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Journal of Fish Biology (2011) 79, 1746–1759
doi:10.1111/j.1095-8649.2011.03108.x, available online at wileyonlinelibrary.com
A new species of gravel-dwelling loach (Ostariophysi:
Nemacheilidae) from the Nepalese Himalayan foothills
K. W. Conway*†, D. R. Edds‡, J. Shrestha§and R. L. Mayden
*Department of Wildlife and Fisheries Sciences and Texas Cooperative Wildlife Collection,
Texas A&M University, College Station, TX 77843, U.S.A., ‡Department of Biological
Sciences, Emporia State University, Emporia, KS 66801-5087, U.S.A., §Nepal Academy of
Science and Technology (NAST), Khumaltar, Lalitpur, Nepal and Department of Biology,
Saint Louis University, 3507 Laclede Ave, St Louis, MO 63103, U.S.A.
(Received 22 November 2010, Accepted 19 August 2011)
Turcinoemacheilus himalaya, new species, is described from the Koshi and Gandaki River basins
of Nepal. The new species is distinguished from its hypothesised congener, Turcinoemacheilus
kosswigi, from the Euphrates, Tigris and Karoun basins of the Middle East, by the presence of
small scales on the posterior half of its body (v. absence of all scales), its shorter caudal peduncle
(caudal peduncle length 12–15% standard length, LSv. 16 – 23), its shorter snout (snout length
28– 36% head length, LHv. 40 – 49) and by features of its colour pattern, including the presence
of small irregularly shaped dark grey markings over the lateral body surface. Turcinoemacheilus
himalaya is known to date only from Nepal. ©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles
Key words: Cobitoidea; Cypriniformes; Physoschistura;Schistura ; taxonomy; Turcinoemacheilus.
INTRODUCTION
Loaches of the family Nemacheilidae Regan are small, benthic freshwater fishes,
widely distributed throughout much of Europe, Asia and the Lake Tana basin of
Ethiopia (Kottelat, 1990). With well over 500 valid species (Eschmeyer, 2010), the
Nemacheilidae is one of the largest families of the Cypriniformes, and many new
species continue to be described annually, particularly from Asia (Kottelat, 2000;
Bohlen & ˆ
Slechtov´
a, 2009; Hadiaty & Kottelat, 2009, 2010; Tan & Kottelat, 2009;
Hu & Zhang, 2010) but also from the Far East and even from Europe (Coad &
Nalbant, 2005; Economidis, 2005; Stoumboudi et al., 2006; Erk’Akan et al., 2007).
Although nemacheilid generic level diversity is highest in the Far East (Prokofiev,
2009), the majority of nemacheilid species inhabit South and Southeast Asia, where
they are particularly abundant in well-oxygenated hill streams (Kottelat, 1990). To
date, surprisingly few nemacheilid loaches have been reported from Nepal. Shrestha
(1981) recorded five nominal species, including Acanthocobitis botia (Hamilton
1822), Nemacheilus corica (Hamilton 1822), Schistura beavani (G¨
unther 1868),
†Author to whom correspondence should be addressed. Tel.: +1 9798452620; email: kevin.conway@
tamu.edu
1746
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles
NEW NEMACHEILID LOACH FROM NEPAL 1747
Schistura rupecula McClelland 1838 and S. r. inglisi Hora 1935. Rajbanshi (1982)
increased the number to seven, with the addition of Schistura savona (Hamilton
1822) and Schistura scaturigina (McClelland 1839), and Edds (1985) added one
more, Schistura devdevi Hora 1935. Recently, Shrestha (2008) listed 12 species
of nemacheilids from Nepal, with the addition of Schistura himachalensis (Menon
1987), Schistura horai (Menon 1952), Schistura multifasciatus (Day 1878), Schistura
prashadi (Hora 1921) and Schistura sikamaiensis (Hora 1921) [Shrestha (2008) made
no mention of S. devdevi ]. A number of Shrestha’s (2008) new records require further
investigation before they should be considered part of the Nepalese ichthyofauna (e.g.
S. prashadi, a species known otherwise only from Nagaland, India; Menon, 1987).
During ongoing investigations of the ichthyofauna inhabiting the hill streams of
the Koshi and Gandaki River basins of Nepal, the authors became aware of a distinc-
tive new nemacheilid loach that appears to belong to Turcinoemacheilus B˘
an˘
arescu
& Nalbant 1964, a genus reported to date only from the Euphrates-Tigris drainage
of the Middle East (B˘
an˘
arescu & Nalbant, 1964; Coad, 1995; Breil & Bohlen, 2001;
Golzarianpour et al., 2009). The purpose of this paper is to provide a formal descrip-
tion for this new species.
