‘ Nemacheilus’ argyrogaster, a new species of loach from southern Laos (Teleostei: Nemacheilidae)
Author
Kottelat, Maurice
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.
text
Zootaxa
2021
2021-02-19
4933
2
277
288
journal article
8053
10.11646/zootaxa.4933.2.6
432bca9f-72a0-4728-aa8a-2092f7c6e2b6
1175-5326
4550320
7ACF38DE-FE95-4FED-9B9D-77096D0463E1
‘Nemacheilus’
argyrogaster
,
new species
(
Figs. 1–2
)
Holotype
.
MHNG 2785.032
, 63.0 mm SL;
Laos
:
Mekong
drainage:
Xe Kong watershed
:
Xe Kong Prov.
: confluence of
Xe Kong
and
Houai Peung
, downstream of
Ban Kaleum
,
15°42’54’’N
106°48’9”E
;
M. Kottelat
et al.,
28 May 2009
.
Paratypes
.
CMK
21663,
26
(
7 males
45.3–55.0 mm SL,
18 females
49.8–76.9 mm
SL, 1 fixed in 95% ethanol)
;
ZRC
61628,
6
,
45.6–61.8 mm
SL; same data as holotype
.—
CMK
21684,
3
,
52.8–66.5 mm
SL;
Xe Kong Prov.
:
Xe Kong
drainage:
Houai Peung
, about
100 m
upstream of confluence with Xe Kong, downstream of Ban Kaleum,
15°42’56’’N
106°48’16”E
;
M. Kottelat
et al.,
28 May 2009
.
Additional materal
(non-types). All from
Laos
: Mekong drainage: Xe Kong watershed.
CMK
15576,
7
, 37.0–
45.6 mm
SL;
Attapeu Prov.
:
Xe Kong
between
Attapeu
and downstream to
Ban Ouk
,
14°44’51’’N
106°43’59”E
;
M. Kottelat
et al.,
20 May 1999
.—
CMK
15616,
1
,
57.3 mm
SL;
Attapeu Prov.
: unnamed creek entering
Xe Kaman
from the north at
Xe Kaman
dam site,
14°57’40’’N
107°9’16”E
;
M. Kottelat
et al.,
21 May 1999
.—
CMK
15650,
6
,
40.6
–63.0 mm SL;
Attapeu Prov.
: rapids on
Xe Kaman
,
14°53’29’’N
107°7’20”E
;
M. Kottelat
et al.,
21 May 1999
.—
CMK
15672,
15
,
41.4–59.4 mm
SL;
Attapeu Prov.
:
Xe Kaman
at
Muang Xaisettha
,
14°48’27’’N
106°55’52”E
;
M. Kottelat
et al.,
22 May 1999
.—
CMK
15687,
3
,
48.5–58.1 mm
SL;
Attapeu Prov.
:
Xe Pian
at
Ban Mai
,
14°42’22’’N
106°29’46”E
;
M. Kottelat
et al.,
22 May 1999
.—
CMK
15723,
6
, 47.0–
60.4 mm
SL;
Xe Kong Prov.
:
Xe Kong
at
Keng Luang
,
15°26’24’’N
106°43’39”E
;
M. Kottelat
et al.,
24 May 1999
.—
CMK
18996,
2
, fixed in 95 % ethanol; same data as CMK 15723
.—
CMK
15775,
2
,
36.5–42.1 mm
SL;
Xe Kong Prov.
:
Xe Namnoy
, rapids about
1 km
upriver of
Tad Feak
waterfall;
135 masl
,
15°14’9’’N
106°44’55”E
;
M. Kottelat
et al.,
25 May 1999
.—
CMK
15819,
6
,
44.9–61.7 mm
SL;
Xe Kong Prov.
:
Houay Pao
, a west side tributary of
Xe Kong
, entering it about
16 km
upstream of
Muang Kaleum
,
15°50’17’’N
106°45’40”E
;
M. Kottelat
et al.,
26 May 1999
.—
CMK
19006,
1
, fixed in 95 % ethanol; same data as CMK 15819
.—
CMK
21455,
1
,
41.2 mm
SL;
Attapeu Prov.
