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Accepted by L. Page: 27 Jul. 2015; published: 25 Aug. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 4006 (3): 551
–
568
www.mapress.com
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zootaxa
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Article
551
http://dx.doi.org/10.11646/zootaxa.4006.3.8
http://zoobank.org/urn:lsid:zoobank.org:pub:CBDC7284-7EF2-49F4-A875-13E79B597E0A
Mahseers genera To r and Neolissochilus (Teleostei: Cyprinidae) from southern
Vietnam
HUY ĐỨC HOÀNG
1,4
, HÙNG MẠNH PHẠM
1
, JEAN-DOMINIQUE DURAND
2
,
NGÂN TRỌNG TRẦN
1
& PHÚC ĐÌNH PHAN
3
1
Việt Nam National University Hồ Chí Minh city-University of Science, Faculty of Biology, 227 Nguyễn Văn Cừ, District 5, Hồ Chí
Minh City, Vietnam. E-mail: hdhuy@hcmus.edu.vn
2
Institut de Recherche pour le Développement (IRD), UMR 5119 ECOSYM, Université
Montpellier II, Place Eugène Bataillon, CC 93, 34095 Montpellier cedex 5, France
3
Research Institute for Aquaculture No 3, 33 Đặng Tất, Nha Trang, Khánh Hòa, Vietnam
4
Corresponding author. E-mail: hdhuy@hcmus.edu.vn; Phone: +84 8 38304377
Abstract
Two new species and two new basin records of mahseers in the genera To r and Neolissochilus are described from the upper
Krong No and middle Đồng Nai drainages of the Langbiang Plateau in southern Vietnam. These new species and new
records are known from streams and rivers in montane mixed pine and evergreen forests between 140 and 1112 m. Their
populations are isolated in the Sre Pok River of the Mekong basin, the middle of the Đồng Nai basin, and the An Lão
River. Both new species are differentiated from their congeners by a combination of the following characters: 23–24 lat-
eral scales, 9–10 predorsal scales, 2/7 or 1/8 pelvic-fin rays, mouth position, median lobe of lower lip, rostral hood, colour
in life and by divergent mitochondrial DNA. Tor mekongensis sp. nov. is differentiated from Tor dongnaiensis sp. nov. by
the number of transverse scale rows (3/1/2 vs. 4/1/2), number of pelvic-fin rays (2/7 vs. 1/8), a blunt rostral hood vs. point-
ed, caudal-fin lobes that are equal vs. unequal, and by mitochondrial DNA (0.7% sequence divergence). Molecular evi-
dence identifies both species as members of the genus Tor and distinct from all congeners sampled (uncorrected sequence
divergences >1.9% for all Tor species for which homologous COI sequences are available). Tor sinensis is recorded in the
Krong No and the Sre Pok rivers, further south of its known distribution. Polymorphism is described in Neolissochilus
stracheyi with a Tor -like morph and a Neolissochilus-like morph.
Key words: Langbiang Plateau, Tor dongnaiensis sp. nov., Tor mekongensis sp. nov., Tor sinensis, Neolissochilus stra-
cheyi, southern Vietnam
Introduction
Mahseer refers to a group of freshwater cyprinid fishes easily distinguishable by relatively large scales on their
body compared to those of other cyprinid fishes (Desai 2003). The members belong to two genera, Tor and
Neolissochilus. These two genera are distinguished by the presence of a continuous labial groove in Tor vs. an
interrupted groove in Neolissochilus, and 10–14 gill rakers on the lower arm of the first gill arch in the former, and
6–9 in the latter (Rainboth 1985).
Tor is distinguished from other cyprinids by having a fleshy median mental lobe (Roberts 1999). There are 14
valid species of Tor in the trans-Himalayan region and southeast Asia (Eschmeyer 2015; Khare et al. 2014; Laskar
et al. 2013; Kottelat 2013; Madhusoodana et al. 2011; Zhou & Cui 1996; Menon 1992). Among these species, six
have been reported in southeast Asia; namely T. ater Roberts 1999, T. lateravittatus Zhou & Cui 1996, T. polylepis
Zhou & Cui 1996, T. sinensis Wu 1977, T. tambra Valenciennes in Cuvier and Valenciennes 1842, and T.
tambroides Bleeker 1854 (Kottelat 2013; Oijen & Loots 2012). However, the taxonomy and systematics of the
genus Tor is not stable. While Eschmeyer listed in 2013 (cited in Khare et al. 2014) 36 species as valid, in 2015
only 14 are maintained. Characters on which the taxonomy is based may be plastic and responsible for taxonomic
confusion (Mohindra et al. 2007). In this context molecular phylogenetic investigations would be useful, but to
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date most available studies have focussed on diversity of To r in India, a region known to be a diversity hotspot for
the genus (Mohindra et al. 2007; Laskar et al. 2013; Sati et al. 2013; Khare et al. 2014). Only one study has
investigated molecular diversity of Tor at a larger geographic scale (Nguyen et al. 2008). In this study, a close
phylogenetic relationship was highlighted between T. putitora and T. macrolepis, suggesting that they belong to the
same species. This was recently confirmed using both genetic and morpho-anatomic information (Khare et al.
2014). In contrast, an important cryptic diversity has been revealed within some nominal species such as Tor
khudree in India and Tor douronensis (which is a synonym of T. tamb r a according Eschmeyer, 2015) in Vietnam,
Malaysia and Sumatra (Nguyen et al. 2008). In T. tam b r a, three lineages were found, but due to their apparent
allopatric distributions, the authors did not make conclusions about the taxonomic status of the lineages. This first
insight into southeast Asian diversity of To r calls for further investigations, especially considering that this
biogeographic area encompasses several major freshwater biodiversity hotspots (De Bruyn et al. 2013).
During field expeditions in the Langbiang Plateau at high-elevations in the Krong No drainage, a catchment of
the Sre Pok River in the Mekong River drainage, and at low-elevations in the middle Đồng Nai drainage, from
2011 to 2015, two new, large species of To r were collected. They are described on the basis of morphological and
molecular characteristics. Additional information on Tor sinensis Wu, 1977, and Neolissochilus stracheyi (Day
1871), including their diagnosis in southern Vietnam, provided.
