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

Malaysian Borneo

Zachary S. Randall1 and Lawrence M. Page1

Homalopteroides avii, new species, is described from the Rajang River basin, Sarawak, Malaysian Borneo. It isdistinguished 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, 20.226.9% 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, 10.110.9% 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.

HOMALOPTEROIDES Fowler, 1905 was recentlyresurrected by Randall and Page (2012) based onthe following combination of characters: dorsal-

fin origin posterior to pelvic-fin origin, #60 lateral-linescales, #30 predorsal scales, and a mouth morphologycharacterized by two thin and widely separated rostralbarbels, thin crescent-shaped upper lip, the absence ofstructures such as a mental pad or lobes between the lateralportions of the lower lip, and a chin that extends anteriorof the lateral portions of the lower lip. The ten valid speciesof Homalopteroides are: H. wassinkii (Bleeker, 1853), H.modestus (Vinciguerra, 1890), H. rupicola (Prashad andMukerji, 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), andH. yuwonoi (Kottelat, 1998) (Randall and Page, 2012).Homaloptera manipurensis Arunkumar, 1999 was recognizedas a species of Homalopteroides by Randall and Page (2012)and as a questionable junior synonym of Homalopteroidesrupicola (Prashad and Mukerji, 1929) by Kottelat (2012). Wefollow 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 northeasternIndia, Myanmar, Thailand, Laos, Cambodia, Vietnam,Peninsular Malaysia, Sumatra, Java, and Borneo. Six specieshave been reported from Borneo: H. wassinkii (Fowler,1905; Weber and Beaufort, 1916; Hora, 1932; Parenti andLim, 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 fromBorneo 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 fromthe tip of the snout to the origin of the pectoral fin,distance between the anus and anal fin is from the analopening to the base of the first anal-fin ray, snout tonostril distance is from the tip of the snout to the anteriorpart of the nostril, nostril to operculum distance is fromthe most posterior margin of the nostril to the hindmostpart of the opercle, internostril width is the narrowestdistance between the nostrils, interorbital width is theleast distance between the edge of the orbits traversing theorbital rim, interrostral width is the narrowest distancebetween the rostral barbels, and inter-lower lip width isthe narrowest distance between the lateral portions of thelower lip. The lateral cephalic stripe extends horizontallyand posteriorly from between the lateral-rostral andmaxillary barbels to the cheek. The terms origin andinsertion refer, respectively, to the anterior and posteriorpoints of fin bases. All fin-ray counts are given as follows:simple rays in Roman numerals followed by branched raysin Arabic numerals, where dorsal fin- and anal fin-raycounts include the last ray split at the base represented byK. The caudal-fin ray count is the total number ofbranched rays. Small scales found just before the dorsal,pectoral, and anal fins are counted as K. Lengths weremeasured to the nearest 0.1 mm using digital calipers andtaken on the left side when possible. All measurements arein millimeters (mm). Measurements of the head arepresented as proportions of head length (HL), and eyelength is presented as a proportion of interorbital width(IO). Head length and measurements of the body are givenas proportions of standard length (SL). One hundred fifty-five individuals were examined representing all species ofHomalopteroides.Institutional abbreviations follow Sabaj Perez (2012),

the abbreviation for alcoholic specimens is alc, and a ?represents lack of data or uncertain locality. Photographswere taken of preserved specimens using a VisionaryDigital (Palmyra, Virginia) with Canon 40D and 5Dcameras at UF and edited using Photoshop CS3. Whencoordinates were unavailable, they were estimated usingmaps 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 9th edition. Species in thisstudy are recognized using the phylogenetic speciesconcept; i.e., a species is the smallest monophyletic groupthat is diagnosable by one or more unique characterstates.

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 entersBatang Balui, just downstream from logging camp, currentswift, 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 localityand date as holotype.

Paratypes.USNM 323878, 1 alc, 39.5 mm SL, East Malaysia,Sarawak, Batang Balui, trib., Jangan Aya, flowing into BatangBesua, pH 7.3, water temperature 25uC, 2.41uN, 113.73uE, L.R. Parenti, K. Luhat, A. Among, 2 August 1991; USNM323879, 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, 3August 1991.

