A New Species of Homalopteroides (Teleostei: Balitoridae) from Sarawak,

Malaysian Borneo
Zachary S. Randall
1
and Lawrence M. Page
1
Homalopteroides avii, new species, is described from the Rajang River basin, Sarawak, Malaysian Borneo. It is
distinguished from all other species of Homalopteroides by a wider gape of 28.533.3% HL vs. 22.324.2% for H.
wassinkii, 20.028.4% for H. modestus, 19.1% for H. rupicola, 19.127.3% for H. smithi, 20.023.2% for H. stephensoni, 20.2
26.9% for H. weberi, 18.426.0% for H. tweediei, 17.3% for H. indochinensis, 17.421.2% for H. nebulosus, and 18.019.0%
for H. yuwonoi. It is further distinguished from species of Homalopteroides by the presence of a lateral cephalic stripe vs.
absence in H. modestus, H. rupicola, H. smithi, H. tweediei, H. nebulosus, and H. yuwonoi; deeper caudal peduncle, 10.1
10.9% SL, vs. 8.49.0% for H. wassinkii, 8.7% for H. rupicola, 7.79.1% for H. smithi, 6.16.6% for H. stephensoni, 6.78.0%
for H. weberi, 8.09.4% for H. tweediei, 9.39.8% for H. nebulosus, 8.7% for H. indochinensis, and 7.3% for H. yuwonoi;
circumpeduncular scale count of 2022 vs. 16 for H. rupicola, H. stephensoni, H. nebulosus, H. indochinensis, and H. yuwonoi,
1416 for H. tweedei, and 1618 for H. smithi and H. weberi; total pelvic-fin ray count of 9 vs. 10 for H. stephensoni, H.
weberi, and H. yuwonoi. The species is relatively large for the genus, reaching 52.9 mm SL.
H
OMALOPTEROIDES Fowler, 1905 was recently
resurrected by Randall and Page (2012) based on
the following combination of characters: dorsal-
fin origin posterior to pelvic-fin origin, #60 lateral-line
scales, #30 predorsal scales, and a mouth morphology
characterized by two thin and widely separated rostral
barbels, thin crescent-shaped upper lip, the absence of
structures such as a mental pad or lobes between the lateral
portions of the lower lip, and a chin that extends anterior
of the lateral portions of the lower lip. The ten valid species
of Homalopteroides are: H. wassinkii (Bleeker, 1853), H.
modestus (Vinciguerra, 1890), H. rupicola (Prashad and
Mukerji, 1929), H. smithi (Hora, 1932), H. stephensoni (Hora,
1932), H. weberi (Hora, 1932), H. tweediei (Herre, 1940), H.
indochinensis (Silas, 1953), H. nebulosus (Alfred, 1969), and
H. yuwonoi (Kottelat, 1998) (Randall and Page, 2012).
Homaloptera manipurensis Arunkumar, 1999 was recognized
as a species of Homalopteroides by Randall and Page (2012)
and as a questionable junior synonym of Homalopteroides
rupicola (Prashad and Mukerji, 1929) by Kottelat (2012). We
follow Kottelat (2012) due to the inaccessibility of speci-
mens and the lack of detail in the type description of H.
manipurensis.
Species of Homalopteroides are found in northeastern
India, Myanmar, Thailand, Laos, Cambodia, Vietnam,
Peninsular Malaysia, Sumatra, Java, and Borneo. Six species
have been reported from Borneo: H. wassinkii (Fowler,
1905; Weber and Beaufort, 1916; Hora, 1932; Parenti and
Lim, 2005), H. stephensoni (Hora, 1932; Roberts, 1989;
Kottelat and Widjanarti, 2005; Parenti and Lim, 2005; Tan,
2009), H. weberi (Hora, 1932; Inger and Chin, 1962), H.
tweediei (Roberts, 1989), H. nebulosus (Roberts, 1989;
Kottelat and Widjanarti, 2005; Parenti and Lim, 2005),
and H. yuwonoi (Kottelat, 1998; Kottelat and Widjanarti,
2005). Recent examination of balitorid loaches from
Borneo at the USNM revealed a new species of Homalopter-
oides described herein.