MATERIALS AND METHODS
Specimens were captured in 1984, 1985, 1986, 1996 and 2008 by seine, electrofisher or
dip net, killed by an overdose of anaesthetic, fixed in 10% formalin solution and preserved
in 70% ethanol. Counts and measurements follow Kottelat (1990) except that the last two
branched rays of the dorsal and anal fins, which share a single pterygiophore, are counted
separately and that values for head length (LH) reported here are equivalent to lateral head
length of that author. All measurements were taken point to point on the left side of speci-
mens with digital calipers and recorded to 0·1 mm. The head of one specimen was dissected,
dehydrated with a graded series of ethanol, critical point dried (Denton DCP-1 critical point
drier; Denton Vacuum; www.denton vacuum.com), placed on an aluminum stub and coated
with gold (Denton Desk IV XLS; Denton Vacuum) for examination with scanning electron
microscopy (SEM) using a Philips XL-20 SEM (Philips Electronics N.V.; www.philips.com).
Five specimens were cleared and stained (c&s) following the protocol of Taylor & Van Dyke
(1985). Meristic counts were obtained from c&s specimens with the aid of a Leica S8AP0
stereomicroscope (www.leica-microsystems.com). Paired fin counts were obtained from the
left side of c&s specimens only. Numbers in parentheses after a count indicate the frequency
of that count. Meristic counts for Turcinoemacheilus kosswigi B˘
an˘
arescu & Nalbant 1964 were
obtained from B˘
an˘
arescu & Nalbant (1964), Breil & Bohlen (2001) and Golzarianpour et al.
(2009). All photographs were taken using a Leica DFC280 mounted on the aforementioned
microscope. Illustrations were adapted from photographs using Abode Illustrator CS.
A number of habitat measurements were obtained at localities sampled between 1984 and
1986. Dissolved oxygen (DO) (ppm), pH and total hardness (mg l−1CaCO3) were determined
with a Hach kit model AL-36B (Hach Co.; www.hach.com). Current speed (m s−1) was mea-
sured with a Gurley pygmy meter no. 625 (Gurley Precision Instruments; www.gurley.com)
at 60% depth at 10 locations across the microhabitat where the species was captured; depth
(cm) was determined with a metre stick. Substratum composition was qualitatively charac-
terized by visually estimating the relative proportion of mud (<0·06 mm diameter), sand
(0·06–2·00 mm), gravel (2–16 mm), pebble (16 – 64 mm), cobble (64 – 256 mm) and boulder
(>256 mm) (Bain, 1999). Vegetation cover was visually estimated as per cent of the substra-
tum covered by filamentous algae, submerged vegetation, emergent vegetation and organic
debris.
Specimens examined are housed at the American Museum of Natural History, New York
(AMNH), Fischsammlung J. Freyhof, Berlin (FSJF), University of Kansas Natural History
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
1748 K. W. CONWAY ET AL.
Museum and Biodiversity Research Center, Lawrence (KU), Oklahoma State University
Department of Zoology Collection of Vertebrates, Stillwater (OSUS), Texas Cooperative
Wildlife Collection, College Station (TCWC), Tribhuvan University Zoology Collection,
Kathmandu (TZC), University of Michigan Museum of Zoology, Michigan (UMMZ), and
the Zoological Institute and Museum, Hamburg (ZMH). Comparative materials examined are
listed in Appendix.
TAXONOMY
TURCINOEMACHEILUS HIMALAYA NEW SPECIES
(FIGS 1 AND 2)
Nemacheilus shebbearei Hora: Edds (1989: 24, 29, 36, 38, 41, 45, 73, 76, 80, 91,
93, 123, 125; 1993: 52, 55, 59), Edds et al. (2002: 546)
Physoschistura elongata Sen & Nalbant in Singh, Sen, Banarescu & Nalbant:
Shrestha (2008: 135, 331, Fig. 112)
Holotype: KU 40558, 46·6 mm standard length LS; Nepal: Bagmati Zone, Sind-
hupalchok District, Indrawati River at Melamchi Township, 27◦4942·5 N; 85◦34
37·1 E, R. L. Mayden, K. W. Conway, K. Khanal & R. Napit 28–29 October 2008.
Paratypes: KU 40279, 1, 33·2mm LS; Nepal: Palpa/Syangja, Kali Gandaki at
Ramdi, 27◦5360·00 N; 83◦3759·88 E, 26 April 1996, D. Edds. KU 40278,
1, 30·8mmLS; Nepal: Chitwan/Nawalparasi, Narayani River at Narayangarh, 27◦
Fig.1. Turcinoemacheilus himalaya, KU 40558, holotype, 46·6 mm standard length; Nepal: Bagmati Zone,
Indrawati River at Melamchi township.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
NEW NEMACHEILID LOACH FROM NEPAL 1749
(a)
(b)
Fig.2. (a)Turcinoemacheilus himalaya, KU 40557, paratype; Nepal: Bagmati Zone, Indrawati River at Melam-
chi township. (b) Turcinoemacheilus kosswigi.
