A synopsis of the South Asian fishes referred to Puntius - Verlag Dr ...
A synopsis of the South Asian fishes referred to Puntius - Verlag Dr ...
A synopsis of the South Asian fishes referred to Puntius - Verlag Dr ...
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Ichthyol. Explor. Freshwaters, Vol. 23, No. 1, pp. 69-95, 11 figs., 4 tabs., June 2012<br />
© 2012 by <strong>Verlag</strong> <strong>Dr</strong>. Friedrich Pfeil, München, Germany – ISSN 0936-9902<br />
A <strong>synopsis</strong> <strong>of</strong> <strong>the</strong> <strong>South</strong> <strong>Asian</strong> <strong>fishes</strong> <strong>referred</strong> <strong>to</strong> <strong>Puntius</strong><br />
(Pisces: Cyprinidae)<br />
Rohan Pethiyagoda*, Madhava Meegaskumbura** and Kalana Maduwage***<br />
The tropical <strong>Asian</strong> cyprinid genus <strong>Puntius</strong>, which contains some 120 valid species, has long been suspected <strong>to</strong> be<br />
polyphyletic. Here, through an examination <strong>of</strong> external morphology, osteology, and analysis <strong>of</strong> 16S ribosomal<br />
RNA and cy<strong>to</strong>chrome b gene fragments from 31 <strong>South</strong> <strong>Asian</strong> species hi<strong>the</strong>r<strong>to</strong> <strong>referred</strong> <strong>to</strong> <strong>Puntius</strong>, we show that<br />
<strong>the</strong>se <strong>fishes</strong> represent at least five lineages recognisable as genera. <strong>Puntius</strong> sensu stric<strong>to</strong> has <strong>the</strong> rostral barbels<br />
absent; last unbranched dorsal-fin ray weak or strong, smooth; and lateral line complete, with 22-28 pored scales.<br />
Sys<strong>to</strong>mus possesses maxillary and rostral barbels; last unbranched dorsal-fin ray stiff (‘osseous’), serrated; and<br />
lateral line complete, with 27-34 scales. Three new genera are proposed: Dawkinsia (type species Leuciscus filamen<strong>to</strong>sus)<br />
is distinguished by lacking rostral barbels; having <strong>the</strong> last unbranched dorsal-fin ray smooth; lateral<br />
line complete, with 18-22 scales; and a juvenile colour pattern that includes three black bars on <strong>the</strong> body. <strong>Dr</strong>avidia<br />
(type species Cirrhinus fasciatus) is distinguished by having both rostral and maxillary barbels present; lateral<br />
line complete, with 18–26 pored scales; dorsal fin with 4 unbranched and 8 branched rays, last unbranched<br />
dorsal-fin ray smooth; infraorbital 3 deep, partly overlapping <strong>the</strong> preoperculum; and free uroneural and postepiphysial<br />
fontanelle absent. Pethia (type species Barbus nigr<strong>of</strong>asciatus) is distinguished by having <strong>the</strong> last unbranched<br />
dorsal-fin ray stiff, serrated; infraorbital 3 deep, partially overlapping preoperculum; rostral barbels absent;<br />
maxillary barbels absent or minute; a black blotch on <strong>the</strong> caudal peduncle; and frequently, black blotches, spots<br />
or bars on <strong>the</strong> side <strong>of</strong> <strong>the</strong> body. The identities <strong>of</strong> <strong>Puntius</strong> sophore and Sys<strong>to</strong>mus immaculatus are clarified through<br />
<strong>the</strong> designation <strong>of</strong> neotypes; a lec<strong>to</strong>type is designated for Neolissochilus bovanicus; and precedence is given <strong>to</strong> <strong>the</strong><br />
spelling bovanicus over bovianicus.<br />
Introduction<br />
<strong>Puntius</strong> has long been recognized as a “catch-all”<br />
genus for a variety <strong>of</strong> small tropical <strong>Asian</strong><br />
cyprinids whose interrelationships are poorly<br />
unders<strong>to</strong>od (Kortmulder, 1972; Schut et al., 1984;<br />
Kottelat & Pethiyagoda, 1989; Kullander & Fang,<br />
2005; Kullander, 2008). The genus, which contains<br />
some 220 nominal species (about 120 <strong>of</strong> <strong>the</strong>se<br />
* Ichthyology Section, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia.<br />
E-mail: rohanpet@gmail.com<br />
** Department <strong>of</strong> Zoology, University <strong>of</strong> Peradeniya, Peradeniya, Sri Lanka. E-mail: madhava_m@mac.com<br />
*** Faculty <strong>of</strong> Medicine, University <strong>of</strong> Peradeniya, Peradeniya, Sri Lanka. E-mail: kalanapm@gmail.com<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
69<br />
considered valid), has seen a significant accretion<br />
<strong>of</strong> new species in recent years. The striking coloration<br />
<strong>of</strong> many species <strong>of</strong> <strong>Puntius</strong> has also led <strong>to</strong><br />
<strong>the</strong>ir popularity as freshwater-aquarium <strong>fishes</strong>;<br />
many species are traded internationally as ornamentals<br />
(Collins et al., 2012). As <strong>the</strong> genus has<br />
hi<strong>the</strong>r<strong>to</strong> been unders<strong>to</strong>od (e. g., Menon, 1999;<br />
Jayaram, 2010), species <strong>of</strong> <strong>Puntius</strong> range widely<br />
in size, from about 20 <strong>to</strong> 500 mm SL; possess 18-
70<br />
47 lateral-line scales; a complete, incomplete or<br />
interrupted lateral line; none, one or two pairs <strong>of</strong><br />
barbels; a weak or strong, serrated or smooth last<br />
unbranched dorsal-fin ray; and a body coloration<br />
composed <strong>of</strong> a spot or blotch at <strong>the</strong> base <strong>of</strong> <strong>the</strong><br />
caudal fin, one or more blotches or bars on <strong>the</strong><br />
body, one or more lateral stripes, or none at all<br />
(Jayaram, 1991). The genus comprised 48 valid<br />
species in <strong>the</strong> <strong>South</strong> <strong>Asian</strong> region as at 1990 (Talwar<br />
& Jhingran, 1991), since when some 30 additional<br />
species have been described.<br />
The first experimental investigation <strong>of</strong> species<br />
groups within <strong>Puntius</strong> appears <strong>to</strong> have been by<br />
Kortmulder (1972). As a result <strong>of</strong> similarities<br />
between <strong>the</strong>ir larvae and juveniles, and through<br />
a number <strong>of</strong> hybridization experiments, he suggested<br />
close relationships between P. filamen<strong>to</strong>sus<br />
and P. arulius; and between P. s<strong>to</strong>liczkanus, P. conchonius,<br />
P. phutunio, P. cumingii and P. nigr<strong>of</strong>asciatus,<br />
concluding that “cumingii, phutunio and gelius<br />
are more or less neotenous descendants from a<br />
nigr<strong>of</strong>asciatus like ances<strong>to</strong>r.” A single experiment<br />
he performed <strong>to</strong> hybridize P. conchonius and<br />
P. filamen<strong>to</strong>sus, however, produced larvae that<br />
died shortly after hatching as a result <strong>of</strong> being<br />
unable <strong>to</strong> aerate <strong>the</strong>ir swim bladders, suggestive<br />
<strong>of</strong> <strong>the</strong>ir belonging <strong>to</strong> different sub-groups within<br />
<strong>Puntius</strong>.<br />
Following on this Taki et al. (1978), from an<br />
osteological study <strong>of</strong> 23 mostly Sou<strong>the</strong>ast <strong>Asian</strong><br />
species <strong>of</strong> <strong>Puntius</strong> s.l., proposed six groups, <strong>of</strong><br />
which two (<strong>the</strong>ir P. conchonius group and <strong>the</strong><br />
P. arulius group) comprised <strong>South</strong> <strong>Asian</strong> species.<br />
In a later study, Shantakumar & Vishwanath<br />
(2006) included also P. sophore, <strong>the</strong> type species<br />
<strong>of</strong> <strong>Puntius</strong>, and showed that this species, <strong>to</strong>ge<strong>the</strong>r<br />
with P. terio and P. chola, possesses a postepiphysial<br />
fontanelle and a smooth, weak last<br />
unbranched dorsal-fin ray. Both <strong>the</strong>se characters<br />
were absent in <strong>the</strong> o<strong>the</strong>r nine species <strong>of</strong> <strong>Puntius</strong><br />
s.l. <strong>the</strong>y examined, which included two members<br />
<strong>of</strong> <strong>the</strong> P. conchonius group <strong>of</strong> Taki et al. (1978).<br />
They also proposed a group not sampled by <strong>the</strong><br />
previous authors, comprising P. sarana, P. jayarami<br />
and P. orphoides, which differed from <strong>Puntius</strong><br />
s. s. and <strong>the</strong> P. conchonius group by possessing<br />
35-36 (vs. 29-33) vertebrae and a free uroneural<br />
(absent in <strong>Puntius</strong> s. s. and <strong>the</strong> P. conchonius<br />
group).<br />
The foregoing work suggests that <strong>Puntius</strong> s. l.<br />
comprises a number <strong>of</strong> species groups that share<br />
various combinations <strong>of</strong> apomorphic characters<br />
and character states. The vast range <strong>of</strong> <strong>the</strong> genus<br />
(all <strong>of</strong> tropical <strong>South</strong> and Sou<strong>the</strong>ast Asia), and <strong>the</strong><br />
large number <strong>of</strong> included species, has made <strong>the</strong><br />
resolution <strong>of</strong> all <strong>the</strong> species groups <strong>of</strong> which<br />
<strong>Puntius</strong> is presently composed a challenge. The<br />
conservative body plan <strong>of</strong> <strong>the</strong> included species<br />
<strong>to</strong>o, has resulted in analyses <strong>of</strong> external morphology<br />
alone (e. g. Pillay, 1951; Jayaram, 1991; De<br />
Silva & Liyanage, 2010) proving <strong>to</strong> be <strong>of</strong> limited<br />
value in identifying species groups. At <strong>the</strong> same<br />
time, given <strong>the</strong> steady accretion <strong>of</strong> new species<br />
<strong>to</strong> <strong>Puntius</strong> s. l., especially from <strong>South</strong> Asia, an<br />
improved understanding <strong>of</strong> <strong>the</strong> species-groups<br />
that comprise it is desirable. Here, based on an<br />
analysis <strong>of</strong> external morphology and osteology,<br />
and a molecular phylogeny constructed from a<br />
mi<strong>to</strong>chondrial 16S ribosomal RNA gene fragment<br />
(~ 540 bp) and a cy<strong>to</strong>chrome b gene fragment<br />
(~ 550 bp) including 30 putative species <strong>of</strong> Indian<br />
and Sri Lankan <strong>Puntius</strong> s. l., we show that at least<br />
five lineages are present within <strong>South</strong> <strong>Asian</strong><br />
<strong>Puntius</strong>, which we recognize as distinct genera.<br />
Material and methods<br />
Specimens <strong>referred</strong> <strong>to</strong> in this study are deposited<br />
in <strong>the</strong> Australian Museum, Sydney (AMS); <strong>the</strong><br />
Natural His<strong>to</strong>ry Museum, London (BMNH); <strong>the</strong><br />
Collection <strong>of</strong> Maurice Kottelat, Cornol (CMK);<br />
<strong>the</strong> Raffles Museum <strong>of</strong> Biodiversity Research,<br />
Singapore (ZRC); <strong>the</strong> collection <strong>of</strong> <strong>the</strong> Wildlife<br />
Heritage Trust <strong>of</strong> Sri Lanka (WHT), now deposited<br />
in <strong>the</strong> National Museum <strong>of</strong> Sri Lanka, Colombo<br />
(NMSL); and <strong>the</strong> Zoological Survey <strong>of</strong><br />
India, Kolkata (ZSI). Abbreviations: SL, standard<br />
length; HL, head length.<br />
Measurements were made with digital Vernier<br />
calipers and recorded <strong>to</strong> <strong>the</strong> nearest 0.05 mm.<br />
<strong>Dr</strong>awings were made using a Motic dissecting<br />
microscope fitted with a camera lucida. Methods<br />
for taking counts and measurements follow<br />
Pethiyagoda & Kottelat (2005a), except that scales<br />
in transverse line on body were counted downwards<br />
and backwards from dorsal-fin origin <strong>to</strong><br />
<strong>the</strong> pelvic-fin origin. Osteological descriptions are<br />
based on alizarin-stained specimens (2 from each<br />
species, following <strong>the</strong> method <strong>of</strong> Taylor & Van<br />
Dyke, 1985) and/or x-ray radiographs. Osteological<br />
nomenclature follows Conway (2011).<br />
Numbers in (square) brackets following a count<br />
indicate <strong>the</strong> frequency <strong>of</strong> that count.<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
‘<strong>South</strong> Asia’ as used here is <strong>the</strong> political unit<br />
previously <strong>referred</strong> <strong>to</strong> as British India, bounded<br />
approximately by Pakistan, Myanmar and Sri<br />
Lanka. ‘Sou<strong>the</strong>ast Asia’ as used here is tropical<br />
continental Asia east <strong>of</strong> <strong>South</strong> Asia.<br />
DNA analysis. DNA was extracted from ethanolpreserved<br />
tissues extracted from 39 species (see<br />
Table 1) using Qiagen tissue extraction kits and<br />
manufacturer’s pro<strong>to</strong>cols. Mi<strong>to</strong>chondrial 16S ri-<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
71<br />
bosomal RNA gene and cy<strong>to</strong>chrome-b gene fragments<br />
were amplified using primers16Sar and<br />
16Sbr (Palumbi, 1996) which amplified ~540 bp<br />
<strong>of</strong> <strong>the</strong> 16S rRNA gene and CB-J-10933 (5'-TATGT-<br />
TCTACCATGAGGACAAATATC-3'), and BSF4<br />
(5'-CTTCTACTGGTTGTCCTCCGATTCA-3')<br />
(Bossuyt & Milinkovitch, 2000), which amplified<br />
~ 550 bp <strong>of</strong> cy<strong>to</strong>chrome-b gene.<br />
PCR conditions were as follows: denaturation<br />
at 95 °C for 40 s, annealing at 45 °C for 40 s and<br />
Table 1. GenBank accession numbers, species and locations <strong>of</strong> <strong>the</strong> 39 species <strong>of</strong> Cyprinidae included in <strong>the</strong> molecular<br />
analysis.<br />
reference no. Genbank accession no. species location country<br />
16s cytb<br />
WHT8818_9 JF793558 JF793592 Garra ceylonensis Homadola Sri Lanka<br />
WHT8819_31i JF793559 JF793593 Garra mullya Chalakudy India<br />
WHT8820_80 JF793560 JF793594 Labeo dussumieri Elahera Sri Lanka<br />
WHT8821_23i JF793561 JF793595 Labeo dussumieri Alleppey India<br />
WHT8822_29i JF793562 JF793596 Osteochilichthys nashii Chalakudy India<br />
WHT8834_28i JF793574 JF793608 Gonoprok<strong>to</strong>pterus jerdoni Srirangapatam India<br />
WHT8851_1i JF793591 JF793625 Gonoprok<strong>to</strong>pterus curmuca Chalakudy India<br />
