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80
Kottelat: Three new loaches from Myanmar
A new genus and three new species of nemacheilid loaches from
northern Irrawaddy drainage, Myanmar (Teleostei: Cypriniformes)
Maurice Kottelat
Abstract. Malihkaia aligera, new genus, new species, is described from the Mali Hka River in the northernmost
Kachin State, Myanmar. It is distinguished by strongly furrowed lips and a type of sexual dimorphism new in
Nemacheilidae: the pectoral n of males is greatly elongated as a ‘wing’, rigid and curled, with the rst branched
ray the longest, and with all branches and sub-branches adjacent and without membranes between them. Schistura
nubigena, new species, is described from the same locality. It is distinguished by its colour pattern that evolves
from 4 broad bars in juveniles about 20 mm SL, to a pattern of 8 bars that become coalescent along the ank
and along the back, leaving a row of pale spots between them. Schistura wanlainensis, new species, is described
from the Mon Lan Chaung, a tributary of the Mali Hka. It is distinguished by its unique colour pattern of a pale
grey to whitish background with 18–32 narrow, regularly shaped black bars, extending from dorsal midline and
reaching downwards to below level of pectoral ns, continuous over back with contralaterals, wider than interspaces
anteriorly, narrower than interspaces on caudal peduncle; some of anterior bars fused at their dorsal extremity.
Key words. Cobitoidei, Schistura, Malihkaia, Mali Hka
RAFFLES BULLETIN OF ZOOLOGY 65: 80–99
Date of publication: 12 May 2017
http://zoobank.org/urn:lsid:zoobank.org:pub:70097B8F-3EF2-4740-97D4-AD2357234011
© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)
Rue des Rauraques 6, 2800 Delémont, Switzerland (permanent address); and Lee Kong
Chian Natural History Museum, National University of Singapore, 2 Conservatory
Drive, Singapore 117377; Email: mkottelat@dplanet.ch
INTRODUCTION
Loaches of the family Nemacheilidae are a very important
component of the streams and creeks in hilly areas of South
and mainland Southeast Asia. Most have small geographic
ranges and are restricted to a single or a few river systems.
Their restricted movements downwards in less favourable
lowland habitats and the active geomorphological processes
(of which river captures at both large and small scales
are probably the most important) are likely responsible
for speciation and high endemism. In the hilly areas of
mainland Southeast Asia, members of the Nemacheilidae
represent about 20% of the fish fauna of any small to
medium drainage. For example, in the upper Nam Theun of
Laos, up to seven species may occur in syntopy (Kottelat,
1998, 2015, unpubl. data). Nemacheilidae can be used as
indicator species in different ways; their requirement for cold
and well-oxygenated water can be used as an indicator of
water quality; their diversity and specialisation for peculiar
microhabitats at a single site can be used as an indicator of
habitat integrity.
Nemacheilid diversity can also be used as an indicator of
the quality of our knowledge of the biodiversity in a given
area, or to evaluate the quality of Environmental Impact
Assessments, which throughout mainland Southeast Asia
are notoriously of very poor quality, incompetent or even
fabricated. A survey of a moderate size drainage in a hilly
area (let’s say about 1,000 km2) that does not include at
least 6–10 nemacheilid species is likely to have been very
supercial or to have missed all the critical habitats and
therefore of limited value (admittedly, safety or access issues
may be limiting factors). If these 6–10 species do not include
species endemic to that drainage, or unnamed species, or
include species otherwise only known from a very distant,
not contiguous, drainage, then this may suggest a capability
problem. The life history, evolution and habitat of many
genera of Sisoridae (Ng & Kottelat, 2016; Vidthayanon et
al., 2009) and Psilorhynchidae (Conway, 2011; Conway &
Britz, 2010, 2015; Conway & Kottelat, 2007, 2010; Conway
& Mayden, 2008) show the same pattern and they have a
similar indicator potential, except that their diversity at a
single locality is much lower.
The largest genus of the family is Schistura, with about 210
named valid species to which should be added a minimum
of about 50 unnamed species that I am already aware of and
that await description (pers. obs., updated from Kottelat,
2012b, 2013). Loaches of the genus Schistura typically
occur in fast owing water of small streams and less often
in other habitats such as large rivers and caves. The genus
has its greatest diversity in Southeast Asia (Irrawaddy,
Salween, Mae Khlong, Chao Phraya, Mekong and Red River
drainages, and drainages in between) from where about 160
species have been described; most are described and gured
in Kottelat (1990, 1998, 2000, 2001) and Freyhof & Serov
(2001). Besides, new species are still regularly described (e.g.,
Bohlen & Šlechtová, 2010; Ou et al., 2011; Plongsesthee et
al., 2011, 2013; Bohlen et al., 2016), especially in Myanmar
Taxonomy & Systematics
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RAFFLES BULLETIN OF ZOOLOGY 2017
(e.g., Bohlen & Šlechtová, 2013a, b; Bohlen et al., 2014;
Chen & Neely, 2012; Kottelat, 2017). The interrelationships
within Schistura have not been thoroughly studied yet but
the accumulating (and still unpublished) morphological,
molecular and distribution data unsurprisingly show that
the genus is paraphyletic.
Irrawaddy is the main river of Myanmar. It ows north to
south from the northernmost tip of the country in Kachin
state, from the divide with the Brahmaputra in India and the
Salween in China. It is formed by the conuence, upriver of
Myitkyina, of two main branches, the eastern N’mai Hka and
the western Mali Hka. N’mai Hka ows between high hills
along its whole length and is still very difcult to access and
to travel. Its drainage is ichthyologically virtually unexplored
in Myanmar (with the apparent exception of the description
of Glyptosternon malaisei by Rendahl & Vestergren (1941)
and a small survey conducted by FFI in July 2015); there
is information on its tributary Taron (or Trung) from the
Chinese part of its course (where it is called Dulong; Chen
et al., 2006a, b, 2012). The Mali Hka also ows in a hilly
area, but in the area of Putao the valley widens to form a
plain, which is more accessible, by land in the dry season
(if safety allows) and by air. There is somewhat more
information on the sh fauna of the Mali Hka around Putao
(Putao Plain or Putao basin in the literature).
The rst data on the sh fauna of the area of Putao were
published by B. L. Chaudhuri in 1919. They were based
on specimens collected in February 1918 by Murray Stuart,
a geologist of the Geological Survey of India. Chaudhuri
recorded 12 species and considered that three of them were
new to science. In fact, with present knowledge, it seems
that at least 10 were unnamed by that time. Later, Mukerji
(1933, 1934) reported on a collection of shes obtained
in the Mali Hka River drainage in March 1930, but these
included only samples from the area of Sunprabum (approx.
