A synopsis of the South Asian fishes referred to Puntius - Verlag Dr ...

Ichthyol. Explor. Freshwaters, Vol. 23, No. 1, pp. 69-95, 11 figs., 4 tabs., June 2012 

© 2012 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902 

A synopsis of the South Asian fishes referred to Puntius 

(Pisces: Cyprinidae) 

Rohan Pethiyagoda*, Madhava Meegaskumbura** and Kalana Maduwage*** 

The tropical Asian cyprinid genus Puntius, which contains some 120 valid species, has long been suspected to be 

polyphyletic. Here, through an examination of external morphology, osteology, and analysis of 16S ribosomal 

RNA and cytochrome b gene fragments from 31 South Asian species hitherto referred to Puntius, we show that 

these fishes represent at least five lineages recognisable as genera. Puntius sensu stricto has the rostral barbels 

absent; last unbranched dorsal-fin ray weak or strong, smooth; and lateral line complete, with 22-28 pored scales. 

Systomus possesses maxillary and rostral barbels; last unbranched dorsal-fin ray stiff (‘osseous’), serrated; and 

lateral line complete, with 27-34 scales. Three new genera are proposed: Dawkinsia (type species Leuciscus filamentosus) 

is distinguished by lacking rostral barbels; having the last unbranched dorsal-fin ray smooth; lateral 

line complete, with 18-22 scales; and a juvenile colour pattern that includes three black bars on the body. Dravidia 

(type species Cirrhinus fasciatus) is distinguished by having both rostral and maxillary barbels present; lateral 

line complete, with 18–26 pored scales; dorsal fin with 4 unbranched and 8 branched rays, last unbranched 

dorsal-fin ray smooth; infraorbital 3 deep, partly overlapping the preoperculum; and free uroneural and postepiphysial 

fontanelle absent. Pethia (type species Barbus nigrofasciatus) is distinguished by having the last unbranched 

dorsal-fin ray stiff, serrated; infraorbital 3 deep, partially overlapping preoperculum; rostral barbels absent; 

maxillary barbels absent or minute; a black blotch on the caudal peduncle; and frequently, black blotches, spots 

or bars on the side of the body. The identities of Puntius sophore and Systomus immaculatus are clarified through 

the designation of neotypes; a lectotype is designated for Neolissochilus bovanicus; and precedence is given to the 

spelling bovanicus over bovianicus. 

Introduction 

Puntius has long been recognized as a “catch-all” 

genus for a variety of small tropical Asian 

cyprinids whose interrelationships are poorly 

understood (Kortmulder, 1972; Schut et al., 1984; 

Kottelat & Pethiyagoda, 1989; Kullander & Fang, 

2005; Kullander, 2008). The genus, which contains 

some 220 nominal species (about 120 of these 

* Ichthyology Section, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. 

E-mail: rohanpet@gmail.com 

** Department of Zoology, University of Peradeniya, Peradeniya, Sri Lanka. E-mail: madhava_m@mac.com 

*** Faculty of Medicine, University of Peradeniya, Peradeniya, Sri Lanka. E-mail: kalanapm@gmail.com 

Ichthyol. Explor. Freshwaters, Vol. 23, No. 1 

69 

considered valid), has seen a significant accretion 

of new species in recent years. The striking coloration 

of many species of Puntius has also led to 

their popularity as freshwater-aquarium fishes; 

many species are traded internationally as ornamentals 

(Collins et al., 2012). As the genus has 

hitherto been understood (e. g., Menon, 1999; 

Jayaram, 2010), species of Puntius range widely 

in size, from about 20 to 500 mm SL; possess 18-

70 

47 lateral-line scales; a complete, incomplete or 

interrupted lateral line; none, one or two pairs of 

barbels; a weak or strong, serrated or smooth last 

unbranched dorsal-fin ray; and a body coloration 

composed of a spot or blotch at the base of the 

caudal fin, one or more blotches or bars on the 

body, one or more lateral stripes, or none at all 

(Jayaram, 1991). The genus comprised 48 valid 

species in the South Asian region as at 1990 (Talwar 

& Jhingran, 1991), since when some 30 additional 

species have been described. 

The first experimental investigation of species 

groups within Puntius appears to have been by 

Kortmulder (1972). As a result of similarities 

between their larvae and juveniles, and through 

a number of hybridization experiments, he suggested 

close relationships between P. filamentosus 

and P. arulius; and between P. stoliczkanus, P. conchonius, 

P. phutunio, P. cumingii and P. nigrofasciatus, 

concluding that “cumingii, phutunio and gelius 

are more or less neotenous descendants from a 

nigrofasciatus like ancestor.” A single experiment 

he performed to hybridize P. conchonius and 

P. filamentosus, however, produced larvae that 

died shortly after hatching as a result of being 

unable to aerate their swim bladders, suggestive 

of their belonging to different sub-groups within 

Puntius. 

Following on this Taki et al. (1978), from an 

osteological study of 23 mostly Southeast Asian 

species of Puntius s.l., proposed six groups, of 

which two (their P. conchonius group and the 

P. arulius group) comprised South Asian species. 

In a later study, Shantakumar & Vishwanath 

(2006) included also P. sophore, the type species 

of Puntius, and showed that this species, together 

with P. terio and P. chola, possesses a postepiphysial 

fontanelle and a smooth, weak last 

unbranched dorsal-fin ray. Both these characters 

were absent in the other nine species of Puntius 

s.l. they examined, which included two members 

of the P. conchonius group of Taki et al. (1978). 

They also proposed a group not sampled by the 

previous authors, comprising P. sarana, P. jayarami 

and P. orphoides, which differed from Puntius 

s. s. and the P. conchonius group by possessing 

35-36 (vs. 29-33) vertebrae and a free uroneural 

(absent in Puntius s. s. and the P. conchonius 

group). 

The foregoing work suggests that Puntius s. l. 

comprises a number of species groups that share 

various combinations of apomorphic characters 

and character states. The vast range of the genus 

(all of tropical South and Southeast Asia), and the 

large number of included species, has made the 

resolution of all the species groups of which 

Puntius is presently composed a challenge. The 

conservative body plan of the included species 

too, has resulted in analyses of external morphology 

alone (e. g. Pillay, 1951; Jayaram, 1991; De 

Silva & Liyanage, 2010) proving to be of limited 

value in identifying species groups. At the same 

time, given the steady accretion of new species 

to Puntius s. l., especially from South Asia, an 

improved understanding of the species-groups 

that comprise it is desirable. Here, based on an 

analysis of external morphology and osteology, 

and a molecular phylogeny constructed from a 

mitochondrial 16S ribosomal RNA gene fragment 

(~ 540 bp) and a cytochrome b gene fragment 

(~ 550 bp) including 30 putative species of Indian 

and Sri Lankan Puntius s. l., we show that at least 

five lineages are present within South Asian 

Puntius, which we recognize as distinct genera. 

Material and methods 

Specimens referred to in this study are deposited 

in the Australian Museum, Sydney (AMS); the 

Natural History Museum, London (BMNH); the 

Collection of Maurice Kottelat, Cornol (CMK); 

the Raffles Museum of Biodiversity Research, 

Singapore (ZRC); the collection of the Wildlife 

Heritage Trust of Sri Lanka (WHT), now deposited 

in the National Museum of Sri Lanka, Colombo 

(NMSL); and the Zoological Survey of 

India, Kolkata (ZSI). Abbreviations: SL, standard 

length; HL, head length. 

Measurements were made with digital Vernier 

calipers and recorded to the nearest 0.05 mm. 

Drawings were made using a Motic dissecting 

microscope fitted with a camera lucida. Methods 

for taking counts and measurements follow 

Pethiyagoda & Kottelat (2005a), except that scales 

in transverse line on body were counted downwards 

and backwards from dorsal-fin origin to 

the pelvic-fin origin. Osteological descriptions are 

based on alizarin-stained specimens (2 from each 

species, following the method of Taylor & Van 

Dyke, 1985) and/or x-ray radiographs. Osteological 

nomenclature follows Conway (2011). 

Numbers in (square) brackets following a count 

indicate the frequency of that count. 

Pethiyagoda et al.: Synopsis of South Asian Puntius

‘South Asia’ as used here is the political unit 

previously referred to as British India, bounded 

approximately by Pakistan, Myanmar and Sri 

Lanka. ‘Southeast Asia’ as used here is tropical 

continental Asia east of South Asia. 

DNA analysis. DNA was extracted from ethanolpreserved 

tissues extracted from 39 species (see 

Table 1) using Qiagen tissue extraction kits and 

manufacturer’s protocols. Mitochondrial 16S ri- 


71 

bosomal RNA gene and cytochrome-b gene fragments 

were amplified using primers16Sar and 

16Sbr (Palumbi, 1996) which amplified ~540 bp 

of the 16S rRNA gene and CB-J-10933 (5'-TATGT- 

TCTACCATGAGGACAAATATC-3'), and BSF4 

(5'-CTTCTACTGGTTGTCCTCCGATTCA-3') 

(Bossuyt & Milinkovitch, 2000), which amplified 

~ 550 bp of cytochrome-b gene. 

PCR conditions were as follows: denaturation 

at 95 °C for 40 s, annealing at 45 °C for 40 s and 

Table 1. GenBank accession numbers, species and locations of the 39 species of Cyprinidae included in the molecular 

analysis. 

reference no. Genbank accession no. species location country 

16s cytb 

WHT8818_9 JF793558 JF793592 Garra ceylonensis Homadola Sri Lanka 

WHT8819_31i JF793559 JF793593 Garra mullya Chalakudy India 

WHT8820_80 JF793560 JF793594 Labeo dussumieri Elahera Sri Lanka 

WHT8821_23i JF793561 JF793595 Labeo dussumieri Alleppey India 

WHT8822_29i JF793562 JF793596 Osteochilichthys nashii Chalakudy India 

WHT8834_28i JF793574 JF793608 Gonoproktopterus jerdoni Srirangapatam India 

WHT8851_1i JF793591 JF793625 Gonoproktopterus curmuca Chalakudy India 

WHT8848_85 JF793588 JF793622 Tor khudree Mawanana Sri Lanka 

WHT8849_AQ1 JF793589 JF793623 Barbonymus schwanenfeldii Aquar. spec. – 

WHT8823_59 JF793563 JF793597 Pethia bandula Galapitamada Sri Lanka 

WHT8838_64 JF793578 JF793612 Pethia nigrofasciata Mawanana Sri Lanka 

WHT8812_1 EU604682.1 EU604674.1 Pethia reval Kelani River Sri Lanka 

WHT8814_46 EU604684.1 EU604676.1 Pethia cumingii Bentota Sri Lanka 

WHT8816_75 EU604686.1 EU604678.1 Pethia melanomaculata Kandalama Sri Lanka 

WHT8850_AQ4 JF793590 JF793624 Pethia conchonius Aquar. spec. – 

WHT8815_6i EU604687.1 EU604679.1 Pethia ticto Boncron India 

WHT8832_20i JF793572 JF793606 Dravidia fasciata Chalakudy India 

WHT8852_40i JF796852 Dawkinsia assimilis Chalakudy India 

WHT8833_27i JF793573 JF793607 Dawkinsia filamentosa Kottayam India 

WHT8843_33 JF793583 JF793617 Dawkinsia singhala Menik River Sri Lanka 

WHT8844_19 JF793584 JF793618 Dawkinsia srilankensis Pallegama Sri Lanka 

WHT8824_48 JF793564 JF793598 Puntius bimaculatus Bentota Sri Lanka 

WHT8825_88 JF793565 JF793599 Puntius cf. bimaculatus Minneriya Sri Lanka 

WHT8846_11 JF793586 JF793620 Puntius titteya Kalu River Sri Lanka 

WHT8847_43 JF793587 JF793621 Puntius cf. titteya Bentota Sri Lanka 

WHT8826_54 JF793566 JF793600 Puntius thermalis Mawanana Sri Lanka 

WHT8827_24i JF793567 JF793601 Puntius chola Boncron India 

WHT8845_217ai JF793585 JF793619 Puntius sophore Boncron India 

WHT8828_40 JF793568 JF793602 Puntius layardi Walawe River Sri Lanka 

WHT8829_69 JF793569 JF793603 Puntius dorsalis Gin River Sri Lanka 

WHT8830_70 JF793570 JF793604 Puntius kelumi Gin River Sri Lanka 

WHT8831_35i JF793571 JF793605 Puntius dorsalis Mamallapuram Sri Lanka 

WHT8837_16i JF793577 JF793611 Puntius mahecola Kottayam India 

WHT8835_21 JF793575 JF793609 Systomus martenstyni Pallegama Sri Lanka 

WHT8826_76 JF793576 JF793610 Systomus sp. “Richmondi” Elahera Sri Lanka 

WHT8842_21i JF793582 JF793616 Systomus sarana Boncron India 

WHT8840_35 JF793580 JF793614 Systomus timbiri Menik River Sri Lanka 

WHT8841_45 JF793581 JF793615 Systomus spilurus Bentota Sri Lanka 

WHT8839_12 JF793579 JF793613 Systomus pleurotaenia Gin River Sri Lanka

72 

extension at 72 °C for 40 s, 35 cycles, with a final 

extension of 72 °C for 5 min for the 16s gene fragment; 