:
Xe Kong
, riffles in gravel banks at
Ban Khanmaknao
,
14°36’46’’N
106°33’13”E
;
M. Kottelat
et al.,
22 May 2009
.—
CMK
21521,
31
,
39.5–58.5 mm
SL; 3, fixed in 95 % ethanol;
Xe Kong Prov.
:
Xe Nam Noy
immediately below
Taad Fek
waterfall, downstream of
Ban Dan
;
115 masl
,
15°14’42’’N
106°45’7”E
;
M. Kottelat
et al.,
25 May 2009
.—
CMK
21570,
1
, 48.0 mm SL;
Xe Kong Prov.
:
Xe Kong
at
Keng Chang
rapids, near
Ban Song Khone
,
15°30’17’’N
106°46’2”E
;
M. Kottelat
et al.,
26 May 2009
.
Diagnosis.
‘Nemacheilus’
argyrogaster
is distinguished from most other species of
Nemacheilidae
in Southeast Asia by having an obvious bold black midlateral stripe separating the yellowish brown dorsal part of the body from the silvery whitish ventral part. A somewhat similar pattern is present only in
N. binotatus
,
N. longistriatus
,
Schistura nandingensis
,
S. rubrimaculata
and
S. pawensis
, from which it is distinguished by having male with a globulous suborbital flap with tubercles along its free, posterior edge; male with pectoral fin with thickened anterior rays, and branched rays 1–4 and the unculiferous pads behind them covered with small tubercles; lips thin, lower lip with narrow median notch; and 9+8 branched caudal-fin rays.
Description.
See
Figures 1–2
for general appearance and
Table 1
for morphometric data of
holotype
and
11 paratypes
. A moderately elongate nemacheilid with body depth gradually increasing up to slightly in front of dorsalfin origin. Behind dorsal fin, body depth decreasing gradually to end of anal-fin base, then almost uniform until caudal-fin base. Dorsal profile with slight concavity between head and body. Head slightly depressed; body slightly compressed anteriorly, to compressed posteriorly. Interorbital area convex. In lateral view, eye flush with dorsal profile of head. Cheeks not swollen. Snout rounded. Depth of caudal peduncle 1.3–1.6 times in its length, depth almost uniform. Very low dorsal and ventral keels on posterior half of caudal peduncle. Dorsal keel continuous with upper margin of caudal fin. Largest recorded size
76.9 mm
SL for female, 55.0 mm SL for male.
TABLE 1.
Morphometric data of type material of
‘Nemacheilus’
argyrogaster
(n=12). Range and mean include holotype data.
holotype |
range |
mean |
Standard lenth (mm) |
63.0 |
47.4–63.0 |
Total lenth (mm) |
76.6 |
57.9–76.6 |
In percent of standard length |
Total length |
121.5 |
120.2–123.3 |
122.3 |
Head length (dorsal) |
22.5 |
20.8–23.8 |
22.1 |
Head length (lateral) |
24.3 |
23.3–25.5 |
24.6 |
Predorsal length |
50.1 |
50.1–53.5 |
51.7 |
Prepelvic length |
50.5 |
50.2–52.9 |
51.6 |
Pre-anus length |
65.9 |
65.9–69.5 |
67.1 |
Pre-anal length |
75.3 |
75.3–79.9 |
78.2 |
Head depth |
13.4 |
12.2–13.5 |
12.9 |
Body depth at dorsal-fin origin |
18.2 |
16.2–20.3 |
18.2 |
Depth of caudal peduncle |
10.9 |
10.6–11.6 |
11.1 |
Length of caudal peduncle |
16.7 |
14.5–16.7 |
15.4 |
Head width |
14.2 |
13.3–16.0 |
14.8 |
Body width at dorsal-fin origin |
14.5 |
13.2–16.5 |
14.8 |
Snout length |
9.9 |
9.9–11.2 |
10.4 |
Eye diameter |
5.0 |
4.5–5.7 |
5.2 |
Interorbital width |
7.3 |
6.2–7.5 |
7.0 |
Length of dorsal fin |
17.3 |
16.9–19.6 |
18.2 |
Length of upper caudal-fin lobe |
21.1 |
20.6–23.6 |
22.0 |
Length of median caudal-fin rays |
13.7 |
12.7–14.5 |
13.7 |
Length of lower caudal-fin lobe |
23.0 |
21.3–24.3 |
22.8 |
Length of anal fin |
16.1 |
15.4–17.1 |
16.5 |
Length of pelvic fin |
15.9 |
15.6–18.1 |
16.6 |
Length of pectoral fin |
18.7 |
17.3–20.5 |
18.9 |
In percent of dorsal head length |
Snout length |
44 |
44–50 |
47 |
Eye diameter |
22 |
21–25 |
23 |
Interorbital width |
33 |