Diagnosis.Member of Homalopteroides as defined by Ran-dall and Page (2012). Homalopteroides avii is distinguishedfrom all other species of Homalopteroides (Table 1) by a widergape (Fig. 3) of 28.533.3% HL. It is further distinguishedby presence of a lateral cephalic stripe, a deeper caudalpeduncle, a circumpeduncular scale count of 2022, a totalpelvic-fin ray count of 9, a total pectoral-fin ray modal countof 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 aregiven in Table 2. Homalopteroides avii is a relatively largespecies for the genus, reaching 52.9 mm SL. The body is

slightly deeper than wide, arched predorsally, tapersposteriorly to the anal-fin insertion, and has a flattenedventral surface. When viewed dorsally the head is conicaland covered with small tubercles. The eyes are ovoid,positioned dorsolaterally midway between the snout tipand edge of the opercle, and smaller in length than theinterorbital width. The nostrils are closer to the snout tipthan to the edge of the opercle. The origin of the dorsal finis posterior to the origin of the pelvic fin and closer to thecaudal-fin base than to the snout tip. The pectoral finreaches slightly past the pelvic-fin origin and usually isslightly longer than the head. The pelvic fin does not reachthe anus and is not doubly branched (i.e., with 3 or 4 distalpoints on one ray vs. 2). The anal fin does not reach thecaudal-fin base. An axillary pelvic lobe is absent. Margins ofthe dorsal and anal fins are straight. The caudal fin is forkedwith rounded lobes; the lower lobe is slightly longer thanthe upper lobe.

The body is scaled; anterior to the anal-fin origin scales aredeeply embedded. Most scales, especially just posterior tothe supraoccipital and at the dorsal-fin origin, have one ormore small nipples (wart-like spinous projections) along thefree borders of the scales; up to five are present on one scale.The total lateral-line pore count is 4345, predorsal scalecount is 2225, circumpeduncular scale count is 2022. Scalecounts above and below the lateral line are 6K8K and 6K7, respectively. The scale count below the lateral line to thepelvic-fin origin is 6K7. Variations in fin counts are givenin Table 2. Dorsal fin rays iii, 7K; anal-fin rays iii, 45K;pectoral-fin rays ivv, 710, iii; pelvic-fin rays ii, 7; totalcaudal-fin ray count 1617.

The mouth (Fig. 2) is wide (28.533.3% HL) and inferiorwith the lower jaw visible. The lips are thin, smooth, andcontinuous around the corners of the mouth. The upper lipis crescent-shaped, and the lower lip is broadly interruptedmedially. The inter-lower lip width is large (20.022.6%HL), and the chin extends far in front of the lateral portionsof the lower lip. Rostral and postlabial grooves are present.Two pairs of rostral barbels are present, and a pair ofmaxillary barbels is at each corner of the mouth. Themedial-rostral barbel reaches just past or at the base of thelateral-rostral barbels. The lateral-rostral barbel reaches alittle more than half the distance to the base of themaxillary barbel. The maxillary barbel reaches horizontallyto a vertical at the anterior orbital rim or to mid-orbit. Thegill membranes are united to the isthmus with a largecentral furrow where the gill membranes meet. The gillopening extends from the level of mid-orbit to the ventralsurface of the body.

Coloration.In 70% ethanol: The general pattern is shownin Figures 1 and 2. In dorsal view, the ground color is creamand mottled brown. There are 56 brown dorsal saddles. The1st saddle is between the supraoccipital and dorsal-fin origin,is irregular, and may consist of up to five blotches; the 2nd

saddle spans most of the pelvic-fin length; the 3rd saddle islocated between the dorsal-fin insertion and anal-fin origin;the 4th saddle spans most of the anal-fin length; and the 5th

saddle is at the caudal-fin base and in most specimens doesnot reach the ventral surface. In two individuals, there is anextra saddle between the 3rd and 4th saddles. A dark brownpreorbital bar extends from between the bases of the medial-and lateral-rostral barbels and shifts laterally to meet theeye. 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 indark brown, and dark brown blotches may be presentdorsomedially. A dark brown bar covers the post-epiphysealfontanelle. Postorbital bars extend to the supraoccipital andto the posterior edge of the opercle in alignment with thelateral line.

The lateral view shares the same ground color pattern asthe dorsal view. A dark brown lateral cephalic stripe extendshorizontally and posteriorly from between the lateral-rostraland maxillary barbels to the posterior edge of the orbitwhere it continues diagonally upward for a small distance.In some individuals, the upward extension of the stripe is anunconnected dark brown blotch. Over the lateral line is asolid brown stripe that contains circular blotches in largerindividuals. Mottled brown blotches sometimes coalesce toform distinct lines above and below the lateral line and mayform large blotches between the pelvic fin and caudal-finbase in larger individuals.

The venter is cream with mottled dark brown blotchesrestricted to a band at the origin of the anal fin. The bases ofthe rostral barbels are dark brown. The upper lip and thelateral aspect of the lower lip are mottled dark brown.