WHT8848_85 JF793588 JF793622 Tor khudree Mawanana Sri Lanka<br />
WHT8849_AQ1 JF793589 JF793623 Barbonymus schwanenfeldii Aquar. spec. –<br />
WHT8823_59 JF793563 JF793597 Pethia bandula Galapitamada Sri Lanka<br />
WHT8838_64 JF793578 JF793612 Pethia nigr<strong>of</strong>asciata Mawanana Sri Lanka<br />
WHT8812_1 EU604682.1 EU604674.1 Pethia reval Kelani River Sri Lanka<br />
WHT8814_46 EU604684.1 EU604676.1 Pethia cumingii Ben<strong>to</strong>ta Sri Lanka<br />
WHT8816_75 EU604686.1 EU604678.1 Pethia melanomaculata Kandalama Sri Lanka<br />
WHT8850_AQ4 JF793590 JF793624 Pethia conchonius Aquar. spec. –<br />
WHT8815_6i EU604687.1 EU604679.1 Pethia tic<strong>to</strong> Boncron India<br />
WHT8832_20i JF793572 JF793606 <strong>Dr</strong>avidia fasciata Chalakudy India<br />
WHT8852_40i JF796852 Dawkinsia assimilis Chalakudy India<br />
WHT8833_27i JF793573 JF793607 Dawkinsia filamen<strong>to</strong>sa Kottayam India<br />
WHT8843_33 JF793583 JF793617 Dawkinsia singhala Menik River Sri Lanka<br />
WHT8844_19 JF793584 JF793618 Dawkinsia srilankensis Pallegama Sri Lanka<br />
WHT8824_48 JF793564 JF793598 <strong>Puntius</strong> bimaculatus Ben<strong>to</strong>ta Sri Lanka<br />
WHT8825_88 JF793565 JF793599 <strong>Puntius</strong> cf. bimaculatus Minneriya Sri Lanka<br />
WHT8846_11 JF793586 JF793620 <strong>Puntius</strong> titteya Kalu River Sri Lanka<br />
WHT8847_43 JF793587 JF793621 <strong>Puntius</strong> cf. titteya Ben<strong>to</strong>ta Sri Lanka<br />
WHT8826_54 JF793566 JF793600 <strong>Puntius</strong> <strong>the</strong>rmalis Mawanana Sri Lanka<br />
WHT8827_24i JF793567 JF793601 <strong>Puntius</strong> chola Boncron India<br />
WHT8845_217ai JF793585 JF793619 <strong>Puntius</strong> sophore Boncron India<br />
WHT8828_40 JF793568 JF793602 <strong>Puntius</strong> layardi Walawe River Sri Lanka<br />
WHT8829_69 JF793569 JF793603 <strong>Puntius</strong> dorsalis Gin River Sri Lanka<br />
WHT8830_70 JF793570 JF793604 <strong>Puntius</strong> kelumi Gin River Sri Lanka<br />
WHT8831_35i JF793571 JF793605 <strong>Puntius</strong> dorsalis Mamallapuram Sri Lanka<br />
WHT8837_16i JF793577 JF793611 <strong>Puntius</strong> mahecola Kottayam India<br />
WHT8835_21 JF793575 JF793609 Sys<strong>to</strong>mus martenstyni Pallegama Sri Lanka<br />
WHT8826_76 JF793576 JF793610 Sys<strong>to</strong>mus sp. “Richmondi” Elahera Sri Lanka<br />
WHT8842_21i JF793582 JF793616 Sys<strong>to</strong>mus sarana Boncron India<br />
WHT8840_35 JF793580 JF793614 Sys<strong>to</strong>mus timbiri Menik River Sri Lanka<br />
WHT8841_45 JF793581 JF793615 Sys<strong>to</strong>mus spilurus Ben<strong>to</strong>ta Sri Lanka<br />
WHT8839_12 JF793579 JF793613 Sys<strong>to</strong>mus pleurotaenia Gin River Sri Lanka
72<br />
extension at 72 °C for 40 s, 35 cycles, with a final<br />
extension <strong>of</strong> 72 °C for 5 min for <strong>the</strong> 16s gene fragment;<br />
denaturation at 95 °C for 40 s, annealing at<br />
45 °C for 50 s and extension at 72 °C for 60 s, 35<br />
cycles, with a final extension <strong>of</strong> 72 °C for 5 min<br />
for <strong>the</strong> cy<strong>to</strong>chrome-b gene fragment. Products<br />
were gel purified and sequenced on an ABI 377<br />
or ABI 3100 au<strong>to</strong>mated sequencer following<br />
manufacturer’s pro<strong>to</strong>cols.<br />
The 16s sequences were aligned using Clustal<br />
X (Jeanmougin et al., 1998) and adjusted by eye<br />
using Se-Al (ver. 2.0a9; Rambaut, 1996). Positions<br />
which were difficult <strong>to</strong> align and in which we had<br />
low confidence in positional homology were<br />
excluded from subsequent analyses, leaving a<br />
<strong>to</strong>tal 411 bp. Cy<strong>to</strong>chrome-b sequences were<br />
aligned using translated amino acid sequences<br />
using Se-Al (ver. 2.0a9; Rambaut, 1996).<br />
Two Garra species (G. ceylonensis and G. mullya)<br />
were used as an outgroup for rooting <strong>the</strong> tree:<br />
<strong>the</strong> data <strong>of</strong> Yang et al. (2010) and Zheng et al.<br />
(2010) suggest that Garra is a closely-related basal<br />
genus in relation <strong>to</strong> most taxa included in this<br />
analysis. The data were analyzed using Bayesian<br />
and Maximum Parsimony (MP) criteria. We used<br />
Bayesian inference as implemented in MrBayes<br />
(Huelsenbeck & Ronquist, 2001) <strong>to</strong> generate a<br />
phylogenetic hypo<strong>the</strong>sis <strong>of</strong> relationships among<br />
<strong>the</strong> taxa and <strong>to</strong> estimate a general time-reversible<br />
model <strong>of</strong> sequence evolution with gamma-distributed<br />
rate variation among sites and a proportion<br />
<strong>of</strong> invariant sites (GTR+I+G). We ran four<br />
Metropolis-Coupled Markov Chain Monte Carlo<br />
(MCMCMC) chains for 2 000 000 generations. The<br />
summed likelihood <strong>of</strong> <strong>the</strong> four chains converged<br />
on a stationary value by 50 000 generations (<strong>the</strong><br />
burn-in time). We used <strong>the</strong> frequencies <strong>of</strong> clades<br />
in trees that were sampled every ten generations<br />
from <strong>the</strong> last 250 000 generations as estimates <strong>of</strong><br />
<strong>the</strong> posterior probabilities <strong>of</strong> those clades (Huelsenbeck<br />
et al., 2001). Uniform priors were used<br />
throughout and branch lengths, <strong>to</strong>pology, and<br />
nucleotide substitution parameters were unconstrained.<br />
For tree searches under a Maximum<br />
Parsimony criterion (all characters unordered and<br />
weighted equally) we used heuristic searches with<br />
TBR branch-swapping and random taxon addition<br />
as implemented in PAUP*4.0b10. A single tree<br />
(tree score 1919 steps) was recorded. A bootstrap<br />
analysis <strong>to</strong> determine node support was also carried<br />
out within a maximum parsimony framework.<br />
The molecular dataset used for this study<br />
can be accessed at: http://purl.org/phylo/treebase/phylows/study/TB2:S12407<br />
and http://<br />
web.mac.com/madhavameegaskumbura.<br />
Results<br />
The original dataset comprised <strong>of</strong> a 552-bp fragment<br />
<strong>of</strong> cyt-b and a 537-bp fragment <strong>of</strong> 16s. After<br />
removing poorly aligned regions <strong>of</strong> <strong>the</strong> 16s fragment,<br />
a 1060-bp fragment remained, which was<br />
used in <strong>the</strong> analyses.<br />
From our Bayesian analysis, we chose <strong>the</strong> tree<br />
having <strong>the</strong> highest likelihood value as our best<br />
tree (Fig. 1). Maximum Parsimony analysis yielded<br />
a single tree (tree length = 1919; not shown<br />
because <strong>the</strong> tree <strong>to</strong>pology from this analysis was<br />
very similar <strong>to</strong> that <strong>of</strong> <strong>the</strong> Bayesian analysis).<br />
The Bayesian analysis recovered five wellsupported<br />
clades within <strong>Puntius</strong> s. l., that we<br />
consider <strong>to</strong> be distinct genera on <strong>the</strong> basis <strong>of</strong><br />
external-morphological and osteological characters<br />
(see Table 2 for genetic distances and below<br />
for diagnoses): <strong>Puntius</strong> s. s., Sys<strong>to</strong>mus, and three<br />
new genera, Dawkinsia, <strong>Dr</strong>avidia and Pethia. The<br />
phylogeny shows that (1) Pethia is a sister group<br />
<strong>of</strong> <strong>Dr</strong>avidia and Dawkinsia; (2) Sys<strong>to</strong>mus is <strong>the</strong><br />
sister group <strong>of</strong> <strong>Puntius</strong>; and (3) [<strong>Puntius</strong> + Sys<strong>to</strong>mus]<br />
is sister <strong>to</strong> [Dawkinsia + <strong>Dr</strong>avidia + Pethia].<br />
Table 2. Percent uncorrected genetic distances between Dawkinsia, <strong>Dr</strong>avidia, Pethia, <strong>Puntius</strong> and Sys<strong>to</strong>mus for <strong>the</strong><br />
combined 16s and cyt-b fragments analysed.<br />
Pethia Dawkinsia <strong>Dr</strong>avidia Sys<strong>to</strong>mus <strong>Puntius</strong><br />
Pethia – 7.3-12.0 13.5-14.2 10.4-15.6 10.4-16.3<br />
Dawkinsia – 12.2-15.6 8.1-15.6 8.4-16.6<br />
<strong>Dr</strong>avidia – 12.7-15.6 13.0-16.6<br />
Sys<strong>to</strong>mus – 9.5-18.1<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
<strong>Puntius</strong> Hamil<strong>to</strong>n, 1822<br />
Type species. Cyprinus sophore Hamil<strong>to</strong>n, 1822<br />
(Fig. 2).<br />
Diagnosis. A genus <strong>of</strong> cyprinid <strong>fishes</strong> differing<br />
from all <strong>South</strong> and Sou<strong>the</strong>ast <strong>Asian</strong> genera <strong>of</strong><br />
Cyprininae by <strong>the</strong> combination <strong>of</strong> <strong>the</strong> following<br />
characters and character states: adult size usually<br />
less than 120 mm SL; maxillary barbels absent<br />
or present, rostral barbels absent; dorsal fin with<br />
3-4 unbranched and 8 branched rays; anal fin<br />
with 3 unbranched and 5 branched rays; last<br />
unbranched dorsal-fin ray weak or strong, apically<br />
segmented, not serrated (Fig. 3c); lateral line<br />
complete, with 22-28 pored scales on <strong>the</strong> body;<br />
free uroneural present (Fig. 3d); gill rakers simple,<br />
acuminate (not branched or laminate); no antrorse<br />
predorsal spinous ray; a post-epiphysial fontanelle<br />
present (Fig. 3b; but see remarks on P. bimaculatus<br />
and P. titteya, below); supraneurals 4;<br />
infraorbital 3 slender (Fig. 3a); 5th cera<strong>to</strong>branchial<br />
narrow; pharyngeal teeth 5 + 3 + 2; 12-14 abdominal<br />
and 14-16 caudal vertebrae; and a colour<br />
pattern that includes a (sometimes faint) black<br />
spot or blotch on <strong>the</strong> caudal peduncle.<br />
Remarks. Based on <strong>the</strong> comparisons made herein<br />
(Fig. 1) and <strong>the</strong> data <strong>of</strong> Shantakumar & Vishwanath<br />
(2006) and Kullander (2008), we include<br />
<strong>the</strong> following <strong>South</strong> <strong>Asian</strong> species in <strong>Puntius</strong>:<br />
P. chola (Hamil<strong>to</strong>n, 1822), P. dorsalis (Jerdon, 1849),<br />
P. kelumi Pethiyagoda, Silva, Maduwage & Meegaskumbura,<br />
2008, P. layardi (Gün<strong>the</strong>r, 1868),<br />
P. mahecola (Valenciennes, in Cuvier & Valenciennes,<br />
1844), P. pugio Kullander, 2008, P. sophore<br />
(Hamil<strong>to</strong>n, 1822), P. terio (Hamil<strong>to</strong>n, 1822) and<br />
P. <strong>the</strong>rmalis (Valenciennes, in Cuvier & Valenciennes,<br />
1844).<br />
Because <strong>the</strong>y share <strong>the</strong> external characters<br />
diagnostic <strong>of</strong> <strong>Puntius</strong>, we tentatively retain <strong>the</strong><br />
following <strong>South</strong> <strong>Asian</strong> species in that genus:<br />
P. amphibius (Valenciennes, in Cuvier & Valenciennes<br />
1842), P. arenatus (Day, 1878), P. burmanicus<br />
(Day, 1878), P. cauveriensis (Hora, 1937),<br />
P. crescentus Yazdani & Singh, 1994, P. kamalika<br />
Silva, Maduwage & Pethiyagoda, 2008, P. madhusoodani<br />
Krishnakumar, Pereira & Radhakrishnan,<br />
2011, P. melanostigma (Day, 1878), P. mudumalaiensis<br />
Menon & Rema Devi, 1992, P. muzaffarpurensis<br />
Srivastava, Verma & Sharma, 1977, P. narayani<br />
(Hora, 1937), P. parrah (Day, 1865), P. sahyadriensis<br />
Silas, 1953, P. vittatus (Day, 1865) and P. waa<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
73<br />
geni (Day, 1872). We also tentatively retain in<br />
<strong>Puntius</strong> P. denisonii (Day, 1865) and P. chalakkudiensis<br />
Menon, Rema Devi & Thobias, 1999, for<br />
which we lacked material for DNA and osteological<br />
analysis: <strong>the</strong>y possess a strikingly different<br />
coloration and mouth shape <strong>to</strong> all o<strong>the</strong>r congeners<br />
(Menon et al., 1999) and are likely <strong>to</strong> warrant<br />
placement in a separate genus in <strong>the</strong> future.<br />
Two species, P. bimaculatus (Bleeker, 1863a)<br />
and P. titteya Deraniyagala, 1929, that differ from<br />
all <strong>the</strong> o<strong>the</strong>r <strong>Puntius</strong> examined by lacking a postepiphysial<br />
fontanelle (Fig. 4b) and possessing only<br />
7 branched dorsal-fin rays, are never<strong>the</strong>less retained<br />
in <strong>Puntius</strong> (see Discussion).<br />
<strong>Puntius</strong> sophore (Hamil<strong>to</strong>n, 1822)<br />
(Fig. 2)<br />
Cyprinus sophore Hamil<strong>to</strong>n, 1822: 310, 389, pl. 19.<br />
Neotype. ZRC 35064, 71.8 mm SL; Bangladesh:<br />
Srimangal [= Sreemangal] <strong>to</strong>wn market, from Hail<br />
Hoar floodplain near Moulvi Bazaar; D. Hoggarth,<br />
Mar 1993.<br />
Topotypes. ZRC 35065-35069, 5, 59.4-80.6 mm<br />
SL, same data as neotype.<br />
Description <strong>of</strong> neotype. General body shape and<br />
appearance are illustrated in Fig. 2a. Morphometric<br />
data <strong>of</strong> neotype and 5 <strong>to</strong>potypes are given in<br />
Table 3; characters <strong>of</strong> <strong>to</strong>potypes, where different<br />
from neotype, are given in paren<strong>the</strong>ses. Body<br />
s<strong>to</strong>ut, standard length about 2.4 times maximum<br />
depth, laterally compressed. Dorsal pr<strong>of</strong>ile <strong>of</strong><br />
head concave above eye; dorsal pr<strong>of</strong>ile <strong>of</strong> body<br />
convex, rising steeply from occipital, almost<br />
horizontal at dorsal-fin origin. Ventral con<strong>to</strong>ur <strong>of</strong><br />
body convex <strong>to</strong> anal-fin insertion, almost horizontal<br />
at pelvic-fin origin. Dorsal and ventral<br />
pr<strong>of</strong>iles <strong>of</strong> caudal peduncle concave <strong>to</strong> terminal<br />
scales on tail base. Caudal peduncle length 1.3<br />
(1.2-1.4) times its depth.<br />
Head short, its length about one-fifth standard<br />
length, somewhat compressed laterally. Eyes<br />
located forward <strong>of</strong> centre, superiorly on head,<br />
just visible in ventral and dorsal views. Nares<br />
dorsolaterally orientated. Mouth U-shaped in<br />
ventral aspect, subterminal, small, angle <strong>of</strong> gape<br />
not reaching below anterior nares in lateral aspect.<br />
Lips fleshy, entire, adnate <strong>to</strong> jaws. Upper lip<br />
overhanging lower one. Barbels absent.