MATERIALS AND METHODS
Measurements and counts follow Hubbs and Lagler (2004)
and Kottelat (1984; see Randall and Page, 2012 for
measurements used from each source), or are self explan-
atory, except for the following: pre-pectoral length is from
the tip of the snout to the origin of the pectoral fin,
distance between the anus and anal fin is from the anal
opening to the base of the first anal-fin ray, snout to
nostril distance is from the tip of the snout to the anterior
part of the nostril, nostril to operculum distance is from
the most posterior margin of the nostril to the hindmost
part of the opercle, internostril width is the narrowest
distance between the nostrils, interorbital width is the
least distance between the edge of the orbits traversing the
orbital rim, interrostral width is the narrowest distance
between the rostral barbels, and inter-lower lip width is
the narrowest distance between the lateral portions of the
lower lip. The lateral cephalic stripe extends horizontally
and posteriorly from between the lateral-rostral and
maxillary barbels to the cheek. The terms origin and
insertion refer, respectively, to the anterior and posterior
points of fin bases. All fin-ray counts are given as follows:
simple rays in Roman numerals followed by branched rays
in Arabic numerals, where dorsal fin- and anal fin-ray
counts include the last ray split at the base represented by
K. The caudal-fin ray count is the total number of
branched rays. Small scales found just before the dorsal,
pectoral, and anal fins are counted as K. Lengths were
measured to the nearest 0.1 mm using digital calipers and
taken on the left side when possible. All measurements are
in millimeters (mm). Measurements of the head are
presented as proportions of head length (HL), and eye
length is presented as a proportion of interorbital width
(IO). Head length and measurements of the body are given
as proportions of standard length (SL). One hundred fifty-
five individuals were examined representing all species of
Homalopteroides.
Institutional abbreviations follow Sabaj Perez (2012),
the abbreviation for alcoholic specimens is alc, and a ?
represents lack of data or uncertain locality. Photographs
were taken of preserved specimens using a Visionary
Digital (Palmyra, Virginia) with Canon 40D and 5D
cameras at UF and edited using Photoshop CS3. When
coordinates were unavailable, they were estimated using
maps and Google earth. Maps were constructed using
1
Florida Museum of Natural History, University of Florida, Dickinson Hall, Gainesville, Florida 32611; E-mail: (ZSR) zrandall@flmnh.ufl.edu;
and (LMP) lpage1@ufl.edu. Send reprint requests to ZSR.
Submitted: 8 May 2013. Accepted: 15 September 2013. Associate Editor: D. Buth.
F 2014 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-13-055
Copeia 2014, No. 1, 160167
ArcMap Version 9.3.1 in ArcGIS 9
th
edition. Species in this
study are recognized using the phylogenetic species
concept; i.e., a species is the smallest monophyletic group
that is diagnosable by one or more unique character
states.
Homalopteroides avii, new species
Figures 1, 2; Table 2
Homaloptera wassinkii: Parenti and Lim, 2005:188 (see remarks).
Homaloptera sp.: Parenti and Lim, 2005:188 (see remarks).
Fig. 1. Dorsal, lateral, and ventral views of Homalopteroides avii, Malaysia, Sarawak, Rajang River basin. (A) Holotype, USNM 323875, 52.9 mm SL,
(B) paratype, USNM 323879, 37.4 mm SL. Scale bars represent 20 mm.
Randall and PageHomalopteroides avii 161
Holotype.USNM 323875, 1 alc, 52.9 mm SL, East Malaysia,
Sarawak, Batang Balui, trib., Long Tow, where it enters
Batang Balui, just downstream from logging camp, current
swift, pH 7.7, water temperature 26uC, 2.41uN, 113.76uE, L.
R. Parenti, A. Among, K. Luhat, A. Luhat, 6 August 1991.
Paratopotype.UF 185293, 1 alc, 50.5 mm SL. Same locality
and date as holotype.
Paratypes.USNM 323878, 1 alc, 39.5 mm SL, East Malaysia,
Sarawak, Batang Balui, trib., Jangan Aya, flowing into Batang
Besua, pH 7.3, water temperature 25uC, 2.41uN, 113.73uE, L.
R. Parenti, K. Luhat, A. Among, 2 August 1991; USNM
323879, 2 alc, 36.837.4 mm SL, East Malaysia, Sarawak,
Batang Balui, trib., Kemtu, pH 7.3, water temperature 24uC,
2.38uN, 113.75uE, L. R. Parenti, K. Luhat, A. Among, 3
August 1991.
