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94
Accepted by L. Page: 17 May 2018; published: 13 Sept. 2018
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4476 (1): 094
–
108
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4476.1.9
http://zoobank.org/urn:lsid:zoobank.org:pub:A0B33B41-1883-4509-A013-4D9165ED9678
Garra dengba, a new species of cyprinid fish (Pisces: Teleostei)
from eastern Tibet, China
SHU-QING DENG
1,2
, LIANG CAO
1,3
& E ZHANG
1
1
Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan 430072, Hubei Province, China
2
University of Chinese Academy of Sciences, Beijing 100049, China
3
Corresponding author. E-mail: caoliang0205@ihb.ac.cn; Phone: +86-027-68780233
Abstract
Garra dengba is here described from the Chayu-Qu, a tributary flowing into the Brahmaputra River, in Chayu County,
eastern Tibet, China. It shares the presence of an incipient proboscis on the snout with G. arupi, G. elongata, G. gravelyi,
G. kalpangi, and G. rotundinasus, but is distinguished from these five species in having, among other features, fewer
branched dorsal- and anal-fin rays and more perforated lateral-line scales. Its validity was also confirmed by a molecular
phylogenetic analysis based on the cytochrome b gene.
Key words: Garra, new species, morphology, cyt b gene, phylogenetic analysis
Introduction
The Chayu-Qu (= river) is one of largest rivers in eastern Tibet, China, rising in Kangri Garpo Range with the
highest peak of 5475 m above sea level. The upper reach of this river is separated by this mountain range into two
tributaries. The eastern one, also known as the Sang-Qu, has a total length of 178 km, a total fall of 4785 m in its
course, and a catchment area of 6780 km
2
. This tributary flows from northwest to southeast to Zhuwagen where it
turns south, passing through the Chayu County seat and then heading towards Xiachayu to join the western
tributary, Kangri Garpo-Qu. The western tributary, with its source in the southwestern side of Kangri Garpo, has a
total length of 170 km, a total fall of 3000 m in its course, and a catchment area of 5376km
2
. These two tributaries
merge to form the Chayu-Qu (or Lohit River) that flows south into India and turns southwest into the Brahmaputra
River basin.
The fish diversity of the Chayu-Qu is not well understood as the area drained by its lower reach is part of the
disputed Himalayan ranges between China and India. Indian authors have done some research regarding the
ichthyofaunal diversity of their claimed Arunachal Pradesh, which is referred to as Zangnan in China, whereas the
lower reach of the Lohit River basin is inadequately sampled (Gurumayum et al. 2016). The fish species diversity
of the Chayu-Qu was the subject of several Chinese authors (Wu et al. 1977; Wu et al. 1981; Wu &Wu 1992;
Zhang et al. 1995). Eight valid species have been documented from this river (Zhu 1989; Chu 1999; Yue 2000).
Among them are one loach (Nemacheilus subfuscus), three cyprinids (Garra kempi, Schizothorax curvilabiatus,
and Schizothorax molesworthi), and four catfishes (Exostoma labiatum, Glaridoglanis andersonii,
Pareuchiloglanis kamengensis, and Pseudecheneis sulcata). However, several species are poorly understood
taxonomically. Nemacheilus subfuscus was synonymized with Schistura scaturigina McClelland 1839 (Kottelat
2012), and Pareuchiloglanis kamengensis was referred to a new genus Creteuchiloglanis Zhou, Li & Thomson
(Zhou et al. 2011). It is also unveiled in the present investigation that specimens previously recognized by Chinese
authors as G. kempi from the Chayu-Qu actually represent an undescribed species, here named as G. dengba.
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A NEW GARRA SPECIES FROM EASTERN TIBET, CHINA
Materials and methods
Measurements were taken point-to-point with a dial caliper connected to a data recording computer and data
recorded to the nearest 0.1 mm. Measurements and counts were made on the left side of individuals when possible
following Zhang & Chen (2002), including counts of the lateral-line scales. Prepectoral, prepelvic, predorsal and
preanal lengths were measured from the tip of snout to the pectoral-, pelvic-, dorsal-, and anal-fin origins,
respectively. Disc width is the distance between the roots of two maxillary barbels; disc length is measured from
the anterior mid-point of the anterior margin to the posterior mid-point of the posterior margin of the mental
adhesive disc. Head width is the distance of the widest dimension with opercles closed; head depth is from the
midline at the occiput vertically downwards to the ventral contour of the breast; pelvic-to-anal distance is from the
pelvic-fin origin to the anal-fin origin. All morphometric data and meristic counts for specimens examined are
summarized in Table1. The number of specimens with a specific meristic count is included in parentheses after the
count. The anus-to-anal distance is given as a percentage of the pelvic-to-anal distance. Measurements of parts of
the head are expressed as proportions of the head length (HL). The head length and measurements of other parts of
the body are expressed as percentages of standard length (SL). The counts of dorsal- and anal-fin rays were taken
from radiographs. Other fin rays were counted by utilizing a dissecting microscope. Vertebral counts including the
Weberian apparatus were taken from radiographs following the method of Roberts (1989). The terms used in this
study for description of the snout morphology and their definitions follow Reid (1985), Zhang & Chen (2002) and
Nebeshwar & Vishwanath (2013).