74<br />
100<br />
100<br />
74<br />
100<br />
71<br />
10 changes<br />
100<br />
100<br />
91<br />
92<br />
Garra ceylonensis (LK)<br />
Garra mullya (IN)<br />
Labeo dussumieri (LK)<br />
Labeo dussumieri (IN)<br />
100<br />
83<br />
91<br />
100<br />
Osteochilichthys nashii (IN)<br />
Gonoprok<strong>to</strong>pterus jerdoni (IN)<br />
Gonoprok<strong>to</strong>pterus curmuca (IN)<br />
Tor khudree (LK)<br />
Barbonymus schwarnenfeldii (AQ)<br />
92<br />
92<br />
92<br />
99<br />
50<br />
100<br />
87<br />
99<br />
59<br />
100<br />
100<br />
100<br />
57<br />
100<br />
92<br />
96<br />
87<br />
100 Pethia bandula (LK)<br />
99<br />
98 Pethia nigr<strong>of</strong>asciata (LK)<br />
100<br />
98<br />
93<br />
100<br />
85<br />
Pethia reval (LK)<br />
Pethia cumingii (LK)<br />
Pethia melanomaculata (LK)<br />
Pethia conchonius (AQ)<br />
Pethia tic<strong>to</strong> (IN)<br />
<strong>Dr</strong>avidia fasciata (IN)<br />
Dawkinsia assimilis (IN)<br />
Dawkinsia filamen<strong>to</strong>sa (IN)<br />
Dawkinsia singhala (LK)<br />
Dawkinsia srilankensis (LK)<br />
60<br />
99<br />
50<br />
64<br />
64<br />
100<br />
84<br />
100<br />
96<br />
100<br />
64<br />
100<br />
100<br />
100<br />
100<br />
<strong>Puntius</strong> mahecola (IN)<br />
100<br />
66<br />
<strong>Puntius</strong> bimaculatus (LK)<br />
<strong>Puntius</strong> cf. bimaculatus (LK)<br />
<strong>Puntius</strong> cf. titteya (LK)<br />
<strong>Puntius</strong> titteya (LK)<br />
<strong>Puntius</strong> <strong>the</strong>rmalis (LK)<br />
<strong>Puntius</strong> chola (IN)<br />
<strong>Puntius</strong> sophore (IN)<br />
<strong>Puntius</strong> dorsalis (LK)<br />
100<br />
65<br />
<strong>Puntius</strong> dorsalis (IN)<br />
<strong>Puntius</strong> kelumi (LK)<br />
<strong>Puntius</strong> layardi (LK)<br />
Sys<strong>to</strong>mus martenstyni (LK)<br />
Sys<strong>to</strong>mus sp. ‘Richmondi’ (LK)<br />
Sys<strong>to</strong>mus sarana (IN)<br />
100<br />
73<br />
100<br />
100<br />
Sys<strong>to</strong>mus timbiri (LK)<br />
Sys<strong>to</strong>mus spilurus (LK)<br />
Sys<strong>to</strong>mus pleurotaenia (LK)<br />
Outgroup<br />
Last unbranched dorsal-fin<br />
ray serrated; infraorbital 3<br />
deep; SL < 80 mm; rostral<br />
barbels absent; post-epiphysial<br />
fontanelle absent.<br />
Last unbranched dorsal-fin<br />
ray smooth; SL < 80 mm;<br />
rostral, maxillary barbels<br />
present; post-epiphysial<br />
fontanelle absent.<br />
Last unbranched dorsal-fin<br />
ray smooth; rostral barbels<br />
absent; 3 black bars on flank<br />
in juveniles; post-epiphysial<br />
fontanelle absent.<br />
Last unbranched dorsal-fin<br />
ray smooth; 7 branched<br />
dorsal-fin rays; rostral<br />
barbels absent; postepiphysial<br />
fontanelle absent.<br />
Last unbranched dorsal-fin<br />
ray smooth; 8 branched<br />
dorsal-fin rays; rostral barbels<br />
absent; post-epiphysial<br />
fontanelle present.<br />
Last unbranched dorsal-fin<br />
ray serrated; rostral, maxillary<br />
barbels present; SL ><br />
80 mm; 27−34 lateral-line<br />
scales; post-epiphysial<br />
fontanelle absent.<br />
Fig. 1. Bayesian phylogram <strong>of</strong> <strong>the</strong> 31 putative species <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong> s. l. analysed, with key shared<br />
morphological and osteological characters for each group. See Table 1 for list <strong>of</strong> material. Provenance: AQ,<br />
aquarium trade; IN, India; LK, Sri Lanka. Posterior probabilities are shown above nodes, parsimony bootstrap<br />
values below nodes. Pho<strong>to</strong>graphs illustrate species within <strong>the</strong> respective genera: from <strong>to</strong>p, Pethia bandula, <strong>Dr</strong>avidia<br />
fasciata, Dawkinsia srilankensis, <strong>Puntius</strong> titteya, <strong>Puntius</strong> dorsalis and Sys<strong>to</strong>mus asoka.<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
Pelvic-fin origin slightly anterior <strong>to</strong> dorsal-fin<br />
origin. Posterior margin <strong>of</strong> dorsal fin slightly<br />
concave, that <strong>of</strong> anal fin straight. Distal margins<br />
<strong>of</strong> pelvic and pec<strong>to</strong>ral fins convex, rounded.<br />
Dorsal-fin with 4 unbranched rays (last one<br />
smooth, strong) and 8 branched rays, its origin<br />
above ninth lateral-line scale. Anal fin with 3<br />
unbranched and 5 branched rays. Pelvic fin with<br />
1 unbranched and 8 branched rays, not reaching<br />
anus. Pec<strong>to</strong>ral fin with one unbranched and 15<br />
branched rays, not reaching pelvic-fin origin.<br />
Caudal fin deeply forked, its lobes pointed, with<br />
1 + 9 + 8 + 1 principal rays.<br />
Lateral-line pored scales on body 24 (23 [2]),<br />
plus 1 on caudal-fin base; 4 1 /2 scales in transverse<br />
line between lateral line and origin <strong>of</strong> dorsal fin,<br />
4 (3 1 /2 [1]) between lateral line and origin <strong>of</strong> pelvic<br />
fin; 1 /2 2/1/2 1 /2 scales in transverse line on caudal<br />
peduncle; 9 predorsal scales (excluding notched<br />
scale at dorsal-fin origin). An axillary scale approximately<br />
one-third length <strong>of</strong> pelvic fin extends<br />
backwards from pelvic-fin origin. Dorsal-fin base<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
a<br />
b<br />
75<br />
Fig. 2. Cyprinus sophore: a, ZRC 35064, neotype, 71.8 mm<br />
SL; Bangladesh: Hail Hoar floodplain; b, copy <strong>of</strong> Hamil<strong>to</strong>n’s<br />
drawing, in <strong>the</strong> collection <strong>of</strong> <strong>the</strong> Linnean Society<br />
<strong>of</strong> London (courtesy <strong>of</strong> Linnean Society <strong>of</strong> London).<br />
Table 3. Standard length (in millimetres) and proportional measurements as a percentage <strong>of</strong> standard length or<br />
head length <strong>of</strong> <strong>the</strong> neotype (ZRC 35064, 71.8 mm SL) and 5 <strong>to</strong>potypes (ZRC 35065-35069, 59.4-80.6 mm SL) <strong>of</strong><br />
<strong>Puntius</strong> sophore.<br />
neotype range<br />
min. max.<br />
Standard length (mm) 71.8 59.4 80.6<br />
In percent SL<br />
Total length 130 127 133<br />
Head length 28.6 27.3 29.7<br />
Predorsal length 52.8 52.7 56.1<br />
Preanal length 76.9 71.2 76.9<br />
Prepelvic length 50.3 43.6 50.5<br />
Caudal-peduncle length 19.8 18.0 20.5<br />
Caudal-peduncle depth 14.9 13.8 15.2<br />
Body depth 42.5 40.2 42.5<br />
Dorsal-fin height 27.3 23.8 30.8<br />
Length <strong>of</strong> stiff portion <strong>of</strong> last unbranched dorsal-fin ray 17.5 17.1 19.9<br />
Pec<strong>to</strong>ral-fin length 18.9 18.4 23.1<br />
Pelvic-fin length 20.6 19.0 22.6<br />
Distance from dorsal-fin origin <strong>to</strong> hypural fold 54.5 54.5 58.2<br />
Post-dorsal distance 36.4 36.4 39.8<br />
Maximum body width 17.3 14.0 17.3<br />
Length <strong>of</strong> dorsal-fin base 18.2 16.1 18.6<br />
Length <strong>of</strong> anal-fin base 13.9 11.4 14.7<br />
In percent HL<br />
Snout length 27 27 30<br />
Post-orbital head length 50 49 53<br />
Eye diameter 32 27 32<br />
Internarial width 26 22 27<br />
Interorbital width 42 38 42
76<br />
IO5<br />
Pop<br />
IO4<br />
a b c d<br />
IO5<br />
IO4<br />
IO3<br />
IO3<br />
So<br />
So<br />
IO2<br />
IO2<br />
IO1<br />
IO1<br />
Pa<br />
a<br />
Pop<br />
b c d<br />
Pef<br />
shea<strong>the</strong>d by 8 (7 [2]) scales, anal-fin base by<br />
6 (7 [3]) scales.<br />
In preservative (Fig. 2a), head and body<br />
brownish olive, lighter below lateral line; lower<br />
body, belly and chest pale cream-yellow. Fin rays<br />
light brown, interradial membrane hyaline. A distinct,<br />
horizontally-elongate black blotch slightly<br />
larger than eye, about 3 scales wide by 2 scales<br />
high, at posterior end <strong>of</strong> caudal peduncle. A smaller<br />
black blotch at base <strong>of</strong> dorsal-fin rays 5-7.<br />
F<br />
F<br />
So<br />
PU3 PU2 CC<br />
PU3 PU2 CC<br />
Fig. 4. <strong>Puntius</strong> bimaculatus, WHT 11029, 40.5 mm SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural;<br />
Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; supraorbital sensory<br />
canal not shown.<br />
So<br />
Sc<br />
Pa<br />
PU3 PU2 CC<br />
PU3 PU2 CC<br />
Fig. 3. <strong>Puntius</strong> sophore, WHT11062, 28.9 mm SL; SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Pef, postepiphysial<br />
fontanelle; Ph, parhypural; Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3;<br />
Sc, supraorbital sensory canal; So, supraorbital; Un, free uroneural.<br />
Ep<br />
Pls<br />
Sys<strong>to</strong>mus McClelland, 1838<br />
Un<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
H1<br />
Ph<br />
Ep<br />
Pls<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
Type species. Sys<strong>to</strong>mus immaculatus M’Clelland,<br />
1839.<br />
H1<br />
Ph<br />
Diagnosis. A genus <strong>of</strong> cyprinid <strong>fishes</strong> differing<br />
from all <strong>South</strong> and Sou<strong>the</strong>ast <strong>Asian</strong> genera <strong>of</strong><br />
Cyprininae by <strong>the</strong> combination <strong>of</strong> <strong>the</strong> following<br />
characters and character states: adult size greater<br />
than 80 mm SL (in <strong>South</strong> <strong>Asian</strong> species); maxillary<br />
and rostral barbels present; dorsal fin with 4 + 8<br />
rays, its last unbranched ray stiff, strongly serrated<br />
(Fig. 5c); lateral line complete, with 27-34<br />
scales on body; gill rakers simple, acuminate (not<br />
branched or laminate); no antrorse predorsal<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
IO5<br />
IO4<br />
Pop<br />
So<br />
IO3 IO2<br />
IO1<br />
a b c d<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
F<br />
Pa<br />
So<br />
Pls<br />
PU3 PU2 CC<br />
PU3 PU2 CC<br />
Fig. 5. Sys<strong>to</strong>mus spilurus, WHT 11028, 61.1 mm SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural;<br />
Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; Un, free uroneural;<br />
supraorbital sensory canal not shown.<br />
spinous ray; free uroneural present (Fig. 5d);<br />
post-epiphysial fontanelle absent (Fig. 5b); 5 supraneurals;<br />
infraorbital 3 slender (Fig. 5a); 14-15<br />
abdominal and 17-19 caudal vertebrae; and colour<br />
pattern including a prominent, horizontallyelongate<br />
oval black blotch on <strong>the</strong> caudal peduncle,<br />
with no prominent bars or blotches on <strong>the</strong><br />
body.<br />
Remarks. Based on <strong>the</strong> comparisons made herein,<br />
and <strong>the</strong> data <strong>of</strong> Shantakumar & Vishwanath<br />
(2006) and Kottelat & Pethiyagoda (1989), we<br />
include <strong>the</strong> following nine <strong>South</strong> <strong>Asian</strong> species<br />
in Sys<strong>to</strong>mus: S. asoka (Kottelat & Pethiyagoda,<br />
1989), S. compressiformis (Cockerell, 1913), S. jayarami<br />
(Vishwanath & Tombi Singh, 1986), S. martenstyni<br />
(Kottelat & Pethiyagoda, 1991), S. orphoides<br />
Valenciennes, in Cuvier & Valenciennes 1842,<br />
S. pleurotaenia (Bleeker, 1863a), S. sarana (Hamil<strong>to</strong>n,<br />
1822), S. spilurus (Gün<strong>the</strong>r, 1868) and S. timbiri<br />
(Deraniyagala, 1963). An apparently undescribed<br />
Sri Lankan species with <strong>the</strong> label-name<br />
‘<strong>Puntius</strong> sp. Richmondi’ <strong>to</strong>o, belongs <strong>to</strong> Sys<strong>to</strong>mus.<br />
Barbus clavatus McClelland, 1844 and B. shanensis<br />
Hora & Mukerji, 1934 have been variously placed<br />
in <strong>Puntius</strong> (e. g., Jayaram, 1991; Talwar & Jhingran,<br />
1991) and Poropuntius (e. g., Roberts, 1998; Jayaram,<br />
2010) by previous authors. We have not<br />
examined specimens <strong>of</strong> <strong>the</strong>se and so refer <strong>the</strong>m<br />
only tentatively <strong>to</strong> Sys<strong>to</strong>mus, though <strong>the</strong>y may<br />
bear a closer relationship <strong>to</strong> <strong>the</strong> ‘<strong>Puntius</strong> binotatus<br />
group’ <strong>of</strong> Sou<strong>the</strong>ast Asia (see Discussion).<br />
Sys<strong>to</strong>mus immaculatus<br />
(Fig. 6)<br />
77<br />
Ep<br />
Un<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
H1<br />
Ph<br />
Sys<strong>to</strong>mus immaculatus M’Clelland, 1839: 284, 380,<br />
pl. 44, fig. 5.<br />
Neotype. AMS B.7920, 187 mm SL; India: Assam;<br />
F. Day.<br />
Topotype. AMS B.7921, 193 mm SL (caudal fin<br />
broken); India: Assam; F. Day.<br />
Description <strong>of</strong> neotype. General body shape and<br />
appearance are illustrated in Figure 6a. Morphometric<br />
data <strong>of</strong> neotype and 1 <strong>to</strong>potype are given<br />
in Table 4; characters <strong>of</strong> <strong>to</strong>potype, where different<br />
from neotype, are given in paren<strong>the</strong>ses. Body<br />
elongate, standard length about 2.8 times body<br />
depth. Dorsal pr<strong>of</strong>ile <strong>of</strong> head rounded; dorsal<br />
body pr<strong>of</strong>ile rising gently <strong>to</strong> dorsal-fin origin<br />
(a post-nuchal hump present), sloping gently<br />
downwards <strong>the</strong>reafter. Dorsal-fin origin slightly<br />
posterior <strong>to</strong> pelvic-fin origin. Ventral body pr<strong>of</strong>ile<br />
curving gently ventrad <strong>to</strong> pelvic-fin base, almost<br />
level from <strong>the</strong>re <strong>to</strong> anal-fin origin, rising steeply<br />
<strong>to</strong> caudal peduncle; caudal peduncle ventral<br />
margin level.<br />
Head relatively long, its length about onefourth<br />
standard length. Eyes located forward <strong>of</strong><br />
centre, about mid-height on head. Mouth terminal;<br />
lips fleshy; lower-lip post-labial groove shallow,<br />
uninterrupted. Cheek and interorbital with<br />
about 10 sensory pores. Two pairs <strong>of</strong> barbels<br />
present; maxillary barbel long, its length slightly
78<br />
less than eye diameter; rostral barbel shorter,<br />
about half length <strong>of</strong> maxillary barbel.<br />
Pelvic-fin origin slightly anterior <strong>to</strong> dorsal-fin<br />
origin. Posterior margins <strong>of</strong> dorsal and anal fins<br />
concave. Distal margins <strong>of</strong> pec<strong>to</strong>ral and pelvic<br />
fins convex, rounded. Dorsal-fin origin above 11th<br />
lateral-line scale. Dorsal fin with 4 unbranched<br />
and 8 branched rays, last unbranched ray stiff,<br />
strongly serrated posteriorly in its proximal twothirds,<br />
weak, segmented in its distal one-third.<br />
Last unbranched dorsal-fin ray longest, when<br />
adpressed not overlapping tip <strong>of</strong> last branched<br />
dorsal-fin ray. Anal fin with 3 unbranched and 5<br />
branched rays. Pelvic fin with 1 unbranched and<br />
8 branched rays. Pec<strong>to</strong>ral fin with 1 unbranched<br />
and 15 branched rays. Caudal fin forked, lobes<br />
<strong>of</strong> equal length, longest rays about twice as long<br />
as median ones. Caudal fin with 8 upper and 7<br />
lower procurrent rays, and 1 + 9 + 8 + 1 principal<br />
rays.<br />
Table 4. Standard length (in millimetres) and proportional<br />
measurements as a percentage <strong>of</strong> standard length<br />
or head length <strong>of</strong> <strong>the</strong> neotype (AMS B.7920) and a <strong>to</strong>potype<br />
(AMS B.7921) <strong>of</strong> Sys<strong>to</strong>mus immaculatus.<br />
B.7920 B.7921<br />
Standard length (mm) 186.0 192.7<br />
Percent standard length<br />
Total length 127 –<br />
Head length 26.1 25.4<br />
Predorsal length 53.2 54.2<br />
Pre-anal length 75.1 72.1<br />
Pre-pelvic length 41.0 48.2<br />
Caudal-peduncle length 19.4 22.3<br />
Caudal-peduncle depth 15.2 14.8<br />
Body depth 35.9 35.6<br />
Dorsal-fin height 22.7 20.4<br />
Length <strong>of</strong> stiff portion <strong>of</strong> last 15.6 14.7<br />
unbranched dorsal-fin ray<br />
Pec<strong>to</strong>ral-fin length 19.2 19.3<br />
Pelvic-fin length 19.7 18.8<br />
Dorsal-hypural distance 55.4 56.4<br />
Post-dorsal distance 38.8 40.1<br />
Maximum body width<br />
Percent head length<br />
15.4 14.5<br />
Snout length 30.1 29.5<br />
Post-orbital head length 53.6 52.2<br />
Head depth 83.1 81.0<br />
Eye diameter 23.0 22.3<br />
Maxillary barbel length 19.3 23.8<br />
Rostral barbel length 8.3 16.8<br />
Internarial width 23.0 23.5<br />
Interorbital width 43.8 42.8<br />
Lateral line present, complete, with 32 (33)<br />
pored scales on body, 2 (3) on caudal-fin base.<br />
Lateral line sloping gently downwards for about<br />
7 scales, level <strong>the</strong>reafter. Flank scales large, <strong>the</strong>ir<br />
central field (middle one-third) free <strong>of</strong> annuli,<br />
with about 15 irregular polygons. Radii radial,<br />
denser anteriorly and posteriorly, less dense in<br />
dorsal and ventral fields. 5 1 /2 scales in transverse<br />