Diagnosis.Member of Homalopteroides as defined by Ran-
dall and Page (2012). Homalopteroides avii is distinguished
from all other species of Homalopteroides (Table 1) by a wider
gape (Fig. 3) of 28.533.3% HL. It is further distinguished
by presence of a lateral cephalic stripe, a deeper caudal
peduncle, a circumpeduncular scale count of 2022, a total
pelvic-fin ray count of 9, a total pectoral-fin ray modal count
of 15, and a total lateral-line count of 4345.
Description.Dorsal, lateral, and ventral views of an adult H.
avii are shown in Figure 1. Measurements and meristics are
given in Table 2. Homalopteroides avii is a relatively large
species for the genus, reaching 52.9 mm SL. The body is
slightly deeper than wide, arched predorsally, tapers
posteriorly to the anal-fin insertion, and has a flattened
ventral surface. When viewed dorsally the head is conical
and covered with small tubercles. The eyes are ovoid,
positioned dorsolaterally midway between the snout tip
and edge of the opercle, and smaller in length than the
interorbital width. The nostrils are closer to the snout tip
than to the edge of the opercle. The origin of the dorsal fin
is posterior to the origin of the pelvic fin and closer to the
caudal-fin base than to the snout tip. The pectoral fin
reaches slightly past the pelvic-fin origin and usually is
slightly longer than the head. The pelvic fin does not reach
the anus and is not doubly branched (i.e., with 3 or 4 distal
points on one ray vs. 2). The anal fin does not reach the
caudal-fin base. An axillary pelvic lobe is absent. Margins of
the dorsal and anal fins are straight. The caudal fin is forked
with rounded lobes; the lower lobe is slightly longer than
the upper lobe.
The body is scaled; anterior to the anal-fin origin scales are
deeply embedded. Most scales, especially just posterior to
the supraoccipital and at the dorsal-fin origin, have one or
more small nipples (wart-like spinous projections) along the
free borders of the scales; up to five are present on one scale.
The total lateral-line pore count is 4345, predorsal scale
count is 2225, circumpeduncular scale count is 2022. Scale
counts above and below the lateral line are 6K8Kand 6K
7, respectively. The scale count below the lateral line to the
pelvic-fin origin is 6K7. Variations in fin counts are given
in Table 2. Dorsal fin rays iii, 7K; anal-fin rays iii, 45K;
pectoral-fin rays ivv, 710, iii; pelvic-fin rays ii, 7; total
caudal-fin ray count 1617.
The mouth (Fig. 2) is wide (28.533.3% HL) and inferior
with the lower jaw visible. The lips are thin, smooth, and
continuous around the corners of the mouth. The upper lip
is crescent-shaped, and the lower lip is broadly interrupted
medially. The inter-lower lip width is large (20.022.6%
HL), and the chin extends far in front of the lateral portions
of the lower lip. Rostral and postlabial grooves are present.
Two pairs of rostral barbels are present, and a pair of
maxillary barbels is at each corner of the mouth. The
medial-rostral barbel reaches just past or at the base of the
lateral-rostral barbels. The lateral-rostral barbel reaches a
little more than half the distance to the base of the
maxillary barbel. The maxillary barbel reaches horizontally
to a vertical at the anterior orbital rim or to mid-orbit. The
gill membranes are united to the isthmus with a large
central furrow where the gill membranes meet. The gill
opening extends from the level of mid-orbit to the ventral
surface of the body.
Coloration.In 70% ethanol: The general pattern is shown
in Figures 1 and 2. In dorsal view, the ground color is cream
and mottled brown. There are 56 brown dorsal saddles. The
1
st
saddle is between the supraoccipital and dorsal-fin origin,
is irregular, and may consist of up to five blotches; the 2
nd
saddle spans most of the pelvic-fin length; the 3
rd
saddle is
located between the dorsal-fin insertion and anal-fin origin;
the 4
th
saddle spans most of the anal-fin length; and the 5
th
saddle is at the caudal-fin base and in most specimens does
not reach the ventral surface. In two individuals, there is an
extra saddle between the 3
rd
and 4
th
saddles. A dark brown
preorbital bar extends from between the bases of the medial-
and lateral-rostral barbels and shifts laterally to meet the
eye. There is a brown mottled blotch between and anterior
Fig. 2. Mouth of Homalopteroides avii, holotype, USNM 323875,
52.9 mm SL.