28–35 |
32 |
In percent of lateral head length |
Snout length |
41 |
39–45 |
42 |
Eye diameter |
20 |
18–23 |
21 |
Interorbital width |
30 |
25–31 |
28 |
Dorsal fin with 4 unbranched and 8½ branched rays; distal margin slightly concave. First branched ray longest. Pectoral fin with 1 unbranched and 10 (2), 11 (9) or 12* (1) branched rays (including small last ray, usually unbranched), rounded, reaching about halfway of distance to pelvic-fin base; rays without filamentous extensions. See below for sexual dimorphism. No axillary pectoral lobe. Pelvic fin with 1 unbranched and 7* (11) or 8 (1) branched rays (including small last ray, usually unbranched); reaching about to or slightly beyond anus, halfway to anal-fin origin; triangular, posterior margin rounded; origin below base of last unbranched dorsal-fin ray to base of 2nd branched ray; axillary lobe present, entirely free, conspicuous. Anus situated about 1.8–2.4 eye diameter in front of anal-fin origin, about at extremity of pelvic fin. Anal fin with 3 unbranched and 5½ branched rays; distal margin straight. Caudal fin with 9+8 branched rays, forked, tip of lobes rounded, subequal, length of upper lobe about 1.5–1.7 times length of median rays.
FIGURE 1.
‘Nemacheilus’
argyrogaster
, MHNG
2785.032, holotype, 63.0 mm SL; Laos: Mekong drainage: Xe Kong watershed.
Body entirely covered by scales. Scales embedded. Lateral line almost complete, with 88–93 pores, missing on last 3–8 scales. Cephalic lateral line system with 6 supraorbital, 4 + 10–11 infraorbital, 9 preoperculo-mandibular and 3 supratemporal pores.
Anterior naris pierced in front side of a pointed flap-like tube. Posterior naris adjacent to anterior one. Mouth U-shaped, gape about 1.5–2 times wider than long (
Fig. 3a
). Lips thin. Upper lip with median notch, with a few wrinkles (especially near corner of mouth). Lower lip continuous, with narrow median notch; median part with 3–7 shallow and narrow sulci, with a few wrinkles at angle of mouth. Processus dentiformis present. Tip of lower jaw not exposed. A shallow median concavity in lower jaw. Inner rostral barbel reaching beyond corner of mouth, almost vertical of anterior margin of eye; outer one reaching below anterior half of eye. Maxillary barbel reaching at most beyond posterior margin of eye. Intestine with a loop behind stomach (
Fig. 3b
). Air bladder without posterior chamber in abdominal cavity. Two halves of bony capsule of anterior chamber of air bladder connected by a short manubrium.
Sexual dimorphism.
Males with suborbital flap, swollen, globulous, with tubercles along free posterior edge (
Fig. 4a
). Pectoral fin of male with modified rays and covered by small tubercles on rays and interradial membranes (
Fig. 4b
). Unbranched and first branched rays thickened. Branched rays 1–4 extended posteriorly over dorsal surface of membrane by a thickened unculiferous pad (sensu
Conway
et al
. 2012
); rays and unculiferous pads covered with densely set tubercles. Tubercles gradually less numerous and pads gradually narrower on posterior rays, only a small row of smaller tubercles along posterior edge of ray 5 and behind. Branches of branched rays with secondary branching near tip, missing on anterior branch of first ray. Membranes between branched rays at most as wide as width of branch; secondary branches adjacent, without membranes. Smallest male with distinct modification of pectoral-fin rays
36.5 mm
SL. Pectoral fin of female without widened ray, membranes wider, secondary branches located more proximally than in male, with membranes between secondary branches. Ripe females deeper bodied.
FIGURE 2.