line between lateral line and origin <strong>of</strong> dorsal fin,<br />
4 between lateral line and origin <strong>of</strong> pelvic fin;<br />
1 /2 3/1/3 scales in transverse line on caudal peduncle;<br />
12 (13) predorsal scales. An axillary scale<br />
approximately one-fourth length <strong>of</strong> pelvic fin<br />
extends backwards from pelvic-fin origin.<br />
Supraneurals 5 (6); 15 abdominal and 17 (18)<br />
caudal vertebrae (Fig. 6b); free uroneural present;<br />
infraorbital 3 slender.<br />
<strong>Dr</strong>avidia, new genus<br />
Type species. Cirrhinus fasciatus Jerdon, 1849:<br />
305.<br />
Diagnosis. <strong>Dr</strong>avidia differs from all o<strong>the</strong>r <strong>South</strong><br />
and Sou<strong>the</strong>ast <strong>Asian</strong> genera <strong>of</strong> Cyprinidae by <strong>the</strong><br />
combination <strong>of</strong> <strong>the</strong> following characters and<br />
character states: size small, usually less than<br />
60 mm SL; rostral and maxillary barbels present;<br />
lateral line complete, with 18-26 pored scales on<br />
body; dorsal fin with 4 unbranched and 8 branched<br />
rays, <strong>the</strong> last unbranched ray weak, smooth<br />
(Fig. 7c); anal fin with 3 unbranched and 5<br />
branched rays; gill rakers simple, acuminate (not<br />
branched or laminate); no antrorse predorsal<br />
spinous ray; infraorbital 3 deep, partly overlapping<br />
preoperculum (Fig. 7a); free uroneural and<br />
post-epiphysial fontanelle absent (Fig. 7 b, d); and<br />
one or two broad, black bars on flank, between<br />
bases <strong>of</strong> dorsal and anal fins.<br />
Remarks. The following nominal species are<br />
<strong>referred</strong> <strong>to</strong> <strong>Dr</strong>avidia: D. afasciata (Jayaram, 1990),<br />
D. fasciata (Jerdon, 1849), D. kannikattiensis (Arunachalam<br />
& Johnson, 2003), D. melanampyx (Day,<br />
1865) and D. pradhani (Tilak, 1973).<br />
Etymology. Named for <strong>the</strong> <strong>Dr</strong>avidian peoples <strong>of</strong><br />
<strong>the</strong> sou<strong>the</strong>rn peninsular India; gender feminine.<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
a<br />
b<br />
Fig. 6. Sys<strong>to</strong>mus immaculatus, AMS B.7920, neotype, 187 mm SL; India: Assam; a, pho<strong>to</strong>graph; b, radiograph.<br />
IO5<br />
IO4<br />
Pop<br />
IO3<br />
So<br />
IO1<br />
IO2<br />
a b c d<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
F<br />
So<br />
Pa<br />
PU3 PU2 CC<br />
Fig. 7. <strong>Dr</strong>avidia fasciata, WHT 11043, 43.0 mm SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural;<br />
Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; supraorbital sensory<br />
canal not shown.<br />
Ep<br />
Pls<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
H1<br />
Ph<br />
79
80<br />
IO5<br />
IO4<br />
Pop<br />
IO3<br />
So<br />
IO1<br />
IO2<br />
a b c d<br />
Dawkinsia, new genus<br />
Type species. Leuciscus filamen<strong>to</strong>sus Valenciennes,<br />
in Cuvier & Valenciennes, 1844.<br />
Diagnosis. Dawkinsia differs from all <strong>South</strong> and<br />
Sou<strong>the</strong>ast <strong>Asian</strong> genera <strong>of</strong> Cyprinidae by <strong>the</strong><br />
combination <strong>of</strong> <strong>the</strong> following characters and<br />
character states: adult size usually 80-120 mm<br />
SL; rostral barbels absent, maxillary barbels<br />
present or absent; last unbranched dorsal-fin ray<br />
smooth (Fig. 8c); 4 unbranched and 8 branched<br />
dorsal-fin rays, 3 unbranched and 5 branched<br />
anal-fin rays; lateral line complete, with 18-22<br />
scales on body; gill rakers simple, acuminate (not<br />
branched or laminate); no antrorse predorsal<br />
spinous ray; free uroneural present (Fig. 8d); 4-5<br />
supraneurals; 15 precaudal and 14-17 caudal<br />
vertebrae; post-epiphysial fontanelle absent<br />
(Fig. 8b); infraorbital 3 slender, not overlapping<br />
preoperculum (Fig. 8a); juvenile (< 50 mm SL)<br />
colour pattern consisting <strong>of</strong> three black bars on<br />
body, persisting <strong>to</strong> adult stage in some species; a<br />
black, horizontally elongate blotch on <strong>the</strong> caudal<br />
peduncle in adults.<br />
Remarks. Based on <strong>the</strong> comparisons made herein,<br />
and <strong>the</strong> data <strong>of</strong> Taki et al. (1978) and Pethiyagoda<br />
& Kottelat (2005a), we include <strong>the</strong> following<br />
nine species in Dawkinsia: D. arulius (Jerdon, 1849),<br />
D. assimilis (Jerdon, 1849), D. exclamatio (Pethiyagoda<br />
& Kottelat, 2005b), D. filamen<strong>to</strong>sa (Valenciennes,<br />
in Cuvier & Valenciennes, 1844), D. rohani<br />
(Rema Devi, Indra & Marcus Knight, 2010), D. ru<br />
F<br />
Pa<br />
So<br />
PU3 PU2 CC<br />
Fig. 8. Dawkinsia filamen<strong>to</strong>sa, WHT 11026, 67.1 mm SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural;<br />
Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; So, supraorbital; Un, free uroneural;<br />
supraorbital sensory canal not shown.<br />
brotinctus (Jerdon, 1849) (see Knight et al., 2011),<br />
D. singhala (Duncker, 1912), D. srilankensis (Senanayake,<br />
1985) and D. tambraparniei (Silas, 1954).<br />
The genus appears <strong>to</strong> be restricted <strong>to</strong> Sri Lanka<br />
and sou<strong>the</strong>rn India.<br />
Etymology. The genus is named for Richard<br />
Dawkins, for his contribution <strong>to</strong> <strong>the</strong> public understanding<br />
<strong>of</strong> science and, in particular, <strong>of</strong> evolutionary<br />
science; gender feminine.<br />
Pethia, new genus<br />
Ep<br />
Pls<br />
Un<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
H1<br />
Ph<br />
Type species. Barbus nigr<strong>of</strong>asciatus Gün<strong>the</strong>r,<br />
1868.<br />
Diagnosis. Pethia differs from all <strong>South</strong> and<br />
Sou<strong>the</strong>ast <strong>Asian</strong> genera <strong>of</strong> Cyprinidae by <strong>the</strong><br />
combination <strong>of</strong> <strong>the</strong> following characters and<br />
character states: small adult size (usually less than<br />
50 mm SL, exceptionally <strong>to</strong> 80 mm SL); rostral<br />
barbels absent; maxillary barbels usually absent,<br />
minute if present; last unbranched dorsal-fin ray<br />
stiff, serrated (Fig. 9c); 3 or 4 unbranched and 8<br />
branched dorsal-fin rays, 3 unbranched and 5<br />
branched anal-fin rays; gill rakers simple, acuminate<br />
(not branched or laminate); no antrorse<br />
predorsal spinous ray; infraorbital 3 deep, partially<br />
overlapping <strong>the</strong> cheek and preoperculum<br />
(Fig. 9a); free uroneural absent (Fig. 9d); 4 supraneurals;<br />
11-13 precaudal and 13-16 caudal<br />
vertebrae; post-epiphysial fontanelle absent<br />
(Fig. 9b); lateral line complete, interrupted or<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
Pop<br />
IO5<br />
IO4<br />
IO3<br />
So<br />
IO2<br />
IO1<br />
a b c d<br />
incomplete (usually incomplete), with 19-24<br />
scales on body in lateral series; and colour pattern<br />
consisting <strong>of</strong> a black blotch on caudal peduncle<br />
and frequently also o<strong>the</strong>r black blotches, spots or<br />
bars on side <strong>of</strong> body.<br />
Remarks. Based on <strong>the</strong> comparisons made herein,<br />
and <strong>the</strong> data <strong>of</strong> Taki et al. (1978), Kullander &<br />
Fang (2005), Shantakumar & Vishwanath (2006),<br />
Kullander (2008) and Kullander & Britz (2008) we<br />
allocate <strong>the</strong> following 23 species <strong>to</strong> Pethia: P. atra<br />
(Linthoingambi & Vishwanath, 2007), P. bandula<br />
(Kottelat & Pethiyagoda, 1991), P. conchonius<br />
(Hamil<strong>to</strong>n, 1822), P. cumingii (Gün<strong>the</strong>r, 1868),<br />
P. didi (Kullander & Fang, 2005), P. erythromycter<br />
(Kullander, 2008), P. gelius (Hamil<strong>to</strong>n, 1822),<br />
P. khugae (Linthoingambi & Vishwanath, 2007),<br />
P. macrogramma (Kullander, 2008), P. manipurensis<br />
(Menon, Rema Devi & Viswanath, 2000), P. melanomaculata<br />
(Deraniyagala, 1956), P. nankyweensis<br />
(Kullander, 2008), P. nigripinnis Knight, Rema<br />
Devi & Arunachalam, 2012, P. nigr<strong>of</strong>asciata (Gün<strong>the</strong>r,<br />
1868), P. padamya (Kullander & Britz, 2008),<br />
P. phutunio (Hamil<strong>to</strong>n, 1822), P. pookodensis (Mercy<br />
& Jacob, 2007), P. punctata (Day, 1865), P. reval<br />
(Meegaskumbura, Silva, Maduwage & Pethiyagoda,<br />
2008), P. shalynius (Yazdani & Talukdar,<br />
1975), P. s<strong>to</strong>liczkana (Day, 1871), P. <strong>the</strong>lys (Kullander,<br />
2008), P. tiantian (Kullander & Fang, 2005)<br />
and P. tic<strong>to</strong> (Hamil<strong>to</strong>n, 1822). Based on characters<br />
provided in <strong>the</strong>ir original descriptions, and as<br />
pointed out by Kullander & Fang (2005), o<strong>the</strong>r<br />
species such as <strong>Puntius</strong> meingangbii Arunkumar<br />
& Tombi Singh, 2003, P. muvattupuzhaensis Jamee-<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
F<br />
Pa<br />
So<br />
Sc<br />
CC<br />
PU3<br />
PU2<br />
Fig. 9. Pethia nigr<strong>of</strong>asciata, WHT 11022, 35.9 mm SL: a, circumorbital series; b, dorsal view <strong>of</strong> orbital region <strong>of</strong><br />
cranium; c, last unbranched dorsal-fin ray; and d, caudal skele<strong>to</strong>n (scale bar 1 mm). Abbreviations: CC, compound<br />
centrum; F, frontal; Ep, epural; H1-6, hypurals 1-6; IO1-5, infraorbitals 1-5; Pa, parietal; Ph, parhypural;<br />
Pls, pleurostyle; Pop, preopercle; PU2, PU3, preural centra 2, 3; Sc, supraorbital sensory canal; So, supraorbital.<br />
81<br />
la Beevi & Ramachandran, 2005, P. narayani (Hora,<br />
1937), P. ornatus Vishwanath & Laisram, 2004,<br />
P. setnai Chhapgar & Sane, 1992, and P. yuensis<br />
Arunkumar & Tombi Singh, 2003 <strong>to</strong>o, likely warrant<br />
placement in Pethia (though <strong>Puntius</strong> narayani<br />
differs from <strong>Puntius</strong> s. s. and Pethia by having<br />
9 branched dorsal-fin rays and 6 branched anal-fin<br />
rays: Hora, 1937).<br />
Etymology. Pethia is <strong>the</strong> generic vernacular name<br />
for small cyprinids in Sinhala; gender feminine.<br />
Discussion<br />
Ep<br />
Pls<br />
H6<br />
H5<br />
H4<br />
H3<br />
H2<br />
H1<br />
Ph<br />
The mi<strong>to</strong>chondrial DNA phylogeny based on<br />
cyt-b and 16s gene fragments (Fig. 1) shows that<br />
<strong>the</strong>re are five well supported clades that can be<br />
recognized as distinct genera also on <strong>the</strong> basis <strong>of</strong><br />
shared external-morphological and osteological<br />
characters and character states within <strong>South</strong> <strong>Asian</strong><br />
<strong>Puntius</strong> as previously unders<strong>to</strong>od. These generic<br />
allocations are also supported by (uncorrected)<br />
genetic distances in <strong>the</strong> range 7.3-18.1 % (Table 2).<br />
Except as mentioned below, we believe <strong>the</strong> present<br />
analysis <strong>to</strong> include all <strong>the</strong> major lineages represented<br />
by <strong>the</strong> <strong>South</strong> <strong>Asian</strong> <strong>fishes</strong> until now included<br />
in <strong>Puntius</strong>. We also comment below on<br />
several species with distinctive characters for<br />
which we do not have tissue for molecular<br />
analysis. By presenting a clearer definition <strong>of</strong><br />
<strong>Puntius</strong> s. s., <strong>the</strong> present study also opens <strong>the</strong> way<br />
for <strong>the</strong> recognition <strong>of</strong> distinct genera for many <strong>of</strong><br />
<strong>the</strong> Sou<strong>the</strong>ast <strong>Asian</strong> <strong>fishes</strong> hi<strong>the</strong>r<strong>to</strong> <strong>referred</strong> <strong>to</strong>
82<br />
<strong>Puntius</strong>. Our results also corroborate <strong>the</strong> phylogenetic<br />
relationships recovered by Bossuyt et al.<br />
(2004) involving some <strong>of</strong> <strong>the</strong> same species and<br />
analysing <strong>the</strong> same ~ 590 bp segment <strong>of</strong> <strong>the</strong> 16S<br />
rRNA gene and a ~ 540 bp segment <strong>of</strong> <strong>the</strong> cyt-b<br />
gene.<br />
None <strong>of</strong> <strong>the</strong> genus-group names hi<strong>the</strong>r<strong>to</strong> in<br />
<strong>the</strong> synonymy <strong>of</strong> <strong>Puntius</strong> have type species potentially<br />
referable <strong>to</strong> <strong>Puntius</strong> s. s., Sys<strong>to</strong>mus s. s.,<br />
Dawkinsia, <strong>Dr</strong>avidia or Pethia. The only contending<br />
genus-group names are Cephalakompsus Herre,<br />
1924a, Mandibularca Herre, 1924a, Ospatulus Herre,<br />
1924b, and Spratellicypris Herre & Myers, 1931,<br />
all <strong>of</strong> which have type species endemic <strong>to</strong> <strong>the</strong><br />
Philippines. These possess both rostral and maxillary<br />
barbels, by which <strong>the</strong>y are immediately<br />
distinguished from <strong>Puntius</strong>, Dawkinsia and Pethia<br />
(which lack rostral barbels). <strong>Puntius</strong> and Sys<strong>to</strong>mus<br />
are in any event not threatened by Cephalakompsus,<br />
Mandibularca, Ospatulus and Spratellicypris, which<br />
are junior. On <strong>the</strong> basis <strong>of</strong> <strong>the</strong> original descriptions<br />
<strong>of</strong> <strong>the</strong>ir type species in Herre (1924a), <strong>Dr</strong>avidia<br />
differs from Cephalakompsus, Ospatulus and Spratellicypris<br />
by having a smooth (vs. serrated) last<br />
unbranched dorsal-fin ray; and from Mandibularca<br />
by its smaller size (up <strong>to</strong> 80 mm <strong>to</strong>tal length,<br />
vs. 220 mm in Mandibularca), having <strong>the</strong> lower<br />
jaw horizontal (vs. strongly curved upwards) and<br />
habitus (see Herre, 1924a: pl. 1).<br />
<strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>. The recognition <strong>of</strong> additional<br />
genera within <strong>Puntius</strong> as previously unders<strong>to</strong>od<br />
makes it imperative that <strong>the</strong> identity <strong>of</strong><br />
P. sophore (Hamil<strong>to</strong>n, 1822), <strong>the</strong> type species <strong>of</strong><br />
<strong>Puntius</strong>, be stabilised. Hamil<strong>to</strong>n’s description and<br />
figure <strong>of</strong> P. sophore do not allow this species <strong>to</strong> be<br />
unambiguously distinguished from o<strong>the</strong>r valid<br />
species, e. g., P. chola, P. dorsalis. No type specimens<br />
<strong>of</strong> <strong>fishes</strong> described by Hamil<strong>to</strong>n have come<br />
<strong>to</strong> light in <strong>the</strong> 190 years since <strong>the</strong> publication <strong>of</strong><br />
this work; nor has <strong>the</strong> existence <strong>of</strong> such types<br />
been alluded <strong>to</strong> in <strong>the</strong> subsequent literature. They<br />
must <strong>the</strong>refore be presumed lost. Hamil<strong>to</strong>n (1822)<br />
did not specify <strong>the</strong> type locality <strong>of</strong> S. sophore in<br />
his account <strong>of</strong> that species, but it can be inferred<br />
from his preceding account <strong>of</strong> <strong>Puntius</strong> that it was<br />
Bengal. We selected ZRC 35064, a 71.8 mm SL<br />
specimen, as neotype because it was collected<br />
within <strong>the</strong> Ganges floodplain in Bengal (now<br />
Bangladesh); it is consistent with <strong>the</strong> original<br />
description (Hamil<strong>to</strong>n, 1822: pl. 19, fig. 86); and<br />
it is accompanied by a series <strong>of</strong> five <strong>to</strong>potypes<br />
that facilitate a more representative description.<br />
The coloration <strong>of</strong> <strong>the</strong> neotype also matches<br />
that deliniated in a copy <strong>of</strong> Hamil<strong>to</strong>n’s figure in<br />
<strong>the</strong> archives <strong>of</strong> <strong>the</strong> Linnean Society <strong>of</strong> London<br />
(Fig. 2b). This illustrates a specimen with 17 lateral-line<br />
scales, 2 unbranched and 8 branched<br />
dorsal-fin rays, 2 unbranched and 6 branched<br />
anal-fin rays, 1 + 7 + 7 + 1 principal caudal-fin rays,<br />
and 1 unbranched and 7 branched pelvic-fin rays.<br />
Although, as pointed out by Silva et al. (2010),<br />
Hamil<strong>to</strong>n’s drawings contain many inaccuracies<br />
in <strong>the</strong>ir detail, <strong>the</strong> neotype matches Hamil<strong>to</strong>n’s<br />
figure in general appearance. Based on <strong>the</strong> neotype<br />
and <strong>to</strong>potypes described herein, P. sophore<br />
