Brevibora cheeya is a new species of cyprinid fish - Raffles Museum ...

THE RAFFLES BULLETIN OF ZOOLOGY 2011 

THE RAFFLES BULLETIN OF ZOOLOGY 2011 59(1): 77–82 

Date of Publication: 28 Feb.2011 

© National University of Singapore 

BREVIBORA CHEEYA, A NEW SPECIES OF CYPRINID FISH 

FROM MALAY PENINSULA AND SUMATRA 

Te-Yu Liao 

Department of Vertebrate Zoology, Swedish Museum of Natural History, SE-104 05 Stockholm, Sweden. 

E-mail: teyu.Liao@nrm.se 

Department of Zoology, Stockholm University, SE-106 91 Stockholm, Sweden 

Heok Hui Tan * 

Raffl es Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore, 

Kent Ridge, Singapore 119260, Republic of Singapore. 

*: corresponding author; e-mail: dbsthh@nus.edu.sg 

ABSTRACT. – Brevibora cheeya is a new species of cyprinid fi sh from the Malay Peninsula and Sumatra. 

It is distinguished from its only congener, B. dorsiocellata, by the complete lateral line, more scales along 

the lateral row and larger size. 

KEY WORDS. – Taxonomy, Brevibora cheeya, Brevibora dorsiocellata, Southeast Asia, Biodiversity. 

INTRODUCTION 

Brevibora is a monotypic genus diagnosed by the large, 

black dorsal-fi n blotch and less than nine predorsal vertebrae 

(Liao et al., 2010). Brevibora dorsiocellata (Fig. 1) is 

characterized amongst others, by having an incomplete lateral 

line (Duncker, 1904). However, variation of the lateral line 

length has been reported on by subsequent workers (Brittan, 

1972; Roberts, 1989). Examination of various populations of 

B. dorsiocellata reveals that some populations have complete 

lateral line while other populations have incomplete lateral 

line. This character is consistent within each population. 

The purpose of this paper is to describe a new species 

of Brevibora with complete lateral line from the Malay 

Peninsula and Sumatra. 

MATERIAL AND METHODS 

Counts were based on radiographs and cleared-and-stained 

(C&S) specimens. Caudal vertebrae referred to those 

vertebrae bearing a haemal-spine, including the compound 

centrum, and the remaining vertebrae were counted as 

abdominal vertebrae, including the vertebrae forming the 

Weberian apparatus. The last two rays of the dorsal fi n 

articulating on the same pterygiophore are counted as one. 

All perforated scales along the lateral line were counted as 

lateral line scales. Scales of lateral row were counted along 

the row of the lateral line, including scales on the caudal 

fi n. Defi nition of lateral line completeness follows that of 

77 

Brittan (1954). According to his defi nition, the lateral line 

is complete when it extends beyond the insertion of last 

anal-fi n ray; if incomplete, the lateral line may end at any 

point before the insertion of last anal-fi n ray. Predorsal 

scales were counted along the dorsum midline from the fi rst 

scale behind the head to the scale in front of the dorsal-fi n 

insertion. Transverse line scales were counted on a projected 

straight line from the origins of the dorsal and pelvic fi ns 

(F-J in Fig. 2). Circumpeduncular scale row was the number 

of longitudinal scale rows around the caudal peduncle. All 

measurements and counts were conducted on the left side 

of the body whenever possible. 

Measurements on specimens were taken point-to-point to 

the nearest 0.1 mm with a pair of digital calipers. Points 

used in measurements are given in Fig. 2. Numbers in 

parentheses used in description refer to number of specimens 

examined. 

Cleared-and-stained specimens were prepared according to 

Taylor and Van Dyke (1985) with slight modifi cations. 

Terminologies of color pattern followed Brittan (1954) unless 

noted. Vernacular group names follow Fang et al. (2009), 

viz. rasborins for the genus Rasbora and related genera. 

Specimens examined are from the Natural History Museum, 

London (BMNH), the Swedish Museum of Natural History 

(NRM) and the Zoological Reference Collection, Raffl es 

Museum of Biodiversity Research, National University of 

Singapore (ZRC).

Liao & Tan – Brevibora cheeya, new cyprinid fi sh from Malaysia 

TAXONOMY 

Brevibora cheeya, new species 

(Figs. 3, 4) 

Material examined. – .Holotype – ZRC 51965, 26.6 mm SL; 

Malaysia: Terengganu, Rantau Abang, 56 km to Kuala Terengganu; 

P.K.L. Ng et al, 16 May.1995. 

