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Lepidocephalichthys eleios, a new loach from Lake Indawgyi basin, Kachin State, Myanmar (Teleostei: Cobitidae).

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RAFFLES BULLETIN OF ZOOLOGY 2017
Lepidocephalichthys eleios, a new loach from Lake Indawgyi basin,
Kachin State, Myanmar (Teleostei: Cobitidae)
Maurice Kottelat
Abstract. Lepidocephalichthys eleios, new species, is described from the basin of Lake Indawgyi in Irrawaddy
drainage in northern Myanmar. It is a small-sized species (largest known specimen 22.8 mm SL) distinguished
from all congeners in having the last two rays of the pectoral n in male adjacent, without membrane between
them, not fused and without dorsal projections (vs. unbranched parts fused and forming a thick rod, with a dorsal
projection in the other species of the genus). Its colour pattern is made of an irregular midlateral row of small spots,
sometimes forming a stripe; a middorsal row of 9–12 irregular saddles; an arched black band at base of principal
caudal-n rays (no ocellus or black spot); caudal n with 2–3 very irregular bars, irregularly connected or fused,
leaving unpigment patches. The species lives in dense vegetation along the shores and in mud and vegetal debris
at the bottom in the marshy lower course of tributaries of the lake and of its outlet. The incorrect use of ‘lamina
circularis’ in recent cobitid literature is discussed.
Key words. Cobitoidei, Irrawaddy River, sexual dimorphism, lamina circularis
RAFFLES BULLETIN OF ZOOLOGY 65: 707–714
Date of publication: 17 November 2017
http://zoobank.org/urn:lsid:zoobank.org:pub:C2F5CC5E-FA76-4357-B181-9B03D80FA398
© 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 genus Lepidocephalichthys typically inhabit
slow owing streams, lakes and marshes, where they mainly
occur on muddy bottom. The genus presently includes 19
valid species (pers. obs., updated from Kottelat, 2012,
2013). It is widely distributed in Southeast and South Asia,
from Sundaland and the Mekong drainage westwards to the
Indus drainage and Sri Lanka. The species from the Malay
Peninsula were revised by Kottelat & Lim (1992); Havird
& Page (2010), and Havird et al. (2010) revised the genus
and described additional species.
Lake Indawgyi is located in the Irrawaddy drainage,
Myanmar, in the hills southwest of Myitkyina. The rst
information on the sh fauna of Lake Indawgyi was published
by Prashad & Mukerji (1929). They recorded 64 species from
the lake itself and its tributaries. Since, a few ichthyologists
visited the lake and this has resulted in the addition of a few
species, either already known from elsewhere in Myanmar
or new to science (e.g., Britz, 2007, 2010; Kullander, 2008,
2012; Kullander & Britz, 2002; Kullander & Fang, 2005;
Ng & Kottelat, 2007, 2009; Kottelat, 2017).
A survey conducted in 2013 by the Forest Department, the
Zoology Department of Myitkyina University and Fauna &
Flora International, and additional observations made in 2014,
brought the number of species from the lake and its basin
to 95 native species (Kottelat, 2015). Safety conditions in
2013 and 2014 restricted very much the areas that could be
visited around the lake; this survey, however, also obtained
what are possibly the rst sh samples from the outlet of the
lake. The outlet (Indaw Chaung) ows through a large area
of marshland, in a deep channel bordered by dense vegetation
and under mats of ‘oating’ vegetation. The present article
describes a new species from the basin of Lake Indawgyi.
MATERIAL AND METHODS
Measurements and counts follow Kottelat (1990) and Kottelat
& Freyhof (2007). Head length is the lateral head length of
Kottelat (1990). Last 2 branched dorsal and anal-n rays
articulating on a single pterygiophore are noted as “1½”.