All fins are hyaline with fine brown blotches or bands. Thedorsal fin has two bands, the pectoral fin has one or twobands, and the pelvic fin has one or two bands. The anal finhas one brown blotch (restricted in one individual to thesecond ray). The caudal fin has two brown bands, a proximalband which may not always reach the superior extent of theupper lobe, and a subdistal V-shaped band that is pointedanteriorly. The lower lobe of the caudal fin is fully pigmentedup to the posterior edge of the subdistal V-shaped band. The

paired and caudal fins are brown at their bases. The dorsal finhas a dark brown blotch at its origin.

Distribution and ecological notes.Homalopteroides avii wascollected from the Balui River and downstream of theRajang River from Kapit, Rajang River basin, Sarawak(Fig. 4). Individuals were collected in streams with slow toswift currents and bottoms of mud, leaf litter, gravel,boulders, and rock outcrops with a pH of 7.37.7 and watertemperatures 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, Nemacheiluskapuasensis, 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 unavailablefor examination, THH9806, collected from downstream ofthe 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 examinedin parentheses.

Gapewidth% HL

Lateral cephalicstripe present(Yes/No)

Caudal-peduncledepth % SL

Circumpeduncularscale count

Totalpelvic-raycount

Total pectoral-ray count(modal)

Total lateral-line porecount

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), cannotpresently be determined. Two other specimens identified byParenti and Lim (2005) as H. wassinkii, FMNH 97441 and68140, were collected from the Mengiong and Putai rivers,Rajang River basin, respectively. FMNH 68140 was examinedin this study and is identifiable as H. cf. nebulosus; FMNH97441 was not examined.

Etymology.Named avii, in memory of Lawrence AviGreenberg (21 July 198226 November 2011). The diagnos-tic lateral cephalic stripe of this species, reminiscent of asmile, is a symbol of Avis gentle disposition and good-hearted nature. He is an inspiration to and missed friend ofthe first author and many others. Latinized as a noun in thegenitive singular.

Table 2. Morphometric measurements and meristic counts for Homalopteroides avii (n = 5). All measurements in mm. Numbers of individuals inparentheses. 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 26.860.94Body depth 19.0 15.517.1 16.360.72Body depth at anus 13.7 13.013.8 13.360.34Body width 17.2 13.917.2 15.561.60Predorsal length 53.9 52.954.3 53.860.62Prepectoral length 20.4 20.321.3 20.860.46Prepelvic length 46.3 45.146.1 45.760.52Preanal length 79.0 77.079.0 78.260.85Pre-anus length 73.5 71.773.4 72.360.75Distance between anus and anal fin 5.1 5.57.1 6.060.75Dorsal-fin base length 12.6 12.013.2 12.460.53Dorsal-fin length 21.5 20.021.0 20.660.42Pectoral-fin base length 9.8 10.811.8 11.160.48Pectoral-fin length 28.3 26.928.0 27.460.47Pelvic-fin base length 6.4 6.56.8 6.660.20Pelvic-fin length 20.6 19.820.4 20.160.30Anal-fin length 15.9 13.314.8 14.360.69Caudal-peduncle length 12.4 12.614.4 13.760.77Caudal-peduncle depth 10.1 10.210.9 10.560.30% of head length

Head width 69.6 59.068.1 63.263.92Head depth 48.4 45.051.4 47.862.66Snout length 43.3 42.845.9 44.561.31Snout to nostril distance 30.1 32.133.5 32.760.63Nostril-to-operculum distance 66.7 65.968.1 66.961.00Internostril width 19.1 19.022.1 20.461.34Length of orbit 21.3 19.924.3 22.062.13Interorbital width 25.7 24.430.1 26.862.38Length of maxillary barbel 12.9 11.313.4 12.460.94Length of lateral rostral barbel 11.9 11.113.0 11.860.81Width of gape 33.3 28.530.7 29.660.90Inter-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

MeristicsDorsal-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 finLateral-line pore at pelvic-fin origin 1518Lateral-line pore at dorsal-fin origin 1820Lateral-line pore at anal-fin origin 3235Circumpeduncle scale count 2022Number of scale rows above / below lateral line 6 K8 K / 6 K7No. scale rows below lateral line to pelvic-fin origin 6 K 7Predorsal scale count 2225

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DISCUSSION

Species of Homalopteroides can be separated by branchedpelvic-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 tothe 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 otherspecies of the H. wassinkii group by the combination of alarger gape width of 28.533.3% HL and a lateral cephalicstripe (description of color pattern on head unavailable forH. wassinkii and H. indochinensis). The H. wassinkii group hasa wider distribution than does the H. stephensoni group (H.stephensoni, H. weberi, and H. yuwonoi), which is restricted toSumatra and Borneo.