26°30′–26°45′N 97°45′E) and none from the middle and
upper stretches. Fishes were also obtained in Putao in 1934
by R. Malaise (in part reported by Rendahl, 1948).
The area remained unaccessible for decades because of
security reasons until the 1990s. Sampling by S. Kullander
and R. Britz in 1998 resulted in the addition of several
species to the list, either already known from elsewhere in
Myanmar or new to science (e.g., Conway & Britz, 2010;
Kullander, 2012; Kullander & Britz, 2002; Kullander &
Fang, 2005; Ng, 2008; Anonymous, 2015).
A survey of the shes in the drainage of the upper Mali
Hka was conducted in November to December 2014 by the
Forest Department, the Zoology Department of Myitkyina
University and Fauna & Flora International (FFI). Safety
conditions in 2014 restricted very much the areas that can
be visited in northern Myanmar and sampling was possible
only in a few areas of Putao Plain. An additional survey in
July 2015 by FFI sampled additional sites on Putao Plain
and also in higher altitude along the Mon Lan Chaung, a
main tributary owing from the water divide between the
Brahmaputra in India (Arunachal Pradesh) and the Irrawaddy
in Myanmar and entering the Mali Hka from the west. These
surveys resulted in the discovery of 10 species apparently
new to science. Three of them are described here; one of
them represents a new genus.
MATERIAL & METHODS
Measurements and counts follow Kottelat (1990) and
Kottelat & Freyhof (2007). Last 2 branched dorsal and
anal-fin rays articulating on a single pterygiophore are
noted as “1½”. Frequency of meristic values are indicated
in parentheses, if more than one value is observed; asterisks
indicate the condition for the holotype. Abbreviations used:
CMK, collection of the author; MHNG, Muséum d’Histoire
Naturelle, Genève; and ZRC, Lee Kong Chian Natural History
Museum, Singapore.
The toponymy of northern Myanmar is extremely labile,
depending of different transcription systems of the languages
of the various ethnic and language groups. The following
alternative spellings of the mentioned places can be seen
on maps and in the literature: Kang Mu Lon (Kaung Mu
Lon), Mali Hka (Malika, Malikah, Malikha), Mon Lan
Chaung (Mone Lan, Mon Laing), Phunganrazi (Phonganrazi,
Phanganrazi, Hponganrazi, Hpon-kan Razi), Wa Sar Dam
(Wasandum, Wasandam, War Zan Dum, Wasondum). Dam
means village and chaung means stream.
Malihkaia, new genus
Type species. Malihkaia aligera, new species (Figs. 1–2).
Diagnosis. Malihkaia is distinguished from all other genera
of nemacheilids by the unique morphology of the lips and
the unique pectoral n sexual dimorphism. Lips thick, with
numerous, closely set, deep furrows (Fig. 3). Upper lip with
a small median notch, with transverse furrows on its whole
length, edge crenulated. Lower lip with a median interruption;
with transverse furrows on its whole length, edge crenulated.
In the male, the pectoral n is strongly falcate (Fig. 2). The
unbranched ray is exible and shaped as in other species
of Nemacheilidae. The rst branched ray is rigid, arched
and curled upwards (Fig. 4). It is about 5 times wider than
the other branched rays. It is attened dorso-ventrally and
very elongate, reaching beyond pelvic-n base. It is rst
branched at about midlength. The posterior main branch is
thicker than the anterior main branch; it is branched again
twice, the third branching about at distal one-sixth of the
ray. The anterior branch is branched again only once, in a
position intermediate to that of branching points 2 and 3
of posterior main branch. There is no membrane between
all the branches and sub-branches, but there is a membrane
around the tip of the ray.
The second and third branched rays are very slender, branched
only at their distal third, the anterior branch is thicker than
the posterior one and it is branched again once; the posterior
branch has no sub-branches. The membrane between the
branches is very narrow. The spaces and relative ‘depth’
82
Kottelat: Three new loaches from Myanmar
Fig. 1. Malihkaia aligera, MHNG 2766.051, holotype, male, 65.7 mm SL; Myanmar: Irrawaddy drainage; Mali Hka River near Putao.
of the branching increase regularly in the following rays.
The fourth and following branched rays have two branches
that are each branched once. There is some variation in the
branching apparently related to size and small injuries; the
above is based on intact ns.
The lower surface of the unbranched ray and the unbranched
part of rst branched ray and the membrane between the
three rst rays are covered by an unculiferous pad (Conway
et al., 2012), thickest under the rst branched ray. There
are also pads under the membranes between the other rays,
but less developed. There also are granulations (unculi?)
on the dorsal surface of the rays and thickened tissue on
the membranes.
In females, the pectoral n is rounded; there are unculiferous
pads under the rays, thickest under the unbranched and rst
branched ones, decreasing in extant and thickness posteriorly.
The unbranched ray is thicker than the following rays and
rigid; dorsally, the membrane between the rst two rays is
thickened. The rst branched ray is thickened but exible,
round in cross-section, thicker than the following rays;
the anterior branch is unbranched. The posterior branch is
branched once, the anterior sub-branch is unbranched and the
posterior one branched again. In the second branched ray,
the anterior branch is unbranched and the posterior branch
is branched again. The remaining rays are all branched twice
as in other nemacheilids. There is some little variation in
the branching, related with small injuries and regrowth; the
above is based on one intact n on each female.
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RAFFLES BULLETIN OF ZOOLOGY 2017
Fig. 2. Malihkaia aligera, CMK 25508, paratypes; a, male, 73.0 mm SL; b, female, 77.2 mm SL; Myanmar: Irrawaddy drainage: Mali
Hka River near Putao.
84
Kottelat: Three new loaches from Myanmar
Fig. 4. Malihkaia aligera, CMK 25508, 73.0 mm SL; left pectoral
n of male; dorsal (above) and ventral (below) view. Extremity of
fourth branched ray damaged.
Fig. 3. Malihkaia aligera, CMK 25508, 73.0 mm SL; ventral
view of head.
In both sexes, in both pelvic and pectoral ns, the anterior
edge of the unbranched ray has a thick tissue cover. In both
sexes, the pelvic n has thickened tissue on membranes
between the unbranched ray and branched rays 1–3, dorsally
and ventrally, also covering part of these rays ventrally.
Additional characters useful to distinguish the genus are:
processus dentiformis present; suborbital ap present in
males; 8½ branched dorsal-n rays; 9+8 branched caudal-n
rays; axillary pelvic lobe present; anus situated about 1.5–2
eye diameter in front of anal n; body entirely scaled; lateral
line complete; air bladder without posterior chamber; body
with 9–12 bars extending from dorsal midline to level of
pectoral n, bars of quite regular width and shape in front
of dorsal fin, less regular posteriorly, much wider than
interspaces; and black mark at caudal-n base made of a
more or less squarish blotch in middle of base, a more or
less triangular blotch over dorsal procurrent rays, fainter
pigments over base of uppermost and lowermost 4–6 principal
rays of caudal n.