162 Copeia 2014, No. 1
to the nares. This blotch and preorbital bar form a cream-
colored Y when viewed anteriorly. The eye is outlined in
dark brown, and dark brown blotches may be present
dorsomedially. A dark brown bar covers the post-epiphyseal
fontanelle. Postorbital bars extend to the supraoccipital and
to the posterior edge of the opercle in alignment with the
lateral line.
The lateral view shares the same ground color pattern as
the dorsal view. A dark brown lateral cephalic stripe extends
horizontally and posteriorly from between the lateral-rostral
and maxillary barbels to the posterior edge of the orbit
where it continues diagonally upward for a small distance.
In some individuals, the upward extension of the stripe is an
unconnected dark brown blotch. Over the lateral line is a
solid brown stripe that contains circular blotches in larger
individuals. Mottled brown blotches sometimes coalesce to
form distinct lines above and below the lateral line and may
form large blotches between the pelvic fin and caudal-fin
base in larger individuals.
The venter is cream with mottled dark brown blotches
restricted to a band at the origin of the anal fin. The bases of
the rostral barbels are dark brown. The upper lip and the
lateral aspect of the lower lip are mottled dark brown.
All fins are hyaline with fine brown blotches or bands. The
dorsal fin has two bands, the pectoral fin has one or two
bands, and the pelvic fin has one or two bands. The anal fin
has one brown blotch (restricted in one individual to the
second ray). The caudal fin has two brown bands, a proximal
band which may not always reach the superior extent of the
upper lobe, and a subdistal V-shaped band that is pointed
anteriorly. The lower lobe of the caudal fin is fully pigmented
up to the posterior edge of the subdistal V-shaped band. The
paired and caudal fins are brown at their bases. The dorsal fin
has a dark brown blotch at its origin.
Distribution and ecological notes.Homalopteroides avii was
collected from the Balui River and downstream of the
Rajang River from Kapit, Rajang River basin, Sarawak
(Fig. 4). Individuals were collected in streams with slow to
swift currents and bottoms of mud, leaf litter, gravel,
boulders, and rock outcrops with a pH of 7.37.7 and water
temperatures of 2426uC (Fig. 5).
Twenty-nine species of fishes were collected at the four sites
where H. avii was collected: Barbonymus collingwoodii, Cyclo-
cheilichthys apogon, Hampala bimaculata, H. macrolepidota,
Leptobarbus melanotaenia, Lobocheilos ovalis, Luciosoma seti-
gerum, Macrochirichthys macrochirus, Nematabramis steindach-
neri, Osteochilus enneaporos, O. kahajanensis, O. sarawakensis,
Paracrossochilus acerus, Rasbora argyrotaenia, R. dusonensis, R.
cf. sumatrana, Systomus banksi, Tor tambra, Nemacheilus
kapuasensis, Homaloptera orthogoniata, Homalopteroides nebu-
losus, Hemibagrus bongan, H. nemurus, Clarias planiceps,
Mastacembelus unicolor, Macrognathus circumcinctus, Glyp-
tothorax major, Doryichthys martensii, and Glossogobius celebius
(Parenti and Lim, 2005).
Remarks.Parenti and Lim (2005) identified H. avii as H.
wassinkii (USNM 323875, 323878, 323879) and H. sp.
(THH9806). Although THH9806 and THH9809, also identi-
fied as H. sp., are deposited at ZRC and currently unavailable
for examination, THH9806, collected from downstream of
the Rajang River from Kapit, Rajang River basin (1u56.259N,
112u52.269E) and shown in figure 10 of Parenti and Lim
(2005), is H. avii. The identity of THH9809, from the Kapit
Table 1. Characters distinguishing Homalopteroides avii from all other species of Homalopteroides. Numbers of individuals examined
in parentheses.