Material and methods
We recorded morphological data from specimens fixed in 10% formalin and stored in 70% ethanol. Specimens
were deposited at the University of Science, Hồ Chí Minh City, Vietnam (UNS), Zoological Reference Collection
of the Lee Kong Chian Natural History Museum, National University of Singapore (ZRC).
Morphometric data were taken to the nearest 0.1 mm with digital calipers. Methods of taking measurements
and counts, and terminology of the mouth structure followed Kottelat and Freyhof (2007). Abbreviations are: SL—
standard length; BD—body depth; HL—head length; LCP—length of caudal peduncle; DCP—depth of caudal
peduncle. We obtained comparative morphological data from museum specimens of Tor and photographs of these
species in life, preserved, and from the literature: T. ater (Roberts 1999), T. laterivittatus (Zhou & Cui 1996;
Roberts 1999: as T. sinensis, Kottelat 2001), T. t a mbra (Cuvier & Valenciennes 1842: as Barbus tambra; Bleeker
1863: as Labeobarbus tambra; Weber & de Beaufort 1916: as Labeobarbus tambra; Roberts 1993; Zhou & Cui
1996; Roberts 1999; Haryono 2006), T. douronensis (Bleeker 1863: as Labeobarbus douronensis; Weber &
Beaufort 1916: as Labeobarbus douronensis; Zhou & Cui 1996; Haryono 2006), T. tambroides (Weber & Beaufort
1916: as Labeobarbus tambroides; Kottelat 2001), T. yingjiangensis (Chen & Yang 2004), Neolissochilus blanci
(Pellegrin & Fang 1940; Rainboth et al. 2012), N. hendersoni and N. soroides (Khaironizam et al. 2015), N.
stracheyi (Rainboth 1985; Roberts 1999: as T. tamb r a), N. benasi (Pellegrin & Chevey 1936: as Crossochilus
benasi; Rainboth 1985; Roberts & Catania 2007). Due to the undiagnosed diversity within the genus, we relied on
examination of topotypic material and/or original species descriptions.
In order to test the generic placement of our specimens, we obtained fin samples from the new species and
compared them to species currently assigned to the genus To r (Sade & Biun 2012; Wibowo et al. 2013; Yadav et al.
2012; Yang et al. 2010; Yang et al. 2015). We analyzed 584 base pairs (bp) of the mitochondrial COI locus. DNA
was extracted using PureLink® Genomic DNA Mini Kit (Life Technologies). Primers used for amplifications and
sequencing for the COI gene was: FishF1, FishF2 and FishR1 (Ward et al. 2005). PCR purification and sequencing
were accomplished by Macrogen (Amsterdam, Netherlands). Sequences were edited with 4Peaks by A. Griekspoor
and Tom Groothuis (mekentosj.com) and deposited in GenBank under accession numbers KT261292–KT261305.
The data were aligned using the MUSCLE option in MEGA v. 6.06 and refined by eye. Uncorrected pairwise
sequence divergence was estimated using the substitution model of Tamura-Nei implemented in MEGA v. 6.06
(Tamura et al. 2013).
Maximum-likelihood phylogenetic analyses were performed in MEGA v. 6.06 with the TrN+I+Γ model,
performed in jModelTest v. 2.1.4. The best likelihood estimate from each of these runs was selected and statistical
support for this topology was obtained by running 1000 bootstrap replicates in MEGA v. 6.06. Phylogenetic trees
were modified by TreeGraph v. 2.0.47. The outgroup Neolissochilus stracheyi (GenBank Accession No.
JQ346166) was chosen based on the proposed close relationship of Neolissochiulus to Tor (Nguyen et al. 2008).
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TWO NEW CYPRINID FISHES FROM SOUTHERN VIETNAM.
TABLE 1. Diagnostic characters for species of Tor.
SL/BD SL/HL LCP/DCP Lateral
scales
Transverse
scale rows
Predorsal
scales
Rays of
dorsal fin
Rays of
anal fin
Rays of
pelvic fin
Mouth position
ng Nai River
(T. dongnaiensis sp. nov.) 3.0−3.5 3.6−4.1 1.5−1.8 23−24 4/1/2 10 4/9 3/6 1/8 sub-terminal
T. tambroides 3.0−3.4 3.2−4.0 1.5 23−26 3/1/
2−3
8−9 4/9−10 3/5−6 2/8 sub-terminal
Krong No River
(T. mekongensis sp. nov.) 3.3−3.6 3.5−3.9 1.5−2.0 23 3/1/2 9 4/9 3/6 2/7 sub-terminal
T. tambra 3.3−4.0 3.3−4.3 1.5 22−24 3/1/2 8−9 4/8−9 3/5−6 1/7−8 sub-terminal
T. sinensis 2.9−3.7 3.1−3.9 1.2−1.3 23−28 3.5−4.5/1/2 8−10 4/7−9.5 3/5.5 1/8−9 sub-terminal
T. laterivittatus 3.4−3.9 3.6−3.9 1.4−1.5 25−27 3.5−4/1/2 9 4/8.5 3/5.5 1/8 sub-terminal
T. polylepis 4.1−4.2 4.1−4.2 1.3−1.5 34−35 5.5/1/3 14 4/9.5 3/5 1/8 sub-terminal
T. ater - - - 30−31 5/1/2 11−12 - - - sub-terminal
T. hemispinus 3.4−4.0 3.7−4.0 1.3−1.4 30−35 5.5/1/
3.5
12 4/9−10 3/5 - terminal
T. putitora 3.9−4.8 3.3−3.8 1.4−2.0 23−28 4.5/1/
2.5 9−11 4/9−10 3/5 9 sub-terminal
T. mosal 3.4−4.0 3.5−4.0 1.4−1.6 23−26 3.5/1/
3.5
8 4/8−9 3/5 0/8−9 terminal
T. yingjiangensis 3.7−3.9 3.0−3.5 0.9−1.3 24−26 4−4.5/1/3−3.5 10 4/9 3/5 1/8−9 terminal
T. tor 3.2−4.5 3.4−4.3 1.3−1.7 22−28 3.5−4.5/1/
2.5−3.5
11−12 3−4/9 2−3/5 0/9 terminal
T. khudree 2.6−3.6 2.5−3.5 1.3−1.6 24−27 4.5/1/2.5 9−10 4/9 2−3/5 1/8 terminal
T. mussullah 3.0−3.1 3.9−4.0 1.5 26−27 4/1/3.5 - - - - sub-terminal
T. kulkarnii 3.2−3.6 3.9−4.3 1.3−1.7 24−26 3.5/1/2.5 10−11 4/9 3/5 0/9 terminal
T. remadevii 3.5−3.9 - 1.1−1.5 27−29 4.5/1/2.5−3 9−11 4/10 1/5 1/8 slight upturned
terminal
…….continued on the next page
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TABLE 1. (Continued)
Lower median
lobe
Upper median
projection
Rostral hood Caudal fin
lobes
Body colour Source
ng Nai River
(T. dongnaiensis sp. nov.)