Etymology. Named for Mali Hka River, where the type
species was discovered. Gender feminine.
Remarks. The deeply furrowed lower lip of Malihkaia aligera
is unique in South and Southeast Asian Nemacheilidae.
Acanthocobitis (including Paracanthocobitis, see Kottelat,
2012b: 74, 2013: 198) has papillated lips (Kottelat, 1990,
2012a). On the upper lip, the papillae are organised in several
rows; on the lower lip there is a broad median interruption
and, on each side, a widened, strongly papillated pad, covered
by papillae. Species of Acanthocobitis also have a very
different colour pattern, made of obliquely organised bars,
usually of irregular shape and organisation. All species of
Acanthocobitis have a small black spot (sometime ocellated)
at the upper extremity of the base of the caudal n. Finally,
species of Acanthocobitis have a longer dorsal n, with
9–18½ branched rays.
In Nemacheilidae, the deeply furrowed lips are otherwise
known only in some species of Labiatophysa, Triplophysa
and Tarimichthys, all endemic to the Tibetan plateau, the
Tarim and adjacent endorheic basins in China, Kazakhstan
and Kyrgyzstan, and adjacent waters. These genera are
distinguished, among others, by the absence of scales (most
species), males with densely set unculi on the side of the
head below the eye and on the dorsal surface of anterior
pectoral-n rays (Prokoev, 2004, 2010).
Cobitiforms have developed a great variety of sexual
dimorphism, but most of it associated with the pectoral ns
and involving thickening and modication of rays (see, e.g.,
Šlechtová et al., 2008 for sexual dimorphism in Cobitis and
related genera; Kottelat & Lim, 1992 for Lepidocephalichthys;
Kottelat & Tan, 2008 for Kottelatlimia and Acantopsis;
Kottelat, 1990 for Indochinese Nemacheilidae; Bohlen &
Šlechtová, 2011 for Pteronemacheilus); sexual dimorphism
may also involve, for example, thickening of parts of the
body (e.g., in Sabanejewia, Kottelat & Freyhof, 2007) or
vertical torsion of the caudal peduncle (in Microcobitis; pers.
obs., unpubl.). The modied pectoral n of male M. aligera
seems unique in Nemacheilidae but is very reminiscent of
what is observed in some Cobitidae, especially species of
Acantopsis. In Acantopsis, the branches of the rst branched
ray are in contact and there is no membrane between them
and the ray is covered by a dense layer of unculi (pers. obs.;
Kottelat & Tan, 2008). In Kottelatlimia, the rst branched ray
is branched only at the tip and may appear as an unbranched
ray (Kottelat & Tan, 2008). A curled pectoral n in males
is observed in Cobitidae in Acantopsis and Pangio (e.g.,
Kottelat & Tan, 2008; Kottelat & Lim, 1993) but has not
been reported in Nemacheilidae before.
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RAFFLES BULLETIN OF ZOOLOGY 2017
Table 1. Morphometric data of type material of Malihkaia aligera (n=6). Range and mean include holotype data.
Holotype Range Mean
Standard length (mm) 65.7 64.9–77.2
Total length (mm) 81.4 80.8–94.5
In percent of standard length
Total length 123.8 122.0–124.6 123.2
Head length (dorsal) 20.5 20.5–23.6 22.1
Head length (lateral) 24.6 23.8–25.9 24.8
Predorsal length 51.3 50.2–54.7 52.0
Prepelvic length 53.1 52.3–54.0 53.0
Pre-anus length 74.5 72.7–76.4 74.7
Pre-anal length 79.2 77.4–80.6 79.2
Head depth 13.9 13.6–14.4 14.0
Body depth at dorsal-n origin 17.2 16.0–19.0 17.1
Depth of caudal peduncle 11.4 10.3–11.9 11.1
Length of caudal peduncle 14.6 14.6–16.8 15.6
Head width 14.9 14.0–16.2 15.0
Body width at dorsal-n origin 11.5 10.5–13.3 11.7
Snout length 9.8 9.7–11.2 10.3
Eye diameter 5.0 4.4–5.1 4.8
Interorbital width 5.8 5.2–5.9 5.6
Length of dorsal n 21.6 17.3–21.6 19.7
Length of upper caudal-n lobe 21.5 20.5–22.7 21.6
Length of median caudal-n rays 16.1 14.1–16.8 16.1
Length of lower caudal-n lobe 25.4 20.7–25.4 23.0
Length of anal n 17.8 16.7–18.2 17.6
Length of pelvic n 18.8 17.6–20.4 18.7
Length of pectoral n 30.5 19.8–32.2 31.0
In percent of dorsal head length
Snout length 48 41–50 46
Eye diameter 24 18–24 22
Interorbital width 28 23–28 25
In percent of lateral head length
Snout length 40 40–44 41
Eye diameter 20 18–20 19
Interorbital width 24 21–24 23
Malihkaia aligera, new species
(Figs. 1, 2)
Holotype. MHNG 2766.051, male, 65.7 mm SL; Myanmar:
Kachin State: Mali Hka River, about 9 km upstream of Kang
Mu Lon; 402 masl; 27°25′54″N 97°27′56″E; M. Kottelat,
Nyein Chan et al., 26 November 2014.
Paratypes. CMK 25508, 2 males, 64.9–73.0, 2 females,
73.1–77.2 mm SL; ZRC 55630, 1 male, 65.9 mm SL; same
data as holotype.
Diagnosis. See generic diagnosis for characters distinguishing
the species from all other nemacheilids.
Description. See Figs. 1 & 2 for general appearance and
Table 1 for morphometric data of holotype and paratypes.
An elongate nemacheilid with body depth increasing until
somewhat in front of dorsal-n origin, then decreasing until
posterior extremity of dorsal-n base. Behind dorsal n, body
depth almost uniform until caudal-n base. Dorsal prole
with a marked hump behind head. Head strongly arched in
lateral view, slightly depressed; body slightly compressed
anteriorly to very compressed on caudal peduncle. Interorbital
area arched. Eye below dorsal prole of head. Head deep,
with high cheek. Cheeks not swollen (in both sexes). Snout
rounded in dorsal and lateral view. Caudal peduncle 1.3–1.5
times longer than deep, of uniform depth. Low dorsal keel
on posterior half of post-dorsal area. Low ventral keel on
posterior half of caudal peduncle. Dorsal keel continuous
with upper margin of caudal n. Largest recorded size 77.2
mm SL.