Mitochondrial cytochrome b was selected for amplification and sequencing as it is the most widely used locus
in fish phylogenetics (Zardoya & Meyer 1996). The genomic DNA was extracted from alcohol-stored fin tissues.
The mitochondrial cyt b gene, around 1067bp (base pair), was amplified by the polymerase chain reaction in 25 µl
reactions containing 1.5 µl dNTPs, 2.5 µl reaction buffer, 0.5 µl of each primer, 0.5 µl of Taq DNA Polymerase, 1
µl template DNA, and 18.5 µl H2O. The PCR reaction followed Yang et al. (2010): 94°C predenaturing (3min),
94°C denaturing (45s), 55°C annealing (45s), 72°C extension (1 min), for 35 cycles and a final 72°C final
extension (7 min). Amplified products were subsequently purified and used for direct cycle sequencing by a
commercial sequencing company. The primer pairs used for amplification and sequencing were LA-cyp
ATGGCAAGCCTACGAA AAAC and HA-cyp TCGGATTACAAGACCGATGCTT (Tang et al. 2010).
The specimens used for molecular analysis and DNA sequences retrieved from GenBank are listed in Table 2.
Labeo stolizkae and Altigena lippa were selected as outgroups following Zheng et al. (2010) and Wang et al.
(2014); 22 congeners of Garra were used as ingroups for phylogenetic analysis. Multiple alignments were prepared
for all sequences using MEGA 5.0 (Tamura 2011). Then DNASP v5 was used to filter the haplotype (Librado
2009). Phylogenetic analyses were performed using MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003) with the
following model: GTR + I + G. The model was selected by Akaike’s information criterion (AIC), implemented in
jModeltest (Darriba 2012). Bayesian analyses were initiated from random starting trees, and four Markov Chain
Monte Carlo (MCMC) simulations were run simultaneously for 2,000,000 replicates, sampling one tree per 100
replicates for each run and the first quarter of the trees were discarded as burnin; the remaining trees from two
independent runs were used to construct a 50% majority rule consensus tree.
Samples of G. dengba were collected in fish investigations during 1973 and 2016 into a tributary flowing into
the Brahmaputra River basin, Chayu-Qu in Chayu County, eastern Tibet, West China. Some of them were
preserved in 10% formalin preservative for morphological examination and the others in 95% ethyl alcohol for
DNA extraction. A total of 33 specimens were used for morphological measurements and 7 specimens for
molecular analyses. All specimens were stored in the Freshwater Fish Museum at the Institute of Hydrobiology
(IHB), Chinese Academy of Sciences, Wuhan City, Hubei Province, China.
Garra dengba, sp. nov.
(Figs. 1–2)
Garra kempi: Wu et al., 1977: 378 (Chayu-Qu, eastern Tibet, West China); Wu & Wu, 1992: 293–294 (Chayu, eastern Tibet,
West China); Yue, 2000: 244–245 (Chayu, eastern Tibet, West China).
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TABLE 1. Morphometric measurements and meristic counts for Garra dengba sp. nov.
Holotype. IHB 2016032616, 108.6 mm SL; West China: eastern Tibet: Chayu County: Chayu-Qu, a tributary
flowing into Brahmaputra River; about 28°26'55''N 97°02'48''E, collected by Liang Cao, March 2016.
Paratypes. West China: eastern Tibet: Chayu County: IHB 2016032611–2615, 5 specimens, 80.6–97.1 mm
SL, Chayu-Qu; about 28°26'55''N 97°02'48''E, other data same as holotype; IHB 2016032617–2622, 6 specimens,
44.9–86.0 mm SL, Chayu-Qu; approximately 28°17'59''N 97°01'3'E, other data same as holotype.