denaturation at 95 °C for 40 s, annealing at 

45 °C for 50 s and extension at 72 °C for 60 s, 35 

cycles, with a final extension of 72 °C for 5 min 

for the cytochrome-b gene fragment. Products 

were gel purified and sequenced on an ABI 377 

or ABI 3100 automated sequencer following 

manufacturer’s protocols. 

The 16s sequences were aligned using Clustal 

X (Jeanmougin et al., 1998) and adjusted by eye 

using Se-Al (ver. 2.0a9; Rambaut, 1996). Positions 

which were difficult to align and in which we had 

low confidence in positional homology were 

excluded from subsequent analyses, leaving a 

total 411 bp. Cytochrome-b sequences were 

aligned using translated amino acid sequences 

using Se-Al (ver. 2.0a9; Rambaut, 1996). 

Two Garra species (G. ceylonensis and G. mullya) 

were used as an outgroup for rooting the tree: 

the data of Yang et al. (2010) and Zheng et al. 

(2010) suggest that Garra is a closely-related basal 

genus in relation to most taxa included in this 

analysis. The data were analyzed using Bayesian 

and Maximum Parsimony (MP) criteria. We used 

Bayesian inference as implemented in MrBayes 

(Huelsenbeck & Ronquist, 2001) to generate a 

phylogenetic hypothesis of relationships among 

the taxa and to estimate a general time-reversible 

model of sequence evolution with gamma-distributed 

rate variation among sites and a proportion 

of invariant sites (GTR+I+G). We ran four 

Metropolis-Coupled Markov Chain Monte Carlo 

(MCMCMC) chains for 2 000 000 generations. The 

summed likelihood of the four chains converged 

on a stationary value by 50 000 generations (the 

burn-in time). We used the frequencies of clades 

in trees that were sampled every ten generations 

from the last 250 000 generations as estimates of 

the posterior probabilities of those clades (Huelsenbeck 

et al., 2001). Uniform priors were used 

throughout and branch lengths, topology, and 

nucleotide substitution parameters were unconstrained. 

For tree searches under a Maximum 

Parsimony criterion (all characters unordered and 

weighted equally) we used heuristic searches with 

TBR branch-swapping and random taxon addition 

as implemented in PAUP*4.0b10. A single tree 

(tree score 1919 steps) was recorded. A bootstrap 

analysis to determine node support was also carried 

out within a maximum parsimony framework. 

The molecular dataset used for this study 

can be accessed at: http://purl.org/phylo/treebase/phylows/study/TB2:S12407 

and http:// 

web.mac.com/madhavameegaskumbura. 

Results 

The original dataset comprised of a 552-bp fragment 

of cyt-b and a 537-bp fragment of 16s. After 

removing poorly aligned regions of the 16s fragment, 

a 1060-bp fragment remained, which was 

used in the analyses. 

From our Bayesian analysis, we chose the tree 

having the highest likelihood value as our best 

tree (Fig. 1). Maximum Parsimony analysis yielded 

a single tree (tree length = 1919; not shown 

because the tree topology from this analysis was 

very similar to that of the Bayesian analysis). 

The Bayesian analysis recovered five wellsupported 

clades within Puntius s. l., that we 

consider to be distinct genera on the basis of 

external-morphological and osteological characters 

(see Table 2 for genetic distances and below 

for diagnoses): Puntius s. s., Systomus, and three 

new genera, Dawkinsia, Dravidia and Pethia. The 

phylogeny shows that (1) Pethia is a sister group 

of Dravidia and Dawkinsia; (2) Systomus is the 

sister group of Puntius; and (3) [Puntius + Systomus] 

is sister to [Dawkinsia + Dravidia + Pethia]. 

Table 2. Percent uncorrected genetic distances between Dawkinsia, Dravidia, Pethia, Puntius and Systomus for the 

combined 16s and cyt-b fragments analysed. 

Pethia Dawkinsia Dravidia Systomus Puntius 

Pethia – 7.3-12.0 13.5-14.2 10.4-15.6 10.4-16.3 

Dawkinsia – 12.2-15.6 8.1-15.6 8.4-16.6 

Dravidia – 12.7-15.6 13.0-16.6 

Systomus – 9.5-18.1 


Puntius Hamilton, 1822 

Type species. Cyprinus sophore Hamilton, 1822 

(Fig. 2). 

Diagnosis. A genus of cyprinid fishes differing 

from all South and Southeast Asian genera of 

Cyprininae by the combination of the following 

characters and character states: adult size usually 

less than 120 mm SL; maxillary barbels absent 

or present, rostral barbels absent; dorsal fin with 

3-4 unbranched and 8 branched rays; anal fin 

with 3 unbranched and 5 branched rays; last 

unbranched dorsal-fin ray weak or strong, apically 

segmented, not serrated (Fig. 3c); lateral line 

complete, with 22-28 pored scales on the body; 

free uroneural present (Fig. 3d); gill rakers simple, 

acuminate (not branched or laminate); no antrorse 

predorsal spinous ray; a post-epiphysial fontanelle 

present (Fig. 3b; but see remarks on P. bimaculatus 

and P. titteya, below); supraneurals 4; 

infraorbital 3 slender (Fig. 3a); 5th ceratobranchial 

narrow; pharyngeal teeth 5 + 3 + 2; 12-14 abdominal 

and 14-16 caudal vertebrae; and a colour 

pattern that includes a (sometimes faint) black 

spot or blotch on the caudal peduncle. 

Remarks. Based on the comparisons made herein 

(Fig. 1) and the data of Shantakumar & Vishwanath 

(2006) and Kullander (2008), we include 

the following South Asian species in Puntius: 

P. chola (Hamilton, 1822), P. dorsalis (Jerdon, 1849), 

P. kelumi Pethiyagoda, Silva, Maduwage & Meegaskumbura, 

2008, P. layardi (Günther, 1868), 

P. mahecola (Valenciennes, in Cuvier & Valenciennes, 

1844), P. pugio Kullander, 2008, P. sophore 

(Hamilton, 1822), P. terio (Hamilton, 1822) and 

P. thermalis (Valenciennes, in Cuvier & Valenciennes, 

1844). 

Because they share the external characters 

diagnostic of Puntius, we tentatively retain the 

following South Asian species in that genus: 

P. amphibius (Valenciennes, in Cuvier & Valenciennes 

1842), P. arenatus (Day, 1878), P. burmanicus 

(Day, 1878), P. cauveriensis (Hora, 1937), 

P. crescentus Yazdani & Singh, 1994, P. kamalika 

Silva, Maduwage & Pethiyagoda, 2008, P. madhusoodani 

Krishnakumar, Pereira & Radhakrishnan, 

2011, P. melanostigma (Day, 1878), P. mudumalaiensis 

Menon & Rema Devi, 1992, P. muzaffarpurensis 

Srivastava, Verma & Sharma, 1977, P. narayani 

(Hora, 1937), P. parrah (Day, 1865), P. sahyadriensis 

Silas, 1953, P. vittatus (Day, 1865) and P. waa 


73 

geni (Day, 1872). We also tentatively retain in 

Puntius P. denisonii (Day, 1865) and P. chalakkudiensis 

Menon, Rema Devi & Thobias, 1999, for 

which we lacked material for DNA and osteological 

analysis: they possess a strikingly different 

coloration and mouth shape to all other congeners 

(Menon et al., 1999) and are likely to warrant 

placement in a separate genus in the future. 

Two species, P. bimaculatus (Bleeker, 1863a) 

and P. titteya Deraniyagala, 1929, that differ from 

all the other Puntius examined by lacking a postepiphysial 

fontanelle (Fig. 4b) and possessing only 

7 branched dorsal-fin rays, are nevertheless retained 

in Puntius (see Discussion). 

Puntius sophore (Hamilton, 1822) 

(Fig. 2) 

Cyprinus sophore Hamilton, 1822: 310, 389, pl. 19. 

Neotype. ZRC 35064, 71.8 mm SL; Bangladesh: 

Srimangal [= Sreemangal] town market, from Hail 

Hoar floodplain near Moulvi Bazaar; D. Hoggarth, 

Mar 1993. 

Topotypes. ZRC 35065-35069, 5, 59.4-80.6 mm 

SL, same data as neotype. 

Description of neotype. General body shape and 

appearance are illustrated in Fig. 2a. Morphometric 

data of neotype and 5 topotypes are given in 

Table 3; characters of topotypes, where different 

from neotype, are given in parentheses. Body 

stout, standard length about 2.4 times maximum 

depth, laterally compressed. Dorsal profile of 

head concave above eye; dorsal profile of body 

convex, rising steeply from occipital, almost 

horizontal at dorsal-fin origin. Ventral contour of 

body convex to anal-fin insertion, almost horizontal 

at pelvic-fin origin. Dorsal and ventral 

profiles of caudal peduncle concave to terminal 

scales on tail base. Caudal peduncle length 1.3 

(1.2-1.4) times its depth. 

Head short, its length about one-fifth standard 

length, somewhat compressed laterally. Eyes 

located forward of centre, superiorly on head, 

just visible in ventral and dorsal views. Nares 

dorsolaterally orientated. Mouth U-shaped in 

ventral aspect, subterminal, small, angle of gape 

not reaching below anterior nares in lateral aspect. 

Lips fleshy, entire, adnate to jaws. Upper lip 

overhanging lower one. Barbels absent.