The last two posterior rays (6 and 7th branched rays) of the
pectoral n, when fused, are called pectoral rod. Terminology
for mouth parts, see Fig. 1; the lower lip is interrupted in
the middle; each half is made of a eshy lobe in medial
position (mental lobe), which ends in a point, may have a
number of indentations on the medial side, and is separated
from the lateral portion by a small notch; the lateral portion
(postlabial ange) is the area between the mental lobe and
the base of the maxillary barbel; the postlabial ange is thin,
extends backwards, its posterior part is free from the throat
and its posterior edge usually lobed or fringed (Fig. 1).
Abbreviations used: CMK, collection of the author; MHNG,
Muséum d’Histoire Naturelle, Genève; and ZRC, Lee Kong
Chian Natural History Museum, Singapore.
Taxonomy & Systematics
708
Kottelat: Lepidocephalicthys eleios
Paratypes. CMK 25643, 13, 16.9–22.8 mm SL (and 3 xed
in 95 % ethanol); same data as holotype. — CMK 24160,
51, 17.2–22.7 mm SL (and 4 xed in 95 % ethanol); ZRC
56629, 5, 20.3–22.0 mm SL; Myanmar: Kachin State: Indaw
Chaung, outlet of Lake Indawgyi; 25°15′14″N 96°25′14″E;
M. Kottelat & Dankhaung H-Hkai, 1 December 2013. —
CMK 24171, 3, 14.8–19.2 mm SL; Myanmar: Kachin State:
Nam Phaung Sin Chaung (stream) near Nam Phaung Chin
village, entering Lake Indawgyi north of Lonton; 25°07′26″N
96°17′26″E; M. Kottelat & Dankhaung H-Hkai, 3 December
2013.
Diagnosis. Lepidocephalichthys eleios is distinguished from
all other species of the genus by lacking any modications
of the pectoral-fin rays in males. Pectoral-fin rays 7–8
are adjacent, with narrow or without membrane between
then; in males, they are separate, not fused and have no
dorsal or ventral projections. Additional characters useful
to distinguish the species from congeners but not unique to
it are: an irregular midlateral row of irregularly set small
spots, sometimes forming a stripe; a middorsal row of 9–12
saddles, reaching downwards to about ¼ of body depth,
sometimes regular, sometimes irregularly shaped and of
unequal density, wider than interspaces; an arched black
band at base of principal caudal-n rays (no ocellus or black
spot at caudal-n base); caudal n with pigments on rays
and membranes forming 2–3 very irregular bars, posterior
Fig. 1. Mouth parts in Lepidocephalichthys berdmorei (CMK
25927, 71.1 mm SL), as example for the genus; ventral view.
Abbreviations: mb, maxillary barbel; ml, mental lobe of lower
lip; pf, posterior ange of lower lip; rbi, inner rostral barbel; rbo,
outer rostral barbel; ul, upper lip.
Fig. 2. Lepidocephalichthys eleios; MHNG 2768.058, holotype, 20.6 mm SL; Myanmar: Nam Phaung Sin Chaung, tributary of Lake
Indawgyi.
Lepidocephalichthys eleios, new species
(Figs. 2, 3)
Holotype. MHNG 2768.058, 20.6 mm SL; Myanmar: Kachin
State: Nam Phaung Sin Chaung (stream) near Nam Phaung
Sin village, about 1.5 km before entering Lake Indawgyi north
of Lonton; 175 masl; 25°07′23″N 96°17′15″E; M. Kottelat,
Nyein Chan & Dankhaung H-Hkai, 7 December 2014.
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RAFFLES BULLETIN OF ZOOLOGY 2017
Table 1. Morphometric data of holotype (H) and 6 paratypes of Lepidocephalichthys eleios. Range and mean include holotype data.