G. dengba (n=33)
Holotype Range Mean SD
Standard length (SL) 108.6 42.9–108.6 70.0 18.03
Morphometric measurements
% SL
Body depth 15.1 15.1–20.2 18.1 1.44
Head length 20.6 18.8–26.5 22.3 1.60
Head width 17.1 16.3–20.1 18.5 0.83
Head depth 11.6 11.6–16.2 13.8 1.12
Caudal-peduncle length 14.7 13.7–17.5 15.7 1.04
Caudal-peduncle depth 10.4 9.9–12.5 11.0 0.62
Dorsal-fin length 17.6 16.1–25.3 19.9 1.89
Pectoral-fin length 17.0 17.0–23.0 19.6 1.68
Ventral-fin length 16.0 15.4–20.9 17.6 1.14
Anal-fin length 15.1 14.1–19.9 16.2 1.43
Predorsal length 47.1 44.2–52.2 47.8 1.86
Prepectoral length 18.1 17.3–24.6 20.8 1.95
Preventral length 49.5 47.5–56.4 51.3 1.93
Preanal length 75.4 72.4–80.8 76.0 1.86
% HL
Snout length 44.9 37.8–51.9 45.5 3.23
Eye diameter 15.2 13.3–28.4 19.7 3.50
Interorbital width 44.0 43.2–53.8 47.8 3.08
Disc length 49.1 38.2–51.0 43.9 2.79
Disc width 64.1 57.4–73.4 65.8 4.89
% caudal-peduncle length
Width of caudal peduncle 70.5 63.1–79.0 70.5 4.65
% pelvic to anal distance
Vent to anal distance 52.5 48.3–59.6 53.8 3.4
Meristic counts
Dorsal-fin rays iii, 6 iii, 6
Anal-fin rays iii, 4 iii, 4
Pectoral-fin rays i,13 i,11–13
Pelvic-fin rays i, 8 i, 8
Lateral-line scales 44 42–44
Scales above lateral line 4 3.5–4.5
Scales below lateral line 3 2.5–3.0
Predorsal scales 15 14–16
Circumpeduncular scales 12 12–14
Vertebrae 42 39–42
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A NEW GARRA SPECIES FROM EASTERN TIBET, CHINA
Non-type materials. West China: eastern Tibet: Chayu County: Chayu-Qu: IHB 73VII0063–0066,
73VII0147, 73VII0169–0171, 73VII0173–0175, 73VII 0224, 73VII 0284, 73VII0300, 14 specimens, 42.9–89.2
mm SL; about 28°36'22''N 96°54'44''E, collected by Wenxuan Cao, July 1973; IHB2016109150–9153, 4
specimens, 50.8–99.4 mm SL, about 28°22'41''N 97°02'20''E, collected by Liang Cao, October 2016;
IHB2016109154–9156, 3 specimens, 61.6–98.2 mm SL, about 28°40'47'' N 96°49'43''E, collected by Liang Cao,
October 2016.
Diagnosis. Garra dengba is distinguishable from all other Asian species of the genus except G. arupi, G.
elongata, G. gravelyi, G. kalpangi, and G. rotundinasus by possessing an incipient (vs. no or prominent) proboscis
on the snout. It is distinct from these five species in having 6 (vs. 7 or 8) branched dorsal-fin rays, 4 (vs. 5)
branched anal-fin rays, 42–44 (vs. 32–39) perforated lateral-line scales, no lateral black stripes on side of body (vs.
present, except in G. kalpangi), body depth 15.1–20.2% of SL (vs. 21.1–25.8% in G. arupi and G. gravelyi), 12–14
(vs. 15–16 in G. arupi and G. kalpangi) circumpeduncular scales, 14–16 (vs. 8–13) predorsal scales, no mid-lateral
black band on flank (vs. present in G. gravelyi and G. rotundinasus), no black spots along dorsal-fin base (vs.
present in G. gravelyi), no black band across dorsal fin (vs. present in G. arupi and G. elongata), no longitudinal
lateral black stripes on flank (vs. present, except in G. kalpangi), and scaled (vs. scaleless in G. elongata) breast.
Description. Morphometric data and meristic counts for type specimens are given in Table 1. Body elongate,
anteriorly cylindrical and posteriorly slightly compressed laterally, with greatest depth at dorsal-fin origin and least
caudal-peduncle depth slightly closer to caudal-fin base than to posterior end of anal-fin base. Dorsal profile of
head somewhat convex; profile of predorsal body almost straight or slightly convex, nearly straight or slightly
concave from dorsal-fin origin to origin of dorsal procurrent caudal-fin rays. Ventral profile of head straight to
oblique; nearly straight and flattened or somewhat convex from pectoral-fin insertion to anal-fin origin; concave
from there to origin of ventral procurrent caudal-fin rays.
Head moderately large and depressed, longer than wide and wider than deep. Eye small, positioned dorso-
laterally in upper half of head with slightly convex and broad interorbital space. Snout moderately rounded when
ventrally viewed, and obtuse when laterally viewed; with weakly-developed proboscis pointed forward and
reflected downward against snout in front of eyes; proboscis quadrate, delineated anteriorly from transverse rostral
lobe and laterally from lachrymal field (or lateral lobe) by ethmoid furrow (or depressed rostral surface). Two pairs
of barbels, shorter than eye diameter; rostral pair anterolaterally, maxillary ones hidden at corners of mouth, shorter
than rostral ones. Rostral cap well-developed, pendulous and greatly crenulated with papillated distal margin,
separated from upper jaw by deep groove, laterally continuous with lower lip around corners of mouth. Upper lip
absent; upper jaw fully covered by rostral fold, with thin horny sheath on cutting edge. Lower lip modified to form
mental adhesive disc. Disc elliptical, shorter than wide; anterior margin modified into transverse, fleshy and
crescentic skin fold covered by numerous minute papillae, anteriorly separated from lower jaw by deep groove
extending along entire length of lower jaw and posteriorly bordered in transverse deep groove with central callous
pad; lateral and posterior margins surrounding central callous pad in shape of half circle, papillated and distally
free; posterior margin reaching beyond vertical of posterior margin of eye.