74 

100 

100 

74 

100 

71 

10 changes 

100 

100 

91 

92 

Garra ceylonensis (LK) 

Garra mullya (IN) 

Labeo dussumieri (LK) 

Labeo dussumieri (IN) 

100 

83 

91 

100 

Osteochilichthys nashii (IN) 

Gonoproktopterus jerdoni (IN) 

Gonoproktopterus curmuca (IN) 

Tor khudree (LK) 

Barbonymus schwarnenfeldii (AQ) 

92 

92 

92 

99 

50 

100 

87 

99 

59 

100 

100 

100 

57 

100 

92 

96 

87 

100 Pethia bandula (LK) 

99 

98 Pethia nigrofasciata (LK) 

100 

98 

93 

100 

85 

Pethia reval (LK) 

Pethia cumingii (LK) 

Pethia melanomaculata (LK) 

Pethia conchonius (AQ) 

Pethia ticto (IN) 

Dravidia fasciata (IN) 

Dawkinsia assimilis (IN) 

Dawkinsia filamentosa (IN) 

Dawkinsia singhala (LK) 

Dawkinsia srilankensis (LK) 

60 

99 

50 

64 

64 

100 

84 

100 

96 

100 

64 

100 

100 

100 

100 

Puntius mahecola (IN) 

100 

66 

Puntius bimaculatus (LK) 

Puntius cf. bimaculatus (LK) 

Puntius cf. titteya (LK) 

Puntius titteya (LK) 

Puntius thermalis (LK) 

Puntius chola (IN) 

Puntius sophore (IN) 

Puntius dorsalis (LK) 

100 

65 

Puntius dorsalis (IN) 

Puntius kelumi (LK) 

Puntius layardi (LK) 

Systomus martenstyni (LK) 

Systomus sp. ‘Richmondi’ (LK) 

Systomus sarana (IN) 

100 

73 

100 

100 

Systomus timbiri (LK) 

Systomus spilurus (LK) 

Systomus pleurotaenia (LK) 

Outgroup 

Last unbranched dorsal-fin 

ray serrated; infraorbital 3 

deep; SL < 80 mm; rostral 

barbels absent; post-epiphysial 

fontanelle absent. 


ray smooth; SL < 80 mm; 

rostral, maxillary barbels 

present; post-epiphysial 



ray smooth; rostral barbels 

absent; 3 black bars on flank 

in juveniles; post-epiphysial 



ray smooth; 7 branched 

dorsal-fin rays; rostral 

barbels absent; postepiphysial 



ray smooth; 8 branched 

dorsal-fin rays; rostral barbels 

absent; post-epiphysial 

fontanelle present. 


ray serrated; rostral, maxillary 

barbels present; SL > 

80 mm; 27−34 lateral-line 

scales; post-epiphysial 


Fig. 1. Bayesian phylogram of the 31 putative species of South Asian Puntius s. l. analysed, with key shared 

morphological and osteological characters for each group. See Table 1 for list of material. Provenance: AQ, 

aquarium trade; IN, India; LK, Sri Lanka. Posterior probabilities are shown above nodes, parsimony bootstrap 

values below nodes. Photographs illustrate species within the respective genera: from top, Pethia bandula, Dravidia 

fasciata, Dawkinsia srilankensis, Puntius titteya, Puntius dorsalis and Systomus asoka. 


Pelvic-fin origin slightly anterior to dorsal-fin 

origin. Posterior margin of dorsal fin slightly 

concave, that of anal fin straight. Distal margins 

of pelvic and pectoral fins convex, rounded. 

Dorsal-fin with 4 unbranched rays (last one 

smooth, strong) and 8 branched rays, its origin 

above ninth lateral-line scale. Anal fin with 3 

unbranched and 5 branched rays. Pelvic fin with 

1 unbranched and 8 branched rays, not reaching 

anus. Pectoral fin with one unbranched and 15 

branched rays, not reaching pelvic-fin origin. 

Caudal fin deeply forked, its lobes pointed, with 

1 + 9 + 8 + 1 principal rays. 

Lateral-line pored scales on body 24 (23 [2]), 

plus 1 on caudal-fin base; 4 1 /2 scales in transverse 

line between lateral line and origin of dorsal fin, 

4 (3 1 /2 [1]) between lateral line and origin of pelvic 

fin; 1 /2 2/1/2 1 /2 scales in transverse line on caudal 

peduncle; 9 predorsal scales (excluding notched 

scale at dorsal-fin origin). An axillary scale approximately 

one-third length of pelvic fin extends 

backwards from pelvic-fin origin. Dorsal-fin base 


a 

b 

75 

Fig. 2. Cyprinus sophore: a, ZRC 35064, neotype, 71.8 mm 

SL; Bangladesh: Hail Hoar floodplain; b, copy of Hamilton’s 

drawing, in the collection of the Linnean Society 

of London (courtesy of Linnean Society of London). 

Table 3. Standard length (in millimetres) and proportional measurements as a percentage of standard length or 

head length of the neotype (ZRC 35064, 71.8 mm SL) and 5 topotypes (ZRC 35065-35069, 59.4-80.6 mm SL) of 

Puntius sophore. 

neotype range 

min. max. 

Standard length (mm) 71.8 59.4 80.6 

In percent SL 

Total length 130 127 133 

Head length 28.6 27.3 29.7 

Predorsal length 52.8 52.7 56.1 

Preanal length 76.9 71.2 76.9 

Prepelvic length 50.3 43.6 50.5 

Caudal-peduncle length 19.8 18.0 20.5 

Caudal-peduncle depth 14.9 13.8 15.2 

Body depth 42.5 40.2 42.5 

Dorsal-fin height 27.3 23.8 30.8 

Length of stiff portion of last unbranched dorsal-fin ray 17.5 17.1 19.9 

Pectoral-fin length 18.9 18.4 23.1 

Pelvic-fin length 20.6 19.0 22.6 

Distance from dorsal-fin origin to hypural fold 54.5 54.5 58.2 

Post-dorsal distance 36.4 36.4 39.8 

Maximum body width 17.3 14.0 17.3 

Length of dorsal-fin base 18.2 16.1 18.6 

Length of anal-fin base 13.9 11.4 14.7 

In percent HL 

Snout length 27 27 30 

Post-orbital head length 50 49 53 

Eye diameter 32 27 32 

Internarial width 26 22 27 

Interorbital width 42 38 42

76 

IO5 

Pop 

IO4 

a b c d 

IO5 

IO4 

IO3 

IO3 

So 

So 

IO2 

IO2 

IO1 

IO1 

Pa 

a 

Pop 

b c d 

Pef 

sheathed by 8 (7 [2]) scales, anal-fin base by 

6 (7 [3]) scales. 

In preservative (Fig. 2a), head and body 

brownish olive, lighter below lateral line; lower 

body, belly and chest pale cream-yellow. Fin rays 

light brown, interradial membrane hyaline. A distinct, 

horizontally-elongate black blotch slightly 

larger than eye, about 3 scales wide by 2 scales 

high, at posterior end of caudal peduncle. A smaller 

black blotch at base of dorsal-fin rays 5-7. 

F 

F 

So 

PU3 PU2 CC 

PU3 PU2 CC 

Fig. 4. Puntius bimaculatus, WHT 11029, 40.5 mm SL: a, circumorbital series; b, dorsal view of orbital region of 

cranium; c, last unbranched dorsal-fin ray; and d, caudal skeleton (scale bar 1 mm). Abbreviations: CC, compound 

centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural; 

Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; supraorbital sensory 

canal not shown. 

So 

Sc 

Pa 

PU3 PU2 CC 

PU3 PU2 CC 

Fig. 3. Puntius sophore, WHT11062, 28.9 mm SL; SL: a, circumorbital series; b, dorsal view of orbital region of 


centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Pef, postepiphysial 

fontanelle; Ph, parhypural; Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; 

Sc, supraorbital sensory canal; So, supraorbital; Un, free uroneural. 

Ep 

Pls 

Systomus McClelland, 1838 

Un 

H6 

H5 

H4 

H3 

H2 

H1 

Ph 

Ep 

Pls 

H6 

H5 

H4 

H3 

H2 

Type species. Systomus immaculatus M’Clelland, 

1839. 

H1 

Ph 

Diagnosis. A genus of cyprinid fishes differing 

from all South and Southeast Asian genera of 

Cyprininae by the combination of the following 

characters and character states: adult size greater 

than 80 mm SL (in South Asian species); maxillary 

and rostral barbels present; dorsal fin with 4 + 8 

rays, its last unbranched ray stiff, strongly serrated 

(Fig. 5c); lateral line complete, with 27-34 

scales on body; gill rakers simple, acuminate (not 

branched or laminate); no antrorse predorsal 


IO5 

IO4 

Pop 

So 

IO3 IO2 

IO1 

a b c d 


F 

Pa 

So 

Pls 

PU3 PU2 CC 

PU3 PU2 CC 

Fig. 5. Systomus spilurus, WHT 11028, 61.1 mm SL: a, circumorbital series; b, dorsal view of orbital region of 



Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; Un, free uroneural; 

supraorbital sensory canal not shown. 

spinous ray; free uroneural present (Fig. 5d); 

post-epiphysial fontanelle absent (Fig. 5b); 5 supraneurals; 

infraorbital 3 slender (Fig. 5a); 14-15 

abdominal and 17-19 caudal vertebrae; and colour 

pattern including a prominent, horizontallyelongate 

oval black blotch on the caudal peduncle, 

with no prominent bars or blotches on the 

body. 

Remarks. Based on the comparisons made herein, 

and the data of Shantakumar & Vishwanath 

(2006) and Kottelat & Pethiyagoda (1989), we 

include the following nine South Asian species 

in Systomus: S. asoka (Kottelat & Pethiyagoda, 

1989), S. compressiformis (Cockerell, 1913), S. jayarami 

(Vishwanath & Tombi Singh, 1986), S. martenstyni 

(Kottelat & Pethiyagoda, 1991), S. orphoides 

Valenciennes, in Cuvier & Valenciennes 1842, 

S. pleurotaenia (Bleeker, 1863a), S. sarana (Hamilton, 

1822), S. spilurus (Günther, 1868) and S. timbiri 

(Deraniyagala, 1963). An apparently undescribed 

Sri Lankan species with the label-name 

‘Puntius sp. Richmondi’ too, belongs to Systomus. 

Barbus clavatus McClelland, 1844 and B. shanensis 

Hora & Mukerji, 1934 have been variously placed 

in Puntius (e. g., Jayaram, 1991; Talwar & Jhingran, 

1991) and Poropuntius (e. g., Roberts, 1998; Jayaram, 

2010) by previous authors. We have not 

examined specimens of these and so refer them 

only tentatively to Systomus, though they may 

bear a closer relationship to the ‘Puntius binotatus 

group’ of Southeast Asia (see Discussion). 

Systomus immaculatus 

(Fig. 6) 

77 

Ep 

Un 

H6 

H5 

H4 

H3 

H2 

H1 

Ph 

Systomus immaculatus M’Clelland, 1839: 284, 380, 

pl. 44, fig. 5. 

Neotype. AMS B.7920, 187 mm SL; India: Assam; 

F. Day. 

Topotype. AMS B.7921, 193 mm SL (caudal fin 

broken); India: Assam; F. Day. 

Description of neotype. General body shape and 

appearance are illustrated in Figure 6a. Morphometric 

data of neotype and 1 topotype are given 

in Table 4; characters of topotype, where different 

from neotype, are given in parentheses. Body 

elongate, standard length about 2.8 times body 

depth. Dorsal profile of head rounded; dorsal 

body profile rising gently to dorsal-fin origin 

(a post-nuchal hump present), sloping gently 

downwards thereafter. Dorsal-fin origin slightly 

posterior to pelvic-fin origin. Ventral body profile 

curving gently ventrad to pelvic-fin base, almost 

level from there to anal-fin origin, rising steeply 

to caudal peduncle; caudal peduncle ventral 

margin level. 

Head relatively long, its length about onefourth 

standard length. Eyes located forward of 

centre, about mid-height on head. Mouth terminal; 

lips fleshy; lower-lip post-labial groove shallow, 

uninterrupted. Cheek and interorbital with 

about 10 sensory pores. Two pairs of barbels 

present; maxillary barbel long, its length slightly

78 

less than eye diameter; rostral barbel shorter, 

about half length of maxillary barbel. 

Pelvic-fin origin slightly anterior to dorsal-fin 

origin. Posterior margins of dorsal and anal fins 

concave. Distal margins of pectoral and pelvic 

fins convex, rounded. Dorsal-fin origin above 11th 

lateral-line scale. Dorsal fin with 4 unbranched 

and 8 branched rays, last unbranched ray stiff, 

strongly serrated posteriorly in its proximal twothirds, 

weak, segmented in its distal one-third. 