‘Nemacheilus’
argyrogaster
, Laos: Mekong drainage: Xe Kong watershed.
a,
CMK 15650, 48.0 mm SL;
b,
CMK 15672, 47.5 mm SL;
c,
CMK 15819, 62.1 mm.
Coloration.
After one month in formalin. Head and body background colour: dorsal half pale yellowish brown, ventral half whitish. A black midlateral stripe from tip of snout through eye to upper extremity of gill opening, then along lateral line to middle of caudal-fin base, ending in a more intense black mark slightly below middle of base of fin, on base of lower rays of upper lobe and three upper rays of lower lobe. A little blackish spot at base of unbranched ray of lower caudal-fin lobe. Faint grey margin on end of caudal peduncle. Narrow black saddles on back, usually regularly shaped; in some specimens irregularly shaped and set or even dissociated into pairs of spots; total 14–19 saddles,
5–6 in
front of dorsal-fin origin, 4–6 along dorsal-fin base and 4–7 behind dorsal fin, last one as a short stripe extending posteriorly to base of unbranched rays of upper lobe of caudal-fin.
All fins hyaline, with a few black pigments on membranes between branches near branching points and on rays (on edges and between segments) near primary and secondary branching points. Dorsal fin with a small patch of blackish pigments at base of each ray. In pectoral fin, pigments denser along posterior edge of unbranched ray and branched rays 1–4, appearing as thin stripes.
In life: body pale yellowish brown above black midlateral stripe, contrasting with silvery white below midlateral stripe.
FIGURE 3.
Nemacheius
argyrogaster
, CMK
21663;
a,
60.5 mm SL; mouth;
b,
52.5 mm SL; digestive tract.
FIGURE 4.
Nemacheius
argyrogaster
, CMK
21663, 49.4 mm SL, male sexual dimorphism;
a,
suborbital flap;
b,
left pectoral fin, dorsal view.
FIGURE 5.
Laos: Xe Kong province.
a,
confluence of Xe Kong river and Houai Pheung stream, type locality of
‘Nemacheilus’
argyrogaster
, CMK
21663;
b,
Houai Pheung stream about 100 m upstream of confluence with Xe Kong, locality of CMK 21684. 26 fish species were collected with
N. argyrogaster
at both sites.
Notes on biology.
A dissected female (CMK 21521,
57.2 mm
SL) had ripe ovaries with eggs
0.8–0.9 mm
diameter.
‘Nemacheilus’
argyrogaster
was collected in stretches of streams with moderate to fast current, with pebble to stone bottom (
Fig. 5
). Females reach a larger size than males, up to
76.9 mm
SL. The largest examined male is 55.0 mm SL and the smallest with developped suborbital flap and modified pectoral-fin rays is
36.5 mm
SL. In the two large samples examined (MHNG 2785.032, CMK 21663, ZRC 61628, n=32; CMK 21521, n=34), the adult male: female sex ratio is about 2:5.
At 5 of 13 sampling sites,
‘N.’
argyrogaster
was collected in syntopy with the botiid
Ambastaia nigrolineata
(Kottelat & Chu)
(
Fig. 6
), whose juveniles have a colour pattern partly similar to that of
‘N.’
argyrogaster
, composed of black midlateral and middorsal stripes.
FIGURE 6.
Ambastaia nigrolineata
, CMK
27801, 34.0 mm SL, juvenile; Laos: Mekong drainage: Nam Gnouang.
Distribution.
‘Nemacheilus’
argyrogaster
is presently known only from the Xe Kong watershed in the Mekong drainage in
Laos
. The species is expected to be present also in
Cambodia
, at least in Srepok and Sesan watersheds.
Etymology.
From the Greek
ἀργύρΕΙΟΣ
(argyreios, silver) and
γαστήρ
(gaster, belly), reference to the striking silvery white belly in life. A noun in apposition.
Remarks.
As mentioned in the diagnosis,
‘N.’
argyrogaster
is distinguished from most other species of
Nemacheilidae
in Southeast Asia in having an obvious bold black midlateral stripe separating the yellowish brown dorsal part of the body from the silvery whitish ventral part. A somewhat similar colour pattern is observed in
N. binotatus
(
Fig. 7
) known from the Chao Phraya and Mae Khlong drainages. The two species are distinguished by the presence in males
‘N.’
argyrogaster
of modified pectoral fins with conspicuous tubercles on anterior rays and wide unculiferous pads (vs. minute tubercles, narrow unculiferous pads), the mid-dorsal series of saddles (vs. a black stripe from top of head to caudal-fin base), and the absence of small vertically elongated spots above and in contact with the midlateral stripe (vs. presence of one, rarely two or three, spots at about level of the tip of the pectoral fin). Data on
N. binotatus
are from
Kottelat (1990)
and re-examination of specimens cited therein.