Dorsal fin with 3 simple and 6 (33) branched rays, last one split to base; last simple ray slightly shorter than
HL; origin slightly nearer to snout tip than to caudal-fin base; distal margin slightly concave. Pectoral fin with 1
simple and 11 (1), 12 (24), or 13 (8) branched rays, reaching about two-thirds distance to pelvic-fin insertion;
shorter or equal to HL. Pelvic fin with 1 unbranched and 8 (33) branched rays, extending beyond midway to anal-
fin origin and surpassing anus, somewhat shorter than HL; inserted closer to anal-fin origin than to anterior end of
pectoral-fin base; positioned vertically at base of first or second branched dorsal-fin ray. Anal fin with 3 simple and
4 (33) branched rays, last one split to base; distal margin slightly concave; origin closer to pelvic-fin insertion than
to caudal-fin base. Caudal fin deeply forked; upper and lower lobes equal in length and shape.
Body scaled; scales moderately sized; scales on chest and belly smaller than those on flank and embedded.
Lateral line complete, horizontal, with 40 (20), 41 (8), or 42 (5) plus 2 scales on caudal-fin base; scale rows above
lateral line 3 (1), 3.5 (9), or 4 (23) and below 2.5 (15) or 3 (18). Predorsal midline scales almost same size as flank
scales and not embedded, regularly or irregularly arranged. Circumpeduncular scale rows 12 (32) or 14 (1).
Axillary scales 2; first one present at base of pelvic fin, and last one reaching beyond base of last pelvic-fin ray.
Anus positioned almost midway from pelvic- to anal-fin origin. Gas bladder bipartite, anterior chamber elliptical
and posterior chamber stick-like, as long as anterior one. Intestine long, forming coils. Vertebrae 39 (3), 40 (16), 41
(12), or 42 (2).
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FIGURE 1. Garra dengba sp. nov., IHB 2016032616, holotype, 108.6 mm SL. Lateral (a), dorsal (b) and ventral (c) views of
body. Scale bar = 1 cm.
FIGURE 2. Garra dengba sp. nov., IHB 2016032616, holotype,108.6 mm SL. (a) Dorsal view of head, and (b) ventral view of
mental adhesive disc. Scale bar = 1 cm.
Coloration. In fresh specimens, head and body yellowish dorsally and laterally, grayish on ventral surface. In
formalin-preserved specimens, head black or dark brown dorsally and laterally. Ground color of body dark or light
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A NEW GARRA SPECIES FROM EASTERN TIBET, CHINA
brown dorsally, becoming yellowish ventrally. Dorsal and caudal fins without dark chromatophores on distal
interradial membranes constituting small black blotches. Dorsal, pectoral, pelvic, and anal fins grayish white with
yellowish base.
Distribution. Known only from the Chayu-Qu (or Lohit River), a tributary flowing into the Brahmaputra
River basin, in Chayu County, eastern Tibet, West China (Fig 3).
FIGURE 3. Collection localities of Garra dengba sp. nov.. Black dots indicate sampling sites.
Habitat and biology. This new species was caught in the mainstem and tributaries of the Chayu-Qu where it
occurs in pebbly- and sandy-bottomed flowing water close to riverbanks (Fig. 4). Juveniles prefer shallow, slow-
running water with pebbly sand substrate along the bank. Macroscopic visual examination of the gonads from some
specimens caught in March showed the gonad maturation at the fourth stage, so indicating that this fish possibly
spawns in spring. It mainly feeds on algae and organic debris as found in the gut content of several specimens
examined.
Etymology. The specific epithet, herein utilized as a noun in apposition, is named after “Dengba”, the Chinese
name of Mishmi people who are not officially recognized as an ethnic group by the Chinese government. In China,
Dengba people are now living in the area between the eastern Himalaya Mountains and western Hengduan
Mountains at an altitude of 1000 meters above sea level. They have a concentrated distribution in Chayu County,
eastern Tibet where the type material of the new species was collected.
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FIGURE 4. Habitat of G. dengba. (a) Hill stream with shallow, slowly running water with pebbly sand substrate along the river
bank; (b) main stem with pebbly or sandy substrate, swiftly flowing water close to riverbanks.
TABLE 2. The mitochondrial cytochrome b sequences analyzed in this study.