Dorsal n with 4 unbranched and 8½ branched rays; distal
margin straight to slightly concave. Second branched ray
longest. In rst branched ray, anterior branch not branched
a second time, shorter than posterior branch. Pectoral n
with 1 unbranched and 10 branched rays, falcate in males,
86
Kottelat: Three new loaches from Myanmar
reaching beyond pelvic-fin base, rounded in females,
reaching about two thirds of distance to pelvic-n base (see
diagnosis of genus for more detailed description of pectoral
n; Fig. 4). Pelvic n with 1 unbranched and 6 (1) or 7
(5*) branched rays, reaching about three-fths of distance
to anal-n origin (not reaching anus); rounded; posterior
margin straight; origin below base of branched dorsal-n
rays 1 to 2. Axillary pelvic lobe present, entirely free. Anus
situated about 1.5–2 eye diameter in front of anal n. Anal
n with 3 unbranched and 5½ branched rays; distal margin
straight. Caudal n with 9+8 branched rays; forked, lobes
rounded, lower lobe slightly longer than upper one; upper
lobe 1.3–1.5 times longer than median rays.
Body entirely scaled except on anterior half of predorsal area
and belly in front of pelvic ns. Lateral line complete, with
89–99 pores. Cephalic lateral line system with 6 supraorbital,
4+10–11 infraorbital, 8–9 preoperculo-mandibular and 3
supratemporal pores.
Anterior nostril pierced in front side of a pointed ap-like
tube. Posterior nostril adjacent to anterior one. Mouth
U-shaped, gape about 1.5 times wider than long (Fig. 3). Lips
thick, with numerous, closely set, deep furrows. Upper lip
with a small median notch, with furrows on its whole length,
edge crenulated. Processus dentiformis present. Lower lip
with a median interruption; with furrows on its whole length,
edge crenulated. Tip of lower jaw not exposed. No median
concavity in lower jaw. Inner rostral barbel reaching beyond
corner of mouth; outer one reaching vertical of middle of
eye. Maxillary barbel reaching beyond posterior margin of
eye. Intestine with a loop posterior to stomach (Fig. 5). Air
bladder without visible posterior chamber.
Sexual dimorphism. Males with suborbital ap; females
with neither suborbital ap nor suborbital slit. Pectoral n
rounded in females, strongly falcate in males. In males,
unbranched and rst branched rays rigid, arched and curled
upwards. First branched ray about 5 times wider than other
branched rays, attened dorso-ventrally, reaching beyond
pelvic-n base; without membrane between the branches
and all sub-branches. Second branched ray slender, branched
only at tip, membrane between branches very narrow. See
diagnosis of genus for more detailed description of pectoral
n of both sexes.
Coloration. In formalin, shortly after fixation. Body
background colour pale yellowish brown, throat and belly
whitish; except otherwise stated, all markings blackish
brown to black. Head black, cheek marmorated, lower side
whitish. Irregularly set and shaped patches of black pigments
on lips, but always a patch on posterior edge of lower lip
and on adjacent part of throat. Body with 9–12 bars (3–4
predorsal, 3 subdorsal, 3–5 postdorsal), extending from dorsal
midline to level of pectoral n (or ventral midline on caudal
peduncle), mostly continuous over back with contralaterals;
bars of quite regular width and shape in front of dorsal n,
less regular posteriorly; much wider than interspaces.
A conspicuous more or less squarish black blotch at middle
of caudal-n base, depth about ⅓–¼ of depth of base of n.
A more or less triangular black blotch over dorsal procurrent
rays, including dorsal midline. Fainter pigments over base
of uppermost and lowermost 4–6 principal rays of caudal
fin, making a slightly elongated, arched mark, variably
contrasted, and variably connected with other two blotches.
Space between upper triangular blotch and last bar on body
somewhat reddish. Axial stripe faint, or not distinct except
on caudal peduncle where darker and wider, combining
with blotch at middle of caudal-n base to make a larger
and more conspicuous blotch.
Dorsal n hyaline, no pigments on membranes, with a small
black spot at base of unbranched rays and middle and distal
parts of last unbranched ray black; pigments on rays at
branching points and on middle of unbranched part of rst
branched rays (and branched rays 2–4 in largest specimens).
Caudal n hyaline, with pigments between segments of
branched rays and near branching points. Anal n hyaline,
with pigments near branching points of branched rays.
Pelvic fin hyaline, with pigments along rays and near
branching points of branched rays. Pectoral fin with
membranes hyalines, pigments on rays, darker along posterior
edge of branched rays 2–10. In males, unbranched and rst
branched rays more or less completely covered by pigments,
on dorsal side, on whole length; identical in females, but
pigments less densely set.
Fig. 5. Malihkaia aligera, MHNG 2766.051, holotype,
65.7 mm SL; digestive tract. Scale bar = 1 mm.
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RAFFLES BULLETIN OF ZOOLOGY 2017
Notes on biology. The two females did not have ripe ovaries.
The stomach contained unidentiable insect remains.
Distribution and habitat. Malihkaia aligera has been
observed only once, in the Mali Hka River near Putao.
It was collected over gravel and cobble, in fast rifes, in
about 60 cm depth (Fig. 6). Other species collected in the
same microhabitat at this locality include Bangana sp.,
Garra sp. (Cyprinidae), Psilorhynchus brachyrhynchus
(Psilorhynchidae), Homalopteroides rupicola (Balitoridae),
Acanthocobitis sp., Schistura malaisei, S. nubigena, S.
sikmaiensis (Nemacheilidae), Batasio procerus (Bagridae),
Amblyceps murraystuarti (Amblycipitidae), Pseudecheneis
brachyurus (Sisoridae) and Mastacembelus armatus
(Mastacembelidae). Schistura malaisei and Homalopteroides
rupicola were the most abundant species.
Etymology. From the Latin aliger (aligera, aligerum),
winged. An adjective.
Remarks. The colour pattern of the six available specimens
of M. aligera is made of very regular bars on body, with
Fig. 6. Type locality of Malihkaia aligera and Schistura nubigena;
Myanmar: Kachin state: Mali Hka River near Kang Mu Lon; 402
masl; 26 November 2014.
Fig. 7. Schistura sikmaiensis, CMK 25560; Myanmar: Irrawaddy drainage: Mula River at Lika; a, female, 61.6 mm SL; b, male, 72.7 mm SL.
very well marked edges, on a pale background. This
general pattern is rare among Southeast Asian nemacheilids.