Gape
width
% HL
Lateral cephalic
stripe present
(Yes/No)
Caudal-
peduncle
depth % SL
Circumpeduncular
scale count
Total
pelvic-ray
count
Total pectoral-
ray count
(modal)
Total lateral-
line pore
count
H. avii
(n 5 5)
28.533.3 Y 10.110.9 2022 9 15 4345
H. wassinkii
(n 5 2)
22.324.2 ? 8.49.0 20 9 18 47
H. modestus
(n 5 69)
20.028.4 N 9.111.5 1820 9 15 3944
H. rupicola
(n 5 1)
19.1 N 8.7 16 89 16 4244
H. smithi
(n 5 21)
19.127.3 N 7.79.1 1618 9 17 3542
H. stephensoni
(n 5 5)
20.023.2 Y 6.16.6 16 10 16 4652
H. weberi
(n 5 42)
20.226.9 Y 6.78.0 1618 10 16 4449
H. tweediei
(n 5 4)
18.426.0 N 8.09.4 1416 9 13 3438
H. indochinensis
(n 5 1)
17.3 ? 8.7 16 9 17 44
H. nebulosus
(n 5 4)
17.421.2 N 9.39.8 16 9 14 3840
H. yuwonoi
(n 5 1)
18.019.0 N 7.3 16 10 16 41
Randall and PageHomalopteroides avii 163
area along the Rajang River (2u0.129N, 112u55.839E), cannot
presently be determined. Two other specimens identified by
Parenti and Lim (2005) as H. wassinkii, FMNH 97441 and
68140, were collected from the Mengiong and Putai rivers,
Rajang River basin, respectively. FMNH 68140 was examined
in this study and is identifiable as H. cf. nebulosus; FMNH
97441 was not examined.
Etymology.Named avii, in memory of Lawrence Avi
Greenberg (21 July 198226 November 2011). The diagnos-
tic lateral cephalic stripe of this species, reminiscent of a
smile, is a symbol of Avis gentle disposition and good-
hearted nature. He is an inspiration to and missed friend of
the first author and many others. Latinized as a noun in the
genitive singular.
Table 2. Morphometric measurements and meristic counts for Homalopteroides avii (n = 5). All measurements in mm. Numbers of individuals in
parentheses. Holotype is represented by a *.
Morphometrics Holotype Range (n = 4) Mean%SD (n = 4)
Standard length 52.9 36.850.5
% of standard length
Head length 25.8 25.928.0 26.860.94
Body depth 19.0 15.517.1 16.360.72
Body depth at anus 13.7 13.013.8 13.360.34
Body width 17.2 13.917.2 15.561.60
Predorsal length 53.9 52.954.3 53.860.62
Prepectoral length 20.4 20.321.3 20.860.46
Prepelvic length 46.3 45.146.1 45.760.52
Preanal length 79.0 77.079.0 78.260.85
Pre-anus length 73.5 71.773.4 72.360.75
Distance between anus and anal fin 5.1 5.57.1 6.060.75
Dorsal-fin base length 12.6 12.013.2 12.460.53
Dorsal-fin length 21.5 20.021.0 20.660.42
Pectoral-fin base length 9.8 10.811.8 11.160.48
Pectoral-fin length 28.3 26.928.0 27.460.47
Pelvic-fin base length 6.4 6.56.8 6.660.20
Pelvic-fin length 20.6 19.820.4 20.160.30
Anal-fin length 15.9 13.314.8 14.360.69
Caudal-peduncle length 12.4 12.614.4 13.760.77
Caudal-peduncle depth 10.1 10.210.9 10.560.30
% of head length
Head width 69.6 59.068.1 63.263.92
Head depth 48.4 45.051.4 47.862.66
Snout length 43.3 42.845.9 44.561.31
Snout to nostril distance 30.1 32.133.5 32.760.63
Nostril-to-operculum distance 66.7 65.968.1 66.961.00
Internostril width 19.1 19.022.1 20.461.34
Length of orbit 21.3 19.924.3 22.062.13
Interorbital width 25.7 24.430.1 26.862.38
Length of maxillary barbel 12.9 11.313.4 12.460.94
Length of lateral rostral barbel 11.9 11.113.0 11.860.81
Width of gape 33.3 28.530.7 29.660.90
Inter-lower lip width 22.6 20.021.4 20.660.67
% of interorbital width
Length of orbit 82.9 78.788.1 82.364.06
Meristics
Dorsal-fin ray count iii, 7K (5)
Pectoral-fin ray count v, 8, ii & v, 8, i (*); v, 9, i & iv, 10, i (2); v, 9, i (1); v, 7, ii & v, 8, i (1)
Pelvic-fin ray count ii, 7 (5)
Anal-fin ray count ii, 4, i (*); i, 5K (3); ii, 5K (1)
Caudal-fin ray count 16 (1); 17 (4*)