Taxon Location Drainage Accession number
Garra dengba1 Chayu, Tibet, China Brahmaputra River MH243429
Garra dengba2 Chayu, Tibet, China Brahmaputra River MH243435
Garra dengba3 Chayu, Tibet, China Brahmaputra River MH243431
Garra dengba4 Chayu, Tibet, China Brahmaputra River MH243434
Garra dengba5 Chayu, Tibet, China Brahmaputra River MH243430
Garra dengba6 Chayu, Tibet, China Brahmaputra River MH243432
Garra dengba7 Chayu, Tibet, China Brahmaputra River MH243433
Garra imberba Xinping, Yunnan, China Red River KC119047
Garra nujiangensis Cangyuan, Yunnan, China Salween River KC119055
Garra findolabium Jinping, Yunnan, China Red River JQ864598
Garra fasciacauda Puer, Yunnan, China Lancang-Jiang JQ864597
Garra tengchongensis Tengchong, Yunnan, China Irrawaddy River JQ864586
Garra dulongensis Dulongjiang,Yunnan, China Irrawaddy River JQ864590
Garra qiaojiensis Yingjiang, Yunnan, China Irrawaddy River JQ864583
Garra salweenica Dehong, Yunnan, China Salween River JQ864593
Garra orientalis Longlin, Guangxi, China Red River JQ864581
Garra mirofrontis Yunxian, Yunnan, China Lancang-Jiang JQ864584
Garra bicornuta Peninsular India Thunga River JX074238
Garra ceylonensis Sri Lanka X074252
Garra cf annandalei Nepal Ganges River JX074253
Garra flavatra Myanmar Sea tributary AP011410
Garra fuliginosa Thailand Chao Phraya River JX074255
Garra gotyla Unknown Brahmaputra River JX074256
Garra kempi India Brahmaputra River JX074243
Garra lamta Nepal Ganges River JX074241
Garra lissorhynchus India Brahmaputra River JX074242
Garra mullya Peninsular India JX074237
Garra nasuta India Brahmaputra River JX074295
Garra spilota Myanmar Irrawaddy River AP011327
Labeo stolizkae Ruili, Yunnan, China Irrawaddy River GU086536
Altigena lippa Mengna, Yunnan, China Lancang-Jiang JX074198
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Sequence variation and molecular phylogeny
A total of seven sequences of cyt b (1067 bp) gene of G. dengba were successfully amplified; 22 sequences of
Garra congeners retrieved from GenBank, along with these seven sequences, were used as ingroups for
phylogenetic analysis. For the 1067 bp cyt b sequences, there were 628 conserved sites, 439 variable sites, 345
parsimony informative sites, and 94 singleton sites. The average frequency of four nucleotides of this new species
was A=29.4%, T=29.8%, C=27.0%, and G= 13.8%; the base composition was A-T rich (59.2%). The seven
sequences of G. dengba had 0.1–0.6% sequence divergence, with an overall intraspecific distance value 0.3%.
Interspecific genetic distance values among sampled species pairs of Garra were 6.4–21.3% (overall mean
distance 13.3%). The species under description had the minimum sequence divergence of 10.3% with G.
dulongensis, and the maximum one of 17.2% with G. lissorhynchys and G. i m b e r b a ; the mean genetic distance
values between it and other sampled congeneric species was 14.1 %.
The Bayesian 50% majority consensus tree based on cyt b gene for this new species and 22 congeneric species
is shown in Fig. 5. From the tree topologies, G. dengba was distantly related to G. k e m p i . It clustered with G.
dulongensis and G. tengchongensis, both from a tributary flowing into the Irrawaddy River basin, to form an
independent lineage with 100% posterior probability. Garra kempi was sister to G. s p i l o t a from the Irrawaddy
River basin in Myanmar; this pair grouped with G. flavatra and G. lissorhynchys to represent a lineage poorly
supported with < 50% posterior probability. The sister group of this lineage included two lineages: the one
including G. dulongensis, G. tengchongensis, and G. dengba and the other whose basal group was occupied by the
species pair G. gotyla and G. qiaojiensis.
TABLE 3. Main diagnostic characters for Garra dengba and five closely associated congeners with an incipient
proboscis on the snout.
Data from: a—Nebeshwar et al (2009); b—Vishwanath & Kosygin ( 2000); c—Kullander & Fang (2004); d—Menon
(1964); e—Zhang (2006); f—Nebeshwar et al (2012).
Discussion
This new species has until now been misidentified as G. kempi by Chinese authors (Wu et al. 1977; Wu & Wu
1992; Yue, 2000; Zhang & Chen 2002). The type locality of G. kempi is in the Siyom River below Dema, Abor
Hills in the Brahmaputra River basin (Hora 1921). It was documented by Menon (1964) and Talwar & Jhingran
(1992) from the Siyom River, and by Vishwanath (1993) from the Leimarkhong stream and tributaries of the Tuival
River of the Chindwin River basin in Manipur, India. Wu et al. (1977) were the first to record this species from the
Chayu-Qu in Chayu County, eastern Tibet, China. Their identification was followed by Wu &Wu (1992) to
document G. kempi from the river and by Zhang et al. (1995) from the Yarlung Zangbo-Jiang (the main
Brahmaputra River in China before it enters into India) in Motuo County. The historical distribution of the species
Characters G. dengba G. arupi G. elongata G. gravelyi G. kalpangi G. rotundinasus
Body depth in % SL 15.1–20.2 21.8–23.6
a
17.4–19.2
b
21.1–25.8
b
18.9–23.8
f
18.5–21.0
e
Branched dorsal-fin rays 6 7
a
7
c
7
d
8
f
8
e
Branched anal-fin rays 4 5
a
5
b
5
d
5
f
5
e
Lateral-line scales 42–44 35–36
a
41–42
b
34–36
b
32–33
f
36–37
e
Circumpeduncular scales 12–14 15–16
a
12
a
12
c
16
f
12
e
Predorsal scales 14–16 11–12
a
13
b
8–9
b
10–11
f
10–11
e
Mid-lateral black band on flank absent absent
a
absent
b
present
d
absent
f
absent
e
Black spots at dorsal-fin base absent absent
a
absent
b
present
d
absent
f
absent
e
Black band across dorsal fin absent present
a
present
b
absent
b
absent
f
absent
e
Lateral stripes on side of body absent present
a
present
b
present
d
absent
f
present
e
Scales on chest present present
a
absent
b
present
d
present
f
present
e
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also included the Daying-Jiang and Longchuan-Jiang, two tributaries discharging into the upper Irrawaddy River
basin in Yunnan, China (Chu & Cui 1989; Chen 1998). Actually, the species under the name of G. kempi from this
river basin was the misidentification of G. tengchongensis (Zhang & Chen 2002). Despite the recognition of the
specimens from the Yarlung Zangbo-Jiang basin in eastern Tibet, China as G. kempi by Chinese researchers, they
had no access to the type specimens of this species. The data utilized by Zhang & Chen (2002) for G. kempi was
from Menon (1964). The updated information about this species is available in both Nebeshwar et al. (2009) and
Nebeshwar & Vishwanath (2017). According to this information, G. k e m p i is a deep-bodied species (depth 20.7–
23.1% of SL) which develops a weakly developed transverse lobe delineated posteriorly by a shallow transverse
groove and no proboscis on the snout; 7–8 and 5 branched dorsal- and anal-fin rays, respectively; and 40–42
perforated lateral-line scales. By contrast, G. dengba has a slender body (depth 15.1–20.2 vs. 20.7–23.1% SL) with
a weakly developed or incipient proboscis on the snout, 6 branched dorsal-fin rays, 4 branched anal-fin rays, and
42–44 perforated lateral-line scales (see Table 1).