Paratypes - ZRC 40196, 10, 20.4-27.3 mm SL; same data as 

holotype. - CMK 8156, 20, 12.9-22.8 mm SL, ZRC 24127, 17, 

13.3-22.3 mm SL; same locality as holotype; M. Kottelat et al., 18 

Mar.1992. - ZRC 51966, 46, BMNH 2010.1.11.1-70, 70, 14.1-25.5 

mm SL; Malaysia: Terengganu, Rantau Abang, 56 km to Kuala 

Terengganu; L. Rüber et al., 28 Feb.2009. 

Additional non-type material. – ZRC 39158, 11 (1 C&S), 11.4-28.0 

mm SL; Indonesia: Sumatra: Jambi: Berbak Nature Reserve, Sungai 

Air Hitam Dalam; H.H. Ng & S.H. Tan, 16-17 Jun.1995. - ZRC 

43116, 1, 19.6 mm SL; Indonesia: Sumatra: Riau: Indragiri, Sungai 

Jakil; H.H.Tan et al., 25 Nov.1996. - ZRC 30911, 2 ex., 30.4-30.9 

mm SL; Sumatra: Banka island, Kampung Jelit; M. Kottelat et 

al., 2 Mar.1993. - ZRC 3206, 406, 10.9-38.8 mm SL; Malaysia: 

Pahang, Tasek Bera; C.C. Lindsey, 29 Mar.1963. - ZRC 18826, 7, 

25.9-31.9 mm SL; Malaysia: Selangor, Tanjung Malim; P.K.L. Ng 

Fig. 1: Brevibora dorsiocellata. ZRC 42313, 22.7 mm SL; 

Indonesia: Sumatra: Jambi: Sungai Alai: 19.5 km to Muara Tebo 

from Muara Bungo. 

Fig. 2: Points used in measurements of rasborins species. 1, 

Standard length (SL): A-N, from tip of upper jaw to end of hypural 

plate. 2, Total length (SL): A-H, from tip of upper jaw to tip of 

upper caudal-fi n lobe. 3, Head length: A-E, from tip of upper jaw 

to posterior edge of opercle. 4, Head depth: at level of posterior 

margin of orbit (D). 5, Orbital diameter: B-D, between horizontal 

margins of osseous orbit. 6, Snout length: A-B, from tip of upper 

jaw to anterior margin of osseous orbit. 7, Interorbital width: 

distance between upper margins (C) of each osseous orbit. 8, 

Predorsal length: A-F, from tip of upper jaw to dorsal-fi n origin. 

9, Preanal length: A-K: from tip of upper jaw to anal-fi n origin. 

10, Prepectoral length: A-I, from tip of upper jaw to pectoral-fi n 

origin. 11, Prepelvic length: A-J, from tip of upper jaw to pelvicfi 

n origin. 12, Dorsal hypural length: F-N, from anterior edge of 

dorsal-fi n insertion to end of hypural plate. 13, Body depth: at level 

of dorsal fi n origin (F). 14, Caudal peduncle length: L-N, from 

end of anal-fi n base to end of hypural plate. 15, Caudal peduncle 

depth: narrowest part of caudal peduncle (G-M). 

78 

et al., 14 Sep.1991. - ZRC 42852, 4, 20.0-21.5 mm SL; Malaysia: 

Johor, Sungei Kayu; H.H. Tan et al., 12 Mar.1998. 

Diagnosis. – Brevibora cheeya is distinguished from B. 

dorsiocellata by the complete lateral line (25-30 perforated 

scales vs. 4-9), more scales along the lateral row (29-32 vs. 

25-27), larger size (up to 39.0 mm SL vs. 23.0 mm SL); 

and larger head and prepectoral length (head length 28.5- 

30.0% SL, vs. 24.4-27.9; prepectoral length 29.6-32.9% 

SL, vs. 25.8-28.8). 

Description. – Refer to Fig. 3 for overall body shape and to 

Table 1 for morphometric data. Body laterally compressed, 

rather elongated. Head short. Snout pointed. Mouth terminal, 

reaching vertical of anterior orbit. Dorsal-fi n origin at highest 

point of body outline. Predorsal contour almost straight, with 

a slight depression on snout. Ventral outline slightly curved 

from tip of snout to the middle of caudal peduncle. Barbel, 

keel and tubercles absent. 