long absent pointed unequal silver grey and yellowish in
sub-adult
Present paper
T. tambroides long present pointed equal silver-bronze and reddish Bleeker 1854,
Haryono 2006 Weber & de
Beaufort 1916
Krong No River
(T. mekongensis sp. nov.)
not long absent blunt equal silver grey Present paper
T. tambra not long absent blunt equal olive, or dark- or slightly olive Cuvier & Valenciennes 1842,
Haryono 2006, Weber & de
Beaufort 1916
T. sinensis long absent pointed equal darkish above and brownish or
bronzy beneath, a dark
longitudinal stripe
Zhou & Cui 1996, Roberts
1999
T. laterivittatus long absent pointed equal dark green to silver, a dark
longitudinal stripe
Zhou & Cui 1996
T. polylepis short absent pointed equal silver, a dark longitudinal
stripe
Zhou & Cui 1996
T. ater short absent pointed equal dark brown, a dark
longitudinal stripe
Roberts 1999
T. hemispinus short absent blunt equal - Chen & Chu 1985
T. putitora varying length absent blunt equal reddish sap-green, light orange
fading to silvery white
Menon 1992
T. mosal long absent blunt equal delicate yellowish shade
below, caudal reddish orange
Desai 2003, Madhusoodana, et
al. 2010
T. yingjiangensis short absent blunt equal - Chen & Yang 2004
T. tor varying length absent blunt equal greyish green Menon 1992
T. khudree varying length absent blunt unequal yellowish white and silver
bluish grey
Menon 1992,
Desai 2003
T. mussullah absent - - equal silver-bronze Madhusoodana, et al. 2010
T. kulkarnii short absent blunt unequal dorsal greyish, lateral silvery Menon 1992
T. remadevii short absent blunt equal dorsal greenish to metallic
black, upper lateral silvery to
white
Madhusoodana, et al.2010
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TWO NEW CYPRINID FISHES FROM SOUTHERN VIETNAM.
Results
Morphological data. Our specimens of mahseers from the Krong No and the Đồng Nai drainages belong to four
lineages. Two of four differed from each other and from all species in Tor by a combination of 15 characters shown
in Table 1. Tor sinensis and Neolissochilus stracheyi also were collected.
Molecular data. The newly collected specimens from the Krong No and Đồng Nai Rivers in Vietnam belong
to four mitochondrial lineages (Fig. 1). Three are embedded within a clade containing all To r species used as
ingroup and one characterized by a COI sequence nearly identical to the outgroup, Neolissochilus stracheyi from
Laos (JQ346166) (Fig. 1). This clade receives ≥72% bootstrap support.
Among the three To r lineages, one consists of five specimens collected in the Krong No River (UNS00614,
UNS00873, UNS00876, UNS00945, ZRC 54629) that has a COI sequence similarity of 99.7% with a specimen
identified as Tor sinensis from Yunnan, China (HM536900). In contrast, the two remaining Tor lineages, one
observed in the Krong No River and another in the Đồng Nai River, present no close relationship with any of the
COI sequences of Tor available in GenBank (Fig. 1). These two lineages have divergences of 0.7% larger than the
divergence observed between T. putitora (JX27226) and T. tor (JN032125), estimated at 0.5% (Table 2).
The newly collected specimens of Neolissochilus from the Krong No River are embedded with N. stracheyi
specimens from Laos (JQ346166) (99.8% similarity). This clade receives 100% bootstrap support. COI sequences
of all morphologies of N. stracheyi found in the Krong No River, including a Neolissochilus-like morph
(Neolissochilus blanci type and golden type) and a Tor-like morph, show 100% similarity (Fig. 1, Table 2).
TABLE 2. Samples used in the phylogenetic analysis of To r with the outgroup Neolissochilus stracheyi and genetic
distances between species.
123456789
1. T. dongnaiensis
sp. nov.
KT261298–KT261301
2. T. tambroides
Sumatra, Indonesia
JQ665787
0.031
3. T. mekongensis
sp. nov.
KT261292
0.007 0.027
4. T. tambra
Sabah, Malaysia
JN646100 (as T. douronensis)
0.042 0.040 0.034
5. T. sinensis
Lam Dong, Dak Lak, Vietnam
KT261293–KT261297
0.039 0.033 0.031 0.037
6. T. sinensis
Yunnan, China
HM536900
0.036 0.030 0.028 0.034 0.003
7. T. putitora
India
JX127226
0.023 0.025 0.019 0.034 0.033 0.030
8. T. tor
India
JN032125
0.025 0.027 0.021 0.036 0.035 0.032 0.005
9. N. stracheyi
Lam Dong, Vietnam
KT261302–KT261305
0.045 0.047 0.041 0.060 0.057 0.054 0.039 0.045
10. N. stracheyi
Laos
JQ346166
0.047 0.049 0.043 0.062 0.059 0.056 0.041 0.048 0.002
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FIGURE 1. Maximum-likelihood tree based on COI mitochondrial gene sequences for species of Tor and for Neolissochilus
stracheyi. Numbers on branches are ML bootstrap values (values > 70% shown).
Taxonomic conclusions. Morphological and molecular data sets provide evidence that the newly collected
specimens from the Krong No River and the Đồng Nai River represent new species of Tor and are described here.