Last unbranched dorsal-fin ray longest, when 

adpressed not overlapping tip of last branched 

dorsal-fin ray. Anal fin with 3 unbranched and 5 

branched rays. Pelvic fin with 1 unbranched and 

8 branched rays. Pectoral fin with 1 unbranched 

and 15 branched rays. Caudal fin forked, lobes 

of equal length, longest rays about twice as long 

as median ones. Caudal fin with 8 upper and 7 

lower procurrent rays, and 1 + 9 + 8 + 1 principal 

rays. 

Table 4. Standard length (in millimetres) and proportional 

measurements as a percentage of standard length 

or head length of the neotype (AMS B.7920) and a topotype 

(AMS B.7921) of Systomus immaculatus. 

B.7920 B.7921 

Standard length (mm) 186.0 192.7 

Percent standard length 

Total length 127 – 

Head length 26.1 25.4 

Predorsal length 53.2 54.2 

Pre-anal length 75.1 72.1 

Pre-pelvic length 41.0 48.2 

Caudal-peduncle length 19.4 22.3 

Caudal-peduncle depth 15.2 14.8 

Body depth 35.9 35.6 

Dorsal-fin height 22.7 20.4 

Length of stiff portion of last 15.6 14.7 

unbranched dorsal-fin ray 

Pectoral-fin length 19.2 19.3 

Pelvic-fin length 19.7 18.8 

Dorsal-hypural distance 55.4 56.4 

Post-dorsal distance 38.8 40.1 

Maximum body width 

Percent head length 

15.4 14.5 

Snout length 30.1 29.5 

Post-orbital head length 53.6 52.2 

Head depth 83.1 81.0 

Eye diameter 23.0 22.3 

Maxillary barbel length 19.3 23.8 

Rostral barbel length 8.3 16.8 

Internarial width 23.0 23.5 

Interorbital width 43.8 42.8 

Lateral line present, complete, with 32 (33) 

pored scales on body, 2 (3) on caudal-fin base. 

Lateral line sloping gently downwards for about 

7 scales, level thereafter. Flank scales large, their 

central field (middle one-third) free of annuli, 

with about 15 irregular polygons. Radii radial, 

denser anteriorly and posteriorly, less dense in 

dorsal and ventral fields. 5 1 /2 scales in transverse 

line between lateral line and origin of dorsal fin, 

4 between lateral line and origin of pelvic fin; 

1 /2 3/1/3 scales in transverse line on caudal peduncle; 

12 (13) predorsal scales. An axillary scale 

approximately one-fourth length of pelvic fin 

extends backwards from pelvic-fin origin. 

Supraneurals 5 (6); 15 abdominal and 17 (18) 

caudal vertebrae (Fig. 6b); free uroneural present; 

infraorbital 3 slender. 

Dravidia, new genus 

Type species. Cirrhinus fasciatus Jerdon, 1849: 

305. 

Diagnosis. Dravidia differs from all other South 

and Southeast Asian genera of Cyprinidae by the 

combination of the following characters and 

character states: size small, usually less than 

60 mm SL; rostral and maxillary barbels present; 

lateral line complete, with 18-26 pored scales on 

body; dorsal fin with 4 unbranched and 8 branched 

rays, the last unbranched ray weak, smooth 

(Fig. 7c); anal fin with 3 unbranched and 5 

branched rays; gill rakers simple, acuminate (not 


spinous ray; infraorbital 3 deep, partly overlapping 

preoperculum (Fig. 7a); free uroneural and 

post-epiphysial fontanelle absent (Fig. 7 b, d); and 

one or two broad, black bars on flank, between 

bases of dorsal and anal fins. 

Remarks. The following nominal species are 

referred to Dravidia: D. afasciata (Jayaram, 1990), 

D. fasciata (Jerdon, 1849), D. kannikattiensis (Arunachalam 

& Johnson, 2003), D. melanampyx (Day, 

1865) and D. pradhani (Tilak, 1973). 

Etymology. Named for the Dravidian peoples of 

the southern peninsular India; gender feminine. 


a 

b 

Fig. 6. Systomus immaculatus, AMS B.7920, neotype, 187 mm SL; India: Assam; a, photograph; b, radiograph. 

IO5 

IO4 

Pop 

IO3 

So 

IO1 

IO2 

a b c d 


F 

So 

Pa 

PU3 PU2 CC 

Fig. 7. Dravidia fasciata, WHT 11043, 43.0 mm SL: a, circumorbital series; b, dorsal view of orbital region of 



Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; supraorbital sensory 

canal not shown. 

Ep 

Pls 

H6 

H5 

H4 

H3 

H2 

H1 

Ph 

79

80 

IO5 

IO4 

Pop 

IO3 

So 

IO1 

IO2 

a b c d 

Dawkinsia, new genus 

Type species. Leuciscus filamentosus Valenciennes, 

in Cuvier & Valenciennes, 1844. 

Diagnosis. Dawkinsia differs from all South and 

Southeast Asian genera of Cyprinidae by the 


character states: adult size usually 80-120 mm 

SL; rostral barbels absent, maxillary barbels 

present or absent; last unbranched dorsal-fin ray 

smooth (Fig. 8c); 4 unbranched and 8 branched 

dorsal-fin rays, 3 unbranched and 5 branched 

anal-fin rays; lateral line complete, with 18-22 

scales on body; gill rakers simple, acuminate (not 


spinous ray; free uroneural present (Fig. 8d); 4-5 

supraneurals; 15 precaudal and 14-17 caudal 

vertebrae; post-epiphysial fontanelle absent 

(Fig. 8b); infraorbital 3 slender, not overlapping 

preoperculum (Fig. 8a); juvenile (< 50 mm SL) 

colour pattern consisting of three black bars on 

body, persisting to adult stage in some species; a 

black, horizontally elongate blotch on the caudal 

peduncle in adults. 


and the data of Taki et al. (1978) and Pethiyagoda 

& Kottelat (2005a), we include the following 

nine species in Dawkinsia: D. arulius (Jerdon, 1849), 

D. assimilis (Jerdon, 1849), D. exclamatio (Pethiyagoda 

& Kottelat, 2005b), D. filamentosa (Valenciennes, 

in Cuvier & Valenciennes, 1844), D. rohani 

(Rema Devi, Indra & Marcus Knight, 2010), D. ru 

F 

Pa 

So 

PU3 PU2 CC 

Fig. 8. Dawkinsia filamentosa, WHT 11026, 67.1 mm SL: a, circumorbital series; b, dorsal view of orbital region of 



Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; Un, free uroneural; 

supraorbital sensory canal not shown. 

brotinctus (Jerdon, 1849) (see Knight et al., 2011), 

D. singhala (Duncker, 1912), D. srilankensis (Senanayake, 

1985) and D. tambraparniei (Silas, 1954). 

The genus appears to be restricted to Sri Lanka 

and southern India. 

Etymology. The genus is named for Richard 

Dawkins, for his contribution to the public understanding 

of science and, in particular, of evolutionary 

science; gender feminine. 

Pethia, new genus 

Ep 

Pls 

Un 

H6 

H5 

H4 

H3 

H2 

H1 

Ph 

Type species. Barbus nigrofasciatus Günther, 

1868. 

Diagnosis. Pethia differs from all South and 

Southeast Asian genera of Cyprinidae by the 


character states: small adult size (usually less than 

50 mm SL, exceptionally to 80 mm SL); rostral 

barbels absent; maxillary barbels usually absent, 

minute if present; last unbranched dorsal-fin ray 

stiff, serrated (Fig. 9c); 3 or 4 unbranched and 8 

branched dorsal-fin rays, 3 unbranched and 5 

branched anal-fin rays; gill rakers simple, acuminate 

(not branched or laminate); no antrorse 

predorsal spinous ray; infraorbital 3 deep, partially 

overlapping the cheek and preoperculum 

(Fig. 9a); free uroneural absent (Fig. 9d); 4 supraneurals; 

11-13 precaudal and 13-16 caudal 

vertebrae; post-epiphysial fontanelle absent 

(Fig. 9b); lateral line complete, interrupted or 


Pop 

IO5 

IO4 

IO3 

So 

IO2 

IO1 

a b c d 

incomplete (usually incomplete), with 19-24 

scales on body in lateral series; and colour pattern 

consisting of a black blotch on caudal peduncle 

and frequently also other black blotches, spots or 

bars on side of body. 


and the data of Taki et al. (1978), Kullander & 

Fang (2005), Shantakumar & Vishwanath (2006), 

Kullander (2008) and Kullander & Britz (2008) we 

allocate the following 23 species to Pethia: P. atra 

(Linthoingambi & Vishwanath, 2007), P. bandula 

(Kottelat & Pethiyagoda, 1991), P. conchonius 

(Hamilton, 1822), P. cumingii (Günther, 1868), 

P. didi (Kullander & Fang, 2005), P. erythromycter 

(Kullander, 2008), P. gelius (Hamilton, 1822), 

P. khugae (Linthoingambi & Vishwanath, 2007), 

P. macrogramma (Kullander, 2008), P. manipurensis 

(Menon, Rema Devi & Viswanath, 2000), P. melanomaculata 

(Deraniyagala, 1956), P. nankyweensis 

(Kullander, 2008), P. nigripinnis Knight, Rema 

Devi & Arunachalam, 2012, P. nigrofasciata (Günther, 

1868), P. padamya (Kullander & Britz, 2008), 

P. phutunio (Hamilton, 1822), P. pookodensis (Mercy 

& Jacob, 2007), P. punctata (Day, 1865), P. reval 

(Meegaskumbura, Silva, Maduwage & Pethiyagoda, 

2008), P. shalynius (Yazdani & Talukdar, 

1975), P. stoliczkana (Day, 1871), P. thelys (Kullander, 

2008), P. tiantian (Kullander & Fang, 2005) 

and P. ticto (Hamilton, 1822). Based on characters 

provided in their original descriptions, and as 

pointed out by Kullander & Fang (2005), other 

species such as Puntius meingangbii Arunkumar 

& Tombi Singh, 2003, P. muvattupuzhaensis Jamee- 


F 

Pa 

So 

Sc 

CC 

PU3 

PU2 

Fig. 9. Pethia nigrofasciata, WHT 11022, 35.9 mm SL: a, circumorbital series; b, dorsal view of orbital region of 



Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; Sc, supraorbital sensory canal; So, supraorbital. 

81 

la Beevi & Ramachandran, 2005, P. narayani (Hora, 

1937), P. ornatus Vishwanath & Laisram, 2004, 

P. setnai Chhapgar & Sane, 1992, and P. yuensis 

Arunkumar & Tombi Singh, 2003 too, likely warrant 

placement in Pethia (though Puntius narayani 

differs from Puntius s. s. and Pethia by having 

9 branched dorsal-fin rays and 6 branched anal-fin 

rays: Hora, 1937). 

Etymology. Pethia is the generic vernacular name 

for small cyprinids in Sinhala; gender feminine. 

Discussion 

Ep 

Pls 

H6 

H5 

H4 

H3 

H2 

H1 

Ph 

The mitochondrial DNA phylogeny based on 

cyt-b and 16s gene fragments (Fig. 1) shows that 

there are five well supported clades that can be 

recognized as distinct genera also on the basis of 

shared external-morphological and osteological 

characters and character states within South Asian 

Puntius as previously understood. These generic 

allocations are also supported by (uncorrected) 

genetic distances in the range 7.3-18.1 % (Table 2). 

Except as mentioned below, we believe the present 

analysis to include all the major lineages represented 

by the South Asian fishes until now included 

in Puntius. We also comment below on 

several species with distinctive characters for 

which we do not have tissue for molecular 

analysis. By presenting a clearer definition of 

Puntius s. s., the present study also opens the way 

for the recognition of distinct genera for many of 

the Southeast Asian fishes hitherto referred to

82 

Puntius. Our results also corroborate the phylogenetic 

relationships recovered by Bossuyt et al. 