Of the seven species of Homalopteroides reported to occurin Borneo (H. wassinkii, H. stephensoni, H. weberi, H. tweediei,H. nebulosus, H. yuwonoi, and H. avii), H. stephensoni, fromthe Mahakam (Hora, 1932), Rajang, Katingan, Barito,Kapuas, and Segama (Tan, 2009) basins, H. weberi, fromthe Akar River (Hora, 1932), H. yuwonoi, from the Kapuasbasin (Kottelat, 1998), and H. avii, from the Rajang basin, areknown only from Borneo. Homalopteroides wassinkii wasoriginally described from Java, and H. tweediei and H.nebulosus were originally described from Peninsular Malay-sia. Some records of species of Homalopteroides identifiedfrom Borneo may be misidentifications. Records of H.wassinkii in Borneo by Fowler (1905; Baram basin), Weberand Beaufort (1916; Kapuas and Mahakam basins), andParenti 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 theRajang basin by Parenti and Lim (2005), and as H. nebulosus

in the Kapuas basin by Roberts (1989) and Kottelat andWidjanarti (2005), and in the Rajang basin by Parenti andLim (2005), warrant further investigation.Homalopteroides avii is known only from the Rajang River

basin of Borneo. Specimens of H. avii were collected fromthe Balui River in 1991 around 40 km SW of the Bakunhydroelectric dam prior to its construction. We are unawareof any recent collections from the Balui River, and thepopulation status of H. avii is unknown. It is likely that H.avii has been extirpated from its type locality. The discoveryof this species 22 years after it was first collected is atestament to the important role natural history museumsand collecting expeditions have for understanding biodi-versity in the face of constantly changing environments dueto 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; UF176377, 10 alc; UF 176408, 2 alc; UF 176438, 8 alc; UF176454, 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; ROM51147, 2 alc; ZRC 41272, 4 alc. Myanmar: TanintharyiRegion: 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/headlength for all species of Homalopteroides.

Randall and PageHomalopteroides avii 165

alc; SU 66428 (paratype of Homaloptera nebulosa), 1 alc; ZRC1759 (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, 3alc; UF 183411, 2 alc; UF 183915, 1 alc; USNM 109821(syntype of Homaloptera smithi), 5 alc. Surathani Prov.: ANSP179981, 2 alc; NIFI 3030, 3 alc. Trang Prov.: ANSP 76851, 3alc; ANSP 76852, 1 alc.

Homalopteroides stephensoni: Indonesia: West Kalimantan:Sungai Pinoh: USNM 230254, 5 alc.

Homalopteroides tweediei: Malaysia: Johore: Mawai: BMNH1938.12.1.132 (paratype of Homaloptera tweediei), 1 alc; SU33012 (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: ANSP68718, 11 alc. East Malaysia: Sarawak: Akar River: BMNH1895.7.2.81 (syntype of Homaloptera weberi), 7 alc. EastMalaysia: Sarawak: BMNH 1933.8.9.7, 1 alc; BMNH1978.9.5.4547, 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, 1alc; UMMZ 238960, 1 alc. Brunei: Darussalam: FMNH 117636,2 alc.

Homalopteroides yuwonoi: Indonesia: Kalimantan Barat: Ka-puas basin: MZB 5938 (holotype of Homaloptera yuwonoi), 1alc.

ACKNOWLEDGMENTS

We would like to thank L. Parenti, A. Among, K. Luhat, andA. Luhat for collecting Homalopteroides avii, and L. Parentiand J. Clayton for providing Figure 5. For specimen loansand access to institutional specimens, we thank M. SabajPerez (ANSP), J. Maclaine (BMNH), M. Rogers (FMNH), S.Suksri (NIFI), H. Lopez-Fernandez (ROM), R. Robins (UF), D.Nelson (UMMZ), J. Williams (USNM), and K. Lim (ZRC). TheU.S. National Science Foundation award (DEB 0845392) toD. 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 AllCypriniformes Species Inventory Project funded by the U.S.National Science Foundation (DEB 1022720).

LITERATURE CITED

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Fig. 5. Habitat of Homalopteroides avii in Sarawak, Balui River, RajangRiver basin, 2 August 1991, 2.41uN, 113.73uE. Photo by Lynne Parenti.

Randall and PageHomalopteroides avii 167

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