4224·12 N; 84◦2554·13 E, 23 April 1996, D. Edds. KU 40280, 9 (2 c&s),
35·9–48·1mmLS; Nepal: Gulmi/Parbat, Gaundi River at Gumti, 28◦354·00 N;
83◦3335·99 E, 19 May 1996, D. Edds. KU 40281, 4, 20·8–39·4mmLS; Nepal:
Tanahun, Seti River at Khairenitar, 28◦159·88 N; 84◦40·13 E, 15 June 1996,
D. Edds. KU 40282, 1, 37·5mmLS; Nepal: Parbat, Lungdi River, purchased from
fishermen fishing 1 h walk north of Parse, c. 28◦100·12 N; 83◦480·00 E,
3 September 1996, D. Edds. KU 40557, 14 (2c&s), 27·3–46·6mmLS; same data
as holotype. OSUS 15908, 4, 24·6–46·3mmLS; Nepal: Gulmi, Kali Gandaki at
confluence with Baandigarh River, 19 August 1984, D. Edds. KU 40568, 11 (1c&s
1SEM), 18·1–52·6mmLS; Nepal: Bagmati Zone, Sindhupalchok District, Melam-
chi River 4·828 km (3 miles) upstream from confluence with Indrawati River, 27◦
5722 N; 85◦3226.6 E, R. L. Mayden et al., 29 October 2008. OSUS 15621, 4,
29·8–34·7mmLS; Nepal: Chitwan, Narayani River at Narayangarh, above irriga-
tion building, 1 April 1984, D. Edds. OSUS 16314, 26, 24·7–46·6mmLS; Nepal:
Gulmi, Kali Gandaki at confluence with Gaundi River near Phulbaari, 28◦349·00
N; 83◦3338·00 E, 20 January 1985, D. Edds. OSUS 16325, 7, 34·6–46·7mm
LS; Nepal: Baglung, Kali Gandaki at Jyaamri Ghat, 22 January 1985, D. Edds.
OSUS 16744, 25, 28·6–53·4mmLS; Nepal: Gulmi, Kali Gandaki at confluence
with Gaundi River, 28◦349·00 N; 83◦3338·00 E, 22 May 1985, D. Edds.
OSUS 17197, 32, 22·7–49·6mmLS; Nepal: Gulmi, Kali Gandaki at confluence
with Gaundi River, 28◦349·00 N; 83◦3338·00 E, 21 May 1985, D. Edds. TCZ
uncat., 3, 36·1–45·1mmLS; same data as holotype.
Diagnosis: Turcinoemacheilus himalaya is distinguished from its only congener,
T. kosswigi, by the presence of small scales on the posterior half of its body (v. scales
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
1750 K. W. CONWAY ET AL.
Table I. Morphometric characters of holotype (KU 40558) and 20 paratypes (KU 40280,
40557, 40568 and TCZ uncat.) of Turcinoemacheilus himalaya n. sp. expressed as a percentage
of standard length (LS)orheadlength(LH)
Holotype Range Mean ±s.d.
LS46·6 27·3–53·6
As percentage of LS
Body depth 11·49·2–11·910·8±0·9
Head length 19·3 18·3–20·719·7±1·0
Dorsal head length 18·2 16·6–19·818·5±1·0
Pre-dorsal length 54·5 53·2–57·055·3±1·2
Pre-pelvic length 50·4 47·8–52·550·3±1·4
Pre-anal length 75·5 75·5–80·178·4±1·6
Snout-anus 62·4 59·0–64·162·1±1·5
Caudal peduncle length 14·6 12·0–15·113·9±0·9
Caudal peduncle depth 7·96·6–8·57·8±0·6
Pectoral fin length 17·2 15·3–18·517·0±0·8
Pelvic fin length 14·8 13·7–16·814·9±0·9
As percentage of LH
Head width 48·9 40·5–48·845·5±2·7
Head depth 36·7 34·2–41·538·4±2·9
Eye diameter 13·3 12·1–18·314·5±1·8
Snout length 36·7 28·1–36·733·9±2·3
Interorbital width 21·1 14·3–22·219·0±2·4
completely absent), its shorter caudal peduncle (caudal peduncle length 12–15%LS
v. 16–23), its shorter snout (snout length 28– 36% LHv. 40 – 49) and the presence of
small irregularly shaped dark grey markings along the lateral side of body (v. absence
of such markings).
Description: general body shape as in Fig. 1. Largest specimen examined was
53·6mmLS(range 27·3–53·6 mm). Morphometric characters are listed in Table I.
Body elongate, circular in cross-section anterior to dorsal-fin origin, becoming
increasingly more ovoid in cross-section posterior to dorsal-fin origin. Body depth
greatest midway between occiput and dorsal-fin origin, gently decreasing in depth
towards caudal-fin base. Dorsal and ventral margins of caudal peduncle with low
fleshy ridge. Head and eye small, snout moderate, rounded in dorsal view. Eye
obliquely shaped, its ventral margin occluded in lateral view by a thick layer of skin.