Interestingly, one of the species collected together with M.
aligera is Schistura sikmaiensis, which also has a regular and
contrasted colour pattern (Fig. 7). A third species collected
at the same locality is S. nubigena (described below), which
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Kottelat: Three new loaches from Myanmar
Fig. 8. Schistura nubigena, MHNG 2766.052, holotype, 33.6 mm SL; Myanmar: Irrawaddy drainage: Mali Hka River near Putao.
has a sharply contrasted colour pattern of 4 bold black bars
on a whitish background. Finally, S. wanlainensis, also
described below, from the same area but at higher altitude
and habitat with higher gradient, also has a very contrasted
colour pattern of very regular bars on a pale background.
Schistura nubigena, new species
(Figs. 8–10)
Holotype. MHNG 2766.052, 33.6 mm SL; Myanmar: Kachin
state: Mali Hka River, about 9 km upstream of Kang Mu
Lon; 402 masl; 27°25′54″N 97°27′56″E; M. Kottelat, Nyein
Chan et al., 26 November 2014.
Paratypes. CMK 25509, 11, 19.5–38.7 mm SL; same data
as holotype.
Diagnosis. Schistura nubigena is distinguished from all other
species of the genus by its unique colour pattern made of
4 black bars in juveniles that evolve into 8 bars, more or
less fused on the ank and the back to leave only 2 series
of pale yellowish spots in the predorsal area and 4 narrow
transverse bands in subdorsal and postdorsal areas. Although
not unique, the following characters help to identify the
species: incomplete lateral line, reaching at most to above
pelvic-n base; two vertically elongated blotches on proximal
extremity of all rays of caudal n (one blotch per lobe);
no observed sexual dimorphism; 8½ branched dorsal rays.
Description. See Figs. 8–10 for general appearance and
Table 2 for morphometric data of holotype and three largest
paratypes. A moderately elongate nemacheilid with body
depth slowly increasing up to about above tip of pectoral n.
Behind dorsal n, body depth decreasing slowly to caudal-n
base. Dorsal prole with a small concavity between head and
body. Head slightly compressed; body slightly compressed
anteriorly to compressed posteriorly. Interorbital area slightly
convex. In lateral view, eye below or ushed with dorsal
prole of head. Cheeks not swollen. Snout rounded in dorsal
and lateral view. Caudal peduncle depth 1.6–2.0 times in its
length, of uniform depth. Low dorsal crest on posterior half
of post-dorsal area. Low ventral crest on entire length of
caudal peduncle. Dorsal crest continuous with upper margin
of caudal n. Largest recorded size 38.7 mm SL.
Dorsal n with 4 unbranched and 8½ branched rays; distal
margin straight to slightly concave. Second branched ray
longest. Pectoral n with 1 unbranched or 8 (3*) and 9 (1)
branched rays (including small last rays, usually unbranched),
rounded, reaching about two thirds of distance to pelvic-n
base. No axillary pectoral lobe. Pelvic n with 1 unbranched
and 6 branched rays; reaching to anus; falcate, posterior
margin rounded; origin below base of fourth unbranched or
rst branched dorsal-n ray. Axillary pelvic lobe present,
entirely free. Anus situated about 1 eye diameter in front
of anal n. Anal n with 3 unbranched and 5½ branched
rays; distal margin straight. Caudal n with 9+8 branched
rays; forked, lobes rounded, lower lobe slightly longer than
upper one.
Body entirely covered by scales, except nape and predorsal
area, and belly in front of anal n. Scales deeply embedded.
Lateral line incomplete, reaching between tip of pectoral n
and pelvic-n origin, with 19–31 pores (number apparently
increasing with increasing size). Cephalic lateral line system
with 6 supraorbital, 4+11 infraorbital, 9 preoperculo-
mandibular and 3 supratemporal pores.
Anterior nostril pierced in front side of a pointed ap-like
tube. Posterior nostril adjacent to anterior one. Mouth
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Fig. 9. Schistura nubigena, CMK 25509, paratypes; Myanmar: Irrawaddy drainage: Mali Hka River near Putao; a, 20.0 mm SL; b, 21.1
mm SL; c, 23.8 mm SL; d, 26.7 mm SL; e, 26.9 mm SL.
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Kottelat: Three new loaches from Myanmar
U-shaped, gape about 2 times wider than long (Fig. 11).
Lips thin but eshy. Upper lip without median notch, with
numerous shallow furrows on whole length, edge nely
crenulated. Processus dentiformis present. Lower lip with
wide median notch; median part with 2–4 sulci, lateral parts
smooth. Tip of lower jaw exposed. No median notch or
concavity in lower jaw. Inner rostral barbel reaching slightly
beyond base of outer one; outer one reaching slightly beyond
base of maxillary barbel. Maxillary barbel reaching almost
vertical of posterior margin of eye. Intestine straight behind
stomach (Fig. 12). Air bladder without free posterior chamber.
Sexual dimorphism. None of the characters associated
with sexual dimorphism in other nemacheilids have been
observed in S. nubigena.
Coloration. After fixation in formalin. Head and body
background colour pale yellowish grey; throat, belly and
lower part of caudal peduncle whitish; except otherwise
stated, all markings dark brown to black. Top of head and
opercle dark brown. A white squarish patch between tip of
snout and nostrils; a few smaller, less sharply marked pale
spots on top of head.
Body with 4 bars (behind head, below dorsal-n origin,
below end of dorsal-n base and on caudal peduncle). Bars
continuous across back with contralaterals, not reaching
down to level of pectoral fin. Bars slightly wider than
interspaces in small specimens (less than about 22 mm SL;
Fig. 9b), becoming wider with increasing size. Bars wider
on dorsal midline and at lower extremity. In many specimens
bars appearing divided vertically with a paler median area
(Fig. 9e). Some or all bars maybe in contact at lower and
upper extremities. In largest specimen, all bars appearing as
vertically split in two (total 8 bars; Fig. 10), and all fused
along ank in a broad milateral stripe, predorsal bars also
merged on back, leaving only two longitudinal rows of 3
yellowish spots. Interspace between subdorsal and postdorsal
bars persisting as 4 narrow transverse band; an additional
one between head and rst ank bar. In most specimens,
Table 2. Morphometric data of holotype and three largest paratypes of Schistura nubigena.