H Range Mean
Standard length [mm] 20.6 18.1–22.8
Total length [mm] 24.9 21.1–27.0
In percent of standard length
Total length 120.6 116.7–123.5
Head length 23.9 22.3–24.9 23.8
Pre-dorsal length 52.9 52.9–56.7 54.9
Pre-pelvic length 50.2 46.9–51.2 49.5
Pre-anal length 72.5 71.2–72.5 72.0
Body depth at dorsal-n origin 14.1 14.0–15.1 14.4
Depth of caudal peduncle 9.4 9.4–10.3 9.9
Length of caudal peduncle 19.4 17.9–23.1 19.7
Snout length 8.8 7.5–8.8 8.3
Eye diameter 3.9 3.9–4.4 4.1
Interorbital width 3.8 3.3–4.6 3.8
Length of pelvic n 13.9 13.6–15.8 14.5
Length of pectoral n 13.2 11.0–17.8 14.5
Length of caudal n 21.0 19.1–23.3 21.0
In percent of head length
Eye diameter 16 16–19 17.5
Snout length 37 32–37 35.1
Interorbital width 16 14–19 16.2
one along distal margin, bars irregularly in contact or fused,
leaving variously shaped unpigmented patches; postlabial
ange very long and narrow (Fig. 4); head scaleless; caudal
n truncate to slightly emarginate.
Description. See Fig. 2 for general appearance and Table 1 for
morphometric data of holotype and 6 paratypes. Specimens
are very small, soft and difcult to handle; measurements
and counts were limited to these specimens but characters
were checked on all paratypes. A moderately elongate
cobitid with body depth increasing posteriorly from tip of
snout up to about half of predorsal distance; behind dorsal
n, body depth decreasing slowly to end of anal-n base,
then uniform until caudal-n base. Dorsal prole continuous
between head and body. Head and body very compressed.
Interorbital area arched. Eye below dorsal prole of head.
Snout pointed. Suborbital spine bid, posterior (inner) prong
strongly curved medially (about a quarter of a circle), length
equal to about half of eye diameter, anterior (outer) prong
shorter, slightly curved. Caudal peduncle depth 1.7–2.3
times in its length, of uniform depth. Largest recorded size
22.8 mm SL.
Dorsal n with 4 unbranched and 6½ branched rays; distal
margin convex; last 2 unbranched rays with space between
them. Second branched ray longest. Pectoral fin with 1
unbranched and 7 branched rays; reaching about halfway
to pelvic-n base; last two branched rays (6 and 7) not
fused to form a pectoral rod but adjacent, with a narrow or
without membrane between them, not thickened, without
dorsal or ventral projection (in both sexes). Pelvic n with
1 unbranched and 6 branched rays; reaching about halfway
to anal-n origin; falcate; posterior margin straight; origin
slightly in front of vertical through dorsal-n origin. Anus
immediately in front of anal-n origin. Anal n with 3
unbranched and 5½ branched rays; distal margin slightly
convex. Caudal n with 7+7 branched rays; truncate to very
slightly emarginate.
Body entirely covered by deeply embedded scales. No
scales on head. No lateral line. Anterior naris at tip of an
(obliquely) truncate tube; posterior naris as a long slit,
adjacent to anterior one. Mouth U-shaped, gape about
twice as wide as long (Fig. 4). Upper lip with smooth edge.
Lower lip with long mental lobe, with one or two short,
thick projections along medial edge (close to base); rest of
mental lobe slender and thin (hardly thicker than postlabial
ange). Postlabial ange starting from tip of mental lobe,
sometimes extending backwards further than tip of lobe) and
with deeply concave posterior edge. Shape of ange variable.
Lobe and ange sometimes together with appearance of at
‘barbel’, especially in dehydrated specimens. Inner rostral
barbel reaching beyond corner of mouth; outer one reaching
beyond posterior margin of eye. Maxillary barbel reaching
beyond posterior margin of eye.