Dorsal-fi n origin opposite or slightly ahead to pelvic-fi n 

origin, tip of last branched ray opposite to anal-fi n origin, 

pointed, posterior outline rather straight. D. ii.6 (11). 

Longest pectoral-fi n ray (1 st or 2 nd branched ray) almost 

reaching pelvic-fi n origin. P. ii.11 (2), ii.12 (8). Pelvic fi n 

triangular, longest ray almost reaching anal-fi n origin. V. ii.8 

(10). Anal-fi n origin closely behind vent, posterior outline 

concave, longest ray not reaching caudal fi n A. iii.5 (11). 

Caudal fi n deeply forked. 

Lateral line complete with 25-30 perforated scales [25 (1), 

26 (2), 27 (5), 28 (1), 29 (1), 30 (1)], last two to six scales 

not perforated. Lateral row with 29-32 scales [29 (3), 30 

(3), 31 (3), 32 (2)], including two scales on base of caudal 

fi n, lateral row descending from 1 st to 4 th scale and rather 

horizontal between 5 th and 7 th , then descending from 8 th 

scale onwards to caudal-fi n base (scale number based on 

holotype). Predorsal scales 10-11 [10 (9), 11 (2)]. Transverse 

Fig. 3: Brevibora cheeya. ZRC 51965, holotype, 26.6 mm 

SL; Malaysia: Terengganu, Rantau Abang, 56 km to Kuala 

Terengganu. 

Fig. 4: Brevibora cheeya. ZRC 39158, 11.4 mm SL; Indonesia: Jambi 

Province: Berbak Nature reserve, Sungai Air Hitam Dalam.

line scales ½ 4/1/3 ½ (11). One scale between lateral-line 

scale row and the pelvic-fi n origin. Circumpeduncular scales 

½ 3/1/1 ½ (11). Pectoral axillary scale rudimentary. Pelvic 

axillary scale well developed, corresponding to 1/3 of pelvic 

fi n in length. 

Predorsal vertebrae 8-9 [8 (10), 9 (1)], abdominal vertebrae 

16 (11), caudal vertebrae 15-16 [15 (10), 16 (1)], total 

vertebrae 31-32 [31 (10), 32 (1)]; pharyngeal teeth in three 

rows, 5, 4, 2; 2, 4, 5 (1), rasborin process present. 

Coloration in preservative. – Dorsum dark brownish, ventral 

gradually lighter. Abdomen yellowish. Dorsum of head dark 

brownish. Tip of snout more densely pigmented than adjacent 

area, exhibiting an appearance of dusky lips. Suborbital 

yellowish. Gill cover rather transparent with brownish 

pigment. Iris whitish, black dorsally and ventrally. Dorsal 

stripe prominent, half of dorsal-scale row in width. Reticulate 

pattern on sides distinct. Melanophores densely scattered 

along distal margin of scales in three or four rows, more 

prominent in dorsal half of body. An axial streak running from 

two scales in front of caudal-fi n base to the upper edge of 

gill opening, fading out anteriorly. Dark lateral stripe absent. 

Supra-anal pigment present as a streak and connected with 

subpeduncle streak, both blackish. A distinct black blotch on 

Fig. 5: Map of Sumatra and Malay Peninsula showing the 

distribution of Brevibora cheeya (square; hollow symbol denotes 

the type locality) and B. dorsiocellata from comparative material 

(triangle; hollow symbol denotes the type locality). 


79 

anterior middle dorsal fi n, intensely pigmented with a defi ned 

outline, oval in shape, covering two unbranched and fi rst four 

branched rays. Other fi ns hyaline, with some melanophores 

scattered on fi n rays. Melanophores most intense on distal 

half of anal fi n rays, similar to a blotch in overall shape. 

Inter-radial membrane of anal fi n transparent. It is not sure 

whether the blotch on anal fi n due to stronger pigmentation 

caused by environmental factors. In the smallest specimen of 

ZRC 39158 (Fig. 4) the distal half of its pelvic fi n rays with 

highly scattered melanophores and pigmentation on its anal 

fi n rays more intense than larger specimens. It is postulated 

that this pigmentation fades out upon maturity. 

Distribution and field notes. – Brevibora cheeya is 

distributed both in the westerly and easterly fl owing river 

basins of Peninsular Malaysia (western: Selangor; eastern: 

Terengganu, Pahang, Johor), Central Sumatra (Jambi, Riau) 

and Banka Island (Fig. 5). Brevibora dorsiocellata appears 

to be restricted to the Muar river basin in Malaysia, but 

this could be due to insuffi cient sampling (based on ZRC 

material and second author pers. obs.); but occurs in Central 

Sumatra, where B. cheeya is also present, but seemingly in 

different sub-basins. The Brevibora dorsiocellata group is 

also present in Borneo, and will be covered in more detail 

in future works. 