Paratype Holotype Paratype Paratype
Standard length (mm) 38.7 33.6 26.2 26.9
Total length (mm) 48.1 40.9 32.2 33.1
In percent of standard length
Total length 124.2 121.8 122.6 123.4
Head length (dorsal) 22.1 21.4 24.0 21.1
Head length (lateral) 23.2 24.6 24.8 23.9
Predorsal length 54.2 54.2 54.7 52.4
Prepelvic length 52.8 51.8 52.6 50.1
Pre-anus length 71.7 69.0 70.9 69.6
Pre-anal length 76.3 73.1 72.5 76.4
Head depth 13.0 11.1 11.2 11.7
Body depth at dorsal-n origin 15.6 16.4 12.6 14.1
Depth of caudal peduncle 11.0 11.4 11.0 9.9
Length of caudal peduncle 21.5 18.4 20.2 18.3
Head width 14.6 14.4 13.2 13.9
Body width at dorsal-n origin 13.7 11.8 9.2 9.6
Snout length 10.0 9.6 9.9 9.2
Eye diameter 4.2 4.2 4.7 4.8
Interorbital width 5.5 6.0 6.9 7.0
Length of dorsal n 21.4 20.4 20.8 20.2
Length of upper caudal-n lobe 22.7 20.6 19.2 19.5
Length of median caudal-n rays 18.2 17.9 15.1 17.7
Length of lower caudal-n lobe 23.4 22.8 19.8 21.3
Length of anal n 22.2 21.0 19.3 20.9
Length of pelvic n 19.9 17.5 17.3 19.3
Length of pectoral n 21.6 20.3 16.0 19.4
In percent of dorsal head length
Snout length 45 45 41 44
Eye diameter 19 20 20 23
Interorbital width 25 28 29 33
In percent of lateral head length
Snout length 43 39 40 39
Eye diameter 18 17 19 20
Interorbital width 24 25 28 29
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Fig. 10. Schistura nubigena, CMK 25509, paratype, 38.7 mm SL; Myanmar: Irrawaddy drainage: Mali Hka River near Putao.
Fig. 11. Schistura nubigena, CMK 25509, paratype. 38.7 mm
SL; mouth.
posteriormost bar more irregular than others, with lower
posterior corner projecting towards ventral midline and
caudal-n base (Fig. 9b–e), and in two specimens appearing
as a separate blotch.
Black marks at caudal-n base appearing as two vertically
elongated deep black blotches, one covering proximal
extremity of all branched rays of lower lobe, and one on all
those of upper lobe (best seen on Fig. 9c). Two posterior
dorsal procurrent rays black. Overimposed on upper extremity
of lower black blotch, a small patch of less dense pigments,
oriented diagonally upwards forwards. In larger specimens,
this last patch may be connected to lower posterior extension
of posterior bar, leaving a pair of large roundish white
blotches (paler than rest of yellowish background colour;
Figs. 9d, e, 10f). A faint inner axial stripe visible in few
specimens.
Dorsal n with hyaline membranes and a dark patch along
anterior half of base, above subdorsal bar; an elongate
patch of black pigments at midlength of last unbranched
ray and near rst branching of branched rays. Caudal n
with hyaline membranes, and pigments along all rays and
between all segments. Anal and pelvic n membranes and
rays hyaline. Pectoral fin membranes and rays hyaline;
in largest specimens (33.6, 38.7 mm SL; Figs. 8, 10), an
elongate patch of pigment near branching of branched rays
1–4, on dorsal side only.
Distribution. Schistura nubigena has been observed only
once, in the Mali Hka River near Putao. It was collected
over gravel and cobble, in fast riffles (Fig. 6) together
with Malihkaia aligera; see under that species for more
information.
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Kottelat: Three new loaches from Myanmar
Fig. 12. Schistura nubigena, MHNG 2766.052, holotype, 33.6 mm
SL; digestive tract. Scale bar = 1 mm.
Etymology. From the Latin adjective nubigenus, -a, -um,
who engenders clouds, a reference to the white spots on the
back resulting from the partial fusion of the bars (nubigena is
also a noun meaning cloud-born, but this is not the meaning
used here).
Remarks. Schistura nubigena has a colour pattern made of
4 black bars in juveniles that evolve into 8 bars, more or
less fused on the ank and the back to leave only 2 series
of pale yellowish spots in the predorsal area and 4 narrow
transverse bands in subdorsal and postdorsal areas. This
colour pattern, and its ontogeny, has not been reported for any
named species of nemacheilid in Southeast and South Asia.
Schistura wanlainensis, new species
(Figs. 13, 14)
Holotype. MHNG 2766.053, female, 84.4 mm SL; Myanmar:
Kachin State: Phunganrazi Wildlife Sanctuary: area of Wa
Sar Dam: Mon Lan Chaung (stream) near Wan Lain Dam
village); 27°29′10″N 97°10′29″E, 850 masl; Nyein Chan,
25 July 2015.
Paratypes. CMK 26417, 5, 79.6–91.8 mm SL; same data as
holotype. — CMK 26584, 1, 81.2 mm SL; CMK 26377, 3,
71.2–78.2 mm SL; ZRC 55631, 1, 74.8 mm SL; Myanmar:
Phunganrazi Wildlife Sanctuary: area of Wa Sar Dam: Mon
Lan Chaung (stream); 27°28′51″N 97°10′35″E, 843 masl;
Nyein Chan, 22 July 2015.
Diagnosis. Schistura wanlainensis is distinguished from
all other species placed in Schistura in Southeast Asia by
its unique colour pattern of a pale yellowish to pale grey
background with 18–32 narrow, regularly shaped black bars,
extending from dorsal midline and reaching downwards
to below level of pectoral n, continuous over back with
contralaterals in most specimens, wider than interspaces
anteriorly, narrower than interspaces on caudal peduncle;
some of anterior bars fused at their dorsal extremity.
Additional characters useful to identify the species (but
not unique) are: black pattern at base of caudal n made
of a vertically elongated blotch at middle of base, and a
smaller blotch at its dorsal and ventral extremities (blotches
contiguous in one specimen); 8½ branched dorsal-n rays;
caudal n forked; caudal peduncle depth 1.2–1.5 times in its
length; very low dorsal and ventral crests on caudal peduncle;
lips thick, eshy; upper lip with median notch; lower lip
with narrow median interruption, with numerous furrows;
processus dentiformis present, feebly marked, very wide
(more than half of mouth gape); largest recorded size 91.8
mm SL; and females with a suborbital slit (males unknown
but expected to have a suborbital ap).
Description. See Figs. 13 & 14 for general appearance and
Table 3 for morphometric data of holotype and paratypes. A
moderately elongate nemacheiline with a massive appearance.
Body depth increasing up to dorsal-n origin. Behind dorsal
n, body depth decreasing slowly until above base of anal
n, then uniform until caudal-n base.
Dorsal prole continuous between head and body. Head
slightly depressed; body slightly compressed anteriorly, very
compressed posteriorly. Interorbital area arched. Eye below
dorsal prole of head. Cheeks not swollen. Snout rounded
in lateral and dorsal views. Caudal peduncle 1.2–1.5 times
longer than deep, of uniform depth. Very low dorsal crest
on posterior half of post-dorsal area. Very low ventral
crest along entire length of caudal peduncle. Dorsal crest
continuous with upper margin of caudal n. Largest recorded
size 91.8 mm SL.