Mouth subterminal, strongly arched, with two pairs of rostral barbels and one pair
of maxillary barbels (Fig. 3). Upper and lower lips fleshy, weakly papilliated. Lower
lip with a triangular-shaped median interruption. Upper and lower jaws covered
by a horny sheath. Processus dentiformis present on upper jaw, weakly developed.
Deep groove between rostral fold and upper lip. Rostral fold not developed into lobes
between barbels. Barbels weakly papilliated. Inner rostral barbel short, reaching to or
slightly past base of outer rostral barbel when extended, outer rostral barbel reaching
slightly past base of maxillary barbel when extended, maxillary barbel reaching to or
slightly past horizontal through centre of eye when extended. Anterior nostril small,
surrounded by a short flap. Posterior nostril larger than anterior, crescent shaped,
narrowly separated from anterodorsal margin of eye by a narrow strip of skin.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
NEW NEMACHEILID LOACH FROM NEPAL 1751
(a)
ORB UL
LL
P
MB
(b)
20 µm
1 mm
LJ
IRB
RC
Fig. 3. (a) Scanning electron micrograph picture of the mouthparts of Turcinoemacheilus himalaya, KU 40568,
paratype, 38·8 mm standard length; Nepal: Bagmati Zone, Melamchi River, upstream from confluence
with Indrawati River. (b) Close up of tastebud on surface of right outer rostral barbel, highlighted (a) ( ).
IRB, inner rostral barbel; LJ, lower jaw; LL, lower lip; MB, maxillary barbel; ORB, outer rostral barbel;
P, sensory pore of mandibular sensory canal; RC, rostral cap; UL, upper lip.
Dorsal fin with iii.7.i(4) or iv.7.i(1) rays. Anal fin with iii.5.i rays. Principal caudal
rays 9+8, dorsal procurrent rays 5(1), 7(3) or 8(1), ventral procurrent rays 3(1), 4(3)
or 5(1). Pelvic fin with i.5.i rays, pectoral fin with i.7.i rays. Paired fins horizontally
placed with large fleshy axillary skin-flap at base. Pelvic-fin origin ranging from
anterior to dorsal-fin origin to slightly posterior to dorsal-fin origin. Skin on ven-
tral surface of unbranched pectoral- and pelvic-fin rays greatly thickened. Dorsal fin
rounded, posterior edge straight. First dorsal-fin pterygiophore strongly bifurcated,
inserted between vertebral centra 16/17(5). Anal fin small, rounded, posterior edge
straight, posteriormost tip not reaching base of caudal fin when depressed. Anal-fin
origin situated closer to caudal-fin base than dorsal-fin origin. First anal-fin pterygio-
phore, strongly bifurcated, inserted between haemal spines of vertebral centra 28/29
(3) or 30/31(2). Caudal fin weakly forked, upper and lower lobes rounded, lower
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
1752 K. W. CONWAY ET AL.
lobe slightly longer than upper lobe. Anus positioned slightly posterior to pelvic
fins; the distance between anus and anal-fin origin much greater than the distance
between anus and pelvic-fin origin.
Total number of vertebrae 42-43, consisting of 21+21(1) 22+20(1), 22+21(2) or
23+20(1) abdominal and caudal vertebrae. Total number of pleural ribs 12(2), 13(2)
or 14(1), borne on vertebral centra 5-16(2), 5-17(2) or 5-18(1). Caudal fin skeleton
with six hypurals. Second (free) uroneural of caudal skeleton and supraneurals pos-
terior to the third absent. Anterior swimbladder chamber bilaterally paired, housed
within a boney capsule formed by lateral process of second vertebral centrum and
outer arm of os suspensorium. Boney capsule with two openings laterally and a large
continuous flange of membrane bone along posteroventral edge. Neurocranium with
large, irregularly shaped postepiphysial fontanelle, bordered by frontal, parietal and
supraorbital bones. Fifth ceratobranchial with a single row of eight to nine small
curved teeth with pointed tips (Fig. 4).