Sexual dimorphism. There is no external sexual dimorphism
except that in some of the largest specimens the pectoral
n is more rigid and expanded laterally (Fig. 2), and in
these specimens the last two branched pectoral-n rays are
adjacent, without membrane between them. These specimens
are probably males: a rigid and laterally expanded pectoral n
is observed in most species of the loach families Cobitidae
and Nemacheilidae and is a character of males. In the
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Kottelat: Lepidocephalicthys eleios
Fig. 3. Lepidocephalichthys eleios, paratypes; Myanmar: Lake Indawgyi basin. a, CMK 21460, 19.8 mm SL; b, CMK 21460, 22.6 mm
SL; c, CMK 24171, 19.2 mm SL; d, CMK 25643, 19.5 mm SL; e, CMK 25643, 22.8 mm SL.
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RAFFLES BULLETIN OF ZOOLOGY 2017
part of n) and on rays, except at joints between segments.
Anal n hyaline, with dark brown pigments along most of
unbranched rays 3–4, most of unbranched part of branched
rays; black marks on membranes near branching points
(pattern mostly visible only with magnication). A dark
spot at base of anterior rays. Pelvic n hyaline, with a few
isolated dots on rays and membranes. Pectoral n hyaline.
In life: body slightly purplish brown, slightly translucent.
Notes on biology. Lepidocephalichthys eleios was observed
at 3 of 12 sites sampled in Lake Indawgyi basin: one in
the outlet of the lake, two in a tributary. These three sites
are characterised by deep water, very slow current, dense
shore vegetation (Figs. 5, 6), large extents of oating mats
of plants (Fig. 7), and bottom made of mud or clay covered
with a layer of vegetal debris. The specimens were obtained
from the leaf litter, from among the dense vegetation or
by scraping with a tray net the underside of the ‘oating’
vegetation. Other species collected in the same micro-habitat
were: Oreichthys sp., Pethia erythromycter, Heteropneustes
other specimens, the ns are softer, and there is a narrow
membrane between the last two rays. The specimens with
a narrow membrane have a somewhat stouter appearance
(not quantied), as observed in females of other species of
Lepidocephalichthys. See further comments under Notes
on biology.
Coloration. After one month in formalin. Head and body
background colour pale yellowish grey; except otherwise
stated, markings darker grey to blackish. Head with stripe
between eye and base of outer rostral barbel, and diffuse
blotches on nape and dorsal surface of snout. Body with
9–12 middorsal saddles (3–5 predorsal, 2–3 subdorsal, 3–6
postdorsal; modally 4+2+5) reaching downwards to about
¼ of body depth, sometimes regular (Figs. 2, 3a, d, e),
sometimes irregularly shaped and of unequal density (Fig.
3b); wider than interspaces. An irregular midlateral row of
irregularly set small spots or irregular blotches, sometimes
forming a stripe; midlateral row or stripe leaving an
unpigmented band between it and middorsal row of saddles.
Otherwise, very few pigments on ank in front of level of
anal n; pigments denser posteriorly, especially on lower
half of caudal peduncle. Three diffuse blotches along ventral
midline: rst one at anal-n origin (on body and n); second
one at base of last anal-n rays; third one at about midlength
of caudal peduncle. Axial streak present.
Black mark at caudal-n base: an arched band (tips pointing
backwards) at base of principal rays, not reaching dorsal and
ventral midlines. No ocellus.
Dorsal n hyaline, with dark brown pigment along most of
unbranched rays 3–4, most of unbranched part of branched
rays; black marks on membranes near branching points. A
dark spot at base of anterior rays. Caudal n hyaline, with
pigments on rays and membranes forming 2–3 irregular
bars, posterior one along distal margin; bars irregularly in
contact or fused, leaving variously shaped unpigment patches;
pigments present on membranes (especially in posterior
Fig. 4. Lepidocephalichthys eleios; CMK 24160, 21.0 mm SL (left) and 20.6 mm SL (right); mouth in ventral (left) and latero-ventral (right)
views. Abbreviations: ml, mental lobe of lower lip; pf, postlabial ange. Arrows indicate tip of mental lobe and tip of medial projection.
Fig. 5. Myanmar: Kachin State: Nam Phaung Sin Chaung,
tributary of Lake Indawgyi, north of Lonton; type locality of
Lepidocephalichthys eleios.