(2004) involving some of the same species and 

analysing the same ~ 590 bp segment of the 16S 

rRNA gene and a ~ 540 bp segment of the cyt-b 

gene. 

None of the genus-group names hitherto in 

the synonymy of Puntius have type species potentially 

referable to Puntius s. s., Systomus s. s., 

Dawkinsia, Dravidia or Pethia. The only contending 

genus-group names are Cephalakompsus Herre, 

1924a, Mandibularca Herre, 1924a, Ospatulus Herre, 

1924b, and Spratellicypris Herre & Myers, 1931, 

all of which have type species endemic to the 

Philippines. These possess both rostral and maxillary 

barbels, by which they are immediately 

distinguished from Puntius, Dawkinsia and Pethia 

(which lack rostral barbels). Puntius and Systomus 

are in any event not threatened by Cephalakompsus, 

Mandibularca, Ospatulus and Spratellicypris, which 

are junior. On the basis of the original descriptions 

of their type species in Herre (1924a), Dravidia 

differs from Cephalakompsus, Ospatulus and Spratellicypris 

by having a smooth (vs. serrated) last 

unbranched dorsal-fin ray; and from Mandibularca 

by its smaller size (up to 80 mm total length, 

vs. 220 mm in Mandibularca), having the lower 

jaw horizontal (vs. strongly curved upwards) and 

habitus (see Herre, 1924a: pl. 1). 

South Asian Puntius. The recognition of additional 

genera within Puntius as previously understood 

makes it imperative that the identity of 

P. sophore (Hamilton, 1822), the type species of 

Puntius, be stabilised. Hamilton’s description and 

figure of P. sophore do not allow this species to be 

unambiguously distinguished from other valid 

species, e. g., P. chola, P. dorsalis. No type specimens 

of fishes described by Hamilton have come 

to light in the 190 years since the publication of 

this work; nor has the existence of such types 

been alluded to in the subsequent literature. They 

must therefore be presumed lost. Hamilton (1822) 

did not specify the type locality of S. sophore in 

his account of that species, but it can be inferred 

from his preceding account of Puntius that it was 

Bengal. We selected ZRC 35064, a 71.8 mm SL 

specimen, as neotype because it was collected 

within the Ganges floodplain in Bengal (now 

Bangladesh); it is consistent with the original 

description (Hamilton, 1822: pl. 19, fig. 86); and 

it is accompanied by a series of five topotypes 

that facilitate a more representative description. 

The coloration of the neotype also matches 

that deliniated in a copy of Hamilton’s figure in 

the archives of the Linnean Society of London 

(Fig. 2b). This illustrates a specimen with 17 lateral-line 

scales, 2 unbranched and 8 branched 

dorsal-fin rays, 2 unbranched and 6 branched 

anal-fin rays, 1 + 7 + 7 + 1 principal caudal-fin rays, 

and 1 unbranched and 7 branched pelvic-fin rays. 

Although, as pointed out by Silva et al. (2010), 

Hamilton’s drawings contain many inaccuracies 

in their detail, the neotype matches Hamilton’s 

figure in general appearance. Based on the neotype 

and topotypes described herein, P. sophore 

may be distinguished from P. chola by the characters 

given in Talwar & Jhingran (1991) and 

Jayaram (2010), and from P. dorsalis by the characters 

given in Pethiyagoda et al. (2008). 

Because of a lack of material, we have been 

unable to assign several other Indian species with 

40 or more lateral-line scales, hitherto in Puntius, 

to any of the above genera: their generic placement 

must depend on future work. Perhaps most 

prominent among these are ‘P.’ deccanensis, ‘P.’ fraseri, 

and P. sharmai, which lack rostral barbels but 

possess a pair of maxillary barbels, a stiff, strongly 

serrated last unbranched dorsal-fin ray, and 

42-47 lateral-line scales, though the first species 

differs from the latter two by having 9 branched 

dorsal-fin rays and 7 branched anal-fin rays, vs. 

8 and 5 in Puntius (Hora & Misra, 1938; Yazdani 

& Babu Rao, 1976; Menon & Rema Devi, 1993). 

Also with a high lateral-line scale count (43) is 

Puntius punjaubensis (Day, 1871), which has no 

barbels and possesses a smooth last unbranched 

dorsal-fin ray: it is almost certainly not a member 

of any of the preceding genera. We note in passing 

that the original spelling ‘punjaubensis’ is not 

a lapsus for ‘punjabensis’, the spelling adopted in 

the current literature (e. g., Jayaram, 1991, 2010; 

Talwar & Jhingran, 1991): Day retained the 

original spelling also in his ‘Fishes of India’ (1878), 

‘Punjaub’ having been an accepted contemporaneous 

spelling. 

Another intriguing taxon is ‘Puntius’ ophicephalus, 

for which Menon (1999) created the 

genus Eechathalakenda. Possessing both maxillary 

and rostral barbels, a smooth last unbranched 

dorsal-fin ray and 40-42 lateral-line scales, this 

species does not fit within any of the preceding 

genera. Menon (1999) diagnosed Eechathalakenda 

from Gonoproktopterus by its “elongated Channa 

like appearance, 7 branched rays in the dorsal fin 

(9 in Gonoproktopterus), a slightly elongated but 


weak, smooth and segmented last undivided ray 

to the dorsal fin, four barbels and 40-42 scales 

along the lateral line”. We tentatively treat Eechathalakenda 

as valid. 

The relationships of ‘P.’ guganio (Hamilton, 

1822) and ‘P.’ nangalensis Jayaram, 1990 (barbels 

absent; last unbranched dorsal-fin ray stiff, serrated; 

lateral line incomplete, with 36-39 and 

31-33 scales respectively, in lateral series), remain 

to be elucidated. Jayaram (1990) considered these 

two species to be closely related. In the absence 

of material for comparison, we retain both species 

in ‘Puntius’ for the present, although they do not 

belong to this or any of the genera discussed 

herein. 

Unlike all the other species of Puntius examined 

in this study, P. bimaculatus (restricted to 

southern India and Sri Lanka) and P. titteya (a Sri 

Lankan endemic) lack a post-epiphysial fontanelle. 

These two species also group separately 

from all the other species of Puntius examined 

(Fig. 1) and share a suite of additional distinguishing 

characters: only 7 (vs. 8) branched dorsal-fin 

rays; 1 /2 3 (vs. 1 /2 4- 1 /2 5) scale rows between dorsalfin 

origin and lateral-line row; a lateral line that 

is in P. titteya incomplete and in P. bimaculatus 

occasionally interrupted or incomplete; and a 

coloration which includes a red lateral stripe – 

unusual in Puntius. Although they appear to 

warrant placement in a separate genus, we refrain 

from doing so as this would render Puntius 

paraphyletic (the two lower clades of Puntius in 

Fig. 1 evidently do not possess a suite of external 

characters that distinguish them). We feel this 

problem is best resolved once a wider sample of 

species is available for analysis. 

South Asian Systomus. Having long been treated 

as a synonym of Puntius (Bleeker, 1863b; Day, 

1878; Banarescu, 1997), Systomus was revived as 

a valid genus by Rainboth (1996), who assigned 

six Cambodian species previously in Puntius to 

it on the basis of the finely-serrated last unbranched 

dorsal-fin ray, possession of rostral and/ 

or maxillary barbels, and gill rakers fewer than 

12 on the first arch. Further species were added 

by subsequent authors (e. g. Ng & Tan, 1999; 

Parenti & Lim, 2005; Kottelat & Tan, 2011; see also 

Roberts & Catania, 2007). Given that some species 

of Systomus sensu Rainboth (1996) are popular 

aquarium fishes (e. g., ‘S.’ johorensis, ‘S.’ lateristriga, 

‘S.’ partipentazona), the genus-name is in 

wide use despite other authors (e. g., Kottelat, 


83 

2000, 2001; Kottelat & Widjanarti, 2005) retaining 

it as a junior synonym of Puntius, because removing 

these six species out of a large, hetero geneous 

genus without resolving the generic identity of 

the remaining species would likely cause confusion. 

This confusion is in part because the type 

species of Systomus, S. immaculatus, considered a 

synonym of S. sarana since Day (1878), is poorly 

known. M’Clelland (1839) characterized Systomus 

as follows: “Intermaxillaries protractile, dorsal 

and anal short, the former opposite to the ventrals 

and preceded by a spinous ray; body elevated 

and marked by two or more distinct dark spots, 

or diffuse spots either on the fins or opercula, 

prominence on the apex of the lower jaw obscure”. 

The first of the eight species he included in the 

first division of the genus (characterized by “depth 

of the body . . . equal to about half the length, and 

the spiny ray of the dorsal . . . serrated behind”) 

was S. immaculatus, subsequently designated as 

type species by Jordan (1919). This species he 

described as having “four cirrhi, a slight golden 

tinge on the opercula, fins dark, thirty-two scales 

on the lateral line and ten in an oblique line from 

the base of the ventrals to the dorsum”. He gave 

the type locality of S. immaculatus as “Small 

streams with sandy bottoms in Assam, and probably 

occasionally in the large rivers” adding that 

S. chrysosomus (a junior synonym of S. sarana) “is 

probably a variety only of this species, which is 

generally distributed throughout the rivers and 

ponds of all parts of India, sometime weighing 

as much as two pounds [0.9 kg]”. 

The type/s of S. immaculatus, however, are 

apparently lost. A search for possible types of this 

species in the collections of BMNH and SMF, the 

only repositories known to contain fish types of 

M’Clelland, proved fruitless (J. Maclaine, BMNH; 

and H. Zetzsche, SMF, pers. comms.). A search 

of the fish collections of ZSI too, failed to reveal 

any cyprinid specimens attributable to M’Clelland 

(RP, pers. obs.). Given the confusion that exists 

with regard to the identity of the genus, it is 

necessary that the identity of S. immaculatus, its 

type species, be clarified through the designation 

of a neotype: we here designate AMS B.7920 

(locality Assam, India), as neotype of Systomus 

immaculatus M’Clelland, 1839. The specimen is 

illustrated in Fig. 6 and its morphometric and 

meristic data given in Table 2, which supplement 

the description and diagnosis of M’Clelland 

(1839).

84 

M’Clelland’s (1839) description of S. immaculatus 

matches that of ‘Puntius’ sarana (type locality 

‘ponds and rivers of Bengal’) (Jayaram, 1991; 

Talwar & Jhingran, 1991). As shown here (Figs. 

1, 6) and corroborated also by Shantakumar & 

Vishwanath (2006), ‘P.’ sarana belongs to a lineage 

distinct from the one that includes Puntius sophore 

(the type species of Puntius): the former possess 

both rostral and maxillary barbels, a stiff, strongly 

serrated last unbranched dorsal-fin ray, a 

complete lateral line with 27-34 scales on the 

body, and a horizontally elongate oval black 

blotch on the caudal peduncle. This adds confidence 

to our selection of a neotype for S. immaculatus 

from among specimens nominally 

identified as ‘Puntius sarana’ from Assam. Systomus 

immaculatus (= S. sarana) may be distinguished 

from the other species here included in 

Systomus based on the characters provided above 

for the neotype and topotype, and those provided 

in the works cited in Remarks under the 

diagnosis of Systomus. 

Barbodes, Barbonymus and Poropuntius resemble 

Systomus in possessing two pairs of barbels 

and a serrated last unbranched dorsal-fin ray. 