Cephalic sensory system well-developed (Fig. 4). Supraorbital, infraorbital, pre-
opercularmandibular and otic sensory canals enclosed in a series of short, tube-like
Pmax
(a) (b)
(c)
Max
lOl
Apal
Kii
iii
i
E
LE
Fr
Pa
Epo
Apto
Asph
Exoc
Soc
Fig. 4. Aspects of the anatomy of Turcinoemacheilus himalaya, paratype, KU 40557, 46·6 mm standard length;
Nepal: Bagmati Zone, Melamchi River, upstream from confluence with Indrawati River. (a) Skull in
dorsal view, anterior to top, cephalic lateral line canal illustrated on left side only. Free sensory canal
ossifications in dark grey, extraethmoid ossifications of right side in light grey. (b) Ceratobranchial 5, left
side in dorsal view. Cartilage grey. (c) Stomach and anterior portion of intestine in ventral view, anterior
to top. Scale bars equal 1 mm. Apal, autopalatine; Asph, autosphenotic; Apto, autopterotic; E, ethmoid
complex; Epo, epioccipital; Exoc, exoccipital; Fr, frontal; K, kinethmoid; Le, lateral ethmoid; Max,
maxilla; Pa, parietal; Pmax, premaxilla; Soc, supraoccipital; i, meseopreautopalatine; ii, 2nd preethmoid;
iii, preautopalatine.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
NEW NEMACHEILID LOACH FROM NEPAL 1753
ossifications. Sensory canal ossifications without contact to underlying dermal ossifi-
cations, excluding parietal portion of supratemporal canal. Lateral line canal incom-
plete, extending as a continuous canal along body side until point below insertion of
last dorsal-fin ray, continued thereafter as a series of short interrupted canals before
terminating completely on body side opposite or slightly posterior to anal-fin origin.
Scales small, cycloid, focus large, surrounded by a small number of circuli, and
four to five weakly developed radii along posterior margin. Scales deeply embedded
in skin and widely separate from each other, present on posterior half of body only.
Stomach large, sac-like, intestine straight, uncoiled (Fig. 4).
Sexual dimorphism: no obvious sexual dimorphism. Males appear more slender
bodied than females. Both sexes exhibit a row of small tubercles along fin membranes
between pectoral rays 1–4, which are better developed in males.
Colouration: In alcohol, body background colour light cream to white. Occiput
dark grey. Dorsal surface of body with a highly variable number of dark grey-brown
saddles (ranging from 5 to 9, often confluent) that extend ventrally to form dark
grey-brown bars, which are often forked ventrally, along lateral side of body. Dorsal
saddles and lateral bars more intensely developed in smaller individuals (c. 20 mm
LS). Posteriormost bar, situated at base of caudal fin, much darker than more anterior
bars in all individuals examined. Entire lateral side of body, excluding area directly
below dorsal procurrent rays and area above ventral procurrent rays, speckled with
small irregularly shaped dark grey markings. Such markings, which are generally
more highly developed in larger specimens (>30 mm LS), are heavily concentrated
along horizontal septum along posteriormost part of caudal peduncle, producing (in
combination with posteriormost body bar) a horizontal T-shaped marking at base of
caudal fin. Ventral surface of body without pigmentation.
Dorsal fin with a small dark spot anteriorly, at base of unbranched rays, and two
weak irregular strips across centre, formed by aggregations of small melanophores
on either side of branching point in branched rays. Caudal fin with two irregular
black vertical bars across centre, formed by aggregations of melanophores on either
side of branching point in branched principal caudal-fin rays. Pectoral fins with weak
scattering of small melanophores over surfaces of 4–5 anteriormost rays. Anal and
pelvic fins immaculate.
In life, background colour light grey-cream dorsally and white ventrally [Fig. 2(a)].
Dorsal saddles, lateral markings on body and markings on fins dark grey. Horizontal
T-shaped marking at base of caudal fin prominent dark grey, highlighted by intense
light areas without pigment dorsally and ventrally.
Distribution, habitat, abundance and associated species: specimens were collected
from large rivers (>10 m width) and medium-sized (5–10 m wide) tributaries of
the Koshi and Gandaki River systems of eastern and central Nepal (Fig. 5). At the
type locality in the Koshi basin (Fig. 6), the Indrawati River is a pristine, fast to tor-
rential flowing river, over a cobble and boulder substratum, with a well-developed
gravel shoreline. At this site, T. himalaya was collected close to shore, together
with juveniles of Psilorhynchus pseudecheneis Menon & Datta 1964 (Psilorhynchi-
dae) and an unidentified species of Schistura McClelland 1838 (Nemacheilidae),
by sifting a dip net or small seine through gravel. Other species collected at the
type locality, in adjacent habitats, included Botia almorhae Gray 1831 (Botiidae);
Barilius vagra (Hamilton 1822), Opsarius bendelisis (Hamilton 1807), Schizotho-
raichthys sp., Schizothorax richardsonii (Gray 1832), Tor putitora (Hamilton 1822)
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
1754 K. W. CONWAY ET AL.
80
30
28
26
30
28
26
82 84 86 88 90
80
100 km
82 84 86 88 90
Fig. 5. Distribution of Turcinoemacheilus himalaya (). Type locality represented with an open symbol ( ).
(Cyprinidae); Myersglanis blythii (Day 1870) and Pseudecheneis cf. crassicauda Ng
& Edds 2005 (Sisoridae).