may be distinguished from P. chola by <strong>the</strong> characters<br />
given in Talwar & Jhingran (1991) and<br />
Jayaram (2010), and from P. dorsalis by <strong>the</strong> characters<br />
given in Pethiyagoda et al. (2008).<br />
Because <strong>of</strong> a lack <strong>of</strong> material, we have been<br />
unable <strong>to</strong> assign several o<strong>the</strong>r Indian species with<br />
40 or more lateral-line scales, hi<strong>the</strong>r<strong>to</strong> in <strong>Puntius</strong>,<br />
<strong>to</strong> any <strong>of</strong> <strong>the</strong> above genera: <strong>the</strong>ir generic placement<br />
must depend on future work. Perhaps most<br />
prominent among <strong>the</strong>se are ‘P.’ deccanensis, ‘P.’ fraseri,<br />
and P. sharmai, which lack rostral barbels but<br />
possess a pair <strong>of</strong> maxillary barbels, a stiff, strongly<br />
serrated last unbranched dorsal-fin ray, and<br />
42-47 lateral-line scales, though <strong>the</strong> first species<br />
differs from <strong>the</strong> latter two by having 9 branched<br />
dorsal-fin rays and 7 branched anal-fin rays, vs.<br />
8 and 5 in <strong>Puntius</strong> (Hora & Misra, 1938; Yazdani<br />
& Babu Rao, 1976; Menon & Rema Devi, 1993).<br />
Also with a high lateral-line scale count (43) is<br />
<strong>Puntius</strong> punjaubensis (Day, 1871), which has no<br />
barbels and possesses a smooth last unbranched<br />
dorsal-fin ray: it is almost certainly not a member<br />
<strong>of</strong> any <strong>of</strong> <strong>the</strong> preceding genera. We note in passing<br />
that <strong>the</strong> original spelling ‘punjaubensis’ is not<br />
a lapsus for ‘punjabensis’, <strong>the</strong> spelling adopted in<br />
<strong>the</strong> current literature (e. g., Jayaram, 1991, 2010;<br />
Talwar & Jhingran, 1991): Day retained <strong>the</strong><br />
original spelling also in his ‘Fishes <strong>of</strong> India’ (1878),<br />
‘Punjaub’ having been an accepted contemporaneous<br />
spelling.<br />
Ano<strong>the</strong>r intriguing taxon is ‘<strong>Puntius</strong>’ ophicephalus,<br />
for which Menon (1999) created <strong>the</strong><br />
genus Eechathalakenda. Possessing both maxillary<br />
and rostral barbels, a smooth last unbranched<br />
dorsal-fin ray and 40-42 lateral-line scales, this<br />
species does not fit within any <strong>of</strong> <strong>the</strong> preceding<br />
genera. Menon (1999) diagnosed Eechathalakenda<br />
from Gonoprok<strong>to</strong>pterus by its “elongated Channa<br />
like appearance, 7 branched rays in <strong>the</strong> dorsal fin<br />
(9 in Gonoprok<strong>to</strong>pterus), a slightly elongated but<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
weak, smooth and segmented last undivided ray<br />
<strong>to</strong> <strong>the</strong> dorsal fin, four barbels and 40-42 scales<br />
along <strong>the</strong> lateral line”. We tentatively treat Eechathalakenda<br />
as valid.<br />
The relationships <strong>of</strong> ‘P.’ guganio (Hamil<strong>to</strong>n,<br />
1822) and ‘P.’ nangalensis Jayaram, 1990 (barbels<br />
absent; last unbranched dorsal-fin ray stiff, serrated;<br />
lateral line incomplete, with 36-39 and<br />
31-33 scales respectively, in lateral series), remain<br />
<strong>to</strong> be elucidated. Jayaram (1990) considered <strong>the</strong>se<br />
two species <strong>to</strong> be closely related. In <strong>the</strong> absence<br />
<strong>of</strong> material for comparison, we retain both species<br />
in ‘<strong>Puntius</strong>’ for <strong>the</strong> present, although <strong>the</strong>y do not<br />
belong <strong>to</strong> this or any <strong>of</strong> <strong>the</strong> genera discussed<br />
herein.<br />
Unlike all <strong>the</strong> o<strong>the</strong>r species <strong>of</strong> <strong>Puntius</strong> examined<br />
in this study, P. bimaculatus (restricted <strong>to</strong><br />
sou<strong>the</strong>rn India and Sri Lanka) and P. titteya (a Sri<br />
Lankan endemic) lack a post-epiphysial fontanelle.<br />
These two species also group separately<br />
from all <strong>the</strong> o<strong>the</strong>r species <strong>of</strong> <strong>Puntius</strong> examined<br />
(Fig. 1) and share a suite <strong>of</strong> additional distinguishing<br />
characters: only 7 (vs. 8) branched dorsal-fin<br />
rays; 1 /2 3 (vs. 1 /2 4- 1 /2 5) scale rows between dorsalfin<br />
origin and lateral-line row; a lateral line that<br />
is in P. titteya incomplete and in P. bimaculatus<br />
occasionally interrupted or incomplete; and a<br />
coloration which includes a red lateral stripe –<br />
unusual in <strong>Puntius</strong>. Although <strong>the</strong>y appear <strong>to</strong><br />
warrant placement in a separate genus, we refrain<br />
from doing so as this would render <strong>Puntius</strong><br />
paraphyletic (<strong>the</strong> two lower clades <strong>of</strong> <strong>Puntius</strong> in<br />
Fig. 1 evidently do not possess a suite <strong>of</strong> external<br />
characters that distinguish <strong>the</strong>m). We feel this<br />
problem is best resolved once a wider sample <strong>of</strong><br />
species is available for analysis.<br />
<strong>South</strong> <strong>Asian</strong> Sys<strong>to</strong>mus. Having long been treated<br />
as a synonym <strong>of</strong> <strong>Puntius</strong> (Bleeker, 1863b; Day,<br />
1878; Banarescu, 1997), Sys<strong>to</strong>mus was revived as<br />
a valid genus by Rainboth (1996), who assigned<br />
six Cambodian species previously in <strong>Puntius</strong> <strong>to</strong><br />
it on <strong>the</strong> basis <strong>of</strong> <strong>the</strong> finely-serrated last unbranched<br />
dorsal-fin ray, possession <strong>of</strong> rostral and/<br />
or maxillary barbels, and gill rakers fewer than<br />
12 on <strong>the</strong> first arch. Fur<strong>the</strong>r species were added<br />
by subsequent authors (e. g. Ng & Tan, 1999;<br />
Parenti & Lim, 2005; Kottelat & Tan, 2011; see also<br />
Roberts & Catania, 2007). Given that some species<br />
<strong>of</strong> Sys<strong>to</strong>mus sensu Rainboth (1996) are popular<br />
aquarium <strong>fishes</strong> (e. g., ‘S.’ johorensis, ‘S.’ lateristriga,<br />
‘S.’ partipentazona), <strong>the</strong> genus-name is in<br />
wide use despite o<strong>the</strong>r authors (e. g., Kottelat,<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
83<br />
2000, 2001; Kottelat & Widjanarti, 2005) retaining<br />
it as a junior synonym <strong>of</strong> <strong>Puntius</strong>, because removing<br />
<strong>the</strong>se six species out <strong>of</strong> a large, hetero geneous<br />
genus without resolving <strong>the</strong> generic identity <strong>of</strong><br />
<strong>the</strong> remaining species would likely cause confusion.<br />
This confusion is in part because <strong>the</strong> type<br />
species <strong>of</strong> Sys<strong>to</strong>mus, S. immaculatus, considered a<br />
synonym <strong>of</strong> S. sarana since Day (1878), is poorly<br />
known. M’Clelland (1839) characterized Sys<strong>to</strong>mus<br />
as follows: “Intermaxillaries protractile, dorsal<br />
and anal short, <strong>the</strong> former opposite <strong>to</strong> <strong>the</strong> ventrals<br />
and preceded by a spinous ray; body elevated<br />
and marked by two or more distinct dark spots,<br />
or diffuse spots ei<strong>the</strong>r on <strong>the</strong> fins or opercula,<br />
prominence on <strong>the</strong> apex <strong>of</strong> <strong>the</strong> lower jaw obscure”.<br />
The first <strong>of</strong> <strong>the</strong> eight species he included in <strong>the</strong><br />
first division <strong>of</strong> <strong>the</strong> genus (characterized by “depth<br />
<strong>of</strong> <strong>the</strong> body . . . equal <strong>to</strong> about half <strong>the</strong> length, and<br />
<strong>the</strong> spiny ray <strong>of</strong> <strong>the</strong> dorsal . . . serrated behind”)<br />
was S. immaculatus, subsequently designated as<br />
type species by Jordan (1919). This species he<br />
described as having “four cirrhi, a slight golden<br />
tinge on <strong>the</strong> opercula, fins dark, thirty-two scales<br />
on <strong>the</strong> lateral line and ten in an oblique line from<br />
<strong>the</strong> base <strong>of</strong> <strong>the</strong> ventrals <strong>to</strong> <strong>the</strong> dorsum”. He gave<br />
<strong>the</strong> type locality <strong>of</strong> S. immaculatus as “Small<br />
streams with sandy bot<strong>to</strong>ms in Assam, and probably<br />
occasionally in <strong>the</strong> large rivers” adding that<br />
S. chrysosomus (a junior synonym <strong>of</strong> S. sarana) “is<br />
probably a variety only <strong>of</strong> this species, which is<br />
generally distributed throughout <strong>the</strong> rivers and<br />
ponds <strong>of</strong> all parts <strong>of</strong> India, sometime weighing<br />
as much as two pounds [0.9 kg]”.<br />
The type/s <strong>of</strong> S. immaculatus, however, are<br />
apparently lost. A search for possible types <strong>of</strong> this<br />
species in <strong>the</strong> collections <strong>of</strong> BMNH and SMF, <strong>the</strong><br />
only reposi<strong>to</strong>ries known <strong>to</strong> contain fish types <strong>of</strong><br />
M’Clelland, proved fruitless (J. Maclaine, BMNH;<br />
and H. Zetzsche, SMF, pers. comms.). A search<br />
<strong>of</strong> <strong>the</strong> fish collections <strong>of</strong> ZSI <strong>to</strong>o, failed <strong>to</strong> reveal<br />
any cyprinid specimens attributable <strong>to</strong> M’Clelland<br />
(RP, pers. obs.). Given <strong>the</strong> confusion that exists<br />
with regard <strong>to</strong> <strong>the</strong> identity <strong>of</strong> <strong>the</strong> genus, it is<br />
necessary that <strong>the</strong> identity <strong>of</strong> S. immaculatus, its<br />
type species, be clarified through <strong>the</strong> designation<br />
<strong>of</strong> a neotype: we here designate AMS B.7920<br />
(locality Assam, India), as neotype <strong>of</strong> Sys<strong>to</strong>mus<br />
immaculatus M’Clelland, 1839. The specimen is<br />
illustrated in Fig. 6 and its morphometric and<br />
meristic data given in Table 2, which supplement<br />
<strong>the</strong> description and diagnosis <strong>of</strong> M’Clelland<br />
(1839).
84<br />
M’Clelland’s (1839) description <strong>of</strong> S. immaculatus<br />
matches that <strong>of</strong> ‘<strong>Puntius</strong>’ sarana (type locality<br />
‘ponds and rivers <strong>of</strong> Bengal’) (Jayaram, 1991;<br />
Talwar & Jhingran, 1991). As shown here (Figs.<br />
1, 6) and corroborated also by Shantakumar &<br />
Vishwanath (2006), ‘P.’ sarana belongs <strong>to</strong> a lineage<br />
distinct from <strong>the</strong> one that includes <strong>Puntius</strong> sophore<br />
(<strong>the</strong> type species <strong>of</strong> <strong>Puntius</strong>): <strong>the</strong> former possess<br />
both rostral and maxillary barbels, a stiff, strongly<br />
serrated last unbranched dorsal-fin ray, a<br />
complete lateral line with 27-34 scales on <strong>the</strong><br />
body, and a horizontally elongate oval black<br />
blotch on <strong>the</strong> caudal peduncle. This adds confidence<br />
<strong>to</strong> our selection <strong>of</strong> a neotype for S. immaculatus<br />
from among specimens nominally<br />
identified as ‘<strong>Puntius</strong> sarana’ from Assam. Sys<strong>to</strong>mus<br />
immaculatus (= S. sarana) may be distinguished<br />
from <strong>the</strong> o<strong>the</strong>r species here included in<br />
Sys<strong>to</strong>mus based on <strong>the</strong> characters provided above<br />
for <strong>the</strong> neotype and <strong>to</strong>potype, and those provided<br />
in <strong>the</strong> works cited in Remarks under <strong>the</strong><br />
diagnosis <strong>of</strong> Sys<strong>to</strong>mus.<br />
Barbodes, Barbonymus and Poropuntius resemble<br />
Sys<strong>to</strong>mus in possessing two pairs <strong>of</strong> barbels<br />
and a serrated last unbranched dorsal-fin ray.<br />
Indeed, Menon (1999) placed S. sarana in Barbodes,<br />
<strong>the</strong> type species <strong>of</strong> which is Barbus maculatus<br />
Valenciennes, in Cuvier & Valenciennes 1842 (see<br />
Kottelat, 1999), which is considered a synonym<br />
<strong>of</strong> B. binotatus Valenciennes, in Cuvier & Valenciennes<br />
1842 (Kottelat, 1999; Pethiyagoda & Kottelat,<br />
2005b). Barbus binotatus, however, differs<br />
from species <strong>of</strong> Sys<strong>to</strong>mus by possessing 23-27<br />
lateral-line scales on body and ½4 scale rows<br />
between <strong>the</strong> dorsal-fin origin and lateral line (vs.<br />
32-33 lateral-line scales and 1 /2 5 scale rows between<br />
<strong>the</strong> dorsal-fin origin and lateral line in<br />
S. immaculatus, <strong>the</strong> type species <strong>of</strong> Sys<strong>to</strong>mus), and<br />
having a distinctive colour pattern (see Kottelat<br />
et al., 1993, pl. 15) not observed in any o<strong>the</strong>r species<br />
here allocated <strong>to</strong> Sys<strong>to</strong>mus s. s. Sys<strong>to</strong>mus differs<br />
from Barbonymus by having fewer than 6 rows <strong>of</strong><br />
scales between <strong>the</strong> dorsal-fin origin and lateral<br />
line (7 or more in Barbonymus); and from Poropuntius<br />
by lacking an accessory pore on <strong>the</strong> canals<br />
<strong>of</strong> <strong>the</strong> lateral-line system (accessory pore present<br />
in Poropuntius: Kottelat et al., 1993; Roberts, 1998;<br />
Kottelat, 2001).<br />
Sou<strong>the</strong>ast <strong>Asian</strong> <strong>Puntius</strong>. The recent literature<br />
refers around 45 species <strong>of</strong> Sou<strong>the</strong>ast <strong>Asian</strong><br />
cyprinids <strong>to</strong> <strong>Puntius</strong>. Only one <strong>of</strong> <strong>the</strong>se, however,<br />
belongs <strong>to</strong> <strong>the</strong> genus as it is now unders<strong>to</strong>od:<br />
P. brevis (Bleeker, 1849). Widespread through<br />
Sou<strong>the</strong>ast Asia (Mekong basin <strong>of</strong> Laos, Thailand<br />
and Cambodia; Chao Phraya basin; Malay peninsula<br />
and Java: Kottelat, 2001) it has <strong>the</strong> last unbranched<br />
dorsal-fin ray smooth, a pair <strong>of</strong> maxillary<br />
(and no rostral) barbels, 23-26 lateral-line<br />
scales and a black spot on <strong>the</strong> caudal peduncle.<br />
Taki et al. (1978) included P. brevis in <strong>the</strong>ir P. binotatus<br />
group, though <strong>the</strong> former species lacks<br />
rostral barbels (present in P. binotatus) and has<br />
<strong>the</strong> last unbranched dorsal-fin ray smooth (vs.<br />
posteriorly serrated).<br />
Only one o<strong>the</strong>r Sou<strong>the</strong>ast <strong>Asian</strong> <strong>Puntius</strong> has<br />
<strong>the</strong> last unbranched dorsal-fin ray smooth, <strong>Puntius</strong><br />
oligolepis (Bleeker, 1853). This species, however,<br />
is distinguished from <strong>Puntius</strong> s.s. by having an<br />
incomplete lateral line, only 17 scales in <strong>the</strong> lateral<br />
series, and 1 /2 3 scale rows between <strong>the</strong> dorsalfin<br />
origin and lateral-line scale row. Its relationships<br />
remain <strong>to</strong> be investigated.<br />
All <strong>the</strong> o<strong>the</strong>r Sou<strong>the</strong>ast <strong>Asian</strong> species presently<br />
<strong>referred</strong> <strong>to</strong> <strong>Puntius</strong> require new generic<br />
assignments. Although this is beyond <strong>the</strong> scope<br />
<strong>of</strong> <strong>the</strong> present work for want <strong>of</strong> material, for<br />
convenience <strong>of</strong> discussion and based on data on<br />
external characters available in <strong>the</strong> literature<br />
(Kottelat et al., 1993; Kottelat, 2001; Rainboth,<br />
1996; Roberts, 1989; Weber & de Beaufort, 1916;<br />
and <strong>the</strong> original species descriptions), we here<br />
refer <strong>the</strong>se <strong>to</strong> a number <strong>of</strong> species groups (see also<br />
Sou<strong>the</strong>ast <strong>Asian</strong> Sys<strong>to</strong>mus, below).<br />
The ‘<strong>Puntius</strong> trifasciatus group’ comprises three<br />
species (P. gemellus Kottelat, 1996, P. johor ensis<br />
(Duncker, 1904), and P. trifasciatus Kottelat, 1996)<br />
distinguished by possessing both rostral and<br />
maxillary barbels, 25-27 scales in <strong>the</strong> lateral line<br />
on body, 7-11 gill rakers on <strong>the</strong> anterior gill arch,<br />
and a juvenile coloration consisting <strong>of</strong> 4-5 black<br />