‘Nemacheilus’
argyrogaster
had at first been misidentified as
N. longistriatus
(
Fig. 8
), a species known from the Mekong drainage upstream of the Khone falls, because of the presence of the midlateral stripe (e.g.,
Baird
et al
. 1999: 70
;
Rainboth
et al
. 2012
: pl. 33 fig. 685). The two species are distinguished by the presence in males
‘N.’
argyrogaster
of modified pectoral fins with conspicuous tubercles on and behind anterior branched rays (vs. pectoral fins not modified, tubercles minute and few, on a single row along posterior margin of branched rays 1–3), suborbital flap well developed (vs. rudimentary), the midlateral stripe very regular and present even in the smallest specimens (vs. small specimens [below about
35 mm
SL] with small bars on the flank, connected with the saddles on the back; the bars then become fainter, restricted to course of lateral line, and connected by a midlateral stripe; vestiges of the bars persist and the stripe is irregular). Further, the general appearance is more compact in
‘N.’
argyrogaster
(compare figs. 1–2 and 8) and it has a somewhat smaller eye (eye diameter 21–25 % of dorsal head length, vs. 24–29).
‘Nemacheilus’
argyrogaster
is recorded (at least up to now) only from the Mekong drainage downstream of Khone falls. Data on
N. longistriatus
are from
Kottelat (1990)
and additional examined material.
FIGURE 7.
Nemacheilus binotatus
, CMK
16044, 34.6 mm SL; Thailand: Chao Phraya drainage: Nan River.
FIGURE 8.
Nemacheilus longistriatus
, Mekong
drainage:
a,
CAS 62547, holotype, 53.2 mm SL; Thailand: Mekong at Chiang Khan;
b,
CMK 5402, paratype, 45.1 mm SL; Thailand: Mekong at Chiang Khan;
c,
CMK 23085, 37.5 mm SL; Laos: Xe Bangfai.
Nemacheilus nandingensis
Zhu & Wang
, described from the Salween drainage in
Yunnan
, also has a yellowish to whitish body with a conspicuous black midlateral stripe and a middorsal series of 16–19 squarish saddles (
Zhu & Wang, 1985
). No topotypical specimen could be examined. From the original description, it has the anus closer to the anal-fin origin (about 1 eye diameter, vs.
1.8–2.4 in
‘N.’
argyrogaster
) and consequently the pelvic fin does not reach the anus (vs. reaches).
Nemacheilus nandingensis
was treated as a synonym of
Pteronemacheilus meridionalis
(Zhu)
by Yang (in
Chu & Chen 1990: 49
) and as a valid species of
Schistura
by
Zhu (1989: 54)
,
Kottelat (2012: 113)
and
Chen (2013: 296)
. A species apparently similar to
N. nandingensis
, or the same species, is present in the Mekong drainage in Xishuangbanna (
Yunnan
) (
Fig. 9
); the single available specimen, a male, has a small triangular suborbital flap and a pectoral fin with no tubercles (the first branched ray is thickened and its branches are not secondarily branched). The anus is positioned about 2 eye diameters in front of the anal-fin origin and the mouth has the characters of
Nemacheilus
(see below) and definitively neither the conspicuous median interruption in the lower lip nor the deep sulci in its median parts typical of
Schistura
s.l.
I consider it to be a species of
Nemacheilus
.
FIGURE 9.
Nemacheilus
aff.
nandingensis
, CMK
23949, 56.6 mm SL; China: Xishuangbanna: Mekong drainage: Nam Xing.