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Kottelat: Lepidocephalicthys eleios
fossilis, Chaudhuria sp., Mastacembelus pantherinus,
Monopterus cuchia, Indostomus paradoxus, Badis corycaeus,
Dario hysginon.
Fourteen of the largest and stoutest specimens guessed
to be females were dissected; either the abdominal cavity
was swollen but empty, or there were unripe ovaries. Two
individuals had what seemed to be one and three ova, about
0.1 mm diameter. All samples were obtained in December;
a distended belly and the absence of ova suggest that
spawning possibly already occured for some individuals.
No specimen smaller than 15 mm SL was observed (mesh
size was about 1 mm and they would have been seen).
Several hours of sampling by up to 5 persons, at different
depths of water and over and in different substrates yielded
hundreds of individuals (of which only a subsample was
preserved), but no specimen larger than 22.8 mm SL was
observed. Admittedly, this does not demonstrate that they
are mature, but there is also no evidence that sexual maturity
is not reached at this size or smaller. The modied (rigid
and curled) pectoral ns of males, the female with swollen
belly (possibly spent), and the absence of small individuals
suggest they are mature or close to.
Distribution. Lepidocephalichthys eleios is presently known
only from Lake Indawgyi basin, Kachin State, Myanmar, in
the Irrawaddy drainage. Because no sampling could be done
in similar habitats downstream of the lake in the Mogaung
watershed, it is premature to conclude that L. eleios is
endemic to the lake and its basin.
Etymology. From the classical Greek έλειος heleios, eleios,
meaning ‘of the marsh’, ‘dwelling in the marsh’. A noun
in apposition.
Remarks. The genus Lepidocephalichthys is distinguished
from all other genera of the family Cobitidae by having
the pectoral-n with 1 unbranched and 7 branched rays,
of which, in males, the last 2 rays are fused, forming a
pectoral rod thicker than the other rays, with a variously
developed projection, at about midlength of the dorsal side,
ranging from a small swelling (pectoral rod swelling) to an
elevated, thin, vertical ange (pectoral rod plate); in one
species there is also a vertical ange along the ventral side
of the ray (Nalbant, 1963; Kottelat & Lim, 1992; Havird &
Page, 2010; Deein et al., 2014). Other characters are: slender
body; absence of lateral line; dorsal n with 3–4 unbranched
and 6½ branched rays; pelvic n with 1 unbranched and 6
branched rays; anal n with 3 unbranched and 5½ branched
rays; caudal n with 14 branched rays; three pairs of barbels
(2 rostral and 1 maxillary); each half of lower lip (Fig. 1)
with a pointed mental lobe separated from postlabial ange
by a small notch, posterior margin of ange and medial edge
of lobe denticulated, crenulated or fringed; anterior naris
at tip of a short tube (not extended into a ‘nasal barbel’).
Lepidocephalichthy eleios is, however, missing the main
character diagnosing Lepidocephalichthys, the presence of
the pectoral rod in males. Sexual dimorphism provides very
efcient characters to identify lineages in Cobitidae (Nalbant,
1963, 1994; Šlechtová et al., 2008; Kottelat & Tan, 2008;
Kottelat, 2012: 138). In specimens of L. eleios identied as
males, the last two pectoral rays are not fused into a rod but
close to each other, without membrane between them, they
are not thickened, and there is neither a dorsal nor a ventral
ange. As is described above, all the other characters used to
diagnose the genus are present in L. eleios and show a similar
appearance as in the other species of Lepidocephalichthys.
The organisation of pectoral-n rays 7 and 8 in close contact
in male L. eleios is considered either as a plesiomorphy or
as precursor stage to the fusion into a pectoral rod, and is
possibly related to an incomplete development resulting from
the small size of the species (developmental trunction), as
observed in many other miniature shes (see, e.g., Britz &
Kottelat, 2003, 2008; Kottelat et al., 2006; Britz & Conway,
2009; Britz et al., 2014).