bars on <strong>the</strong> body, changing <strong>to</strong> 3-6 black stripes<br />
on <strong>the</strong> body as adults (Kottelat, 1996); <strong>the</strong>se characters<br />
combine <strong>to</strong> distinguish <strong>the</strong> members <strong>of</strong> this<br />
group from <strong>Puntius</strong>, Sys<strong>to</strong>mus, Pethia and <strong>Dr</strong>avidia.<br />
The members <strong>of</strong> <strong>the</strong> P. trifasciatus group<br />
appear <strong>to</strong> belong <strong>to</strong> a lineage distinct also from<br />
<strong>the</strong> only o<strong>the</strong>r similarly-coloured species <strong>of</strong> Sou<strong>the</strong>ast<br />
<strong>Asian</strong> ‘<strong>Puntius</strong>’, P. lineatus (Duncker, 1904),<br />
which possess no or only a single pair <strong>of</strong> rudimentary<br />
barbels and <strong>the</strong> adults <strong>of</strong> which have <strong>the</strong><br />
lower lip forming a complete post-labial groove,<br />
distinguishing it from all <strong>Puntius</strong> (in ei<strong>the</strong>r its<br />
previous or present conception). Kottelat (1996)<br />
proposed that it be placed in a separate genus (for<br />
<strong>the</strong> present <strong>referred</strong> <strong>to</strong> as <strong>the</strong> ‘P. lineatus’ group).<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
Sou<strong>the</strong>ast <strong>Asian</strong> Sys<strong>to</strong>mus. Of <strong>the</strong> five species<br />
<strong>referred</strong> <strong>to</strong> Sys<strong>to</strong>mus by Rainboth (1996) only<br />
S. orphoides appears <strong>to</strong> be a Sys<strong>to</strong>mus as <strong>the</strong> genus<br />
is now unders<strong>to</strong>od. The o<strong>the</strong>r four included species<br />
possess characters that suggest <strong>the</strong>y belong<br />
<strong>to</strong> different lineages: ‘S.’ aurotaeniatus (Tirant 1885)<br />
lacks rostral barbels and has a coloration consisting<br />
<strong>of</strong> black spots at <strong>the</strong> dorsal-fin origin and on<br />
<strong>the</strong> side <strong>of</strong> <strong>the</strong> body; ‘S.’ johorensis (Duncker, 1904)<br />
lacks rostral barbels and has a striking colour<br />
pattern consisting <strong>of</strong> six black lateral bars; ‘S.’ partipentazona<br />
(Fowler, 1934a) lacks rostral barbels,<br />
has <strong>the</strong> lateral line incomplete and a colour pattern<br />
consisting <strong>of</strong> five black lateral bars including<br />
one across <strong>the</strong> dorsal fin; and ‘S.’ binotatus has a<br />
dark spot at <strong>the</strong> dorsal-fin origin, fewer scale rows<br />
on <strong>the</strong> body (23-27, vs. 27-34 in Sys<strong>to</strong>mus) and<br />
caudal peduncle (12 circumpedun cular scale<br />
rows, vs. 16) and a different head and mouth<br />
shape. Among Sou<strong>the</strong>ast <strong>Asian</strong> cyprinids, only<br />
S. orphoides and Barbus jacobusboehlkei Fowler, 1958<br />
appear <strong>to</strong> belong <strong>to</strong> Sys<strong>to</strong>mus s. s. We here place<br />
<strong>the</strong> remaining species referable <strong>to</strong> Sys<strong>to</strong>mus sensu<br />
Rainboth (1996) tentatively in two species groups<br />
that are, however, unlikely <strong>to</strong> be monophyletic<br />
(e. g., see Kottelat, 1992).<br />
Members <strong>of</strong> <strong>the</strong> ‘<strong>Puntius</strong> binotatus group’ possess<br />
both rostral and maxillary barbels, and a last<br />
unbranched dorsal-fin ray <strong>the</strong> posterior margin<br />
<strong>of</strong> which is more or less strongly serrated. Roberts<br />
(1989) regarded P. binotatus <strong>to</strong> be “<strong>the</strong> most<br />
widely distributed and perhaps most variable<br />
species <strong>of</strong> <strong>Puntius</strong> in sou<strong>the</strong>ast Asia”, noting that<br />
it “occurs from sea level <strong>to</strong> at least 2,000 m . . . and<br />
on small islands inhabited by few o<strong>the</strong>r freshwater<br />
<strong>fishes</strong>.” Close examination will likely show at<br />
least some <strong>of</strong> <strong>the</strong> 13 nominal species from time<br />
<strong>to</strong> time in its synonymy (see Roberts, 1989: 60-61)<br />
<strong>to</strong> be valid, closely-related species, as are P. banksi<br />
Herre, 1940 (see Kottelat & Lim, 1996: 232), and<br />
P. sealei (Herre, 1933) (see Inger & Chin, 1990: 73).<br />
Additionally, as pointed out by Roberts (1989),<br />
P. binotatus is regarded as <strong>the</strong> “parent s<strong>to</strong>ck” for<br />
<strong>the</strong> endemic ‘<strong>Puntius</strong>’ species <strong>of</strong> Lake Lanao and<br />
<strong>the</strong> Lanao Plateau <strong>of</strong> Mindanao in <strong>the</strong> Philippines:<br />
P. amarus (Herre, 1924a), P. disa (Herre, 1932),<br />
P. flavifuscus (Herre, 1924a), P. herrei (Fowler,<br />
1934b), P. joaquinae Wood, 1968, P. ka<strong>to</strong>lo (Herre,<br />
1924a), P. lanaoensis (Herre 1924a), P. lindog (Herre<br />
1924a), P. manalak (Herre, 1924a) and P. sirang<br />
(Herre, 1932). The Lanao ‘<strong>Puntius</strong>’ share <strong>the</strong>se<br />
characters also with o<strong>the</strong>r ‘<strong>Puntius</strong>’ from Mindanao<br />
(P. cataractae (Fowler, 1934b), P. tumba<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
85<br />
(Herre, 1924a) and P. umalii Wood, 1968), as well<br />
as those from Lake Manguao on Palawan (P. manguaoensis<br />
(Day, 1914) and P. ban<strong>to</strong>lanensis (Day,<br />
1914)) and Mindoro (P. hemictenus (Jordan & Richardson,<br />
1908)). Ranging from about 6-24 cm<br />
and possessing 22-32 lateral-line scales most <strong>of</strong><br />
<strong>the</strong>se species show a coloration that includes a<br />
hazy dark stripe on <strong>the</strong> side <strong>of</strong> <strong>the</strong> body, in P. binotatus<br />
accompanied by a dark spot on <strong>the</strong> caudal<br />
peduncle and dorsal-fin origin, accompanied in<br />
some cases by numerous o<strong>the</strong>r dark spots and/<br />
or blotches on <strong>the</strong> body (see Kottelat et al., 1993,<br />
pl. 15). ‘Sys<strong>to</strong>mus’ dunckeri (Ahl, 1929), ‘S.’ everetti<br />
(Boulenger, 1894), ‘S.’ ku chingensis (Herre, 1940),<br />
‘S.’ lateristriga (Valen ciennes, in Cuvier & Valenciennes,<br />
1842), ‘S.’ xou thos Kottelat & Tan, 2011,<br />
‘<strong>Puntius</strong>’ rhombeus Kottelat, 2001 and ‘P.’ semifasciolatus<br />
(Gün<strong>the</strong>r, 1868) <strong>to</strong>o, are apparently members<br />
<strong>of</strong> this group. The genus name Barbodes is<br />
available for <strong>the</strong> ‘<strong>Puntius</strong> binotatus group’, with<br />
Cephalakompsus, Mandibularca, Ospatulus and<br />
Spratellicypris as synonyms.<br />
The remaining Sou<strong>the</strong>ast <strong>Asian</strong> ‘Sys<strong>to</strong>mus’<br />
sensu Rainboth (1996) we place in <strong>the</strong> ‘<strong>Puntius</strong><br />
tetrazona group’ because <strong>the</strong>y possess a posteriorly<br />
serrated last unbranched dorsal-fin ray,<br />
maxillary barbels, and a striking coloration that<br />
includes 3-6 black bars and/or blotches on <strong>the</strong><br />
body; <strong>the</strong> lateral line may be complete or incomplete,<br />
and rostral barbels present or absent. These<br />
species share a strikingly barred or blotched<br />
coloration, which resembles that <strong>of</strong> some species<br />
<strong>of</strong> Pethia (e. g. P. nigr<strong>of</strong>asciata) and <strong>Dr</strong>avidia (e. g.<br />
D. fasciata), as does <strong>the</strong>ir small size. They differ<br />
from <strong>the</strong> former genus, however, in possessing a<br />
complete lateral line and at least maxillary barbels;<br />
and <strong>the</strong> latter in possessing a serrated posterior<br />
edge <strong>to</strong> <strong>the</strong> last unbranched dorsal-fin ray. Additionally,<br />
at least some <strong>of</strong> <strong>the</strong> included species<br />
have a slender infraorbital 3 (Taki et al., 1978),<br />
which serves <strong>to</strong> distinguish <strong>the</strong>m from species <strong>of</strong><br />
Pethia. The <strong>Puntius</strong> tetrazona group includes<br />
‘S.’ anchisporus (Vaillant, 1902), ‘S.’ endecanalis<br />
(Roberts, 1989), ‘S.’ foerschi (Kottelat, 1982), ‘S.’ hexazona<br />
(Weber & de Beaufort, 1912), ‘S.’ partipentazona,<br />
‘S.’ penta zona (Boulenger, 1894), ‘S.’ pulcher<br />
(Rendahl, 1922), ‘S.’ rhomboocellatus (Koumans,<br />
1940) and ‘S.’ te trazona (Bleeker, 1855). Kottelat’s<br />
(1992) observation that ‘tiger barbs’ belong <strong>to</strong> “at<br />
least two lineages which are not obviously related”<br />
is supported also by <strong>the</strong> data <strong>of</strong> Taki et al.<br />
(1978), whose ‘tetrazona group’ (‘S.’ tetrazona and<br />
‘S.’ partipentazona) lacks rostral barbels whereas
86<br />
a<br />
b<br />
Fig. 10. Barbus bovanicus, AMS B.7829, lec<strong>to</strong>type, 95.1 mm SL; India: Bhavani River, Tamil Nadu; a, pho<strong>to</strong>graph;<br />
b, radiograph.<br />
<strong>the</strong>ir ‘pentazona group’ (‘S.’ everetti, ‘S.’ lateristriga,<br />
‘S.’ pentazona and ‘S.’ rhomboocellatus) possesses<br />
<strong>the</strong>m; Taki et al. (1978) also distinguished <strong>the</strong> two<br />
groups by <strong>the</strong>ir colour patterns.<br />
<strong>Dr</strong>avidia. Menon (1999) chose <strong>the</strong> junior name<br />
Labeo melanampyx Day, 1865 over its senior subjective<br />
synonym, Cirrhinus fasciatus Jerdon, 1849 (<strong>the</strong><br />
type species <strong>of</strong> <strong>Dr</strong>avidia) citing Art. 23(b) <strong>of</strong> ICZN<br />
(1985), on <strong>the</strong> grounds that <strong>the</strong> former was “a<br />
favourite aquarium fish and several aquarium<br />
journals mention this species as <strong>Puntius</strong> melanampyx.<br />
A change is not considered desirable.”<br />
However, Art. 23(b) <strong>of</strong> ICZN (1985) requires that<br />
such a case is <strong>to</strong> be <strong>referred</strong> <strong>to</strong> <strong>the</strong> Commission<br />
for a ruling, which has not been done in <strong>the</strong> 13<br />
years following Menon (1999). As such, we follow<br />
Arts. 23.1 and 22.9.3 <strong>of</strong> ICZN (1999) and use <strong>the</strong><br />
senior synonym C. fasciatus, which anyway is also<br />
<strong>the</strong> name in prevailing usage (e. g., Silas, 1956;<br />
Jayaram, 1991; Talwar & Jhingran, 1991; Arunachalam,<br />
2000; Bhat, 2005).<br />
Three additional <strong>South</strong> <strong>Asian</strong> species included<br />
in <strong>Puntius</strong> by previous authors (e. g.,<br />
Jayaram, 1991; Talwar & Jhingran, 1991) bear two<br />
pairs <strong>of</strong> barbels and a smooth last unbranched<br />
dorsal-fin ray: ‘P.’ bovanicus, ‘P.’ carnaticus and<br />
‘P.’ jerdoni. Barbus bovanicus (AMS B.7829, syntype,<br />
95.1 mm SL, here designated lec<strong>to</strong>type; Fig. 10)<br />
has 9 branched dorsal-fin rays, 23 lateral-line<br />
scales on body, eye in upper half <strong>of</strong> head, thick,<br />
fleshy lips, and <strong>the</strong> lower lip present medially: it<br />
is apparently a Neolissochilus. It fur<strong>the</strong>r differs<br />
from all species <strong>of</strong> <strong>Puntius</strong>, Sys<strong>to</strong>mus, Dawkinsia,<br />
Pethia and <strong>Dr</strong>avidia in possessing 22 (vs. 13-19)<br />
caudal vertebrae and 6 (vs. 3-5) supraneurals. We<br />
note in passing that <strong>the</strong> spellings bovanicus and<br />
bovianicus were published simultaneously, in plate<br />
138 fig. 1, and on page 566, <strong>of</strong> Day (1878), both <strong>of</strong><br />
which were released on 1 December 1878 (see<br />
Whitehead & Talwar, 1976: 54 and references<br />
<strong>the</strong>rein). In keeping with subsequent usage, as<br />
first reviser we give B. bovanicus precedence over<br />
B. bovianicus.<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
a<br />
b<br />
Fig. 11. Barbus jerdoni, AMS B.7935, syntype, 179.3 mm SL; India: Canara [Karnataka].<br />
Pethiyagoda & Kottelat (2005b) noted that <strong>the</strong><br />
figure <strong>of</strong> Barbus carnaticus in Day (1878: pl. 137)<br />
“possibly illustrates a species <strong>of</strong> Neolissochilus”.<br />
Jerdon’s (1849) description <strong>of</strong> B. carnaticus is<br />
uninformative, and <strong>the</strong>re is no known surviving<br />
type material. As a result, subsequent authors<br />
have followed Day’s (1878: 563, pl. 137) conception<br />
<strong>of</strong> <strong>the</strong> species. Two <strong>of</strong> Day’s specimens in<br />
<strong>the</strong> AMS collection (B.7963, 103.3 mm SL; B.7626,<br />
230 mm SL) have 28 and 29 lateral-line scales on<br />
<strong>the</strong> body plus 2 on <strong>the</strong> base <strong>of</strong> <strong>the</strong> caudal fin;<br />
1 /2 5 + 1 + 4 scales in transverse line from dorsal-fin<br />
origin <strong>to</strong> pelvic-fin origin; 4 smooth, unbranched<br />
and 8 branched dorsal-fin rays; 3 unbranched and<br />
5 branched anal-fin rays; both rostral and maxillary<br />
barbels; and <strong>the</strong> anal fin, when adpressed,<br />
reaching just beyond <strong>the</strong> hypural fold. This is<br />
additionally a large cyprinid, reported <strong>to</strong> reach a<br />
<strong>to</strong>tal length <strong>of</strong> 60 cm (Talwar & Jhingran, 1991).<br />
It differs from Neolissochilus, however, by having<br />
<strong>the</strong> last unbranched dorsal-fin ray stiff (vs. weak<br />
and segmented: Rainboth, 1985) and <strong>the</strong> lower<br />
jaw exposed medially, which characters distin-<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
87<br />
guish it also from Hypsibarbus and Gonoprok<strong>to</strong>perus,<br />
genera it o<strong>the</strong>rwise resembles superficially.<br />
Its generic allocation remains uncertain.<br />
We hesitate <strong>to</strong> follow Menon (1999) in allocating<br />
Barbus jerdoni <strong>to</strong> Gonoprok<strong>to</strong>pterus (for which<br />
he incorrectly used Hypselobarbus: see Jayaram,<br />
1997). Although <strong>the</strong> AMS syntype <strong>of</strong> G. jerdoni<br />
(B.7935, 179.3 mm SL; Fig. 11) has 9 branched<br />
dorsal-fin rays, <strong>the</strong> last unbranched ray weak,<br />
segmented; rostral and maxillary barbels; and<br />
6 + 1 + 3 scales in transverse line between dorsalfin<br />
origin and pelvic-fin origin, which are consistent<br />
with Gonoprok<strong>to</strong>pterus as presently unders<strong>to</strong>od<br />
(Talwar & Jhingran, 1991; Jayaram, 1997), it possesses<br />
25 lateral-line scales, substantially lower<br />
than <strong>the</strong> 31-45 lateral-line scales observed in<br />
o<strong>the</strong>r species <strong>of</strong> this genus. Its generic placement<br />
<strong>to</strong>o deserves fur<strong>the</strong>r investigation.<br />
Dawkinsia. Arguing against <strong>the</strong> monophyly <strong>of</strong><br />
<strong>the</strong> ‘filamen<strong>to</strong>sus complex’ <strong>of</strong> Jayaram (1991), who<br />
did not point <strong>to</strong> any putative synapomorphies,<br />
Pethiyagoda & Kottelat (2005a) proposed <strong>the</strong>
88<br />
‘<strong>Puntius</strong> filamen<strong>to</strong>sus group’ for those species <strong>of</strong><br />
<strong>Puntius</strong> s. l., <strong>the</strong> juveniles <strong>of</strong> which possess a<br />
unique shared colour pattern <strong>of</strong> three black bars<br />
on <strong>the</strong> side. This was demonstrated first by De<br />
Silva et al. (1981) in a comparative study <strong>of</strong> <strong>the</strong><br />
on<strong>to</strong>geny <strong>of</strong> Dawkinsia singhala and <strong>Dr</strong>avidia fasciata,<br />
since when <strong>the</strong> genus has expanded considerably<br />
(Pethiyagoda & Kottelat, 2005a; Rema Devi<br />
et al., 2010; Marcus Knight et al., 2011). Figure 1<br />
suggests that <strong>the</strong> general coloration <strong>of</strong> <strong>the</strong> sistergenera<br />
Dawkinsia, <strong>Dr</strong>avidia and Pethia derive from<br />
a common ancestry which, taken <strong>to</strong>ge<strong>the</strong>r with<br />
apparent relationships within <strong>the</strong> P. tetrazona<br />
group and P. trifasciatus group, suggests that<br />
coloration could be a relatively conservative<br />
character useful in identifying relationships<br />
among <strong>the</strong>se <strong>fishes</strong> (see also Kortmulder, 1986;<br />
Kortmulder & van der Poll, 1981).<br />
Pethia. Pethia (as <strong>the</strong> ‘<strong>Puntius</strong> conchonius group’)<br />
has been a group <strong>of</strong> <strong>fishes</strong> long recognized as<br />
having a separate identity within <strong>Puntius</strong> as previously<br />