Dorsal n with 4 unbranched and 8½ branched rays; distal
margin straight to slightly concave. Second branched ray
longest. Pectoral n with 1 unbranched and 11 branched rays
(two specimens with an additional small, unbranched ray),
rounded, reaching about halfway of distance to pelvic-n
base; rays without lamentous extensions. No proper axillary
pectoral lobe, but swelling dorsally at base of n. Pelvic n
with 1 unbranched and 7 branched rays; reaching about two
thirds of distance to anus; posterior edge rounded, second or
third ray longest; origin below base of branched dorsal-n
rays 2–3. Axillary pelvic lobe present, free, conspicuous.
Anus situated about 2 eye diameters in front of anal n,
hidden under a protuberant, massive genital papilla (at least
in females before spawning). Anal n with 3 unbranched and
5½ branched rays; distal margin slightly concave. Caudal n
with 9+8 branched rays; dorsal and ventral procurrent rays
cannot be counted; forked, upper lobe 1.4–1.8 times longer
than median rays, lobes rounded, of equal length.
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Fig. 13. Schistura wanlainensis, MHNG 2766.053, holotype, 84.4 mm SL; Myanmar: Irrawaddy drainage: Mon Lan Chaung near Wan
Lam Dam.
Body entirely scaled, except around lateral line pores 1–5 and
belly between pectoral ns. Scales deeply embedded and not
densely set in anterior half of body. Lateral line complete,
with 90–106 pores (difcult to count with accuracy). Many
lateral line scales with 2 auxillary pores, above and below
main pore. Cephalic lateral line system with 6 supraorbital,
4+11 infraorbital, 8–10 preoperculo-mandibular and 3
supratemporal pores.
Anterior nostril pierced in front side of a pointed ap-like
tube. Posterior nostril adjacent to anterior one. Mouth arched,
gape about two times wider than long (Fig. 15). Lips thick,
eshy. Upper lip with median notch, with furrows along
whole length, edge crenulated. Lower lip with narrow median
interruption; median part without sulci, lateral part with
numerous furrows on whole length. Processus dentiformis
present, feebly marked, very wide (more than half of mouth
gape); no sharp edge on upper jaw. Tip of lower jaw
exposed. A median concavity in lower jaw. Inner rostral
barbel not reaching corner of mouth; outer one reaching
base of maxillary barbel. Maxillary barbel reaching vertical
of posterior margin of eye. In single dissected specimen,
intestine with a large loop behind stomach (Fig. 16). No
apparent posterior chamber of air bladder.
Sexual dimorphism. All specimens apparently females;
presence of eggs, about 0.6 mm diameter observed in four
specimens 78.2–91.8 mm SL; most slender specimen (85.6
mm SL) indeterminate, remaining specimens not checked
but all with distended belly. All specimens have a short slit
in a position that would correspond to posterior edge of
suborbital ap in species in which males have a suborbital
ap (compare Figs. 17 & 18); in species in which such a
slit is present in females, the males usually have a suborbital
ap. No tubercles or unculi observed.
Coloration. After two months in formalin. Head and body
background colour pale yellowish to pale grey; throat and
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Kottelat: Three new loaches from Myanmar
Fig. 14. Schistura wanlainensis, CMK 26417, paratypes; Myanmar: Kachin state: Irrawaddy drainage: Mon Lan Chaung near Wan Lam
Dam; a, 85.6 mm SL; b, 91.8 mm SL.
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posteriorly by a band of black and red pigments on upper
branched ray. Lower blotch elongated, slanted towards lower
edge of caudal n, on base of principal unbranched ray and
lower branched rays 1–3, continued posteriorly by a band of
black and red pigments on lower branched ray. No visible
inner axial stripe.
Dorsal fin with hyaline membranes. All branched rays
black except distal extremity; proximal third paler in some
specimens, greyish in one (holotype). Last unbranched
ray black on posterior half, orange on anterior half. Other
unbranched rays black. Membrane between last unbranched
and rst branched ray orange in distal area; membranes
between branches of rst branched ray orange; distalmost
part of membrane between rst two branched rays yellowish.
Caudal n with hyaline membranes in most specimens, pale
orange in some. Rays blackish brown. One or two irregular
and incomplete vertical rows of spots corresponding to
Table 3. Morphometric data of type material of Schistura wanlainensis (n=11). Range and mean include holotype data.
Holotype Range Mean
Standard length (mm) 84.4 71.2–91.8
Total length (mm) 100.0 95.9–108.8
In percent of standard length
Total length 118.4 116.0–121.5 119.3
Head length (dorsal) 17.8 18.5–21.0 19.8
Head length (lateral) 21.4 21.4–23.2 22.1
Predorsal length 51.0 50.0–54.7 51.9
Prepelvic length 52.7 52.7–56.7 54.3
Pre-anus length 73.9 73.1–76.4 75.0
Pre-anal length 79.5 77.0–80.2 79.2
Head depth 12.4 12.4–14.2 13.0
Body depth at dorsal-n origin 23.1 19.6–24.9 22.6
Depth of caudal peduncle 11.6 10.7–12.5 11.5
Length of caudal peduncle 14.9 14.6–17.6 15.8
Head width 14.3 14.3–16.0 15.1
Body width at dorsal-n origin 17.2 14.7–21.0 17.3
Snout length 8.8 8.8–10.2 9.5
Eye diameter 3.1 2.4–3.7 3.1
Interorbital width 5.7 5.5–6.1 5.8
Length of dorsal n 16.6 15.6–19.7 18.1
Length of upper caudal-n lobe 16.8–16.8 21.1
Length of median caudal-n rays 10.7 10.0–13.4 10.6
Length of lower caudal-n lobe 19.9 17.5–21.4 19.8
Length of anal n 15.1 14.1–17.7 15.8
Length of pelvic n 16.6 15.1–17.8 16.6
Length of pectoral n 18.1 16.4–19.8 18.6
In percent of dorsal head length
Snout length 50 46–49 48
Eye diameter 17 12–17 16
Interorbital width 32 27–33 29
In percent of lateral head length
Snout length 41 40–45 43
Eye diameter 14 11–15 14
Interorbital width 27 25–28 26
belly whitish to pale yellowish; darker on back; head darker
grey in most specimens. Except otherwise stated, markings
blackish brown to black. Head with vermiculated pattern on
top, snout, suborbital area and opercle. Body with 18–32 bars,
extending from dorsal midline and reaching downwards to
below level of pectoral n (on belly in largest specimens
but not reaching ventral midline); continuous over back
with contralaterals in most specimens; bars of regular shape,
wider than interspaces anteriorly, narrower than interspaces
on caudal peduncle; some of anterior bars fused at their
upper extremity.