Lepidocephalichthys eleios cannot be a member of the
other genera of Cobitidae as it does not have the characters
distinguishing them from Lepidocephalichthys either. To
mention only the genera present on mainland Southeast Asia:
the lamina circularis in males of Cobitis and Misgurnus; the
serrated edge of the 2nd ray of the pectoral-n in males of
Kottelatlimia; the strongly elongated body and snout and
very ornamented lips with large conical and laciniated
projections along the edges of Acantopsis; the slender to
anguilliform body, the backward position of the dorsal n
and the curled pectoral n in males of Pangio; the deep and
uniformly dark brown body, small eye and poorly distinct or
missing mental lobe of Lepidocephalus; the four barbel-like
projections of the lower lip and the keeled and vertically
distorted caudal peduncle in males of Microcobitis; and the
ridges of papillae along the barbel and lips and the slender
translucent to whitish body of Acanthopsoides.
Fig. 6. Myanmar: Kachin State: marshes along Indaw Chaung,
outlet of Lake Indawgyi (a); mats of oating vegetation are moved
by farmers (b); locality and habitat of Lepidocephalichthys eleios
(CMK 24160).
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RAFFLES BULLETIN OF ZOOLOGY 2017
Kottelat & Lim (1992) described sexual dimorphism in the
Sundaic and some Indochinese species of Lepidocephalichthys.
Havird & Page (2010) reported sexual dimorphism in
most species of Lepidocephalichthys, but with a confused
terminology. The main problem is their misunderstanding of
the ‘lamina circularis’ present in some genera of Cobitidae.
They commented (p. 137) on the fused last two pectoral-
n rays: “[t]his modication is referred to as the lamina
circularis and is formed by a fusion and hardening of the
innermost (seventh and eighth) pectoral rays. In mature males
of other cobitid genera, the lamina circularis is formed by
different pectoral rays (usually the second) or is absent”.
To refer to just any sexually dimorphic modication of the
pectoral rays in Cobitidae as ‘lamina circularis’ is erroneous
as the structures compared are not homologous (a structure
on n rays 7 and 8 vs. a structure on ray 2). There is an
extensive literature on cobitids and their sexual dimorphism
by authors in Europe, China, Japan, and Korea, spanning
over about 140 years (e.g., Bacescu, 1962; Nalbant, 1963;
Šlechtová et al., 2008). Lamina circularis means circular
lamella. The sexual dimorphism in Cobitis was rst described
by Canestrini (1871) as a swelling at the base of the rst
branched pectoral-n ray; it was later recognised as a circular,
‘scale-shaped’ projection of the ray itself and became known
as Canestrini scale and later as lamina circularis, which is
more appropriate because the structure is not a scale. The
lamina circularis is a laminar, usually but not always circular,
posterior projection of the rst (proximal-most) segment of
the dorsal hemitrich of the rst branched pectoral-n ray
(see, e.g., Nalbant, 1963: 356; Kottelat & Freyhof, 2007: 301,
g. 61). A second lamina circularis is present on the second
branched ray in some species. Obviously, the lamina circularis
of males of Cobitis and Misgurnus is not homologous with
the fused pectoral-n rays 7–8 of male Lepidocephalichthys,
with the thickened pectoral-n rays 1–2 of male Theriodes,
with the curled pectoral ns with thickened pectoral-n
ray 2 of male Pangio, Lepidocephalus, Acantopsis and
Acanthopsoides, with the modied pectoral-n rays 5–8 of
male Neoeucirrhichthys; or with the blade-like formation or
‘saw’ on pectoral-n ray 2 of male Kottelatlimia (Kottelat
& Lim, 1992; Kottelat & Tan, 2008; Šlechtová et al., 2008;
Kottelat, 2012; Deein et al., 2014).