Lateral-line pore count 4144 + 21 on caudal fin
Lateral-line pore at pelvic-fin origin 1518
Lateral-line pore at dorsal-fin origin 1820
Lateral-line pore at anal-fin origin 3235
Circumpeduncle scale count 2022
Number of scale rows above / below lateral line 6 K8 K / 6 K7
No. scale rows below lateral line to pelvic-fin origin 6 K 7
Predorsal scale count 2225
164 Copeia 2014, No. 1
DISCUSSION
Species of Homalopteroides can be separated by branched
pelvic-fin ray count into two groups, the H. wassinkii group
(67 branched pelvic-fin rays) and the H. stephensoni group
(8 branched pelvic-fin rays). Homalopteroides avii belongs to
the H. wassinkii group along with H. wassinkii, H. modestus,
H. rupicola, H. smithi, H. tweediei, H. indochinensis, and H.
nebulosus. Homalopteroides avii is distinguished from other
species of the H. wassinkii group by the combination of a
larger gape width of 28.533.3% HL and a lateral cephalic
stripe (description of color pattern on head unavailable for
H. wassinkii and H. indochinensis). The H. wassinkii group has
a wider distribution than does the H. stephensoni group (H.
stephensoni, H. weberi, and H. yuwonoi), which is restricted to
Sumatra and Borneo.
Of the seven species of Homalopteroides reported to occur
in Borneo (H. wassinkii, H. stephensoni, H. weberi, H. tweediei,
H. nebulosus, H. yuwonoi, and H. avii), H. stephensoni, from
the Mahakam (Hora, 1932), Rajang, Katingan, Barito,
Kapuas, and Segama (Tan, 2009) basins, H. weberi, from
the Akar River (Hora, 1932), H. yuwonoi, from the Kapuas
basin (Kottelat, 1998), and H. avii, from the Rajang basin, are
known only from Borneo. Homalopteroides wassinkii was
originally described from Java, and H. tweediei and H.
nebulosus were originally described from Peninsular Malay-
sia. Some records of species of Homalopteroides identified
from Borneo may be misidentifications. Records of H.
wassinkii in Borneo by Fowler (1905; Baram basin), Weber
and Beaufort (1916; Kapuas and Mahakam basins), and
Parenti and Lim (2005; Rajang basin) were misidentifica-
tions of H. weberi (Randall and Page, 2012:335), H.
stephensoni (Hora, 1932:281), and H. avii (this study),
respectively. The specimens from Borneo identified as H.
tweediei in the Kapuas basin by Roberts (1989) and in the
Rajang basin by Parenti and Lim (2005), and as H. nebulosus
in the Kapuas basin by Roberts (1989) and Kottelat and
Widjanarti (2005), and in the Rajang basin by Parenti and
Lim (2005), warrant further investigation.
Homalopteroides avii is known only from the Rajang River
basin of Borneo. Specimens of H. avii were collected from
the Balui River in 1991 around 40 km SW of the Bakun
hydroelectric dam prior to its construction. We are unaware
of any recent collections from the Balui River, and the
population status of H. avii is unknown. It is likely that H.
avii has been extirpated from its type locality. The discovery
of this species 22 years after it was first collected is a
testament to the important role natural history museums
and collecting expeditions have for understanding biodi-
versity in the face of constantly changing environments due
to anthropogenic influence.
MATERIAL EXAMINED
Homalopteroides indochinensis: Vietnam: Indo-China (? Ton-
kin): BMNH 1933-8-19-50 (holotype, unique), 1 alc.
Homalopteroides modestus: Thailand: Kanchanaburi Prov.:
Mae Khlong basin: ANSP 179826, 5 alc; NIFI 4508, 1 alc;
NIFI 4517, 1 alc; UF 172926, 1 alc; UF 173067, 1 alc; UF
176377, 10 alc; UF 176408, 2 alc; UF 176438, 8 alc; UF
176454, 4 alc; UF 176544, 1 alc; UF 176557, 8 alc;
UF 181080, 5 alc; UF 181141, 1 alc; UF 181160, 9 alc;
ZRC 53385, 1 alc; ZRC 53386, 1 alc. Thailand: Tak Prov.:
Salween basin: NIFI 3786, 1 alc; NIFI 4514, 1 alc; ROM
51147, 2 alc; ZRC 41272, 4 alc. Myanmar: Tanintharyi
Region: Tenasserim basin: ZRC 22889, 1 alc. Myanmar: (?)