Indeed, Menon (1999) placed S. sarana in Barbodes, 

the type species of which is Barbus maculatus 

Valenciennes, in Cuvier & Valenciennes 1842 (see 

Kottelat, 1999), which is considered a synonym 

of B. binotatus Valenciennes, in Cuvier & Valenciennes 

1842 (Kottelat, 1999; Pethiyagoda & Kottelat, 

2005b). Barbus binotatus, however, differs 

from species of Systomus by possessing 23-27 

lateral-line scales on body and ½4 scale rows 

between the dorsal-fin origin and lateral line (vs. 

32-33 lateral-line scales and 1 /2 5 scale rows between 

the dorsal-fin origin and lateral line in 

S. immaculatus, the type species of Systomus), and 

having a distinctive colour pattern (see Kottelat 

et al., 1993, pl. 15) not observed in any other species 

here allocated to Systomus s. s. Systomus differs 

from Barbonymus by having fewer than 6 rows of 

scales between the dorsal-fin origin and lateral 

line (7 or more in Barbonymus); and from Poropuntius 

by lacking an accessory pore on the canals 

of the lateral-line system (accessory pore present 

in Poropuntius: Kottelat et al., 1993; Roberts, 1998; 

Kottelat, 2001). 

Southeast Asian Puntius. The recent literature 

refers around 45 species of Southeast Asian 

cyprinids to Puntius. Only one of these, however, 

belongs to the genus as it is now understood: 

P. brevis (Bleeker, 1849). Widespread through 

Southeast Asia (Mekong basin of Laos, Thailand 

and Cambodia; Chao Phraya basin; Malay peninsula 

and Java: Kottelat, 2001) it has the last unbranched 

dorsal-fin ray smooth, a pair of maxillary 

(and no rostral) barbels, 23-26 lateral-line 

scales and a black spot on the caudal peduncle. 

Taki et al. (1978) included P. brevis in their P. binotatus 

group, though the former species lacks 

rostral barbels (present in P. binotatus) and has 

the last unbranched dorsal-fin ray smooth (vs. 

posteriorly serrated). 

Only one other Southeast Asian Puntius has 

the last unbranched dorsal-fin ray smooth, Puntius 

oligolepis (Bleeker, 1853). This species, however, 

is distinguished from Puntius s.s. by having an 

incomplete lateral line, only 17 scales in the lateral 

series, and 1 /2 3 scale rows between the dorsalfin 

origin and lateral-line scale row. Its relationships 

remain to be investigated. 

All the other Southeast Asian species presently 

referred to Puntius require new generic 

assignments. Although this is beyond the scope 

of the present work for want of material, for 

convenience of discussion and based on data on 

external characters available in the literature 

(Kottelat et al., 1993; Kottelat, 2001; Rainboth, 

1996; Roberts, 1989; Weber & de Beaufort, 1916; 

and the original species descriptions), we here 

refer these to a number of species groups (see also 

Southeast Asian Systomus, below). 

The ‘Puntius trifasciatus group’ comprises three 

species (P. gemellus Kottelat, 1996, P. johor ensis 

(Duncker, 1904), and P. trifasciatus Kottelat, 1996) 

distinguished by possessing both rostral and 

maxillary barbels, 25-27 scales in the lateral line 

on body, 7-11 gill rakers on the anterior gill arch, 

and a juvenile coloration consisting of 4-5 black 

bars on the body, changing to 3-6 black stripes 

on the body as adults (Kottelat, 1996); these characters 

combine to distinguish the members of this 

group from Puntius, Systomus, Pethia and Dravidia. 

The members of the P. trifasciatus group 

appear to belong to a lineage distinct also from 

the only other similarly-coloured species of Southeast 

Asian ‘Puntius’, P. lineatus (Duncker, 1904), 

which possess no or only a single pair of rudimentary 

barbels and the adults of which have the 

lower lip forming a complete post-labial groove, 

distinguishing it from all Puntius (in either its 

previous or present conception). Kottelat (1996) 

proposed that it be placed in a separate genus (for 

the present referred to as the ‘P. lineatus’ group). 


Southeast Asian Systomus. Of the five species 

referred to Systomus by Rainboth (1996) only 

S. orphoides appears to be a Systomus as the genus 

is now understood. The other four included species 

possess characters that suggest they belong 

to different lineages: ‘S.’ aurotaeniatus (Tirant 1885) 

lacks rostral barbels and has a coloration consisting 

of black spots at the dorsal-fin origin and on 

the side of the body; ‘S.’ johorensis (Duncker, 1904) 

lacks rostral barbels and has a striking colour 

pattern consisting of six black lateral bars; ‘S.’ partipentazona 

(Fowler, 1934a) lacks rostral barbels, 

has the lateral line incomplete and a colour pattern 

consisting of five black lateral bars including 

one across the dorsal fin; and ‘S.’ binotatus has a 

dark spot at the dorsal-fin origin, fewer scale rows 

on the body (23-27, vs. 27-34 in Systomus) and 

caudal peduncle (12 circumpedun cular scale 

rows, vs. 16) and a different head and mouth 

shape. Among Southeast Asian cyprinids, only 

S. orphoides and Barbus jacobusboehlkei Fowler, 1958 

appear to belong to Systomus s. s. We here place 

the remaining species referable to Systomus sensu 

Rainboth (1996) tentatively in two species groups 

that are, however, unlikely to be monophyletic 

(e. g., see Kottelat, 1992). 

Members of the ‘Puntius binotatus group’ possess 

both rostral and maxillary barbels, and a last 

unbranched dorsal-fin ray the posterior margin 

of which is more or less strongly serrated. Roberts 

(1989) regarded P. binotatus to be “the most 

widely distributed and perhaps most variable 

species of Puntius in southeast Asia”, noting that 

it “occurs from sea level to at least 2,000 m . . . and 

on small islands inhabited by few other freshwater 

fishes.” Close examination will likely show at 

least some of the 13 nominal species from time 

to time in its synonymy (see Roberts, 1989: 60-61) 

to be valid, closely-related species, as are P. banksi 

Herre, 1940 (see Kottelat & Lim, 1996: 232), and 

P. sealei (Herre, 1933) (see Inger & Chin, 1990: 73). 

Additionally, as pointed out by Roberts (1989), 

P. binotatus is regarded as the “parent stock” for 

the endemic ‘Puntius’ species of Lake Lanao and 

the Lanao Plateau of Mindanao in the Philippines: 

P. amarus (Herre, 1924a), P. disa (Herre, 1932), 

P. flavifuscus (Herre, 1924a), P. herrei (Fowler, 

1934b), P. joaquinae Wood, 1968, P. katolo (Herre, 

1924a), P. lanaoensis (Herre 1924a), P. lindog (Herre 

1924a), P. manalak (Herre, 1924a) and P. sirang 

(Herre, 1932). The Lanao ‘Puntius’ share these 

characters also with other ‘Puntius’ from Mindanao 

(P. cataractae (Fowler, 1934b), P. tumba 


85 

(Herre, 1924a) and P. umalii Wood, 1968), as well 

as those from Lake Manguao on Palawan (P. manguaoensis 

(Day, 1914) and P. bantolanensis (Day, 

1914)) and Mindoro (P. hemictenus (Jordan & Richardson, 

1908)). Ranging from about 6-24 cm 

and possessing 22-32 lateral-line scales most of 

these species show a coloration that includes a 

hazy dark stripe on the side of the body, in P. binotatus 

accompanied by a dark spot on the caudal 

peduncle and dorsal-fin origin, accompanied in 

some cases by numerous other dark spots and/ 

or blotches on the body (see Kottelat et al., 1993, 

pl. 15). ‘Systomus’ dunckeri (Ahl, 1929), ‘S.’ everetti 

(Boulenger, 1894), ‘S.’ ku chingensis (Herre, 1940), 

‘S.’ lateristriga (Valen ciennes, in Cuvier & Valenciennes, 

1842), ‘S.’ xou thos Kottelat & Tan, 2011, 

‘Puntius’ rhombeus Kottelat, 2001 and ‘P.’ semifasciolatus 

(Günther, 1868) too, are apparently members 

of this group. The genus name Barbodes is 

available for the ‘Puntius binotatus group’, with 

Cephalakompsus, Mandibularca, Ospatulus and 

Spratellicypris as synonyms. 

The remaining Southeast Asian ‘Systomus’ 

sensu Rainboth (1996) we place in the ‘Puntius 

tetrazona group’ because they possess a posteriorly 

serrated last unbranched dorsal-fin ray, 

maxillary barbels, and a striking coloration that 

includes 3-6 black bars and/or blotches on the 

body; the lateral line may be complete or incomplete, 

and rostral barbels present or absent. These 

species share a strikingly barred or blotched 

coloration, which resembles that of some species 

of Pethia (e. g. P. nigrofasciata) and Dravidia (e. g. 

D. fasciata), as does their small size. They differ 

from the former genus, however, in possessing a 

complete lateral line and at least maxillary barbels; 

and the latter in possessing a serrated posterior 

edge to the last unbranched dorsal-fin ray. Additionally, 

at least some of the included species 

have a slender infraorbital 3 (Taki et al., 1978), 

which serves to distinguish them from species of 

Pethia. The Puntius tetrazona group includes 

‘S.’ anchisporus (Vaillant, 1902), ‘S.’ endecanalis 

(Roberts, 1989), ‘S.’ foerschi (Kottelat, 1982), ‘S.’ hexazona 

(Weber & de Beaufort, 1912), ‘S.’ partipentazona, 

‘S.’ penta zona (Boulenger, 1894), ‘S.’ pulcher 

(Rendahl, 1922), ‘S.’ rhomboocellatus (Koumans, 

1940) and ‘S.’ te trazona (Bleeker, 1855). Kottelat’s 

(1992) observation that ‘tiger barbs’ belong to “at 

least two lineages which are not obviously related” 

is supported also by the data of Taki et al. 

(1978), whose ‘tetrazona group’ (‘S.’ tetrazona and 

‘S.’ partipentazona) lacks rostral barbels whereas

86 

a 

b 

Fig. 10. Barbus bovanicus, AMS B.7829, lectotype, 95.1 mm SL; India: Bhavani River, Tamil Nadu; a, photograph; 

b, radiograph. 

their ‘pentazona group’ (‘S.’ everetti, ‘S.’ lateristriga, 

‘S.’ pentazona and ‘S.’ rhomboocellatus) possesses 

them; Taki et al. (1978) also distinguished the two 

groups by their colour patterns. 

Dravidia. Menon (1999) chose the junior name 

Labeo melanampyx Day, 1865 over its senior subjective 

synonym, Cirrhinus fasciatus Jerdon, 1849 (the 

type species of Dravidia) citing Art. 23(b) of ICZN 

(1985), on the grounds that the former was “a 

favourite aquarium fish and several aquarium 

journals mention this species as Puntius melanampyx. 

A change is not considered desirable.” 

However, Art. 23(b) of ICZN (1985) requires that 

such a case is to be referred to the Commission 

for a ruling, which has not been done in the 13 

years following Menon (1999). As such, we follow 

Arts. 23.1 and 22.9.3 of ICZN (1999) and use the 

senior synonym C. fasciatus, which anyway is also 

the name in prevailing usage (e. g., Silas, 1956; 

Jayaram, 1991; Talwar & Jhingran, 1991; Arunachalam, 

2000; Bhat, 2005). 

Three additional South Asian species included 

in Puntius by previous authors (e. g., 

Jayaram, 1991; Talwar & Jhingran, 1991) bear two 

pairs of barbels and a smooth last unbranched 

dorsal-fin ray: ‘P.’ bovanicus, ‘P.’ carnaticus and 

‘P.’ jerdoni. Barbus bovanicus (AMS B.7829, syntype, 

95.1 mm SL, here designated lectotype; Fig. 10) 

has 9 branched dorsal-fin rays, 23 lateral-line 

scales on body, eye in upper half of head, thick, 

fleshy lips, and the lower lip present medially: it 

is apparently a Neolissochilus. It further differs 

from all species of Puntius, Systomus, Dawkinsia, 

Pethia and Dravidia in possessing 22 (vs. 13-19) 

caudal vertebrae and 6 (vs. 3-5) supraneurals. We 

note in passing that the spellings bovanicus and 

bovianicus were published simultaneously, in plate 

138 fig. 1, and on page 566, of Day (1878), both of 

which were released on 1 December 1878 (see 

Whitehead & Talwar, 1976: 54 and references 

therein). In keeping with subsequent usage, as 

first reviser we give B. bovanicus precedence over 

B. bovianicus. 


a 

b 

Fig. 11. Barbus jerdoni, AMS B.7935, syntype, 179.3 mm SL; India: Canara [Karnataka]. 