Two other species of Lepidocephalichthys have been observed
in Lake Indawgyi basin, L. berdmorei and L. alkaia. In
both, the last 2 pectoral-n rays are fused into a rod in adult
males. Lepidocephalichthys berdmorei is a large species (up
to 78 mm SL) found in small streams with moderate to fast
current, usually with pebble to stone bottom. It has a stout
body, yellowish brown colouration with numerous small black
spots, a midlateral row of irregular small black blotches, a
black blotch on base of branched caudal-n rays 3–7; the
caudal n has 3–6 thin black bars; the last two pectoral-n
rays are fused in a thick cylindrical rod. Lepidocephalichthys
alkaia reaches up to 61 mm SL and is found in more sluggish
streams. It has a stout body, a pale yellowish background
colour, a midlateral row of blotches more or less contiguous
to form a midlateral stripe, continued by black pigments on
median caudal-n rays and membranes inbetween, a black
blotch on base of branched rays 3–7 of caudal n base, and
the caudal n has 5–8 thin irregular vertical rows of black
spots; the last two pectoral-n rays are fused into a rod with
a low dorsal swelling, highest at about midlength.
Material used for comparison. Lepidocephalichthys alkaia:
CMK 24128, 50; CMK 25618, 33; Myanmar: Lake Indawgyi
basin. — CMK 14649, 7; Thailand: Salween drainage. L.
berdmorei: CMK 24152, 2; CMK 24190, 1; Myanmar: Lake
Indawgyi basin. — CMK 25430, 1; Myanmar: Tenasserim.
— CMK 14648, 1; Thailand: Salween drainage. — CMK
25927, 8; CMK 25941, 3; Laos: Mekong drainage. L. cf.
goalparensis: CMK 26462, 1; CMK 26518, 1; Myanmar:
Irrawaddy drainage: Putao. L. cf. kranos: CMK 12646,
4; Laos: Mekong drainage. L. furcatus: CMK 16516,
5; Thailand: Tapi drainage. L. guntea: CMK 5429, 1;
India: Karnataka. L. hasselti: CMK 24894, 5; Myanmar:
Tenasserim. — CMK 13249, 47; Laos: Mekong drainage.
CMK 7316, 1; Indonesia: Sumatra: Siak drainage. — CMK
20580, 1; Indonesia: Sumatra: Musi drainage. CMK 8263,
21; Malaysia: Kelantan. — CMK 5124, 24; Thailand: Sai
Buri. L. jonklaasi: CMK 12190, 29; Sri Lanka. L. lorentzi:
CMK 6967, 2; Indonesia: Borneo: Kapuas drainage. L.
micropogon: CMK 26657, 2; Myanmar: Irrawaddy drainage:
Myitkyina. — CMK 25065, 1; CMK 25230, 1; Myanmar:
Tenasserim. L. thermalis: CMK 7059, 10; Sri Lanka. —
CMK 9340, 12; India: Kerala. L. tomaculum: CMK 8033, 6
paratypes; Malaysia: Selangor. — CMK 11273, 7; Indonesia:
Sumatra: Batang Hari. — CMK 11927, 4; Indonesia: Bintan.
L. zeppelini: CMK 7968, 2; CMK 23025, 2; Laos: Mekong
drainage. — CMK 10749, 5; Thailand: Mekong drainage.
ACKNOWLEDGEMENTS
Nature and Wildlife Conservation Division, Forest Department
gave permission to survey in Indawgyi Wildlife Sanctuary.
The material of the new species was obtained from surveys
conducted by Fauna & Flora International (FFI), with
nancial support from the Leona M. and Harry B. Helmsley
Charitable Trust and the Critical Ecosystem Partnership Fund.
For help in organising the work and logistics I thank Zau
Lunn and Frank Momberg (FFI). For their help in the eld
I thank Nyein Chan, Dankhaung H-Hkai (Fauna & Flora
International), Kyi Oo, Soe Myint, Nay Myo Tun (Indawgyi
Wildlife Sanctuary) and Khin Maung Thein (Department of
Fisheries). I thank two anonymous reviewers for reading and
commenting on the manuscript.
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