The type locality in Malaysia is in a coastal heath forest 

(Fig. 6). This habitat consists mainly of Melaleuca 

(Myrtaceae) stands on sandy and peaty substrate. The 

waters are tannin stained, slow fl owing, and acidic (pH 

4.5, as measured in February 2009). Syntopic species 

recorded from the type locality include: Boraras maculatus, 

Cyclocheilichthys apogon, Osteochilus spilurus, Parachela 

maculicauda, P. oxygastroides, Rasbora einthovenii, R. 

trilineata, Systomus johorensis, Trigonopoma gracile, 

T. pauciperforatum (Cyprinidae), Lepidocephalichthys 

furcatus, Pangio alcoides, P. semicincta (Cobitidae), 

Kryptopterus macrocephalus, Ompok leiacanthus, Wallago 

leerii (Siluridae), Hemibagrus nemurus, Pseudomystus 

leiacanthus (Bagridae), Clarias batrachus, C. meladerma 

(Clariidae), Parakysis verrucosa (Akysidae), Aplocheilus 

panchax (Aplocheilidae), Hemirhamphodon pogonognathus 

(Hemiramphidae), Monopterus albus (Synbranchidae), Nandus 

Fig. 6: Type locality of Brevibora cheeya at Rantau Abang, 

Terengganu, Malaysia (February 2009).

Table 1: Morphometry of Brevibora cheeya. SD = standard deviation. Measurements of B. dorsiocellata are from 12 specimens including two of paralectotypes BMNH 1913.5.24.13-14 and ten 

of ZRC 42313. Differences in range of variation are indicated in bold. 

Brevibora cheeya Brevibora dorsiocellate 


Holotype Paratype (n=10) n=12 

range mean SD range mean SD 

Standard length (SL; in mm) 26.6 20.4-27.3 17.0-22.1 

In % of SL 

total length 137.3 137.1-143.0 139.2 1.7 137.2-144.7 141 (n=10) 2.6 

head length 28.5 28.5-30.0 29.3 0.5 24.4-27.9 26.3 1.0 

body deapth 26.5 24.4-26.7 25.6 0.7 25.8-28.4 27.0 (n=10) 0.8 

caudal peduncular depth 11.9 11.7-13.0 12.3 0.5 10.5-13.6 11.8 0.5 

caudal peduncular length 18.5 16.6-18.5 17.3 0.6 17.3-20.7 18.5 0.8 

predorsal length 72.3 70.2-73.9 72.3 1.1 70.6-76.8 73.6 1.3 

preanal length 51.2 51.2-55.2 53.5 1.1 48.8-56.0 53.5 1.3 

80 

prepectoral length 29.6 29.6-32.9 31 0.8 25.8-28.8 27.7 0.8 

prepelvic length 53.8 52.7-54.6 53.7 0.6 51.8-57.9 54.4 1.3 

dorsal-hypural length 50.8 49.3-52.2 50.7 1.1 51.0-55.4 52.7 1.4 

length of dorsal fi n 25.8 26.2-29.3 27.8 1.0 

length of anal fi n 16.5 16.5-18.8 17.7 (n=7) 0.9 

In % of head length (HL) 

head depth 63.5 59.5-63.9 61.8 1.3 60.3-65.5 62 1.1 

snout length 32.4 28.3-32.8 30.5 1.6 23.4-30.9 28.3 1.7 

orbital diameter 35.1 33.3-40.0 37.2 2.2 34.5-37.7 36 1.1 

interorbital width 35.1 33.8-41.0 37.7 2.1 29.8-43.4 37.3 2.3

nebulosus (Nandidae), Pristolepis grooti (Pristolepididae), 

Belontia hasseltii, Betta imbellis, B. waseri, Luciocephalus 

pulcher, Parosphromenus paludicola, Trichopodus leerii, T. 

trichopterus, Trichopsis vittata (Osphronemidae), Channa 

bankanensis, C. lucius and C. striata (Channidae) (Kottelat 

et al., 1992; unpublished data). 