Specimens of Tor sinensis and Neolissochilus stracheyi represent new records for the drainages.
Tor dongnaiensis sp. nov.
Holotype: UNS00859, 240 mm SL; middle Đồng Nai drainage: Cát Tiên National Park, Lâm Đồng Province,
Vietnam (11°26’33.32” N 107°26’4.01” E, 162 m), 7 March 2014, Hoàng Đức Huy, Phạm Mạnh Hùng and Trần
Trọng Ngân (Fig. 2).
Paratypes (all same locality as holotype): ZRC 54628, 114 mm SL, 11 March 2014; UNS00861, 167 mm SL,
12 March 2014; UNS00862, 201 mm SL, 9 April 2014; ZRC 54627, 243 mm SL, 26 October 2013; UNS00880,
167 mm SL, 1 June 2014; UNS00888, 189 mm SL, 23 October 2014; UNS00889, 411 mm SL, 23 October 2014
(11°39’20.8” N 107°16’04.7” E, 195 m).
Diagnosis. Tor dongnaiensis most closely resembles T. tambroides but differs in having a yellow to grey vs.
silver-bronze and reddish body, transverse scale rows 4/1/2 vs. 3/1/2, predorsal scales 10 vs. 8-9, pelvic-fin rays 1/
8 vs. 2/8, median projection in upper lip absent vs. present, and caudal fin lobes unequal vs. equal. Tor
dongnaiensis differs from T. a t er in having lateral scales 23–24 vs. 30–31, transverse scale rows 4/1/2 vs. 5/1/2,
predorsal scales 10 vs. 11–12, median lobe in lower lip long vs. short, stripe along side of body absent vs. present.
Tor dongnaiensis differs from T. laterivittatus and T. sinensis in having transverse scale rows 4/1/2 vs. 3–4/1/2,
stripe along side of body absent vs. present. Tor dongnaiensis differs from T. yingjiangensis in having deeper body
28.3–33.6 vs. 25.5–27.3% SL, caudal peduncle length 18.4–22.3 vs. 11.3–14.8% SL, and median lobe in lower lip
long vs. short. Tor dongnaiensis differs from T. t a mbra in having transverse scale rows 4/1/2 vs. 3/1/2, predorsal
scales 10 vs. 8–9, lips thickened vs. normal, and median lobe in lower lip long vs. short.
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TWO NEW CYPRINID FISHES FROM SOUTHERN VIETNAM.
FIGURE 2. Type localities of Tor mekongensis (black star) and T. dongnaiensis (white star), and collection localities of T.
sinensis (black triangle), and Neolissochilus stracheyi (white cross).
Description. General appearance in Figure 3; meristic and morphometric data of specimens given in Table 3.
Head conical, longer than deep, its depth 1.3–1.5 times in HL. Snout slightly pointed. Mouth subterminal and
oblique, its posterior edge not extending to vertical line of orbit. Rostral hood straight and pointed. Lips thick
fleshy; upper lip rolled backwards and upwards without median projection; median lobe of lower lip long, more or
less square and reaching an imaginary line between the corners of the mouth (Fig. 4e, f). Rostral and maxillary
barbels almost equal in length and both longer than eye diameter.
Body elongate, moderately compressed, its depth about 3.0–3.5 times in SL; caudal peduncle about 1.5–1.8
times longer than deep. Dorsal body profile convex, ventral profile rounded. Lateral line complete, 23–24 scales;
10 predorsal scales; 4/1/2 scales in transverse row anterior to pelvic-fin insertion.
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TABLE 3. Morphometric and meristic characters of Tor dongnaiensis sp. nov. and Tor mekongensis sp. nov.
Tor dongnaiensis Tor mekongensis
Holotype Range Mean±SD N Holotype Range Mean±SD N
SL (mm) 240 11.4–41.1 21.6±8.9 8 337 7.6–33.7 16.4±15.0 3
Morphometrics
% SL
Head length 24.2 24.2–28.1 25.8±1.4 8 25.5 25.5–28.9 27.8±2.0 3
Depth of body 28.3 28.3–33.6 31.1±1.7 8 27.6 27.6–30.2 28.9±1.3 3
Body width at dorsal-fin origin 15.4 14.9–20.1 16.4±1.7 8 13.6 13.6–15.8 14.4±1.2 3
Predorsal length 51.5 50.8–53.5 51.7±1 8 50.4 50.4–52.2 51.3±0.9 3
Prepectoral length 25.1 23.5–27.4 25.6±1.2 8 24.6 24.6–27.7 26.2±1.5 3
Prepelvic length 51.7 46.9–54.5 50.9±2.2 8 48.4 48.4–52.8 50.8±2.3 3
Preanal length 75.0 74.1–85.2 76.5±3.7 8 74.2 74.2–76.3 74.9±1.2 3
Distance between pectoral- and pelvic-fin origins 23.8 21.1–24 22.5±1.1 8 20.5 20.1–21.1 20.6±0.5 3
Distance between pelvic- and anal-fin origins 20.4 17.5–22.6 20.1±1.6 8 20.5 19.5–20.5 19.9±0.5 3
Depth of caudal peduncle 11.9 11.3–13.5 12.3±0.6 8 11.3 11.3–13.2 12.3±1.0 3
Length of caudal peduncle 19.6 18.4–22.3 20.3±1.3 8 21.4 18.9–21.4 20.0±1.3 3
Length of dorsal fin 22.1 17.5–26.3 23.3±2.6 8 21.4 21.4–27.0 24.9±3.1 3
Length of dorsal-fin base 12.9 12.9–15.8 14.3±0.9 8 13.6 13.6–15.1 14.4±0.7 3
Length of pectoral fin 20.8 19.7–23.5 21.7±1.2 8 20.2 20.2–22.4 21.5±1.2 3
Length of pelvic fin 19.2 18.9–20.9 19.5±0.7 8 17.5 17.5–18.9 18.3±0.7 3
Length of anal fin 19.6 19.3–22.2 20.5±1.2 8 21.5 20.1–22.4 21.3±1.1 3
Length of anal-fin base 7.1 7.1–8.6 7.7±0.5 8 6.8 6.8–20.4 11.6±7.6 3
% HL
Head depth 72.4 68.2–79 72±3.9 8 65.1 65.1–69.6 67.6±2.3 3
Head depth at eye 55.2 50–58.5 54.2±2.9 8 53.5 52.2–54.5 53.4±1.2 3
Maximum head width 59.5 53.1–77.3 58.8±7.7 8 58.1 52.2–58.1 55.3±3.0 3
Snout length 39.7 33.3–39.7 36.1±2.5 8 32.6 30.4–32.6 31.6±1.1 3
Interorbital width 43.1 31.3–49.2 39.6±5.3 8 44.2 28.3–44.2 36.3±8.0 3
Eye diameter 19.0 15.9–22.2 19.4±1.9 8 17.4 17.4–26.1 22.1±4.4 3
Mouth width 25.9 22.4–31.9 26.3±3.2 8 25.6 25.6–27.3 26.3±0.9 3
Counts
Lateral-line scales 24 23–24 23.4±0.5 8 23 23 23.0±0.0 3
Scales between lateral line and origin of dorsal fin 4 4 4.0±0.0 8 3 3 3.0±0.0 3
Scales between lateral line and origin of pelvic fin 2 2 2.0±0.0 8 2 2 2.0±0.0 3
Scales between lateral line and origin of anal fin 3 3 3.0±0.0 8 3 3 3.0±0.0 3
Circumpeduncular scales 12 12 12.0±0.0 8 12 12 12.0±0.0 3
Predorsal scales 10 10 10.0±0.0 8 9 9 9.0±0.0 3
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TWO NEW CYPRINID FISHES FROM SOUTHERN VIETNAM.