Pethiyagoda & Kottelat (2005b) noted that the 

figure of Barbus carnaticus in Day (1878: pl. 137) 

“possibly illustrates a species of Neolissochilus”. 

Jerdon’s (1849) description of B. carnaticus is 

uninformative, and there is no known surviving 

type material. As a result, subsequent authors 

have followed Day’s (1878: 563, pl. 137) conception 

of the species. Two of Day’s specimens in 

the AMS collection (B.7963, 103.3 mm SL; B.7626, 

230 mm SL) have 28 and 29 lateral-line scales on 

the body plus 2 on the base of the caudal fin; 

1 /2 5 + 1 + 4 scales in transverse line from dorsal-fin 

origin to pelvic-fin origin; 4 smooth, unbranched 

and 8 branched dorsal-fin rays; 3 unbranched and 

5 branched anal-fin rays; both rostral and maxillary 

barbels; and the anal fin, when adpressed, 

reaching just beyond the hypural fold. This is 

additionally a large cyprinid, reported to reach a 

total length of 60 cm (Talwar & Jhingran, 1991). 

It differs from Neolissochilus, however, by having 

the last unbranched dorsal-fin ray stiff (vs. weak 

and segmented: Rainboth, 1985) and the lower 

jaw exposed medially, which characters distin- 


87 

guish it also from Hypsibarbus and Gonoproktoperus, 

genera it otherwise resembles superficially. 

Its generic allocation remains uncertain. 

We hesitate to follow Menon (1999) in allocating 

Barbus jerdoni to Gonoproktopterus (for which 

he incorrectly used Hypselobarbus: see Jayaram, 

1997). Although the AMS syntype of G. jerdoni 

(B.7935, 179.3 mm SL; Fig. 11) has 9 branched 

dorsal-fin rays, the last unbranched ray weak, 

segmented; rostral and maxillary barbels; and 

6 + 1 + 3 scales in transverse line between dorsalfin 

origin and pelvic-fin origin, which are consistent 

with Gonoproktopterus as presently understood 

(Talwar & Jhingran, 1991; Jayaram, 1997), it possesses 

25 lateral-line scales, substantially lower 

than the 31-45 lateral-line scales observed in 

other species of this genus. Its generic placement 

too deserves further investigation. 

Dawkinsia. Arguing against the monophyly of 

the ‘filamentosus complex’ of Jayaram (1991), who 

did not point to any putative synapomorphies, 

Pethiyagoda & Kottelat (2005a) proposed the

88 

‘Puntius filamentosus group’ for those species of 

Puntius s. l., the juveniles of which possess a 

unique shared colour pattern of three black bars 

on the side. This was demonstrated first by De 

Silva et al. (1981) in a comparative study of the 

ontogeny of Dawkinsia singhala and Dravidia fasciata, 

since when the genus has expanded considerably 

(Pethiyagoda & Kottelat, 2005a; Rema Devi 

et al., 2010; Marcus Knight et al., 2011). Figure 1 

suggests that the general coloration of the sistergenera 

Dawkinsia, Dravidia and Pethia derive from 

a common ancestry which, taken together with 

apparent relationships within the P. tetrazona 

group and P. trifasciatus group, suggests that 

coloration could be a relatively conservative 

character useful in identifying relationships 

among these fishes (see also Kortmulder, 1986; 

Kortmulder & van der Poll, 1981). 

Pethia. Pethia (as the ‘Puntius conchonius group’) 

has been a group of fishes long recognized as 

having a separate identity within Puntius as previously 

understood. Following on the earlier work 

of Kortmulder (1972), and the osteological studies 

of Taki et al. (1978), Kullander & Fang (2005) 

described the ‘Puntius conchonius group’ as being 

characterized by small adult size, broad infraorbital 

3, absence of rostral barbels, minute or absent 

maxillary barbels, a stiff and serrated last unbranched 

dorsal fin ray, an often abbreviated 

lateral line, a colour pattern including a black 

blotch or spot anteriorly on the side (absent in 

P. conchonius) and a black blotch on the caudal 

peduncle, and a broad 5th ceratobranchial characterized 

by a pointed dorsal tip, spaced moderate-sized 

teeth, and absence of dorsal angle. Our 

results agree broadly with this characterization 

except that the 5th ceratobranchial is slender in 

P. punctata, and with an obtuse tip in P. nigrofasciata, 

P. reval and P. cumingii. 

Distribution. The present results suggest a 

greater degree of generic endemism in South 

Asian cyprinids than previously suspected. Several 

cyprinid genera are already known to be 

endemic to the peninsula of India, including 

Betadevario, Horalabiosa, Gonoproktopterus, Lepidopygopsis, 

Parapsilorhynchus and Rohtee. To these 

are now added Dawkinsia and Dravidia, which are 

restricted to Sri Lanka and the southern region 

of the Indian peninsula. Puntius s. s. too, turns out 

to be largely restricted to South Asia, only a single 

species, P. brevis, extending its range through 

Indochina to Java. Pethia too, appears mostly 

restricted to South Asia, with only P. stoliczkanus 

extending its range to Laos (Kottelat, 2001) and 

Thailand (Smith, 1945, as Puntius stoliczkae). Likewise, 

Systomus s. s. is represented in Southeast 

Asia only by S. jacobusboehlkei and S. orphoides. 

Character states. Some of the character states we 

report here are variable between closely-related 

species and even within species. Silva et al. (2008), 

for example, reported that one of the specimens 

of Puntius kamalika they stained possessed a free 

uroneural while another did not, and that free 

uroneurals were altogether absent in its sister 

species, P. mahecola. Similarly, while free uroneurals 

are present in each of the two specimens of 

P. sophore and P. chola cleared and stained by us, 

Shantakumar & Vishwanath (2006) reported them 

to be absent in both these species. Although we 

have included this character in the generic diagnoses 

above, it is important to note that it may 

be variable. 

While the deep infraorbital 3 is synapomorphic 

in Pethia, this character state occurs also in some 

species of Puntius as understood here, including 

P. bimaculatus, P. kelumi, P. mahecola and P. titteya. 

Similarly, character states that are synapomorphic 

in other cyprinine genera sometimes present 

themselves as apomorphies in individual species 

of Puntius as presently defined. For example, a 

few species retained in Puntius possess numerous 

rows of sensory papillae on the head, apparently 

an apomorphy shared by the closely-related 

species-group P. cauveriensis, P. dorsalis, P. kelumi 

and P. layardi (Pethiyagoda et al., 2008), but not 

other species of Puntius. Among Asian cyprinine 

genera, this character-state occurs also in Cyclocheilichthys, 

Eirmotus, Neobarynotus and Oreichthys 

(Kottelat, 1996; Tan & Kottelat, 2008; Schäfer, 

2009). Likewise, some species of Dawkinsia develop 

nuptial tubercles on the snout and cheeks 

(Pethiyagoda & Kottelat, 2005a), a character that 

occurs also in many other cyprinid genera. The 

dense tuberculation of the body in sexually mature 

males of Puntius kelumi, however (Pethiyagoda 

et al., 2008), while not reported from any other 

member of the Puntius, occurs in several cyprinid 

lineages. 

Nevertheless, the five genera into which the 

South Asian species previously in Puntius are 

now allocated are distinguishable from one another 

by reliable external characters: Puntius, 

smooth last unbranched dorsal-fin ray, no rostral 


arbels, no bars on flank in juveniles; Systomus, 

serrated last unbranched dorsal-fin ray, rostral 

barbels present; Dravidia, smooth last unbranched 

dorsal-fin ray, rostral barbels present; Dawkinsia, 

smooth last unbranched dorsal-fin ray, rostral 

barbels absent, black bars on flank in juveniles; 

Pethia, serrated last unbranched dorsal-fin ray, 

rostral barbels absent. All these species possess 

fewer than 40 lateral-line scales. 

A key to South Asian Puntius s. l. 

The following key allows generic allocation of 

South Asian fishes included in Puntius by Jayaram 

(1991, 2010), Talwar & Jhingran (1991), Menon 

(1999) and authors of new species from the region 

from 1990-2010. Generic names in quotes indicate 

provisional assignment pending future revision; 

such species are unlikely members of the stated 

genera. 

1. – 18-39 scales in lateral line (or if incomplete, 

in lateral series) on body. 

........................................................................2 

– 40-47 scales in lateral line on body. 

......................................................................10 

2. – Last unbranched dorsal-fin ray serrated 

posteriorly. 

........................................................................3 

– Last unbranched dorsal-fin ray smooth. 

........................................................................5 

3. – Lateral line complete; a black blotch on 

caudal peduncle; adults exceed 80 mm SL. 

.......................................................... Systomus 

– Lateral line abbreviated; maximum SL 

< 80 mm. 

........................................................................4 

4. – 19-24 scales in lateral series; one or more 

black spots, blotches or bars on body. 

................................................................Pethia 

– 31-39 scales in lateral series; no black markings 

on body. 

................‘Puntius’ guganio, ‘P.’ nangalensis 

5. – Juveniles (< 50 mm SL) with two or more 

black bars on body. 

........................................................................6 

– No black bars on body at any stage. 

........................................................................7 


89 

6. – Both rostral and maxillary barbels present. 

........................................................... Dravidia 

– Rostral barbels absent. 

.........................................................Dawkinsia 

7. – 9 branched dorsal-fin rays. 

........................................................................8 

– 7 or 8 branched dorsal-fin rays. 

........................................................................9 

8. – Last unbranched dorsal-fin ray stout, heavily 

ossified. 

...............................‘Neolissochilus’ bovanicus 

– Last unbranched dorsal-fin ray weak, segmented. 

..............................‘Gonoproktopterus’ jerdoni 

9. – Rostral barbels present; when adpressed, 

anal fin reaching beyond hypural notch. 

........................ ‘Gonoproktopterus’ carnaticus 

– Rostral barbels absent; when adpressed, 

anal fin falling short of hypural notch. 

.............................................................Puntius 

10. – Both rostral and maxillary barbels present. 

........................... Eechathalakenda ophicephala 

– Rostral barbels absent. 

......................................................................11 

11. – Last unbranched dorsal-fin ray serrated; 

maxillary barbels present. 

......................................................................12 

– Last unbranched dorsal-fin ray smooth; 

maxillary barbels absent. 

....................................‘Puntius’ punjaubensis 

12. – Lateral line complete; 9 branched dorsal-fin 

rays; 7 branched anal-fin rays. 

...................................... ‘Puntius’ deccanensis 

– Lateral line incomplete; 8 branched dorsalfin 

rays; 5 branched anal-fin rays. 

......................... ‘Puntius’ sharmai, ‘P.’ fraseri 

Material examined. Dawkinsia arulius, BMNH 1980.11. 

25:73-74, 2, 88.0-90.9 mm SL; BMNH 89.2.1.664-5, 2, 

67.1-67.6 mm SL; India: Tamil Nadu: Cauvery River at 

Coorg. 

Dawkinsia assimilis, WHT 6250, 5, topotypes, 43.3- 

53.2 mm SL; India: Karnataka: Netravati River, 42 km 

from Madikeri on Mangalore-Mysore Road. 