Etymology. – Cheeya and Beiya, are two Chinese deities 

who hunt ghosts for Yama; “chee” and “bei” mean seven 

and eight, respectively, and “ya” is an honorable title for a 

respected person. Cheeya is tall and Beiya is short; in allusion 

to its relatively larger size as compared to B. dorsiocellata. 

A noun in apposition. 

DISCUSSION 

Brevibora dorsiocellata was described by Duncker (1904) 

as a member of Rasbora, based on specimens from Muar 

(north-western corner of the southern state of Johor in 

Malaysia). This species is a popular aquarium fi sh well-known 

for the conspicuous black blotch on its dorsal fi n. Duncker 

(1904) mentioned that B. dorsiocellata has an incomplete 

lateral line with only eight perforated scales. However, in 

his drawing (Duncker, 1904), the lateral line is complete. 

Weber & Beaufort (1916) noticed this discrepancy and after 

examination of the types sent by Duncker, they stated that 

the drawing is erroneous. In addition, the drawing did not 

show the unique lateral line pattern shared by Brevibora and 

Kottelatia. Meinken (1951) described a subspecies of Rasbora 

dorsiocellata as R. dorsiocellata macrophthalma, also with a 

black blotch on the dorsal fi n and an incomplete lateral line, 

which was subsequently synonymized with B. dorsiocellata 

by Brittan (1954). The holotype of B. macrophthalma is 

missing (Eschmeyer & Fricke, 2009), so no direct comparison 

can be made. Moreover, the locality is vague and listed 

only as Malay Archipelago, as it had been obtained from 

the aquarium trade. Since B. macrophthalma also possesses 

an incomplete lateral line, it would not affect the validity 

of B. cheeya. We follow Brittan’s opinion in treating B. 

macrophthalma as a synonym of B. dorsiocellata. 

Brittan (1954) considered variation in length of lateral line 

as an intra-species variation in B. dorsiocellata. However, 

this conclusion was based on only fi ve specimens from three 

localities. Apparently the sample size is insuffi cient and, 

indeed, no variation was observed within any population 

by Brittan (1954). Roberts (1989) also noted the difference 

in lateral line length, but he adopted the interpretation of 

Brittan without additional comments. In material of B. 

dorsiocellata from Malay Peninsula and Sumatra, we found 

that lateral line length (complete or incomplete) is a stable 

character within a population and locality, and hence the 

lateral line completeness is a good character for species 

diagnosis within this genus. Brittan (1954) reported that two 

specimens of B. dorsiocellata (s. l.) from Pahang (SU 31196) 

possess incomplete lateral line of 30 and 33 mm SL, one 

with 20 pores on the left and 28 on the right, the other with 

24 pores on both sides. Although we did not examine these 

two specimens, it is likely that these two specimens are B. 


81 

cheeya. In some rasborins with complete lateral line, the last 

few scales are usually not perforated, commonly observed 

in the Rasbora daniconius and R. trifasciata species groups 

(Brittan, 1954), and this is probably the reason why Brittan 

(1954) made a unique defi nition of lateral line completeness 

(see MATERIAL AND METHODS). 

Kottelat & Vidthayanon (1993) provided a list of 47 miniature 

fi sh species from South and Southeast Asia according to the 

criterion proposed by Weitzman & Vari (1988), being less 

than 20 mm SL at female maturity or unknown maturity 

but never surpassing 26 mm SL in the wild. Brevibora 

dorsiocellata (sensu lato) was not included in their list, and 

none of the subsequent authors considered this species as 

a miniature fi sh. However, after this present description of 

B. cheeya, B. dorsiocellata (less than 23 mm SL) meets the 

criterion of a miniature fi sh. 

A black blotch in the middle of dorsal fi n is a conspicuous 

character distinguishing Brevibora from the other rasborins. 

Among species of rasborins, Rasbora atridorsalis and R. 

dorsinotata also possess a black blotch on the dorsal fi n 

(Kottelat & Chu, 1987), but the blotch is at the tip rather 

than in the middle of the fi n. 