In the Gandaki basin, specimens were collected (during each season of the year)
in well-oxygenated, shallow riffles and runs with moderate to swift flow over clean,
mainly pebble and cobble substratum (Table II). In 1984–1986, 140 specimens were
Fig. 6. Indrawati River at Melamchi township. Type locality of Turcinoemacheilus himalaya.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
NEW NEMACHEILID LOACH FROM NEPAL 1755
Table II. Habitat characteristics at sites of 16 collections of Turcinoemacheilus himalaya in
Nepal’s Gandaki River drainage in 1984–1986
Environmental factor Mean ±s.d. Range
Physicochemistry
Temperature (◦C) 18·2±1·2 12–25
Dissolved oxygen (mg l−1)11·1±0·5 8–15
pH 8·1±0·17·5–8·5
Total hardness (mg l−1) 146·0±5·4 103–171
Current velocity (m s−1)0·5±0·10·2–0·9
Water depth (cm) 27·6±4·2 13–66
Substratum composition
Silt (<0·06 mm) 9·7±3·3 0–40
Sand (0·06–2 mm) 17·9±4·7 0–53
Gravel (2–16 mm) 17·2±3·8 0–50
Pebble (16–64 mm) 22·1±5·0 0–80
Cobble (64–256 mm) 26·8±4·5 0–70
Boulder (>256 mm) 6·2±2·1 0–35
Vegetation cover (%)
Filamentous algae 11·5±3·1 0–30
Submerged <0·1±<0·1 0–5
Emergent <0·1±<0·1 0–5
Debris 1·2±0·9 0–15
captured in 16 of 156 collections at 11 of 81 sites sampled throughout the basin, with
one to 32 fish captured per collection (mode =1, median =4 individuals). Syntopic
species included Aspidoparia jaya (Hamilton 1822), Aspidoparia morar (Hamilton
1822), Bangana dero (Hamilton 1822), Barilius barila (Hamilton 1822), B. shacra
(Hamilton 1822), B. vagra,Chagunius chagunio (Hamilton 1822), Crossocheilus
latius (Hamilton 1822), Cyprinion semiplotum (McClelland 1839), Garra annan-
dalei Hora 1921, Garra gotyla (Gray 1830), Garra lamta (Hamilton 1822), Neolis-
sochilus dukai (Day 1878), Neolissochilus hexagonolepis (McClelland 1839), Opsar-
ius barna (Hamilton 1822), O. bendelisis,Puntius conchonius (Hamilton 1822),
Puntius sophore (Hamilton 1822), Raiamas bola (Hamilton 1822), Schizothorax
progastus (McClelland 1839), T. putitora,Tor tor (Hamilton 1822) (Cyprinidae);
N. corica,Schistura. cf. scaturigina,S. beavani,S. rupecula (Nemacheilidae), Botia
lohachata Chaudhuri 1912 (Botiidae); Psilorhynchus nepalensis Conway & May-
den 2008 (Psilorhynchidae); Amblyceps mangois (Hamilton 1822) (Amblycipiti-
dae); Glyptothorax kashmirensis Hora 1923, Glyptothorax pectinopterus (McClel-
land 1842), Glyptothorax trilineatus Blyth 1860, M. blythii,Pseudecheneis serracula
Ng & Edds 2005 (Sisoridae); and Mastacembelus armatus (Lac´
ep`
ede 1800) (Mas-
tacembelidae). Other species collected in different habitats at these sites included
Chela cachius (Hamilton 1822), Devario devario (Hamilton 1822), Labeo bata
(Hamilton 1822), Labeo boga (Hamilton 1822), Labeo pangusia (Hamilton 1822),
Puntius guganio (Hamilton 1822), Puntius ticto (Hamilton 1822), S. richardsonii
(Cyprinidae); Balitora brucei Gray 1830 (Balitoridae); A. botia (Nemacheilidae);
Mystus vittatus (Bloch 1794) (Bagridae); Glyptothorax cavia (Hamilton 1822), Glyp-
tothorax garhwali Tilak 1969, Glyptothorax indicus Talwar in Talwar & Jhingran
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
1756 K. W. CONWAY ET AL.
1991 (Sisoridae); Xenentodon cancila (Hamilton 1822) (Belonidae); Pseudambassis
baculis (Hamilton 1822) and Pseudambassis ranga (Hamilton 1822) (Ambassidae).
In 1996, 19 individuals were captured in six collections at 56 sites sampled, with one
to nine individuals per collection. Other species not previously collected at Gandaki
basin sites of T. himalaya occurrence included Anguilla bengalensis (Gray 1831)
(Anguillidae); Semiplotus modestus Day 1870 (Cyprinidae); Glyptothorax gracilis
(G¨
unther 1864), Glyptothorax telchitta (Hamilton 1822) and Pseudecheneis eddsi
Ng 2006 (Sisoridae).
Etymology: From the Sanskrit (Hima-Alaya), meaning ‘adobe of snow’, in
reference to the Himalayan Mountain Range to which the species is endemic.