Garra dengba is characterized by having a weakly developed or incipient proboscis on the snout. This
character is shared with four congeners: G. elongata, G. gravelyi, G. kalpangi, and G. rotundinasus (Nebeshwar et
al. 2012). It is also present in G. arupi, a new species first described by Nebeshwar et al. (2009) from the upper
Brahmaputra River basin. In its original description, the species was referred to as a species without a proboscis on
the snout, but the accompanying image 3 (page 199) clearly illustrated an incipient proboscis with several tubercles
along the anterior margin before the nostrils, forwarded-pointed and reflected downward against the snout,
separated anteriorly from the tuberculated rostral field or transverse lobe and laterally from the infraorbital field by
a shallow ethmoid furrow (Reid 1985). Undoubtedly, G. arupi has an incipient proboscis in common with G.
dengba, G. elongata, G. gravelyi, G. kalpangi, and G. rotundinasus. A prominent proboscis on the snout can be
found in G. bispinosa, G. qiaojiensis, and G. orientalis (Zhang 2005: fig. 2) and also in G. arunachalensis, G.
birostris, G. cornigera, G. gotyla, G. koladynensis, G. quadratirostris, and G. trilobata (Nebeshwar & Vishwanath
2017: fig. 4). The proboscis of this kind is distinct, heavily tuberculated and more or less anteriorly free. See the
diagnosis and Table 3 for morphological differences among G. dengba and other five congeneric species with a
weakly-developed proboscis on the snout.
Garra rupecula was originally described by McClelland (1838) from the Mishmi Hills in the Brahmaputra
River basin. Menon (1964) designated it to the species group having a dark band across the dorsal fin, a W-shaped
band in the caudal fin, and naked chest and belly. Other species that can be put under this group are G.
lissorhynchus, G. ahboyai, G. nambulica, and G. paralissorhynchus (Vishwananth & Joyshree 2005) and G.
dampaensis (Lalronunga et al. 2013). It appears that G. rupecula, like all these species, possesses 4 branched anal-
fin rays. However, the description of the species by Menon (1964) was based on specimens collected from Manipur
valley (in the Chindwin River basin) and its distribution in this basin has been ruled out (Vishwanath &
Linthoingambi 2008). Garra rupecula is currently known only by McClelland’s (1838) very brief statement
(Lalronunga et al. 2013). According to its original description, it has 16 circumpeduncular scales, 35 perforated
lateral-line scales, and no scales on the chest, thus differing from the species under description.
Garra dengba has 4 branched anal-fin rays instead of 5 in most currently recognized species of Garra, or even
genera of the Labeonini. This character was found in other eight species: G. manipurensis by Vishwanath &
Sarojnalini (1988), G. nambulica by Vishwanath & Joyshree (2005), G. abhoyai, G. paralissorhynchus, and G.
lissorhynchus by Nebeshwar et al. (2012), G. dampaensis by Lalronunga et al. (2013), G. t y a o by Arunachalam et
al. (2014) and G. namyaensis by Shangningam & Vishwanath (2012) (see Table 4). This new species differs from
these eight species in having an incipient proboscis on the snout (vs. absent), and 42–44 (vs. 31–37) lateral-line
perforated scales; from all of them except G. namyaensis in having 12–14 (vs. 15–16) circumpeduncular scales;
from all of them except G. lissorhynchus in having 14–16 (vs. 18–29 in G. abhoyai or 8–12 in G. manipurensis, G.
paralissorhynchus and G. t y a o ) predorsal scales; from all of them except G. abhoyai and G. nambulica in having
rostral lobes (vs. absent); and from all of them except G. manipurensis in the absence (vs. presence) of a W-shaped
band on the caudal fin. It further differs from G. t y a o in having a shallower (vs. deeper) body (depth 15.1–20.2 vs.