Comparative material. – Brevibora dorsiocellata: 

BMNH1913.5.24.13-14, 2 paralectotypes, 17.0 & 19.8 

mm SL; Malaysia: Johore. - ZRC 2315, 3 paralectotypes, 

14.4-15.7 mm SL; Malaysia: Negeri Sembilan, Kuala 

Jelai. - BMNH 1994.12.16.237, 1, 21.6 mm SL; Indonesia: 

Kalimantan Tengah: Sungai Sebangau: 1 km downstream 

from Keram Benkari (10km South of Palangka Raya). - 

ZRC 42313, 11 (out of 67), up to 23.0 mm SL; Indonesia: 

Sumatra: Jambi: Sungai Alai: 19.5 km to Muara Tebo from 

Muara Bungo. ZRC 38494, 2, 9.6-17.8 mm SL; Indonesia: 

Sumatra: Jambi: Danau Pinang, Sungai Pijoan. - ZRC 38580, 

21, 13.5-19.9 mm SL; Indonesia: Sumatra: Jambi: Danau 

Kamining. - ZRC 38563, 33, 10.8-20.9 mm SL; Indonesia: 

Sumatra: Jambi: Sungai Alai. NRM 57237, 1 (C&S), 19.8 

mm SL; Aquarium. 

ACKNOWLEDGEMENTS 

The fi rst author is grateful to Erik Åhlander for translating the 

German literature; Kelvin K. P. Lim for his assistance during 

his stay in the Raffl es Museum of Biodiversity Research; 

two anonymous reviewers for their constructive suggestions. 

Part of the research was conducted in the Raffl es Museum 

of Biodiversity Research, supported by a grant to the fi rst 

author from the Riksmusei Vänner. The second author 

thanks Lukas Rüber, Sebastien Lavoué, Norsham Yaacob 

and Dennis Yong, for assistance and logistic support during 

the Malaysian peat swamp survey in 2009. Material collected 

and used for research by the second author is funded by 

various research grants including RMBR, R-154-000-318- 

112, R-264-001-004-272; and the National Geographic grant 

8509-08 and Natural Environment Research Council grant 

NE/F003749/1 to Lukas Rüber.


LITERATURE CITED 

Brittan, M.R., 1954. A revision of the Indo-Malayan fresh-water 

fi sh genus Rasbora. Monographs of the Institute of Science 

and Technology, Manila. 224 pp. 

Duncker, G., 1904. Die Fische der malayischen Halbinsel. 

Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 

21: 133–207. 

Eschmeyer, W. N. & R. Fricke (eds.), 2009. Catalog of Fishes 

electronic version (version of 9 September 2009). Available 

via http://research.calacadcmy.org/ichthyology/catalog/ 

fi shcatsearch.html. 

Fang, F., M. Noren, T.Y. Liao, M. Källersjö & S.O. Kullander, 2009. 

Molecular phylogenetic interrelationships of the south Asian 

cyprinid genera Danio, Devario, and Microrasbora (Teleostei, 

Cyprinidae, Danioninae). Zoologica Scripta, 38: 237–256. 

Kottelat, M. & X.L. Chu, 1987. Two new species of Rasbora 

Bleeker, 1860 from southern Yunnan and northern Thailand. 

Spixiana, 10: 313–318. 

Kottelat, M. & C. Vidthayanon, 1993. Boraras micros, a new genus 

and species of minute freshwater fi sh from Thailand (Teleostei: 

Cyprinidae). Ichthyological Exploration of Freshwaters, 4: 

161–176. 

82 

Kottelat, M., P. K. L. Ng & K. K. P. Lim, 1992. Recent collections 

of freshwater fish from Terengganu, Peninsular Malaysia. 

Malayan Naturalist, 46: 7–12. 

Liao, T.Y., S.O. Kullander & F. Fang, 2010. A phylogenetic analysis 

of the cyprinid fi sh genus Rasbora (Teleostei: Cyprinidae). 

Zoologica Scripta, 39: 155–176. 

Meinken, H., 1951. Mitteilungen der Fischbestimmungsstelle des 

VDA. Aquarien und Terrarien Zeitschrift, 4: 119–120. 

Roberts, T. R., 1989. The freshwater fi shes of western Borneo 

(Kalimantan Barat, Indonesia). Memoirs of the California 

Academy of Sciences, 14: 1–10. 

Taylor, W.R. & G.C. Van Dyke, 1985. Revised procedures for 

staining and clearing small fi shes and other vertebrates for 

bone and cartilage study. Cybium, 9: 107–119. 

Weber, M. & L.F. de Beaufort, 1916. The fishes of the Indo- 

Australian Archipelago. III. Ostariophysi: II Cyprinoidea, 

Apodes, Synbranchi. Brill, Leiden. 455 pp. 

Weitzman, S.H. & R.P. Vari, 1988. Miniaturization in South 

American freshwater fishes; an overview and discussion. 

Proceedings of the Biological Society of Washington, 101: 

444–465.

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