DISCUSSION
B˘
an˘
arescu & Nalbant (1964) provided a very brief diagnosis for Turcinoemacheilus
in their review of the loaches of Turkey: ‘K¨
orper niedrig, gestreckt und dick,
vollst¨
andig schuppenlos. Die Seitenlinie sehr kurz. After¨
offnung weit nach vorn
gedr¨
angt, n¨
aher der Bauchflossenbasis als dem Ansatz der Afterflosse gelegen. Ansatz
der R¨
uckenflosse hinter dem Hinterrand der Bauchflossen. Schwanzflosse schwach
eingebuchtet; alle Flossen sind klein und mehr oder weniger abgerundet.’ [Body nar-
row, elongate and thick, entirely scaleless. The lateral line very short. Anus displaced
far anteriorly, closer to the pelvic fins than to the anterior base of anal fin. Anterior
base of dorsal fin behind posterior tip of pelvic fins. Caudal fin slightly emarginated;
all fins small and more or less rounded.]. Turcinoemacheilus himalaya exhibits all
these diagnostic characteristics except for the presence of scales along the posterior
half of the body, which serves to distinguish the new species from its only congener,
T. kosswigi. The two species of Turcinoemacheilus are further distinguished by a
number of morphometric characters, including caudal peduncle length (12–15% LS
in T. himalaya v. 16–23 in T. kosswigi ) and snout length (28–36% LHin T. himalaya
v. 40–49 in T. kosswigi ), and by differences in pigmentation (entire lateral side of
body speckled with small irregularly shaped dark grey markings in T. himalaya v.
absence of such markings in T. kosswigi ).
Previous authors have considered specimens of T. himalaya to belong to either
N. shebbearei (McClelland 1839) (Edds, 1989, 1993; Edds et al., 2002) or
P. elongata (Shrestha, 2008). Turcinoemacheilus himalaya is easily distinguished
from N. shebbearei (currently placed in the synonymy of S. scaturigina, following
Menon, 1987) and P. elongata by the position of its anus, which is situated close
behind the pelvic fins (v. situated closer to the anal fin than pelvic fins in both
N. shebbearei and P. elongata ; Hora, 1935; Singh et al., 1982). Turcinoemacheilus
himalaya is further distinguished from N. shebbearei by the position of the pelvic
fins in relation to the dorsal fin (pelvic fins inserted anterior or directly opposite
dorsal-fin origin in T. himalaya v. pelvic fins inserted posterior to dorsal-fin origin
in N. shebbearei ) and by the absence (v. presence) of a sexually dimorphic subor-
bital flap in males (Hora, 1935). It is further distinguished from P. elongata by the
absence (v. presence) of the posterior chamber of the swimbladder and the absence
(v. presence) of an intestinal coil (Singh et al., 1982). Turcinoemacheilus himalaya
is distinguished from all other members of the Nemacheilidae reported from Nepal
to date (Shrestha, 1981, 2008) by the anterior position of its anus.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759
NEW NEMACHEILID LOACH FROM NEPAL 1757
Turcinoemacheilus himalaya appears to be endemic to Nepal and is known, to
date, only from the upper reaches of the Koshi River basin and the hills and Inner
Tarai of the Gandaki River basin. Like its congener, it appears to be relatively rare,
found in only 22 of 212 collections in the Gandaki system in 1984–1986 and 1996,
much less common than most other loaches in Nepal, and is typically present in low
abundance at sites where it is found. This small benthic fish appears to be sensitive to
pollution, as it was absent in sampling from areas where it had previously occurred in
the Narayani River, Nepal, following dumping of paper mill effluent that decreased
dissolved oxygen and increased alkalinity, silt substratum and plant fibre detritus
(Edds et al., 2002). This raises concern for conservation of the species, given the
deterioration of Nepal’s riverine ecosystems (Edds, 2007).
The discovery of T. himalaya is noteworthy not only because it represents a new
generic record for Nepal but also because it represents the first record of Turci-
noemacheilus outside of the Tigris-Euphrates drainage. Given the specific habitat
requirements of Turcinoemacheilus (i.e. gravel beds with a strong current in the
upper reaches of rivers; Breil & Bohlen, 2001; Golzarianpour et al., 2009), it is
possible that additional populations of Turcinoemacheilus await discovery in remote
regions of Iran, Afghanistan, Pakistan and India.