FIGURE 3. Tor dongnaiensis: (a) juvenile paratype (ZRC 54628, 114 mm SL), (b) sub-adult paratype (UNS00880, 167 mm
SL), (c) adult holotype (UNS00859, 240 mm SL) in life, and (d) in preservative; lateral views. Scale 10 mm.
Dorsal with 4 simple and 9 branched rays, last simple ray smooth; dorsal-fin origin inserted approximately
opposite to pelvic-fin origin; distal margin slightly concave. Pectoral fin pointed with 1 simple and 16 branched
rays. Pelvic fin pointed, with 1 simple and 8 branched rays; axillary scale present. Anus immediately in front of
anal fin. Anal fin pointed with 3 simple and 6 branched rays. Caudal deeply forked with 10+9 principal rays, 9+8
being branched and upper lobe smaller than lower one.
Colour in life. Head dark yellowish turquoise on back, yellowish turquoise around orbit and on side, white on
opercula and lower jaw. Body dark grey on back, light yellow above lateral line, grey white on belly. Scales light
yellow to silver white, black at scale bases. All fins dark grey with yellow tinge on the branched rays. In sub-adult
specimens, scales and fins light yellow but pectoral fin and pelvic fin pinkish orange on coastal rays. In juvenile
specimens, scales bluish silver; dorsal and caudal fins greenish yellow, dark bold on rays; pectoral fins pinkish
orange; pelvic and anal fins yellow with orange at coastal rays and hyaline on distal parts (Fig. 3a, b, c).
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Colour in preservative. Body including head dark grey on back. Opercula becoming black at base. Lower half
of body greyish white. All fins grey to dark grey (Fig. 3d).
Etymology. Specific epithet is in reference to the type locality of Đồng Nai drainage.
Suggested common name. Cá ngựa xám Đồng Nai (Vietnamese), Dongnai mahseer (English).
Tor mekongensis sp. nov.
Holotype: UNS00877, 337 mm SL; upper Ea Krong No drainage: upper Mekong basin in montane evergreen
forest in Bidoup-Núi Bà National Park, Lâm Đồng Province, Vietnam (12°16’23.68” N 108°26’30.17” E, 672 m),
24 June 2014, Hoàng Đức Huy, Phạm Mạnh Hùng and Trần Trọng Ngân (Fig. 2).
Paratypes (all same locality as holotype): UNS00878, 79.5 mm SL, 24 June 2014; UNS00879, 76 mm SL, 24
June 2014.
FIGURE 4. Head of Tor sinensis UNS00873, 205 mm SL (a, c) front view, (b, d) ventral view; T. dongnaiensis holotype
UNS00859, 240 mm SL (e) front view, (f) ventral view; T. mekongensis holotype UNS00877, 337 mm SL (g) front view, (h)
ventral view; Neolissochilus stracheyi Tor-like morph UNS00949, 428 mm SL (i) front view, (k) ventral view; Neolissochilus
stracheyi Neolissochilus-like morph UNS00872, 308 mm SL (l) front view, (m) ventral view. Scale 10 mm.
Diagnosis. Tor mekongensis is similar to T. t a mbra but differs genetically. Tor mekongensis differs from T.
sinensis in having lateral scales 23 vs. 23–28, pelvic-fin rays 2/7 vs. 1/8–9, median lobe short vs. long, rostral hood
rounded and blunt vs. prolonged into a lobe. Tor mekongensis differs from T. dongnaiensis in having predorsal
scales 9 vs. 10, median lobe short vs. long, pelvic-fin rays 2/7 vs. 1/8, upper lip thin vs. thick, colour in life silver
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grey vs. yellowish, lobes of caudal fin nearly equal vs. unequal. Tor mekongensis differs from T. tambroides in
having lateral scales 23 vs. 23–26, median lobe short vs. long, upper lip without median projection vs. median
projection present. Tor mekongensis differs from T. ater in having lateral scales 23 vs. 30–31, scales in transverse
row 3/1/2 vs. 5/1/2, predorsal scales 9 vs. 11–12, stripe on side of body absent vs. present.
Description. General appearance in Figure 5; meristic and morphometric data of specimens given in Table 3.
Head longer than deep, its depth about 1.4–1.5 times in HL and its dorsal profile slightly convex. Snout blunt.