Dawkinsia exclamatio, WHT 6255, holotype, 70.4 mm 

SL; WHT 6249, 3 paratypes, 62.3-71.9 mm SL; WHT 

11056,1, 63.5 mm SL, c&s; WHT 11076, 1, 70.4 mm SL, 

c&s; India: Kerala: Kallada River at Thenmalai.

90 

Dawkinsia filamentosa: MNHN 3908, lectotype, 

82.7 mm SL; India: Kerala: Alleppey. – AMS B.7869, 

94.9 mm SL; India: Madras. – MNHN 3908, 2 paralectotypes, 

76.4-85.4 mm SL; India: Alleppey. – WHT 

11026, 1, 67.1 mm SL, c&s; WHT 11040, 1, 63.0 mm SL, 

c&s; India: Kerala: Chalakkudy. 

Dawkinsia singhala: ZMH 364, lectotype, 24.0 mm 

SL; ZMH 365, 2, paralectotypes, 54.1-63.3 mm SL; WHT 

6245, 4, 54.1-63.3 mm SL; Sri Lanka: Gin River at Wakwella. 

– WHT 11037, 1, 62.1 mm SL, c&s; WHT 11042, 

1, 55.0 mm SL, c&s; Sri Lanka: Morawaka. 

Dawkinsia srilankensis, BMNH 1976.2.10.2, holotype, 

79.0 mm SL; BMNH 1976.2.10.2, 1, paratype, 63.3 mm 

SL; WHT 11053, 1, 61.0 mm SL, c&s; WHT 11063, 1, 

60.7 mm SL, c&s; Sri Lanka; 3 miles N of Pallegama. 

Dawkinsia tambraparniei, ZSI 735, 1, paratype, 

68.7 mm SL, India: Tamil Nadu: tributary of Tambraparni 

River 7 km from Kalladi Kuruchi; ZSI-SRS 4452, 

2, 52.9-64.6 mm SL; India: Tamil Nadu: Tirunelveli. 

Dravidia fasciata, WHT 11043, 1, 43 mm SL, c&s; WHT 

11034, 1, 39.1 mm SL, c&s; India: Kerala: Chalakudy. 

Gonoproktopterus carnaticus AMS B.7963, 1, 103.3 mm 

SL; AMS B.7626, 1, 225 mm SL; India: Bhavani River, 

Tamil Nadu, F. Day. 

Gonoproktopterus jerdoni, AMS B.7935, 1, syntype, 

179.3 mm SL, India: Canara [Karnatake], F. Day. 

Gonoproktopterus kolus AMS B.7518, 1, 153.9 mm SL, 

India: Deccan, F. Day. 

Gonoproktopterus lithopidos AMS B.8374, 1, syntype, 

282 mm SL, India: Canara, F. Day. 

Neolissochilus bovanicus, AMS B.7829, lectotype, 

95.1 mm SL, India: Bhavani River, Tamil Nadu, F. Day. 

Neolissochilus wynaadensis AMS B.7989, 1, syntype, 

137.1 mm SL,India: Wynaad, F. Day. 

Pethia bandula, ZRC 38483, holotype, 34.8 mm SL; 

CMK 7146, 7, paratypes, 29.6-38.5 mm SL; Sri Lanka: 

Galapitamada. 

Pethia conchonius, AMS IA.6883, 41.8 mm SL; aquarium 

specimen. – AMS IA.6518, 46.5 mm SL; aquarium 

specimen. 

Pethia cumingii, BMNH 1859.10.19.101, lectotype, 

33.4 mm SL; BMNH 1859.5.31.47-48, 3, paralectotypes; 

Sri Lanka. – WHT 7685, 1, 37.0 mm SL, c&s; WHT 11047, 

1, 35.6 mm SL, c&s; Sri Lanka: Mawanana. 

Pethia melanomaculata, WHT 1681, 2, 37.0-38.7 mm 

SL; WHT 218, 2, 30.6-31.8 mm SL; WHT 11066, 1, 

62.8 mm SL, c&s; WHT 11068, 1, 37.8 mm SL, c&s; Sri 

Lanka: Mahawelia basin: Hasalaka. 

Pethia nigrofasciata, BMNH 1859.10.19.99-100, 2, 

syntypes, 32.1-45.5 mm SL; Sri Lanka. – WHT 7677, 3, 

41.0-44.4 mm SL; Kelani River at Kitulgala. – WHT 

11022, 1, 35.9 mm SL, c&s; WHT 11023, 1, 35.6 mm SL, 

c&s; Sri Lanka, Mawanana. 

Pethia punctata, WHT 7716, 3, 35.1-37.0 mm SL; 

WHT 11071, 1, 29.3 mm SL, c&s; WHT 11077, 1, 28.7 mm 

SL, c&s; India: Kerala: Calicut. – WHT 7715, 4, 29.7- 

34.2 mm SL; India: Kerala: Thenmalai. 

Pethia reval, WHT 677, holotype, 30.5 mm SL; WHT 

7536, 3, 28.8-36.6 mm SL; Sri Lanka: Kelani Basin: 

Labugama. – WHT 11033, 1, 25.6 mm SL, c&s; WHT 

11044, 1, 28.3 mm SL, c&s; Sri Lanka: Ingiriya. 

Pethia stoliczkana, AMS B.7542, 1, 34.7 mm SL; India: 

West Bengal: Darjeeling. 

Pethia ticto, WHT 7718, 5, 30.8-34.6 mm SL; India: 

West Bengal: Nadia. – AMS B.7974, 2, 49.5-58.6 mm SL; 

India: Ganjam. – WHT 11054, 1, 37.0 mm SL, c&s; WHT 

11065, 1, 37.4 mm SL, c&s; India: W Bengal: Nadia. 

Puntius amphibius, MNHN 73, lectotype, 92.5 mm 

SL; MNHN 2005-0006, 1, paralectotype, 74.3 mm SL; 

India: Maharashtra: Bombay. 

Puntius bimaculatus, WHT 79, 2, 37.0-46.2 mm SL; 

Sri Lanka: Pallegama. – WHT 1065, 8, 40.2-46.0; Sri 

Lanka: Mee Oya [River]. 

Puntius brevis, AMS I.43487-006, 43.2 mm SL; Cambodia: 

Tonle Sap River, 3-5 km downstream of Kompong 

Chhnang. 

Puntius chola,AMS B.7887, 89.3 mm SL; India: Madras. 

Puntius everetti, AMS I.30463-001, 3, 32.5-39.4 mm 

SL; Malaya. 

Puntius denisonii, AMS B.7913, 1, 88 mm SL; India: 

Travancore Hills. 

Puntius dorsalis, WHT 2153, 4, 68.4-115.3 mm SL; 

India: Tamil Nadu: Mamallapuram. – AMS B.7634, 

112.4 mm SL; India: Madras. – WHT 1900, 6, 93.3-133.5; 

Sri Lanka: Minneriya. – WHT 7646, 2, 56.7-65.0 mm SL, 

c&s; Sri Lanka: Tissamaharamaya. 

Puntius kamalika, WHT 7639, holotype, 50.2 mm SL; 

WHT 7640, 11, paratypes, 39.3-55.4 mm SL; Sri Lanka, 

Kalu River at Walandure near Kuruwita. 

Puntius kelumi, WHT 7629, holotype, 70.4 mm; WHT 

7630, 4, 57.2-68.6 mm SL; Sri Lanka: Kalu River: 

Madakada Aranya, near Ingiriya. – WHT 7645, 2, 55.1- 

63.2 mm SL, c&s; Sri Lanka: Homadola. 

Puntius layardi: BMNH 1853.12.27.5, syntypes, 2, 

92.5-112.7 mm SL; Sri Lanka. 

Puntius mahecola, MNHN 3896, lectotype, 50.7 mm 

SL; MNHN 2005-0007, paralectotypes, 5, 48.7-56.5 mm 

SL; India: Kerala: Mahe. – WHT 7681, 2, 56.8-66.8 mm 

SL, c&s; India: Kerala: Kottayam. 

Puntius partipentazona, AMS I.43747-007, 32.1 mm 

SL; Cambodia: Tonle Sap River. 

Puntius sealei, AMS I.22131-006, 9, 45.8-62.4 mm SL; 

Northwest Borneo: Kembayung River. 

Puntius sophore, ZRC 35064, neotype, 71.8 mm SL; 

ZRC 35065-35069, 5, 59.4-80.6 mm SL; Bangladesh: 

Srimangal, from Hail Hoar floodplain near Moulvi 

Bazaar. – WHT 11060, 1, 36.6 mm SL, c&s; WHT 11062, 

1, 28.9 mm SL, c&s; India: West Bengal: Boncron. 

Puntius thermalis, WHT 158, 4, 74.6-76.0 mm SL; 

WHT 11027, 1, 55.8 mm SL, c&s; WHT 11035, 1, 48.9 mm 

SL, c&s; Sri Lanka: Attidiya. 

Puntius titteya, WHT 8013, 1, 32.1 mm SL, c&s; WHT 

8014, 1, 44.2 mm SL, c&s; Sri Lanka: Kelani River near 

Ginigathena. 

Puntius vittatus, WHT 11036, 1, 27.5 mm SL, c&s; 

WHT 11038, 1, 28.5 mm SL, c&s; Sri Lanka: Navinna. 

Systomus immaculatus, AMS B.7920, neotype, 


186.9 mm SL; AMS B.7921, 1, topotype, 192.7 mm SL; 

India: Assam. 

Systomus martenstyni, ZRC 38482, holotype, 

136.0 mm SL; CMK 7168, 11, 67.2-116.8 mm SL; Sri 

Lanka: 2 km N of Pallegama. 

Systomus orphoides, AMS I.43487-062, 2, 106-136 mm 

SL; Cambodia: Tonle Sap River, 3-5 km downstream 

of Kompong Chhnang. 

Systomus pleurotaenia, WHT 11031, 1, 66.2 mm SL, 

c&s; Sri Lanka: Akuressa. 

Systomus sp. ‘Richmondi’ [manuscript name for an 

as yet undescribed species], WHT 9861, 6, 121.5- 

127.0 mm SL; Sri Lanka: Elahera. 

Systomus spilurus, WHT 11028, 1, 61.1 mm SL; Sri 

Lanka, Kuruwita. 

Systomus timbiri, NMSL FF801, holotype, 178 mm 

SL; Sri Lanka: Timbirigasmankada. – WHT 11107, 5, 

127-200 mm SL; Sri Lanka: Walawe River. 

Acknowledgements 

We are grateful to the Department of Wildlife Conservation, 

Sri Lanka, for permitting export of tissue samples 

for molecular analysis; Kelum Manamendra-Arachchi 

and Mohomed Bahir for assisting with the collection of 

material in Sri Lanka and India; Andrew Rao for supplying 

samples from West Bengal; and Sudath Nanayakkara 

for hospitality at the WHT field station at 

Agarapatana, Sri Lanka. MM thanks Christopher 

S. Schnei der for access to laboratory facilities at Boston 

University, and RP thanks Mark McGrouther and 

Amanda Hay (AMS) for laboratory facilities and permission 

to access material in his care and Kelvin Lim (ZRC) 

for the loan of material; James Maclaine (BMNH) and 

Horst Zetzsche (SMF) for information on types; Gina 

Douglas (Linnean Society of London) for permission to 

reproduce Fig. 2b; Johnny Jensen for the photograph of 

D. fasciata in Fig. 1; and Maurice Kottelat, Kevin Conway, 

Shrimati K. Rema Devi and Sri W. Vishwanath for 

discussion and literature. Finally, it is a pleasure to 

thank Maurice Kottelat and an anonymous reviewer 

for valuable comments and criticism that helped substantially 

to improve the manuscript. 

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Received 14 April 2011 

Revised 4 March 2012 

Accepted 6 May 2012

96

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