Schistura rubrimaculata
Bohlen & Šlechtová
(from the
Arakan
range in
Rakhine
,
Myanmar
) and
S. pawensis
Bohlen & Šlechtová
(from
Irrawaddy
drainage in
Myanmar
) too, have a colour pattern that includes a midlateral stripe and a pale belly, but the back is dark brown and the males lack modified pectoral-fin rays. Besides,
S. rubrimaculata
has up to 6 saddles on the back and a maximum known size of
28 mm
SL, and
S. pawensis
has 6–7 weakly marked saddles on the back, no suborbital flap, 7–8 + 7 branched caudal-fin rays (data from Bohlen & Šlechtová, 2013).
Specimens of a
Nemacheilus
superficially similar to
N. masyae
Smith
and
N. pallidus
Kottelat
, (
Fig. 10
) were also collected in the Xe Kong watershed and confused in the field with
‘N.’
argyrogaster
. The two are sympatric but not syntopic.
Nemacheilus
aff.
pallidus
was observed only in the Xe Pian (a tributary of Xe Kong). It too, has a midlateral stripe, thinner than in
‘N.’
argyrogaster
, including in juveniles, without indication that it would be derived from a row of coalescent blotches; there is a conspicuous black spot at the base of the caudal fin, a black spot on the anteriormost branched rays of the dorsal fin at about ¼ of their length, and rows of spots on the dorsal and caudal fins; the upper lobe of the caudal fin is distinctly longer than the lower one, the pectoral-fin rays have long projections, and there are numerous small tubercles on the middle and posterior part of the body. Males also have a suborbital flap, with a few minute tubercles, and there are a few minute tubercles along the anterior 2 branched pectoral-fin rays.
It is not clear to which species these specimens belong, and in fact they may represent a distinct, unnamed species. There is variability in both
N. masyae
and
N. pallidus
as described by
Kottelat (1990: 55
, 63), who identified all his material from the Mekong drainage as
N. pallidus
. Since then, numerous and better preserved specimens have become available. This material allows us to better understand variability within the two species. It appears that
N. pallidus
is restricted to the Chao Phraya drainage; the specimens from the Mekong drainage in
Kottelat (1990
[including fig. 36b from Nam Mun], 2001 [fig. 257 shows a fish from Nam Mang]) are not
N. pallidus
(in prep.).
FIGURE 10.
Nemacheilus
aff.
pallidus
, CMK
15687, 53.5 mm SL; Laos: Mekong drainage: Xe Kong watershed, Xe Pian.
Page
et al
. (2020)
treated
N. pallidus
as a synonym of
N. masyae
, based on their interpretation of the colour pattern and morphometric characters that had been considered diagnostic by
Kottelat (1990)
, supplemented by a molecular analysis. Because they did not see differences in morphometry and because their interpretation of elements of colour pattern placed the sequenced specimens of their
N. pallidus
into their
N. masyae
clade, making it non-monophyletic, they concluded that
N. pallidus
is a junior synonym of
N. masyae
. However, their morphometric and colour pattern data are not presented in a way that allows comparison by colour pattern or by geographic origin, or with the data presented in
Kottelat (1990)
. The morphometric data of all their
N. pallidus
and
N. masyae
are lumped in Table 2 or divided into two groups in a PCA, and there is no discussion of colour pattern variability, except for the statement “the putative differences [...] do not hold up when specimens are examined from across the range of the putative species (
Fig. 7
)” (p. 399). In fact, their figure 7 shows material only from peninsular
Thailand
, the Mae Khlong drainage and the upper Chao Phraya drainage, which is only a fraction of the range; it does not show specimens from the Mekong drainage and south-eastern
Thailand
. Further, the usefulness of the tree is limited because the geographical origin of some samples is not mentioned in the list of examined material.
For these reasons, I am unable to follow the conclusions of
Page
et al
. (2020)
. Their analysis may show that the characters listed to diagnose
N. pallidus
and
N. masyae
are insufficient (or not detailed enough), but it shows neither that the two are synonyms, nor that a single species is involved. There are differences in colour pattern between specimens of
N. pallidus
and
N. masyae
in the figures in
Kottelat (1990
: figs. 29a, 36) and, despite their statement, Page
et al
.’s (2020) figure 7d of a topotype of
N. pallidus
shows a fish very distinct from their
N. masyae
from Peninsular
Thailand
(their figures 7a–b) and Mae Khlong drainage (their figure 7c) (compare size, shape, position and numbers of bars and saddles, and body shape). As mentioned above,
N. pallidus
from the Mekong in
Kottelat (1990
: fig. 36b) is misidentified.