Rostral hood rounded and blunt. Mouth subterminal and oblique, its posterior edge not extending to vertical line of
orbit. Lips normal fleshy, median lobe of lower lip developed, but does not reach the line connecting the corners of
the mouth (Fig. 4g, h). Upper lip not prolonged into a lobe. Rostral and maxillary barbels almost equal in length
and both longer than eye diameter.
Body elongate, moderately compressed, body depth about 3.3–3.6 times in SL; caudal peduncle slender and
long, about 1.5–2.0 times longer than deep. Lateral line complete, 23 scales; 9 predorsal scales; 3/1/2 scales in
transverse row anterior to pelvic-fin insertion.
FIGURE 5. Tor mekongensis: (a) juvenile paratype (UNS00879, 76 mm SL) in life, (b) adult holotype (UNS00877, 337 mm
SL) in life, and (c) in preservative; lateral views. Scale 10 mm.
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Dorsal with 4 simple and 9 branched rays, last simple ray smooth; dorsal-fin origin inserted slightly opposite to
pelvic-fin origin; distal margin slightly concave. Pectoral fin pointed with 1 simple and 15 branched rays. Pelvic fin
pointed, with 2 simple and 7 branched rays; axillary scale present. Anus immediately in front of anal fin. Anal fin
pointed with 3 simple and 6 branched rays. Caudal fin deeply forked with 10+9 principal rays, 9+8 being branched.
Colour in life. Head dark on back, white on lower jaw. Body dark on back, grey on side, snowy white on belly.
Scales silver grey. Dorsal and caudal fins grey on ray, yellow tinge distally. Pelvic fin and anal fin grey to dark grey,
orange tinge distally. Pectoral fin pinkish on origin, grey on rays (Fig. 5a, b).
Colour in preservative. Similar to that of fresh condition except noted below. Body including head dark grey
on back. Lower half of body greyish white. Dorsal fin, pectoral fin, pelvic fin and anal fin grey to blackish. Caudal
fin grey on rays and pale on margin (Fig. 5c).
Etymology. Specific epithet is in reference to the type locality of Mekong drainage.
Suggested common names. Cá ngựa xám Mê Kông (Vietnamese), Mekong mahseer (English).
Tor sinensis Wu 1977
Specimens Examined: upper Ea Krong No drainage: upper Mekong basin in montane evergreen forest in Bidoup-
Núi Bà National Park, Lâm Đồng Province, Vietnam (12°16’23.68” N 108°26’30.17” E, 672 m): UNS00873, 205
mm SL, 25 June 2014; UNS00614, 213 mm SL, 11 March 2012 (12°15’9.37” N 108°26’23.18” E, 650 m); ZRC
54629, 214 mm SL, 12 March 2012; UNS00874, 133 mm SL, 24 June 2014; ZRC 54630, 97 mm SL, 25 June 2014
by Hoàng Đức Huy, Phạm Mạnh Hùng and Trần Trọng Ngân; upper Mekong basin in Vietnam (12°58’02.5” N
107°32’58.2” E): UNS00876, 98 mm SL, 25 June 2014; the Sre Pok River: upper Mekong basin in Dak Lak
Province, Vietnam (12°58’02.5” N 107°32’58.2” E): UNS00945, 300 mm SL, 27 December 2014 by Phan Đình
Phúc (Fig. 2).
FIGURE 6. Tor sinensis Wu 1977: (a) juvenile (UNS00876, 98 mm SL), (b) sub-adult (UNS00873, 205 mm SL) in life; lateral
views. Scale 10 mm.
Diagnosis. Tor sinensis differs from T. ater in having lateral scales 24–25 vs. 30–31, scales in transverse row 3/
1/2 vs. 5/1/2, predorsal scales 10 vs. 11–12, median lobe in lower lip long vs. short, stripe along side of body absent
vs. present. Tor sinensis differs from T. mekongensis in having lateral scales 23–28 vs. 23, pelvic-fin rays 1/8–9 vs.
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2/7, median lobe long vs. short, rostral hood prolonged into a lobe vs. rounded and blunt. Tor sinensis differs from
T. dongnaiensis in having transverse scale rows 3/1/2 vs. 4/1/2, dorsal-fin rays 4/8 vs. 4/9, lobes of caudal fin
nearly equal vs. unequal. Tor sinensis differs from T. t a mbroides in having SL/BD 3.6–4.0 vs. 3.0–3.4, lateral
scales 24–25 vs. 24–26, median projection in upper lip present vs. absent. Tor sinensis differs from T.
yingjiangensis in having colour in life silver-grey vs. yellow, SL/HL 3.3–3.9 vs. 3.0–3.5. Tor sinensis differs from
T. t a m bra in Java in lateral scales 24–25 vs. 22–24, predorsal scales 10 vs. 8–9, median lobe in lower lip long vs.
short.
Colour in life. Head dark on back, white on lower jaw. Body dark on back, yellow to grey-silver on side,
snowy white on abdomen. A slate-grey longitudinal stripe run from posterior of head to the caudal base in adult
specimen (UNS00945), similar to T. laterivittatus, but other specimens without a longitudinal stripe. Scales silver
with black at bases. Dorsal fin yellow grey, pelvic fin and anal fin dark grey. Pectoral fin pinkish on base, and black
on rays. Caudal fin almost dark grey with yellow and pink tinge distally. Juvenile specimens light silver on scales
and fins hyaline. Their dorsal fins green-yellow, dark grey on rays; distal margin concave. Pectoral fins pinkish
orange and pelvic fin orange on coastal rays. Anal fins pinkish yellow on distal parts. Caudal fin greenish yellow,
dark bold on rays (Fig. 6a, b).
