drainage, Guangxi Province.
Pseudocrossocheilus liuchengensis: IHB 82VII1521–3, 82VII1528, 4, 68.0–81.0 mm SL, Pearl River drainage,
Guangxi Province. Pseudogyrinocheilus prochilus: IHB 87V051–3, 3, 116.8–162.3 mm SL, Wu-Jiang of upperY-
angtze River drainage at Guiding County, Guizhou Province; IHB 79III0068, 185.6 mm SL, upper Yangtze River
drainage at Yibing City, Sichuan Province. Qianlabeo striatus: IHB 800858 (holotype), 800933, 800879, 8000892,
800917, 5, 57.2–67.5 mm SL, Beipan-Jiang of Pearl River drainage at An’shun County, Guizhou Province. Recto-
ris luxiensis: IHB 87IV0023, 87IV0032, 87IV0018, 87IV0020, 4, 94.0–135.1 mm SL, upper Yangtze River drain-
age at Dachang County, Sichuan Province. Rectoris posehensis: IHB 75IV1068-73, 81XII0281, 7, 94.5–153.4 mm
SL, Pearl River drainage at Longzhou County, Guangxi Province. Semilabeo notabilis: IHB 75IV1050, 81XII0204,
81XII 0281, 4, 101.9–144.1 mm SL, Pearl River at Longzhou County, Guangxi Province; IHB 86082394, 167.1
mm SL, Hongshui-He of Pearl River drainage in Bama County ,Guangxi Province. Sinilabeo hummali: IHB
2003051 (holotype), 120.0 mm SL, Jialing-Jiang of upper Yangtze River drainage at Chongqing City. Sinocros-
socheilus guizhouensis: IHB 6650415–6, 6650503–5, 6650508 (lectotype), 6, 53.0–76.0 mm SL, Wu-Jiang of
upper Yangtze drainage at Zunyi City, Guizhou Province.
Acknowledgments
This work was supported by a grant from the National Natural Sciences Foundation of China (NSFC No.
30970323). We are indebted to J. Zhou (Guangxi Institute of Fisheries, Nanling) and J.H. Lan (Fishery Bureau of
Du’an County, Guangxi) for their help with the field survey.
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