21.2–29.6 % of SL), six (vs. seven) branched dorsal-fin rays; further from G. paralissorhynchus in having a
shallower (vs. deeper) body (depth 15.1–20.2 vs. 23.7–26.7 % of SL) and an anteriorly moved anus (vent-anal
distance 48.3–59.6 vs. 31.7–35.2 % of ventral-anal distance); further from G. manipurensis in possessing six (vs.
seven) branched dorsal-fin rays and a scaled (vs. scaleless) chest; further from G. lissorhynchus in lacking W-
shaped band on the caudal fin.
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FIGURE 5. Bayesian inference tree derived from cyt b gene for 23 species of Garra. Nodal numbers are posterior probability
values larger than 50%.
Kullander & Fang (2004) also found that five species of Garra from the Rakhine Yoma, southern Myanmar
possessed three or four branched anal-fin rays: G. propulvinus, G. vittatula, G. r a k h i n i c a , G. f l a v a t r a , and G.
poecilura. Garra dengba is distinct from these five species in having, among other features, no rostral lobe (vs.
present), a weakly developed proboscis (vs. absent), a shallower (vs. deeper) body (depth 15.1–20.2 vs 21.9–30.0%
of SL), more perforated lateral-line scales (42–44 vs. less than 35), more predorsal scales (14–16 vs. no more than
12), and fewer circumpeducular scales (12–14 vs. 16).
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TAB LE 4 . Main diagnostic characters for Garra dengba and other congeners with four branched anal-fin rays.
G. dengba
G. abhoyai
G. dampaensis
G. lissorhynchus
G. manipurensis
G. nambulica
G. namyaensis
G. paralissorhynchus
G. tyao
Body depth (% SL)
15.1–20.2
17.6–18.7a
20.1–22.7c
18.1–20.0d
20–21.2f
16.4–19.5g
19.0–21.0h
23.7–26.7b
21.2–29.6j
Vent-anal distance
% pelvic-anal distance
48.3–59.6
38.2–46.5a
15.9–19.6c
37.3–40.2d
24–26.4f
34.0–43.4g
28.0–33.3h
31.7–35.2b
about 30%j
Dorsal-fin rays
iii, 6
iii, 6a
ii,6c
iii, 6d
ii, 7f
ii,6g
ii,6.5h
iii, 6b
ii–iii, 7j
Anal-fin rays
iii, 4
iii, 4a
iii, 4c
iii, 4d
ii, 4f
ii,4g
ii,4–4.5h
iii, 4b
ii–ii, 3–4j
Lateral line scales
42–44
34–36b
27–29c
34–35d
34–35b
36–37g
31h
32–33d
31j
Predorsal scales
14–16
18–29b
10–11c
14–15d
10–11f
16–29g
13h
11–12i
8–10j
Circumpeduncular scales
12–14
16a
16c
16d
16f
16g
14h
16c
15–16j
Rostral lobe
absent
absenta
presentc
presentd
presentb
absentg
presenth
presentb
presentj
W-shaped black band
on caudal fin
absent
presenta
presentc
presentd
absentb
presentg
presenth
presenti
presentj
Dark band across dorsal fin
absent
presenta
presentc
presentd
absentf
presentg
presenth
presenti
presentj
A dark spot at upper
extremity gill-opening
absent
absenta
presentc
presente
absentk
presentg
presenth
presenti
presentj
Proboscis
present
absenta
absentc
absente
absentf
absentg
absenth
absenti
absentj
Scales on chest and belly
present;
absenta
presentc
absente
absent on chest
present on bellyf
absentg
presenth
absenti
presentj
Data from: a-Vishwanath & Linthoingambi (2008); b-Nebeshwar et al (2012); c-Lalronunga et al (2013); d-Nebeshwar et al (2009); e-Menon (1964); f-Vishwanath & Sarojnalini (1988);
g-Vishwanath & Joyshree (2005); h-Shangningam & Vishwanath (2012); i-Vishwanath & Shanta (2005); j-Arunachalam et al (2014); k-Nebeshwar, & Vishwanath (2017)
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TABLE 5. Genetic distances of cyt b computed by MEGA among 23 species.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
1. G. dengba
2. G. bico rnuta 0.142
3. G. ceylonensis 0.165
0.126
4. G. cf. annandalei 0.131
0.103
0.130
5. G. dulongensis 0.103
0.140
0.157
0.114
6. G. findolabium 0.159
0.169
0.165
0.128
0.148
7. G. flavatra 0.169
0.171
0.182
0.158
0.173
0.183
8. G. fuliginosa 0.119
0.133
0.143
0.108
0.132
0.156
0.168
9. G. gotyl a 0.156
0.140
0.139
0.129
0.131
0.144
0.155
0.111
10. G. imberba 0.172
0.174
0.189
0.147
0.164
0.129
0.194
0.149
0.158
11. G. kempi 0.126
0.139
0.157
0.112
0.124
0.131
0.167
0.115
0.129
0.129
12. G. lamta 0.128
0.136
0.148
0.098
0.124
0.144
0.163
0.092
0.119
0.156
0.107
13. G. lissorhynchys 0.172
0.159
0.184
0.166
0.159
0.187
0.116
0.161
0.157
0.213
0.183
0.173
14. G. mirofrontis 0.120
0.139
0.127
0.114
0.123
0.142
0.155
0.075
0.111
0.148
0.131
0.096 0.171