Kayin State: (?) Salween basin: BMNH 1893.2.16.50 (para-
lectotype of Helgia modesta), 1 alc.
Homalopteroides nebulosus: Malaysia: Kelantan: Sok River:
BMNH 1967.11.15.15 (paratype of Homaloptera nebulosa), 1
Fig. 3. Scatterplot of gape width/head
length for all species of Homalopteroides.
Randall and PageHomalopteroides avii 165
alc; SU 66428 (paratype of Homaloptera nebulosa), 1 alc; ZRC
1759 (paratype of Homaloptera nebulosa), 1 alc; ZRC 2020
(holotype of Homaloptera nebulosa), 1 alc.
Homalopteroides rupicola: Myanmar: Myitkyna District: San-
kha River: SU 28726 (paratype of Chopraia rupicola), 1 alc.
Homalopteroides smithi: Thailand: Nakhon Srithammarat:
Ban Kiriwong: BMNH 1934.12.18.34, 1 alc; UF 183330, 3
alc; UF 183411, 2 alc; UF 183915, 1 alc; USNM 109821
(syntype of Homaloptera smithi), 5 alc. Surathani Prov.: ANSP
179981, 2 alc; NIFI 3030, 3 alc. Trang Prov.: ANSP 76851, 3
alc; ANSP 76852, 1 alc.
Homalopteroides stephensoni: Indonesia: West Kalimantan:
Sungai Pinoh: USNM 230254, 5 alc.
Homalopteroides tweediei: Malaysia: Johore: Mawai: BMNH
1938.12.1.132 (paratype of Homaloptera tweediei), 1 alc; SU
33012 (holotype of Homaloptera tweediei), 1 alc; SU 33013
(paratypes of Homaloptera tweediei), 2 alc.
Homalopteroides wassinkii: Indonesia: Java: Buitenzorg:
Tjampea: BMNH 1866.5.2.52, 1 alc. Indonesia: Java: Lab.
Binnenvisscherij (fishery): UMMZ 155660, 1 alc.
Homalopteroides weberi: East Malaysia: Baram River: ANSP
68718, 11 alc. East Malaysia: Sarawak: Akar River: BMNH
1895.7.2.81 (syntype of Homaloptera weberi), 7 alc. East
Malaysia: Sarawak: BMNH 1933.8.9.7, 1 alc; BMNH
1978.9.5.4547, 3 alc; ROM 70456, 3 alc; ROM 70458, 3 alc;
ROM 70464, 3 alc; ROM 70466, 3 alc; ROM 82115, 1 alc;
ROM 82131, 3 alc. East Malaysia: Sabah: FMNH 99366, 1
alc; UMMZ 238960, 1 alc. Brunei: Darussalam: FMNH 117636,
2 alc.
Homalopteroides yuwonoi: Indonesia: Kalimantan Barat: Ka-
puas basin: MZB 5938 (holotype of Homaloptera yuwonoi), 1
alc.
ACKNOWLEDGMENTS
We would like to thank L. Parenti, A. Among, K. Luhat, and
A. Luhat for collecting Homalopteroides avii, and L. Parenti
and J. Clayton for providing Figure 5. For specimen loans
and access to institutional specimens, we thank M. Sabaj
Perez (ANSP), J. Maclaine (BMNH), M. Rogers (FMNH), S.
Suksri (NIFI), H. Lo pez-Fernandez (ROM), R. Robins (UF), D.
Nelson (UMMZ), J. Williams (USNM), and K. Lim (ZRC). The
U.S. National Science Foundation award (DEB 0845392) to
D. Reed provided the Visionary Digital (Palmyra, Virginia)
Fig. 4. Distribution of specimens examined (black dots), site at Kapit of Parenti and Lim 2005 (gray dot), and Bakun Dam (asterisk).
166 Copeia 2014, No. 1
System. Funding for this study was provided by the All
Cypriniformes Species Inventory Project funded by the U.S.
National Science Foundation (DEB 1022720).
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Fig. 5. Habitat of Homalopteroides avii in Sarawak, Balui River, Rajang
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Randall and PageHomalopteroides avii 167