Neolissochilus stracheyi (Day 1871)
Specimens examined: upper Ea Krong No drainage: upper Mekong basin in montane evergreen forest in Bidoup-
Núi Bà National Park, Lâm Đồng Province, Vietnam (12°14’57.47” N 108°40’31.41” E, 1101 m): UNS00603, 193
mm SL, 13 March 2012; UNS00604, 256 mm SL, 19 February 2012 (12°15’36.6” N 108°30’44.5” E, 856 m);
UNS00865–871, 212–322 mm SL, 8 March 2013 (12°15’5.25” N 108°38’32.48” E, 1025 m); UNS00863–864,
160–163 mm SL, 5 March 2013 (12°14’57.47” N 108°40’31.41” E, 1101 m); UNS00872, 308 mm SL, 25 June
2014 (12°16’23.68” N 108°26’30.17” E, 672 m); UNS00948–949, 287–428 mm SL, 22–24 January 2015
(12°16’48.63” N 108°31’40.85” E, 754 m); UNS00950, 534 mm SL, 22 January 2015 (12°16’47.88” N
108°30’20.75” E, 720 m); the An Lão River, Bình Định Province, Vietnam (14°40’30.6” N 108°54’13.4” E, 547
m): UNS00768, UNS00770, UNS00870, 113–130 mm SL, 21 January 2013 (Fig. 2).
Diagnosis. Neolissochilus stracheyi differs from N. hendersoni and N. soroides in lateral stripe present vs.
absent. Neolissochilus stracheyi differs from N. benasi in lateral scales 24–28 vs. 31–32, transverse scale rows 3/1/
2, vs. 4/1/3. Neolissochilus stracheyi in this study is similar to N. blanci from Laos (Pellegrin & Fang 1940,
Rainboth et al. 2012) in living colour, morphometric and meristic characters.
From 13 March 2012 to 22 January 2015 a sample of 17 fresh N. stracheyi was collected from the upper Krong
No River in Bidoup-Núi Bà National Park. The following data were obtained: Neolissochilus-like morph: 16
individuals, 113–534 mm SL with moderately developed lips and without a submentum on the lower lip (Fig. 4l,
m). Among this morph, there were two body-colour types: bronze with a slate-grey lateral stripe (Fig. 7a, b, e) and
golden lacking a lateral stripe and becoming progressively lighter to silvery white below (Fig. 7c). The lateral
stripe appears in preserved specimens of both body colour types. Individuals of both types have the caudal fin
deeply forked, but the lateral stripe type has a convex distal margin on each lobe of the caudal fin, and the golden
type has a straight distal margin on each lobe.
Tor-like morph: 1 individual, 428 mm (UNS00949): rostral hood straight and pointed. Lips thick, fleshy; upper
lip rolled backwards and upwards without median projection; median lobe of lower lip broad, thick and long as the
most highly developed mentum in species of the genus To r. Body-colour bronze with faint slate-grey lateral stripe
(Figs. 4i, k and 7d).
Ecology. All specimens of the new species and new records were found in the Ea Krong No drainage,
consisting of montane mixed pine and evergreen forest (Fig. 8c). Our survey also recorded the occurrence of N.
stracheyi in the An Lão River at ~500 m, in the Krong Ana River at ~ 431m elevation, Chư Yang Sin National Park
(12°29’50.9” N 108°20’29.6” E) and the Đồng Nai River at ~ 650 m elevation, Tà Đùng Nature Reserve
(11°45’10.4” N 108°00’3.0” E). While N. stracheyi occurs along the main river and in streams between 596–1112
m, T. mekongensis and T. tambra occur in lower elevations between 596–850 m. Water conditions where N.
stracheyi occur were 17−26
°C, with pH 6.5−7.66, DO 79−88.6%, conductivity 10–34 µS.cm
-1
and flow velocity
0.17−0.77 m/s. Water conditions where Tor occur were 21−26
°C, with pH 7−7.66, DO 79−80.6%, conductivity
32−34 µS.cm
-1
and flow velocity 0.20−0.77 m/s.
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FIGURE 7. Neolissochilus stracheyi: (a) sub-adult (UNS00864, 163 mm SL) in life, (b) adult (UNS00866, 238 mm SL) in
life, (c) golden adult (UNS00948, 287 mm SL) in life, (d) To r -like morph adult (UNS00949, 428 mm SL) in life, (e) female
adult (UNS00950, 534 mm SL) in life; lateral views. Scale 10 mm.
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TWO NEW CYPRINID FISHES FROM SOUTHERN VIETNAM.
FIGURE 8. Mahseers in the wild: (a) juvenile To r at night in shallow area near stream bank; (b) shoal of Neolissochilus
stracheyi in slow-flowing pool, high elevation main river with riparian vegetation at type locality of Tor mekongensis; (c)
Krong No River (12°15’9.37” N 108°26’23.18” E, 650 m); (d) Đồng Nai River (11°39’20.8” N 107°16’04.7” E, 195 m).
Mahseers of the Krong No River occurred in lotic habitats of main streams, particularly in deep pools. In the
early wet season (June−July), mature individuals move to the depositional zone in moderate flowing pools (5–7 m
deep and 20–50 m wide) with detritus and fallen leaves for gonadal development (villagers, pers. com.). The
juveniles often inhabit shallow areas with mixed silt and sand, slower flow and submergent plants. They share this
habitat with Poropuntius sp., Hampala macrolepidota, and Onychostoma krongnoensis. During wet season surveys
(October 2011, May 2013 and June 2014), many juvenile fishes were observed (Fig. 8a).
Mahseers of Tor and Neolissochilus usually roam in shoals of 20–30 individuals in swift currents or in slow
flowing pools (2–3 m deep) for foraging (Fig. 8b). They feed on plant matter, fallen fruits and insects from riparian
vegetation (Rainboth 1996). Villagers claim that mahseers and carps are often found inside the underwater rock
caves or trunks of large trees in the stream bank.
All specimens of T. dongnaiensis were found in the middle Đồng Nai drainage between 140−200 m (Fig. 8d).
Water conditions were 23−26
°C, pH 7−7.66, DO 75−81.7%, conductivity 33−40 µS.cm
-1
and flow velocity
0.05−1.3 m/s. This species shares habitat with Labeo spp., Gyrinocheilus aymonieri, Cosmocheilus harmandi,
Hampala macrolepidota, Hypsibarbus spp., Wallago attu, Bagarius yarrelli, and Hemibagrus spp.
Discussion
With two new species described here, a total of 16 species of Tor are now recognised. The number of Tor species in
the Mekong River basin is 8.
Tor ‘tambroides’ has been described by Mai et al. (1992) in the Đồng Nai River; however, the illustration and
morphological characters depicted by these authors agrees with the description of T. dongnaiensis. It is probable
that the individual depicted as T. ‘tambroides’ in the Đồng Nai River was confused with T. dongnaiensis, which in
turn questions the presence of T. tambroides in Vietnam.