unders<strong>to</strong>od. Following on <strong>the</strong> earlier work<br />
<strong>of</strong> Kortmulder (1972), and <strong>the</strong> osteological studies<br />
<strong>of</strong> Taki et al. (1978), Kullander & Fang (2005)<br />
described <strong>the</strong> ‘<strong>Puntius</strong> conchonius group’ as being<br />
characterized by small adult size, broad infraorbital<br />
3, absence <strong>of</strong> rostral barbels, minute or absent<br />
maxillary barbels, a stiff and serrated last unbranched<br />
dorsal fin ray, an <strong>of</strong>ten abbreviated<br />
lateral line, a colour pattern including a black<br />
blotch or spot anteriorly on <strong>the</strong> side (absent in<br />
P. conchonius) and a black blotch on <strong>the</strong> caudal<br />
peduncle, and a broad 5th cera<strong>to</strong>branchial characterized<br />
by a pointed dorsal tip, spaced moderate-sized<br />
teeth, and absence <strong>of</strong> dorsal angle. Our<br />
results agree broadly with this characterization<br />
except that <strong>the</strong> 5th cera<strong>to</strong>branchial is slender in<br />
P. punctata, and with an obtuse tip in P. nigr<strong>of</strong>asciata,<br />
P. reval and P. cumingii.<br />
Distribution. The present results suggest a<br />
greater degree <strong>of</strong> generic endemism in <strong>South</strong><br />
<strong>Asian</strong> cyprinids than previously suspected. Several<br />
cyprinid genera are already known <strong>to</strong> be<br />
endemic <strong>to</strong> <strong>the</strong> peninsula <strong>of</strong> India, including<br />
Betadevario, Horalabiosa, Gonoprok<strong>to</strong>pterus, Lepidopygopsis,<br />
Parapsilorhynchus and Rohtee. To <strong>the</strong>se<br />
are now added Dawkinsia and <strong>Dr</strong>avidia, which are<br />
restricted <strong>to</strong> Sri Lanka and <strong>the</strong> sou<strong>the</strong>rn region<br />
<strong>of</strong> <strong>the</strong> Indian peninsula. <strong>Puntius</strong> s. s. <strong>to</strong>o, turns out<br />
<strong>to</strong> be largely restricted <strong>to</strong> <strong>South</strong> Asia, only a single<br />
species, P. brevis, extending its range through<br />
Indochina <strong>to</strong> Java. Pethia <strong>to</strong>o, appears mostly<br />
restricted <strong>to</strong> <strong>South</strong> Asia, with only P. s<strong>to</strong>liczkanus<br />
extending its range <strong>to</strong> Laos (Kottelat, 2001) and<br />
Thailand (Smith, 1945, as <strong>Puntius</strong> s<strong>to</strong>liczkae). Likewise,<br />
Sys<strong>to</strong>mus s. s. is represented in Sou<strong>the</strong>ast<br />
Asia only by S. jacobusboehlkei and S. orphoides.<br />
Character states. Some <strong>of</strong> <strong>the</strong> character states we<br />
report here are variable between closely-related<br />
species and even within species. Silva et al. (2008),<br />
for example, reported that one <strong>of</strong> <strong>the</strong> specimens<br />
<strong>of</strong> <strong>Puntius</strong> kamalika <strong>the</strong>y stained possessed a free<br />
uroneural while ano<strong>the</strong>r did not, and that free<br />
uroneurals were al<strong>to</strong>ge<strong>the</strong>r absent in its sister<br />
species, P. mahecola. Similarly, while free uroneurals<br />
are present in each <strong>of</strong> <strong>the</strong> two specimens <strong>of</strong><br />
P. sophore and P. chola cleared and stained by us,<br />
Shantakumar & Vishwanath (2006) reported <strong>the</strong>m<br />
<strong>to</strong> be absent in both <strong>the</strong>se species. Although we<br />
have included this character in <strong>the</strong> generic diagnoses<br />
above, it is important <strong>to</strong> note that it may<br />
be variable.<br />
While <strong>the</strong> deep infraorbital 3 is synapomorphic<br />
in Pethia, this character state occurs also in some<br />
species <strong>of</strong> <strong>Puntius</strong> as unders<strong>to</strong>od here, including<br />
P. bimaculatus, P. kelumi, P. mahecola and P. titteya.<br />
Similarly, character states that are synapomorphic<br />
in o<strong>the</strong>r cyprinine genera sometimes present<br />
<strong>the</strong>mselves as apomorphies in individual species<br />
<strong>of</strong> <strong>Puntius</strong> as presently defined. For example, a<br />
few species retained in <strong>Puntius</strong> possess numerous<br />
rows <strong>of</strong> sensory papillae on <strong>the</strong> head, apparently<br />
an apomorphy shared by <strong>the</strong> closely-related<br />
species-group P. cauveriensis, P. dorsalis, P. kelumi<br />
and P. layardi (Pethiyagoda et al., 2008), but not<br />
o<strong>the</strong>r species <strong>of</strong> <strong>Puntius</strong>. Among <strong>Asian</strong> cyprinine<br />
genera, this character-state occurs also in Cyclocheilichthys,<br />
Eirmotus, Neobarynotus and Oreichthys<br />
(Kottelat, 1996; Tan & Kottelat, 2008; Schäfer,<br />
2009). Likewise, some species <strong>of</strong> Dawkinsia develop<br />
nuptial tubercles on <strong>the</strong> snout and cheeks<br />
(Pethiyagoda & Kottelat, 2005a), a character that<br />
occurs also in many o<strong>the</strong>r cyprinid genera. The<br />
dense tuberculation <strong>of</strong> <strong>the</strong> body in sexually mature<br />
males <strong>of</strong> <strong>Puntius</strong> kelumi, however (Pethiyagoda<br />
et al., 2008), while not reported from any o<strong>the</strong>r<br />
member <strong>of</strong> <strong>the</strong> <strong>Puntius</strong>, occurs in several cyprinid<br />
lineages.<br />
Never<strong>the</strong>less, <strong>the</strong> five genera in<strong>to</strong> which <strong>the</strong><br />
<strong>South</strong> <strong>Asian</strong> species previously in <strong>Puntius</strong> are<br />
now allocated are distinguishable from one ano<strong>the</strong>r<br />
by reliable external characters: <strong>Puntius</strong>,<br />
smooth last unbranched dorsal-fin ray, no rostral<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
arbels, no bars on flank in juveniles; Sys<strong>to</strong>mus,<br />
serrated last unbranched dorsal-fin ray, rostral<br />
barbels present; <strong>Dr</strong>avidia, smooth last unbranched<br />
dorsal-fin ray, rostral barbels present; Dawkinsia,<br />
smooth last unbranched dorsal-fin ray, rostral<br />
barbels absent, black bars on flank in juveniles;<br />
Pethia, serrated last unbranched dorsal-fin ray,<br />
rostral barbels absent. All <strong>the</strong>se species possess<br />
fewer than 40 lateral-line scales.<br />
A key <strong>to</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong> s. l.<br />
The following key allows generic allocation <strong>of</strong><br />
<strong>South</strong> <strong>Asian</strong> <strong>fishes</strong> included in <strong>Puntius</strong> by Jayaram<br />
(1991, 2010), Talwar & Jhingran (1991), Menon<br />
(1999) and authors <strong>of</strong> new species from <strong>the</strong> region<br />
from 1990-2010. Generic names in quotes indicate<br />
provisional assignment pending future revision;<br />
such species are unlikely members <strong>of</strong> <strong>the</strong> stated<br />
genera.<br />
1. – 18-39 scales in lateral line (or if incomplete,<br />
in lateral series) on body.<br />
........................................................................2<br />
– 40-47 scales in lateral line on body.<br />
......................................................................10<br />
2. – Last unbranched dorsal-fin ray serrated<br />
posteriorly.<br />
........................................................................3<br />
– Last unbranched dorsal-fin ray smooth.<br />
........................................................................5<br />
3. – Lateral line complete; a black blotch on<br />
caudal peduncle; adults exceed 80 mm SL.<br />
.......................................................... Sys<strong>to</strong>mus<br />
– Lateral line abbreviated; maximum SL<br />
< 80 mm.<br />
........................................................................4<br />
4. – 19-24 scales in lateral series; one or more<br />
black spots, blotches or bars on body.<br />
................................................................Pethia<br />
– 31-39 scales in lateral series; no black markings<br />
on body.<br />
................‘<strong>Puntius</strong>’ guganio, ‘P.’ nangalensis<br />
5. – Juveniles (< 50 mm SL) with two or more<br />
black bars on body.<br />
........................................................................6<br />
– No black bars on body at any stage.<br />
........................................................................7<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
89<br />
6. – Both rostral and maxillary barbels present.<br />
........................................................... <strong>Dr</strong>avidia<br />
– Rostral barbels absent.<br />
.........................................................Dawkinsia<br />
7. – 9 branched dorsal-fin rays.<br />
........................................................................8<br />
– 7 or 8 branched dorsal-fin rays.<br />
........................................................................9<br />
8. – Last unbranched dorsal-fin ray s<strong>to</strong>ut, heavily<br />
ossified.<br />
...............................‘Neolissochilus’ bovanicus<br />
– Last unbranched dorsal-fin ray weak, segmented.<br />
..............................‘Gonoprok<strong>to</strong>pterus’ jerdoni<br />
9. – Rostral barbels present; when adpressed,<br />
anal fin reaching beyond hypural notch.<br />
........................ ‘Gonoprok<strong>to</strong>pterus’ carnaticus<br />
– Rostral barbels absent; when adpressed,<br />
anal fin falling short <strong>of</strong> hypural notch.<br />
.............................................................<strong>Puntius</strong><br />
10. – Both rostral and maxillary barbels present.<br />
........................... Eechathalakenda ophicephala<br />
– Rostral barbels absent.<br />
......................................................................11<br />
11. – Last unbranched dorsal-fin ray serrated;<br />
maxillary barbels present.<br />
......................................................................12<br />
– Last unbranched dorsal-fin ray smooth;<br />
maxillary barbels absent.<br />
....................................‘<strong>Puntius</strong>’ punjaubensis<br />
12. – Lateral line complete; 9 branched dorsal-fin<br />
rays; 7 branched anal-fin rays.<br />
...................................... ‘<strong>Puntius</strong>’ deccanensis<br />
– Lateral line incomplete; 8 branched dorsalfin<br />
rays; 5 branched anal-fin rays.<br />
......................... ‘<strong>Puntius</strong>’ sharmai, ‘P.’ fraseri<br />
Material examined. Dawkinsia arulius, BMNH 1980.11.<br />
25:73-74, 2, 88.0-90.9 mm SL; BMNH 89.2.1.664-5, 2,<br />
67.1-67.6 mm SL; India: Tamil Nadu: Cauvery River at<br />
Coorg.<br />
Dawkinsia assimilis, WHT 6250, 5, <strong>to</strong>potypes, 43.3-<br />
53.2 mm SL; India: Karnataka: Netravati River, 42 km<br />
from Madikeri on Mangalore-Mysore Road.<br />
Dawkinsia exclamatio, WHT 6255, holotype, 70.4 mm<br />
SL; WHT 6249, 3 paratypes, 62.3-71.9 mm SL; WHT<br />
11056,1, 63.5 mm SL, c&s; WHT 11076, 1, 70.4 mm SL,<br />
c&s; India: Kerala: Kallada River at Thenmalai.
90<br />
Dawkinsia filamen<strong>to</strong>sa: MNHN 3908, lec<strong>to</strong>type,<br />
82.7 mm SL; India: Kerala: Alleppey. – AMS B.7869,<br />
94.9 mm SL; India: Madras. – MNHN 3908, 2 paralec<strong>to</strong>types,<br />
76.4-85.4 mm SL; India: Alleppey. – WHT<br />
11026, 1, 67.1 mm SL, c&s; WHT 11040, 1, 63.0 mm SL,<br />
c&s; India: Kerala: Chalakkudy.<br />
Dawkinsia singhala: ZMH 364, lec<strong>to</strong>type, 24.0 mm<br />
SL; ZMH 365, 2, paralec<strong>to</strong>types, 54.1-63.3 mm SL; WHT<br />
6245, 4, 54.1-63.3 mm SL; Sri Lanka: Gin River at Wakwella.<br />
– WHT 11037, 1, 62.1 mm SL, c&s; WHT 11042,<br />
1, 55.0 mm SL, c&s; Sri Lanka: Morawaka.<br />
Dawkinsia srilankensis, BMNH 1976.2.10.2, holotype,<br />
79.0 mm SL; BMNH 1976.2.10.2, 1, paratype, 63.3 mm<br />
SL; WHT 11053, 1, 61.0 mm SL, c&s; WHT 11063, 1,<br />
60.7 mm SL, c&s; Sri Lanka; 3 miles N <strong>of</strong> Pallegama.<br />
Dawkinsia tambraparniei, ZSI 735, 1, paratype,<br />
68.7 mm SL, India: Tamil Nadu: tributary <strong>of</strong> Tambraparni<br />
River 7 km from Kalladi Kuruchi; ZSI-SRS 4452,<br />
2, 52.9-64.6 mm SL; India: Tamil Nadu: Tirunelveli.<br />
<strong>Dr</strong>avidia fasciata, WHT 11043, 1, 43 mm SL, c&s; WHT<br />
11034, 1, 39.1 mm SL, c&s; India: Kerala: Chalakudy.<br />
Gonoprok<strong>to</strong>pterus carnaticus AMS B.7963, 1, 103.3 mm<br />
SL; AMS B.7626, 1, 225 mm SL; India: Bhavani River,<br />
Tamil Nadu, F. Day.<br />
Gonoprok<strong>to</strong>pterus jerdoni, AMS B.7935, 1, syntype,<br />
179.3 mm SL, India: Canara [Karnatake], F. Day.<br />
Gonoprok<strong>to</strong>pterus kolus AMS B.7518, 1, 153.9 mm SL,<br />
India: Deccan, F. Day.<br />
Gonoprok<strong>to</strong>pterus lithopidos AMS B.8374, 1, syntype,<br />
282 mm SL, India: Canara, F. Day.<br />
Neolissochilus bovanicus, AMS B.7829, lec<strong>to</strong>type,<br />
95.1 mm SL, India: Bhavani River, Tamil Nadu, F. Day.<br />
Neolissochilus wynaadensis AMS B.7989, 1, syntype,<br />
137.1 mm SL,India: Wynaad, F. Day.<br />
Pethia bandula, ZRC 38483, holotype, 34.8 mm SL;<br />
CMK 7146, 7, paratypes, 29.6-38.5 mm SL; Sri Lanka:<br />
Galapitamada.<br />
Pethia conchonius, AMS IA.6883, 41.8 mm SL; aquarium<br />
specimen. – AMS IA.6518, 46.5 mm SL; aquarium<br />
specimen.<br />
Pethia cumingii, BMNH 1859.10.19.101, lec<strong>to</strong>type,<br />
33.4 mm SL; BMNH 1859.5.31.47-48, 3, paralec<strong>to</strong>types;<br />
Sri Lanka. – WHT 7685, 1, 37.0 mm SL, c&s; WHT 11047,<br />
1, 35.6 mm SL, c&s; Sri Lanka: Mawanana.<br />
Pethia melanomaculata, WHT 1681, 2, 37.0-38.7 mm<br />
SL; WHT 218, 2, 30.6-31.8 mm SL; WHT 11066, 1,<br />
62.8 mm SL, c&s; WHT 11068, 1, 37.8 mm SL, c&s; Sri<br />
Lanka: Mahawelia basin: Hasalaka.<br />
Pethia nigr<strong>of</strong>asciata, BMNH 1859.10.19.99-100, 2,<br />
syntypes, 32.1-45.5 mm SL; Sri Lanka. – WHT 7677, 3,<br />
41.0-44.4 mm SL; Kelani River at Kitulgala. – WHT<br />
11022, 1, 35.9 mm SL, c&s; WHT 11023, 1, 35.6 mm SL,<br />
c&s; Sri Lanka, Mawanana.<br />
Pethia punctata, WHT 7716, 3, 35.1-37.0 mm SL;<br />
WHT 11071, 1, 29.3 mm SL, c&s; WHT 11077, 1, 28.7 mm<br />
SL, c&s; India: Kerala: Calicut. – WHT 7715, 4, 29.7-<br />
34.2 mm SL; India: Kerala: Thenmalai.<br />
Pethia reval, WHT 677, holotype, 30.5 mm SL; WHT<br />
7536, 3, 28.8-36.6 mm SL; Sri Lanka: Kelani Basin:<br />
Labugama. – WHT 11033, 1, 25.6 mm SL, c&s; WHT<br />
11044, 1, 28.3 mm SL, c&s; Sri Lanka: Ingiriya.<br />
Pethia s<strong>to</strong>liczkana, AMS B.7542, 1, 34.7 mm SL; India:<br />
West Bengal: Darjeeling.<br />
Pethia tic<strong>to</strong>, WHT 7718, 5, 30.8-34.6 mm SL; India:<br />
West Bengal: Nadia. – AMS B.7974, 2, 49.5-58.6 mm SL;<br />
India: Ganjam. – WHT 11054, 1, 37.0 mm SL, c&s; WHT<br />
11065, 1, 37.4 mm SL, c&s; India: W Bengal: Nadia.<br />
<strong>Puntius</strong> amphibius, MNHN 73, lec<strong>to</strong>type, 92.5 mm<br />
SL; MNHN 2005-0006, 1, paralec<strong>to</strong>type, 74.3 mm SL;<br />
India: Maharashtra: Bombay.<br />
<strong>Puntius</strong> bimaculatus, WHT 79, 2, 37.0-46.2 mm SL;<br />
Sri Lanka: Pallegama. – WHT 1065, 8, 40.2-46.0; Sri<br />
Lanka: Mee Oya [River].<br />
<strong>Puntius</strong> brevis, AMS I.43487-006, 43.2 mm SL; Cambodia:<br />
Tonle Sap River, 3-5 km downstream <strong>of</strong> Kompong<br />
Chhnang.<br />
<strong>Puntius</strong> chola,AMS B.7887, 89.3 mm SL; India: Madras.<br />
<strong>Puntius</strong> everetti, AMS I.30463-001, 3, 32.5-39.4 mm<br />
SL; Malaya.<br />
<strong>Puntius</strong> denisonii, AMS B.7913, 1, 88 mm SL; India:<br />
Travancore Hills.<br />
<strong>Puntius</strong> dorsalis, WHT 2153, 4, 68.4-115.3 mm SL;<br />
India: Tamil Nadu: Mamallapuram. – AMS B.7634,<br />
112.4 mm SL; India: Madras. – WHT 1900, 6, 93.3-133.5;<br />
Sri Lanka: Minneriya. – WHT 7646, 2, 56.7-65.0 mm SL,<br />
c&s; Sri Lanka: Tissamaharamaya.<br />
<strong>Puntius</strong> kamalika, WHT 7639, holotype, 50.2 mm SL;<br />
WHT 7640, 11, paratypes, 39.3-55.4 mm SL; Sri Lanka,<br />
Kalu River at Walandure near Kuruwita.<br />
<strong>Puntius</strong> kelumi, WHT 7629, holotype, 70.4 mm; WHT<br />
7630, 4, 57.2-68.6 mm SL; Sri Lanka: Kalu River:<br />
Madakada Aranya, near Ingiriya. – WHT 7645, 2, 55.1-<br />
63.2 mm SL, c&s; Sri Lanka: Homadola.<br />
<strong>Puntius</strong> layardi: BMNH 1853.12.27.5, syntypes, 2,<br />
92.5-112.7 mm SL; Sri Lanka.<br />
<strong>Puntius</strong> mahecola, MNHN 3896, lec<strong>to</strong>type, 50.7 mm<br />
SL; MNHN 2005-0007, paralec<strong>to</strong>types, 5, 48.7-56.5 mm<br />
SL; India: Kerala: Mahe. – WHT 7681, 2, 56.8-66.8 mm<br />