15. G. mullya 0.147
0.133
0.094
0.134
0.137
0.159
0.160
0.138
0.140
0.162
0.150
0.142 0.166
0.130
16. G. nasuta 0.136
0.147
0.147
0.115
0.120
0.155
0.170
0.101
0.118
0.158
0.126
0.077 0.176
0.105
0.142
17. G. nujiangensis 0.170
0.170
0.168
0.140
0.161
0.111
0.196
0.159
0.168
0.111
0.154
0.157 0.212
0.149
0.165
0.167
18. G. orientalis 0.114
0.134
0.126
0.101
0.122
0.146
0.160
0.073
0.104
0.143
0.120
0.089 0.165
0.065
0.129
0.109
0.145
19. G. qiaojiensis 0.144
0.148
0.147
0.129
0.128
0.142
0.153
0.116
0.075
0.166
0.122
0.112 0.169
0.104
0.127
0.113
0.159
0.104
20. G. salweenica 0.118
0.127
0.124
0.109
0.112
0.146
0.161
0.068
0.110
0.158
0.118
0.093 0.165
0.064
0.139
0.094
0.154
0.070
0.109
21. G. spilota 0.133
0.150
0.151
0.128
0.137
0.143
0.176
0.133
0.126
0.151
0.102
0.130 0.181
0.122
0.145
0.126
0.151
0.128
0.124
0.120
22. G. tengchongensis
0.119
0.158
0.153
0.116
0.122
0.151
0.180
0.128
0.133
0.181
0.154
0.121 0.187
0.128
0.142
0.123
0.155
0.121
0.138
0.125
0.143
23. G. fasciacauda 0.160
0.166
0.176
0.136
0.153
0.147
0.198
0.138
0.158
0.147
0.154
0.144 0.191
0.146
0.167
0.142
0.149
0.139
0.152
0.145
0.159
0.152
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Garra dengba is so far known only from the Chayu-Qu, a tributary discharging into the Brahmaputra River
basin where a total of 15 species from Garra have been identified as valid (Nebeshwar & Vishwanath 2017).
Among them, seven species have a snout with a proboscis and a transverse lobe in common with G. dengba,
namely G. arunachalensis, G. birostris, G. gotyla, G. kalpangii, G. n a s u t a , G. quadratirostris, and G. tamangi. The
new species is distinct from G. arunachalensis, G. birostris, G. g o t y l a , and G. quadratirostris, in terms of
Nebeshwar & Vishwanath (2013), in possessing more lateral scales (42–44 vs. 33–37), a more anteriorly positioned
anus (vent to anal distance 48.3–59.6 % of pelvic to anal distance vs. 19.0–44.0%), and a shallower (vs. deeper)
body (depth 15.1–20.2% of SL vs. 20.3–28.2%). Garra nasuta is a poorly understood species which is known only
by its original description (Nebeshwar & Vishwanath 2013). It is a pit between the nares and also likely a proboscis
with an anteriorly truncate margin and a transverse lobe. The first character can easily distinguish G. n a s ut a from G.
dengba. Judged from the illustration of McClelland (1838), reproduced as fig. 9 in Nebeshwar & Vishwanath
(2013), G. n a s u t a has a prominent proboscis on the snout instead of an incipient one in G. dengba. The new species
is distinguished from G. tamangi, according to Gurumayum et al. (2016), by having more perforated lateral-line
scales (42–44 vs. 33–34), fewer branched dorsal-fin rays (6 vs. 8–9), more predorsal scales (14–16 vs. 10–11) and
a shallower (vs. deeper) body (depth 15.1–20.2 vs. 20.7–22.3% SL).
The recognition of G. dengba as a valid species is corroborated by its marked sequence divergence from
sampled congeners (Table 5), and its monophyletic nature recovered in the phylogenetic trees inferred from cyt b
(Fig. 5). This new species had a significant sequence divergence from sampled congeners (10.3–17.2%). The BI
trees produced in this study clearly showed that G. dengba grouped with G. tengchongensis, G. dulongensis into a
monophyletic lineage where G. dengba and G. dulongensis were paired species sister to G. tengchongensis, and its
monophyly was well supported with 100% posterior probability. Given G. dulongensis and G. tengchongensis are
regarded as valid, the species status of G. dengba is warranted.
Acknowledgements
We would like to express our greatest gratitude to Yufan Wang (Zhejiang Forest Resource Monitoring Center),
Feng Lin (Guangxi Normal University), and Hao Jiang and Wei Ding (Huazhong Agricultural University) for
collecting specimens. Our grateful thanks also go to Waikhom Vishwanath (Manipur University, India), Seven O.
Kullander (Swedish Museum of Natural History) and Prachya Musikasinthorn (Kasetsart University) for providing
us with relevant literature. This study was supported by a grant from Chengdu Engineering Corporation Limited of
China.
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