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Among Mekong Tor species considered valid, T. tambra is treated as the senior synonym of T. douronensis
(Roberts 1993, 1999; Kottelat 2001, 2013, Eschmeyer, 2015). A molecular phylogeny (Nguyen et al. 2008) of
some Tor species included specimens identified as T. tambra and revealed an important cryptic diversity consisting
of three evolutionary lineages including one in the Mekong River. Kottelat (2013) acknowledged this finding but
did not draw taxonomic and nomenclatural conclusions because no morphological information or materials from
Java were provided in the Nguyen et al.’s (2008) study. However, it is certain, considering the level of divergence
and the phylogenetic relationships that these different lineages correspond to different species. Tor tambra was
described from Java, and the lineage identified in the Mekong River could correspond to T. s inenis.
Neolissochilus stracheyi specimens in the Krong No River exhibit marked variation in body colour and mouth
morphology, including the lips and presence or absence of a fleshy median lobe on the lower lip. Zhou & Cui 1996
recorded an undescribed specimen similar to N. stracheyi with a median lobe of the lower lip in the upper Mekong
River in Yunnan, China. Later, Roberts (1999) recorded the specimens of the Nam Theun watershed in Laos
assigned to T. tambra as having tuberculation covering virtually the entire side of the face. Numerous vs. few
tubercles on the side of the head are distinguishing characteristics separating Neolissochilus from To r (Rainboth
1985). The specimens of Zhou & Cui (1996) and Roberts (1999) could correspond to the Tor-like morph of N.
stracheyi in this study. The variation is interpreted here as intraspecific polymorphism as described by Roberts &
Khaironizam (2008) and Khaironizam et al. (2015). Recognition of polymorphism involving the mouth of N.
stracheyi contributes to a better understanding of polymorphism in Neolissochilus, not only in the Malay Peninsula
(Roberts & Khaironizam 2008, Khaironizam et al. 2015), but also in the southeast Asia mainland.
Conservation status. The large mahseers, T. mekongensis, T. sinensis and N. stracheyi are indigenous fishes of
the Langbiang Plateau in Ea Krong No drainage. During the dry season, when river water levels are low (December
to April), local people frequently harvest these species using gill nets and cast nets. The main river of the Ea Krong
No drainage is approximately 85 km long and runs inside Bidoup-Núi Bà National Park and Chư Yang Sin
National Park. Mahseers are found generally within the protected area. Tor mekongensis probably is distributed in
the Sre Pok River and mainstream Mekong River where the fishes are threatened by development of dams. Given
the available information, we suggest the species should be considered Data Deficient following IUCN’s Red List
categories (IUCN 2014, version 11).
Tor dongnaiensis is recorded from the mainstream Đồng Nai River, where it is harvested by local fishers. As a
result of the development of hydropower dams, including Đồng Nai 3 (11°52’20” N 107°53’23” E), Đồng Nai 4
(11°53’11” N 107°43’37” E), and Đồng Nai 5 (11°47’51” N 107°32’57” E), and sand exploitation activities, this
species is facing a high threat. Given the available information, we suggest the species should be considered Near
Threatened following IUCN’s Red List categories (IUCN 2014, version 11).
Comparative material examined
Neolissochilus benasi: Vietnam, Lao Cai province: Muong Hum (N. 25.338–35.339)
Neolissochilus benasi: Vietnam, Lai Chau province (H.01.59.01.01–H.01.59.01.03)
Neolissochilus blanci: Vietnam, Lai Chau province (H.01.59.03.01–H.01.59.03)
Neolissochilus blanci: Laos, Ban Nam Khueng (39-203.205.)
Tor ater: Laos, Nam Theun (ZRC 40356, ZRC 40357)
Tor douronensis: Indonesia, Java (MNHN 3826)
Tor douronensis: China, Yunnan: Mengla County (KIZ 863532, KIZ 736028)
Tor kulkarnii: India, Maharashtra, Deolali, Darna River (ZSI No.FF 2710, ZSI No. 2710)
Tor laterivittatus: China, Yunnan: Mengla County (KIZ 8840041)
Tor polylepis: China, Yunnan: Mengla Country (KIZ 863563), Yunnan: Xishuanbanna (KIZ 59179)
T. remadevii: India, Chambakkad, Pambar River (ZSI (WGRS) CLT. No. V/F 13119a)
Tor sinensis: Laos, Nam Theun (ZRC 40352-40355, ZRC 40358)
Tor tambra: Indonesia, Java, Buitenzorg (RMNH 2289)
Tor tambra: Laos, Nam Theun (ZRC 40347, ZRC 40348, ZRC 40349)
Tor tambro i d e s : Indonesia, Sumatra (RMNH 2620)
Tor tambro i d e s : China, Yunnan: Mengla County (KIZ 7890575)
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Tor tor: China, Yunnan: Mengla County (KIZ 863535, KIZ 736015)
Tor polylepis: China, Yunnan: Mengla Country (KIZ 863563), Yunnan: Xishuanbanna (KIZ 59179)
Tor yingjiangensis: China, Yunnan, Yingjiang River (KIZ 164401, KIZ 704404, KIZ 764236)
Acknowledgements
Boards and staffs of Bidoup Núi Bà National Park and Cát Tiên National Park kindly facilitated surveys and issued
permissions. Hoàng Đức Khoa, Đưng Gur Ha Bình, K’Jim and several K’Ho people from Đạ Long, Đưng Ia Giêng
villages and Đạ Tẻh district assisted with field work. Hoàng Trọng Khiêm assisted with lab work. Prachya
Musikasinthorn, Norfatimah Mohamed Yunus, and Chirachai Nonpayom provided access to reference literature.
Anonymous peer reviewers scrutinized and commented on the manuscript. This research was funded by the
Vietnam National Foundation for Science and Technology Development (NAFOSTED) under Grant No. 106.15-
2012.100 and the “Fondation Total” (fondation.total.com) through the “DECODIV” project. For all of this
assistance we are most grateful.
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