Black pattern at base of caudal n made of a vertically
elongated blotch at middle of base, and a smaller blotch at
its dorsal and ventral extremities; blotches contiguous in one
specimen. Median blotch occupying about median half of n
base; with a median constriction. Upper blotch elongated,
slanted forwards upwards, at base of principal unbranched ray
and upper branched rays 1–3; in some specimens continued
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accumulation of pigments at primary and secondary branching
points. Black stripes along principal unbranched ray of each
lobe (see above, black pattern at caudal-n base), surrounded
by pale-orange to reddish area.
Anal n with hyaline membranes. Branched rays with one
or two rows of black spots corresponding to accumulation of
black pigments at primary and secondary branching points.
Last unbranched ray orange, as well (in some specimens)
as distal extremity of membrane between last unbranched
and rst branched rays. Blackish areas adjacent to lower
extremity of ank bars.
Pelvic n with hyaline membranes. Branched rays with one
or two rows of black spots corresponding to accumulation of
black pigments at primary and secondary branching points.
Unbranched ray orange (and in one specimen, also distal
extremity of rst branched ray), as well (in some specimens)
as distal extremity of membrane between unbranched and
rst branched rays.
Pectoral n with hyaline membranes. Rays black; in some
specimens, unbranched ray orange as well as anterior half of
rst branched ray. In all branched rays, posterior branch more
densely pigmented than anterior branch. Membrane between
unbranched and rst branched rays orange, at least distally.
Notes on biology. A dissected female (CMK 26584, 81.2
mm SL) with distended belly had ovaries lled with a large
number (probably > 1000) of eggs 0.6 mm in diameter. Its
stomach contained unidentied green vegetal material.
Distribution and habitat. Schistura wanlainensis is presently
known only from the Mon Lan Chaung in the area of Wa
Sar Dam, in Phunganrazi Wildlife Sanctuary (Fig. 19). The
Mon Lan Chaung becomes known as Mula Chaung on Putao
Plain and is a tributary of the Mali Hka, which it enters from
the west on Putao Plain.
At the places of sampling, the Mon Lan Chaung is a large
river about 50 m wide, with very fast water owing over
a stone and boulders bottom. Other species collected in
the same microhabitat at these localities include Garra
cf. dulongensis, Garra sp., Neolissochilus compressus,
Oreinus meridionalis, Semiplotus cirrhosus (Cyprinidae),
Acanthocobitis sp., Schistura malaisei (Nemacheilidae),
Amblyceps murraystuarti (Amblycipitidae), Glyptothorax
sp., Oreoglanis sp. (Sisoridae) and Mastacembelus armatus
(Mastacembelidae). Schistura malaisei was the most abundant
species.
Etymology. Named for Wan Lain Dam village where the
sh were collected, and whose inhabitants have established
a sh conservation zone. An adjective.
Fig. 16. Schistura wanlainensis, CMK 26584, 81.2 mm SL; digestive
track. Scale bar = 10 mm.
Fig. 17. Schistura wanlainensis, CMK 26417, 84.3 mm SL, female;
note suborbital slit.
Fig. 15. Schistura wanlainensis, CMK 26417, 84.3 mm SL; mouth.
Arrows indicate lateral extent of processus dentiformis.
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Fig. 18. Schistura sikmaiensis, CMK 25560; a, female, 61.6 mm
SL, with suborbital slit; b, male, 72.7 mm SL, with suborbital ap.
Fig. 19. Type locality of Schistura wanlainensis; Myanmar: Kachin
state: Mon Lan Chaung near Wan Lain Dam, 850 masl; 25 July
2015. Photograph by Zau Lunn.
Remarks. Schistura sikmaiensis was diagnosed from all
other species of Schistura in Southeast Asia by the absence
of processus dentiformis (Rendahl, 1948: 27; Kottelat, 1990:
208). Examination of additional material suggests that the
processus dentiformis could also be described as present
but feebly marked and very wide, as in S. wanlainensis.
Schistura sikmaiensis (of which S. putaoensis is a synonym)
is recorded from a few localities in the upper Irrawaddy in
Myanmar, China and India (Kottelat, 1990: 208) but it was
not found in the Mon Lan Chaung and its tributaries in the
hills. It was observed only on Putao Plain (see Fig. 7). The
two species share a number of characters, including the
pale background colour, the bars on the anterior part of the
body narrower than those on the posterior part and often
fused in pairs dorsally, a similar black pattern at the base
of the caudal n, and the presence in females of a slit in a
position that corresponds to the posterior edge of suborbital
ap of males.
Schistura wanlainensis is distinguished from S. sikmaiensis
by its general appearance (compare Figs. 13, 14 & 7). For
example, the more slender caudal peduncle whose depth
is 1.8–2.1 times in the depth of the body (vs. 1.3–1.6) and
1.2–1.5 times in its own length (vs. 1.3–1.7), a pointed
snout (vs. rounded), a somewhat shorter head (lateral length
21.4–23.2 % SL vs. 22.3–24.6). Besides, S. wanlainensis
has more (18–32 vs. 13–21) and narrower bars, the bars
on the caudal peduncle narrower than the interspaces (vs.
wider than the interspaces), and the bars on the anterior part
of the body not much narrower than the posterior ones and
not all regularly fused in pairs dorsally (vs. much narrower
and regularly fused). In S. wanlainensis, the bars are of
uniform intensity of black. In S. sikmaiensis, the edge of
the bars is clearly darker than the inner and this is gradually
more marked from the posterior to the anterior bars, with
some bars becoming vertically split and the number of bars
increasing with size.
Comparison material. Schistura sikmaiensis: CMK 25507,
1, 54.5 mm SL; Myanmar: Kachin state: Mali Hka River,
about 9 km upstream of Kang Mu Lon. — CMK 25560, 10,
54.5–77.9 mm SL; Myanmar: Kachin state: Mula River at
Lika village, about 0.8 km downriver of Mulashidi.
ACKNOWLEDGEMENTS
This work partly results from surveys conducted with or by
Fauna & Flora International (FFI), with nancial support
from the Leona M. and Harry B. Helmsley Charitable Trust,
the Critical Ecosystem Partnership Fund, and the Global
Environment Facility – Satoyama Project. Department of
Fisheries gave permission to conduct the surveys. For their
help in the eld in 2014, I thank Nyein Chan and Dankhaung
H-Hkai (FFI). Zau Lunn (FFI) organised the logistics and
provided Figure 19. The July 2015 material was obtained
by Nyein Chan, Zay Yar Aung (FFI) and villagers from
Wa Sar Dam. I thank Jörg Bohlen and Kevin Conway for
reading and commenting on the manuscript.
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