Taking into account the variability in
N. masyae
and
N. pallidus
as understood in
Kottelat (1990)
and mentioned above, and the non-monophyly retrieved in Page
et al
.’s tree, the hypothesis that more than two species are involved should have been investigated. Preliminary observations (unpublished) suggest that
N. pallidus
is restricted to the Chao Phraya drainage (widely spaced narrow saddles, midlateral row of spots, stout caudal peduncle), that
N. masyae
is possibly restricted to the Malay Peninsula (bolder saddles and midlateral blotches, slenderer caudal peduncle), that the populations from the Mae Khlong are potentially distinct from both
N. pallidus
and
N. masyae
and possibly close to populations from south-eastern
Thailand
(midlateral row of blotches fusing into a stripe with growth or age), and that 2 or 3 species are present in the Mekong drainage.
Diagnostic characters of
Nemacheilus
sensu
Kottelat (1990: 43)
are the forked to deeply forked caudal fin; the lips thin, usually smooth, the lower lip continuous along its anterior margin or with a slight median incision; the suborbital flap usually present; and, in most species, the pale background body colour.
Kottelat (1990: 43)
identified three groups of species in
Nemacheilus
and a few species that could not be placed in any group. While
‘N.’
argyrogaster
shares the characters diagnostic of
Nemacheilus
, it does not fit in any of these groups. It lacks the fimbriate posterior margin of the pectoral fin and the tubercles on scales of body seen in group A (e.g.,
N. masyae
,
N. pallidus
); it lacks the acuminate scales on the caudal peduncle and lower lip with deep folds and sulci of group B (e.g.,
N. selangoricus
,
N. spiniferus
); and it has pectoral-fin modifications unknown in group C (e.g.,
N. binotatus
). This suggests that
‘N.’
argyrogaster
might not be closely related to the other
Nemacheilus
, for which reason it is only tentatively placed in that genus.
Comparison material.
List not comprehensive.
Nemacheilus binotatus
: CMK 1768
, 6,
32.8–39.8 mm
SL;
Thailand
:
Chiang Mai Province
: Mae Nam Ping in Chiang Dao gorges.
N. longistriatus
: CMK
5402,
4 paratypes
, 36.0–
46.5 mm
SL;
Thailand
,
Loei Province
:
Mekong
mainstream between
Chiang Khan
to
70 km
downstream.—
CMK
13704,
22
,
30.6–38.8 mm
SL
;
Laos
:
Savannakhet Province
:
Xe Bang Hiang
, about
6 km
downstream of
Xepon.
—
CMK
13777,
7
,
32.4–37.3 mm
SL
;
Laos
:
Savannakhet Province
:
Xe Bang Hiang
at
Ban Tat Hai Xe.
—
CMK
15950,
1
,
67.2 mm
SL
;
Laos
:
Bolikhamsai Province
:
Nam Xao
, a small tributary of
Nam Ngiep.
—
CMK
19312,
7
,
29.8-36.5 mm
SL
;
Laos
:
Khammouan Province
:
Xe Bangfai
at
Keng Mou Man.
—
CMK
23085,
56
,
29.7–46.6 mm
SL
;
Laos
:
Savannakhet Province
:
Xe Bangfai
drainage:
Xe Noy
at
Keng Boua
.
N. nandingensis
:
CMK
23949, 1,
56.6 mm
SL;
China
:
Yunnan
:
Xishuangbanna
,
Nam Xing River
.
N.
cf.
pallidus
: CMK
21246, 21,
46.7–53.8 mm
SL;
Attapeu Prov.
:
Xe Pian
, about
2 km
upstream of Ban Mai.—
CMK
21280,
3
,
43.7–47.1 mm
SL
;
Attapeu Prov.
:
Xe Pian
, about
3 km
downstream of Ban Mai.—
CMK
21330,
14
,
40.1–54.8 mm
SL
;
Attapeu Prov.
: Houai Pin, a tributary of Xe Pian, entering it about
5 km
downstream of Ban Mai.
N. zonatus
:
CMK
19373, 13,
25.1–30.2 mm
SL;
Laos
:
Khammouan Province
:
Xe Bangfai
at
Ban Tha Ponsaoe
.