The authors thank R. Thiel and I. Eidus (ZMH) for access to radiographs and images of
the type series of T. kosswigi, and A. Bentley (KU), J. Freyhof (FSJF), T. Echelle (OSUS),
D. Nelson (UMMZ) and B. Brown (AMNH) for access to museum specimens under their
care. The authors are particularly grateful to J. Bohlen and J. Freyhof for sharing information
and images of T. kosswigi, R. Britz for translating German text and examining specimens of
T. kosswigi, M. Kottelat, J. Bohlen and R. Britz for discussions about nemacheilid genera,
G. W. Lange (St Louis, MO) for photographing the holotype (Fig. 1), A. Conway for help
in collecting morphometrics and D. Siegel (Saint Louis University) and D. Sever (South-
eastern Louisiana University) for access to, and help with, SEM facilities (equipment time
on which was funded via NSF DEB-0809831 to D. Sever). The authors are particularly
grateful to J. Freyhof for valuable comments and suggestions on an earlier version of this
manuscript. Finally, the authors thank those who provided field assistance in Nepal, including
A. B. Gurung, K. Khanal and R. Napit, and the Nepalese Department of National Parks and
Wildlife Conservation for granting permits. This research was supported by funds from the
National Geographic Society Committee for Research and Exploration, Wildlife Conserva-
tion International, the Explorers Club, the Fulbright program, Oklahoma State University,
the University of Kansas Natural History Museum, the Emporia State University Faculty
Research and Creativity Committee (all to D.R.E.), the National Science Foundation (NSF
DEB-0808446 to R.L.M. and KWC and EF 0431326 to R.L.M.) and Texas Agrilife Research
(to K.W.C.). This is publication number 1418 of the Texas Cooperative Wildlife Collection.
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APPENDIX. COMPARATIVE MATERIAL EXAMINED
Acanthocobitis botia: KU 40367, 1, 40 mm standard length, LS; Nepal: Mechi,
Jhapa, Mechi River at Karkarvitta, at bridge crossing at border with India, 26◦38
45·50 N; 88◦944·30 E. KU 40408, 1, 40·5mmLS; Nepal: Mechi, Jhapa, Biring
River at highway, 26◦3830·60 N; 87◦5614·30 E. KU 40433, 1, 30·2mmLS;
Nepal: Kosi, Morang, Lohandra River at Belbari town, 26◦3946·40 N; 87◦24
29·30 E. KU 40459, 40, 37·7–72·5mmLS; Nepal: Kosi: Sunsari, Koshi River
diversion 1 km north of highway around 2 km before bridge crossing Koshi, 26◦
3614·50 N; 87◦126·80 E. KU 40489, 9, 40·8–52·7mm LS; Nepal: Kosi,
Sunsari: Sunsari River at Inarua, 26◦3638·60;87
◦738·80 E. KU 40507, 8,
32·4–63·1mmLS; Nepal: Bheri, Bardiya, main stem of Karnali River downstream
of Chisapani, 28◦3759·60 N; 81◦1630·40 E.
Barbatula barbatula: AMNH 78212, 2, 67·0–83·0mmLS; Sweden: Uppland,
Solna, stream by Ulriksdal. TCWC 14520.01, 74, 25.0-80.0 mm Ls; Poland: Lubrzanka
River, near Kielce.
Micronemacheilus taeniatus: AMNH 227982, 1, 83·0mmLS; Vietnam: Ha Tinh
Province, Huong Son District, Rao An Region, 18◦220·0 N; 105◦130·0 E.
Schistura devdevi : KU 40378, 13, 24·4–27·4mmLS; Nepal: Jhapa, Mechi River
at Karkarvitta, at bridge crossing at border with India, 26◦3845·50 N; 88◦9
44·30. UMMZ 244566, 12, 22·0–32·0mmLS; India: West Bengal, Relli River and
tributaries at Relli, Tista River basin, 27◦350·00 N; 88◦3139·00 E. UMMZ
244570, 1, 22·0mmLS; India: West Bengal, Pala River, tributary of Relli River,
after Relli town on Kalimpang-Relli road, Tista River basin, 27◦242·00 N; 88◦32
9·00 E. UMMZ 244573, 13, 26·0–46·0mmLS; India: West Bengal, Rishi Khola
River at Rishi (on West Bengal and Sikkim border), Tista River basin, 27◦956·00
N; 88◦387·00 E.
Schistura savona: KU 40379, 3, 20·0–26·5mmLS; Nepal: Jhapa, Mechi River
at Karkarvitta, at bridge crossing at border with India, 26◦3845·50 N; 88◦9
44·30 E. KU 40389, 15, 21·7–30·1mmLS; Nepal: Jhapa, Palia River (tributary of
the Mechi) at bridge crossing highway west of Karkarvitta, 26◦392·70 N; 88◦8
13·20 E.
Schistura sp.: AMNH 211268, 2, 57·1–65·9mmLS; Vietnam: Ha Giang Province,
Yen minh district, du gia stream.
Turcinoemacheilus kosswigi : ZMH 1884, 1, holotype (images and radiographs
examined only); Turkey, Tigris basin, Kapozik Kadun, Hakkari. ZMH 1885, 5,
paratypes (images and radiographs examined only); same data as holotype. FSJF
2454, 4 ex., not measured; Turkey: Adıyaman province, Upper River G¨
oksu, 5km
northeast of G¨
olbas¸ı, 37◦500·00 N, 37◦410·00 E.
©2011 The Authors
Journal of Fish Biology ©2011 The Fisheries Society of the British Isles, Journal of Fish Biology 2011, 79, 1746– 1759