SL, c&s; India: Kerala: Kottayam.<br />
<strong>Puntius</strong> partipentazona, AMS I.43747-007, 32.1 mm<br />
SL; Cambodia: Tonle Sap River.<br />
<strong>Puntius</strong> sealei, AMS I.22131-006, 9, 45.8-62.4 mm SL;<br />
Northwest Borneo: Kembayung River.<br />
<strong>Puntius</strong> sophore, ZRC 35064, neotype, 71.8 mm SL;<br />
ZRC 35065-35069, 5, 59.4-80.6 mm SL; Bangladesh:<br />
Srimangal, from Hail Hoar floodplain near Moulvi<br />
Bazaar. – WHT 11060, 1, 36.6 mm SL, c&s; WHT 11062,<br />
1, 28.9 mm SL, c&s; India: West Bengal: Boncron.<br />
<strong>Puntius</strong> <strong>the</strong>rmalis, WHT 158, 4, 74.6-76.0 mm SL;<br />
WHT 11027, 1, 55.8 mm SL, c&s; WHT 11035, 1, 48.9 mm<br />
SL, c&s; Sri Lanka: Attidiya.<br />
<strong>Puntius</strong> titteya, WHT 8013, 1, 32.1 mm SL, c&s; WHT<br />
8014, 1, 44.2 mm SL, c&s; Sri Lanka: Kelani River near<br />
Giniga<strong>the</strong>na.<br />
<strong>Puntius</strong> vittatus, WHT 11036, 1, 27.5 mm SL, c&s;<br />
WHT 11038, 1, 28.5 mm SL, c&s; Sri Lanka: Navinna.<br />
Sys<strong>to</strong>mus immaculatus, AMS B.7920, neotype,<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
186.9 mm SL; AMS B.7921, 1, <strong>to</strong>potype, 192.7 mm SL;<br />
India: Assam.<br />
Sys<strong>to</strong>mus martenstyni, ZRC 38482, holotype,<br />
136.0 mm SL; CMK 7168, 11, 67.2-116.8 mm SL; Sri<br />
Lanka: 2 km N <strong>of</strong> Pallegama.<br />
Sys<strong>to</strong>mus orphoides, AMS I.43487-062, 2, 106-136 mm<br />
SL; Cambodia: Tonle Sap River, 3-5 km downstream<br />
<strong>of</strong> Kompong Chhnang.<br />
Sys<strong>to</strong>mus pleurotaenia, WHT 11031, 1, 66.2 mm SL,<br />
c&s; Sri Lanka: Akuressa.<br />
Sys<strong>to</strong>mus sp. ‘Richmondi’ [manuscript name for an<br />
as yet undescribed species], WHT 9861, 6, 121.5-<br />
127.0 mm SL; Sri Lanka: Elahera.<br />
Sys<strong>to</strong>mus spilurus, WHT 11028, 1, 61.1 mm SL; Sri<br />
Lanka, Kuruwita.<br />
Sys<strong>to</strong>mus timbiri, NMSL FF801, holotype, 178 mm<br />
SL; Sri Lanka: Timbirigasmankada. – WHT 11107, 5,<br />
127-200 mm SL; Sri Lanka: Walawe River.<br />
Acknowledgements<br />
We are grateful <strong>to</strong> <strong>the</strong> Department <strong>of</strong> Wildlife Conservation,<br />
Sri Lanka, for permitting export <strong>of</strong> tissue samples<br />
for molecular analysis; Kelum Manamendra-Arachchi<br />
and Mohomed Bahir for assisting with <strong>the</strong> collection <strong>of</strong><br />
material in Sri Lanka and India; Andrew Rao for supplying<br />
samples from West Bengal; and Sudath Nanayakkara<br />
for hospitality at <strong>the</strong> WHT field station at<br />
Agarapatana, Sri Lanka. MM thanks Chris<strong>to</strong>pher<br />
S. Schnei der for access <strong>to</strong> labora<strong>to</strong>ry facilities at Bos<strong>to</strong>n<br />
University, and RP thanks Mark McGrou<strong>the</strong>r and<br />
Amanda Hay (AMS) for labora<strong>to</strong>ry facilities and permission<br />
<strong>to</strong> access material in his care and Kelvin Lim (ZRC)<br />
for <strong>the</strong> loan <strong>of</strong> material; James Maclaine (BMNH) and<br />
Horst Zetzsche (SMF) for information on types; Gina<br />
Douglas (Linnean Society <strong>of</strong> London) for permission <strong>to</strong><br />
reproduce Fig. 2b; Johnny Jensen for <strong>the</strong> pho<strong>to</strong>graph <strong>of</strong><br />
D. fasciata in Fig. 1; and Maurice Kottelat, Kevin Conway,<br />
Shrimati K. Rema Devi and Sri W. Vishwanath for<br />
discussion and literature. Finally, it is a pleasure <strong>to</strong><br />
thank Maurice Kottelat and an anonymous reviewer<br />
for valuable comments and criticism that helped substantially<br />
<strong>to</strong> improve <strong>the</strong> manuscript.<br />
Literature cited<br />
Ahl, E. 1929. Übersicht über die lebend eingeführten<br />
asiatischen Arten der Gattung Barbus. Teil 3. Das<br />
Aquarium, 1929: 165-169.<br />
Arunachalam, M. 2000. Assemblage structure <strong>of</strong> stream<br />
<strong>fishes</strong> in <strong>the</strong> Western Ghats (India). Hydrobiologia,<br />
430: 1-31.<br />
Arunachalam, M. & J. A. Johnson. 2003. A new species<br />
<strong>of</strong> <strong>Puntius</strong> Hamil<strong>to</strong>n (Pisces: Cyprinidae) from<br />
Kalakad Mundanthurai Tiger Reserve, Tamil Nadu,<br />
India. Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry Society,<br />
99 (2002 [2003]): 474-480.<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
91<br />
Arunkumar, L. & H. T. Tombi Singh. 2003. Two new<br />
species <strong>of</strong> puntiid fish from <strong>the</strong> Yu River system <strong>of</strong><br />
Manipur. Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry<br />
Society, 99: 481-487.<br />
Banarescu, P. 1997. The status <strong>of</strong> some nominal genera<br />
<strong>of</strong> Eurasian Cyprinidae (Osteichthyes, Cypriniformes).<br />
Revue Roumaine de Biologie, Série de<br />
Biologie Animale, 42: 19-30.<br />
Bhat, A. 2005. Ecomorphological correlates in tropical<br />
stream <strong>fishes</strong> <strong>of</strong> sou<strong>the</strong>rn India. Environmental<br />
Biology <strong>of</strong> Fishes, 73: 211-225.<br />
Bleeker, P. 1849. Bijdrage <strong>to</strong>t de kennis der ichthyologische<br />
fauna van Midden- en Oost-Java, met beschrijving<br />
van eenige nieuwe species. Verhandelingen<br />
van het Bataviaasch Genootschap van Kunsten<br />
en Wetenschappen, 23 (12): 1-23.<br />
— 1853. Diagnostische beschrijvingen van nieuwe <strong>of</strong><br />
weinig bekende vischsoorten van Sumatra. Tiental<br />
V-X. Natuurkundig Tijdschrift voor Nederlandsch<br />
Indië, 4: 243-302.<br />
— 1855. Nalezingen op de vischfauna van Sumatra.<br />
Visschen van Lahat en Sibogha. Natuurkundig<br />
Tijdschrift voor Nederlandsch Indië, 9: 257-280.<br />
— 1863a. Description de quelques espèces nouvelles<br />
de Cyprinoïdes du Ceylon. Verslagen en Mededeelingen<br />
der Koninklijke Akademie van Wetenschappen.<br />
Afdeling Natuurkunde, 15: 239-253.<br />
— 1863b-1864. Atlas ichthyologique des Indes Orientales<br />
Néêrlandaises. Tome III. Cyprins. Müller,<br />
Amsterdam, 1863: pp. 1-48, pls. 102-132, 1864: pp.<br />
49-150, pls. 133-144.<br />
Bossuyt, F., M. Meegaskumbura, N. Beenaerts, D. J.<br />
Gower, R. Pethiyagoda, K. Roelants, A. Mannaert,<br />
M. Wilkinson, M. M. Bahir, K. Manamendra-<br />
Arachchi, P. K. L. Ng, C. J. Schneider, O. V. Oommen<br />
& M. C. Milinkovitch. 2004. Local endemism<br />
within <strong>the</strong> Western Ghats-Sri Lanka Biodiversity<br />
Hotspot. Science, 306: 479-481.<br />
Bossuyt, F. & M. C. Milinkovitch. 2000. Convergent<br />
adaptive radiation in Madagascan and <strong>Asian</strong> ranid<br />
frogs reveal covariation between larval and adult<br />
traits. Proceedings <strong>of</strong> <strong>the</strong> National Academy <strong>of</strong><br />
Sciences <strong>of</strong> <strong>the</strong> United States <strong>of</strong> America, 97: 6585-<br />
6590.<br />
Boulenger, G. A. 1894. Descriptions <strong>of</strong> new freshwater<br />
<strong>fishes</strong> from Borneo. Annals and Magazine <strong>of</strong> Natural<br />
His<strong>to</strong>ry, Series 6, 13 (75): 245-251.<br />
Chhapgar, B. F. & S. R. Sane. 1992. A new fish <strong>of</strong> <strong>the</strong><br />
genus <strong>Puntius</strong> Hamil<strong>to</strong>n (Ostariophysi: Cyprinidae)<br />
from Goa. Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry<br />
Society, 89: 357-359.<br />
Cockerell, T. D. A. 1913. Observations on fish scales.<br />
Bulletin <strong>of</strong> <strong>the</strong> Bureau <strong>of</strong> Fisheries, 32: 117-174, pls.<br />
32-40.<br />
Collins, R. A., K. F. Armstrong, R. Meier, Y. Yi, S. D. J.<br />
Brown, R. H. Cruickshank, S. Keeling & C. Johns<strong>to</strong>n.<br />
2012. Barcoding and border biosecurity: identifying<br />
cyprinid <strong>fishes</strong> in <strong>the</strong> aquarium trade. PLoS ONE,<br />
7 (1): e28381. doi:10.1371/journal.pone.0028381
92<br />
Conway, K. W. 2011. Osteology <strong>of</strong> <strong>the</strong> <strong>South</strong> <strong>Asian</strong><br />
genus Psilorhynchus McClelland, 1839 Teleostei:<br />
Ostariophysi: Psilorhynchidae), with investigation<br />
<strong>of</strong> its phylogenetic relationships within <strong>the</strong> order<br />
Cypriniformes. Zoological Journal <strong>of</strong> <strong>the</strong> Linnean<br />
Society, 163: 150-154.<br />
Cuvier, G. & A. Valenciennes. 1842. His<strong>to</strong>ire naturelle<br />
des poissons. Tome seizième. Bertrand, Paris,<br />
xx + 472 pp., pls. 456-487.<br />
Cuvier, G. & A. Valenciennes. 1844. His<strong>to</strong>ire naturelle<br />
des poissons. Tome dix-septième. Bertrand, Paris,<br />
xxiii + 497 pp., pls. 487-519.<br />
Day, A. L. 1914. Two new cyprinoid <strong>fishes</strong> <strong>of</strong> <strong>the</strong> genus<br />
Barbus from Lake Manguao, Palawan, P. I. Philippine<br />
Journal <strong>of</strong> Science, 9 (Sect. D, 2): 187-192,<br />
pl. 1.<br />
Day, F. 1865. On <strong>the</strong> <strong>fishes</strong> <strong>of</strong> Cochin, on <strong>the</strong> Malabar<br />
Coast <strong>of</strong> India. Part II. Anacanthini. Proceedings <strong>of</strong><br />
<strong>the</strong> Zoological Society <strong>of</strong> London, 1865: 286-318.<br />
— 1871. Monograph <strong>of</strong> Indian Cyprinidae (part 1-3).<br />
Journal and Proceedings <strong>of</strong> <strong>the</strong> Asiatic Society <strong>of</strong><br />
Bengal, 40: 95-142, 277-367, 337-367, pls. 9, 21-23.<br />
— 1872. Monograph <strong>of</strong> Indian Cyprinidae (parts 4-6).<br />
Journal and Proceedings <strong>of</strong> <strong>the</strong> Asiatic Society <strong>of</strong><br />
Bengal, 41: 1-29, 171-198; 318-326, pl. 1.<br />
— 1875-78. The <strong>fishes</strong> <strong>of</strong> India; being a natural his<strong>to</strong>ry<br />
<strong>of</strong> <strong>the</strong> <strong>fishes</strong> known <strong>to</strong> inhabit <strong>the</strong> seas and<br />
fresh waters <strong>of</strong> India, Burma, and Ceylon. Quaritsch,<br />
London, (1875): 1-168, pls. 1-40; (1876): 169-368,<br />
pls. 41-78; (1877): 369-552, pls. 79-138; (1878): i-xx,<br />
553-778, pls. 139-195.<br />
De Silva, M. P. K. S. K. & N. P. P. Liyanage. 2010. A<br />
multivariate approach for developing a dicho<strong>to</strong>mous<br />
key for identification and differentiation <strong>of</strong><br />
<strong>Puntius</strong> (Osteichthyes: Cyprinidae) species in Sri<br />
Lanka. Journal <strong>of</strong> <strong>the</strong> National Science Foundation<br />
<strong>of</strong> Sri Lanka, 38: 15-27.<br />
De Silva, S. S., K. Kortmulder & P. Maitipe. 1981. The<br />
identity <strong>of</strong> <strong>Puntius</strong> melanampyx singhala (Duncker,<br />
1911) (Pisces, Cyprinidae). Ne<strong>the</strong>rlands Journal <strong>of</strong><br />
Zoology, 31: 777-785.<br />
Deraniyagala, P. E. P. 1929. Two new freshwater <strong>fishes</strong>.<br />
Ceylon Journal <strong>of</strong> Science (B), 15: 73-77, pls. 21-22.<br />
— 1956. Two new subspecies and one new species <strong>of</strong><br />
cyprinoid <strong>fishes</strong> from Ceylon. Proceedings <strong>of</strong> <strong>the</strong><br />
12th Annual Sessions <strong>of</strong> <strong>the</strong> Ceylon Association for<br />
<strong>the</strong> Advancement <strong>of</strong> Science, 1: 34-35.<br />
— 1963. A new cyprinid <strong>Puntius</strong> timbiri from Ceylon.<br />
Spolia Zeylanica, 30: 63-64, pl. 1.<br />
Duncker, G. 1904. Die Fische der malayischen Halbinsel.<br />
Jahrbuch der Hamburgischen Wissenschaftlichen<br />
Anstalten, 2. Beiheft, Mitteilungen aus dem Naturhis<strong>to</strong>rischen<br />
Museum in Hamburg, 21: 133-207,<br />
2 pls.<br />
Duncker, G. 1912. Die Süsswasserfische Ceylons. Jahrbuch<br />
der Hamburgischen Wissenschaftlichen Anstalten,<br />
2. Beiheft, Mitteilungen aus dem Naturhis<strong>to</strong>rischen<br />
Museum in Hamburg, 29 (1911 [1912]):<br />
241-272.<br />
Fowler, H. W. 1934a. Zoological results <strong>of</strong> <strong>the</strong> third De<br />
Schauensee Siamese Expedition, part 5: additional<br />
<strong>fishes</strong>. Proceedings <strong>of</strong> <strong>the</strong> Academy <strong>of</strong> Natural Sciences<br />
<strong>of</strong> Philadelphia, 86: 335-352.<br />
— 1934b. Descriptions <strong>of</strong> new <strong>fishes</strong> obtained 1907 <strong>to</strong><br />
1910, chiefly in <strong>the</strong> Philippine Islands and adjacent<br />
seas. Proceedings <strong>of</strong> <strong>the</strong> Academy <strong>of</strong> Natural Sciences<br />
<strong>of</strong> Philadelphia, 85: 233-367.<br />
— 1958. Some new taxonomic names <strong>of</strong> fishlike vertebrates.<br />
Notulae Naturae (Philadelphia), 310:<br />
1-16.<br />
Gün<strong>the</strong>r, A. 1868. Catalogue <strong>of</strong> <strong>the</strong> <strong>fishes</strong> in <strong>the</strong> British<br />
Museum, vol. 7. British Museum, London. i-xx +<br />
1-512 pp.<br />
Hamil<strong>to</strong>n, F. 1822. An account <strong>of</strong> <strong>the</strong> <strong>fishes</strong> found in<br />
<strong>the</strong> river Ganges and its branches. Constable, Edinburgh<br />
& London, vii + 405 pp, 39 pls.<br />
Herre, A. W. C. T. 1924a. Distribution <strong>of</strong> <strong>the</strong> true freshwater<br />
<strong>fishes</strong> in <strong>the</strong> Philippines. I. The Philippine<br />
Cyprinidae. Philippine Journal <strong>of</strong> Science, 24: 249-<br />
307, pls. 1-2.<br />
— 1924b. The distribution <strong>of</strong> true fresh-water <strong>fishes</strong><br />
in <strong>the</strong> Philippines and its significance. Pp. 1561-1570<br />
in: G. Lightfoot (ed.), Proceedings <strong>of</strong> <strong>the</strong> Pan-Pacific<br />
Science Congress, Australia, 1923. Vol. 2. Melbourne.<br />
— 1932. Five new Philippine <strong>fishes</strong>. Copeia, 1932:<br />
139-142.<br />
— 1940. New species <strong>of</strong> <strong>fishes</strong> from <strong>the</strong> Malay Peninsula<br />
and Borneo. Bulletin <strong>of</strong> <strong>the</strong> Raffles Museum,<br />
16: 5-26, pl. 1-20.<br />
Herre, A. W. C. T. & G. S. Myers. 1931. Fishes from<br />
sou<strong>the</strong>astern China and Hainan. Lingnan Science<br />
Journal, 10: 233-254.<br />
Hora, S. L. 1937. Notes on <strong>fishes</strong> in <strong>the</strong> Indian Museum,<br />
XXVIII. On three collections <strong>of</strong> fish from Mysore<br />
and Coorg, south India. Records <strong>of</strong> <strong>the</strong> Indian<br />
Museum, 39: 5-28.<br />
— 1942. Fishes <strong>of</strong> Poona, Part 2. Journal <strong>of</strong> <strong>the</strong> Bombay<br />
Natural His<strong>to</strong>ry Society, 43: 218-225.<br />
Hora, S. L. & K. S. Misra. 1938. Fish <strong>of</strong> Deolali, Part III.<br />
Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry Society, 40:<br />
20-38, pls. 1-3.<br />
Hora, S. L. & D. D. Mukerji. 1934. Notes on <strong>fishes</strong> in <strong>the</strong><br />
Indian Museum. XXIII. On a collection <strong>of</strong> fish from<br />
<strong>the</strong> S. Shan States, Burma. Records <strong>of</strong> <strong>the</strong> Indian<br />
Museum, 36: 353-370.<br />
Huelsenbeck, J. P. & F. Ronquist. 2001. MRBAYES:<br />
Bayesian inference <strong>of</strong> phylogenetic trees. Bioinformatics,<br />
17: 754-755.<br />
Huelsenbeck, J. P., F. Ronquist, R. Neilsen & J. P. Bollback.<br />
2001. Bayesian inference <strong>of</strong> phylogeny and its<br />
impact on evolutionary biology. Science, 294: 2310-<br />
2314.<br />
ICZN [International Commission <strong>of</strong> Zoological Nomenclature].<br />
1985. International Code <strong>of</strong> Zoological<br />
Nomenclature [3rd ed.]. International Trust for<br />
Zoological Nomenclature, London, 338 pp.<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
— 1999. International Code <strong>of</strong> Zoological Nomenclature<br />
[4th ed.]. International Trust for Zoological<br />
Nomenclature, London, 306 pp.<br />
Inger, R. F. & P. K. Chin. 1962. The freshwater <strong>fishes</strong> <strong>of</strong><br />
North Borneo. Fieldiana: Zoology, 45: 1-268.<br />
Jameela Beevi, K. S. & A. Ramachandran. 2005. A new<br />
species <strong>of</strong> <strong>Puntius</strong> (Cyprinidae, Cyprininae) from<br />
Kerala, India. Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry<br />
Society, 102: 83-85.<br />
Jayaram, K. C. 1990. Two new species <strong>of</strong> <strong>the</strong> genus<br />
<strong>Puntius</strong> Hamil<strong>to</strong>n (Pisces: Cyprinidae) from India.<br />
Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry Society, 87:<br />
106-109.<br />
— 1991. Revision <strong>of</strong> <strong>the</strong> genus <strong>Puntius</strong> Hamil<strong>to</strong>n from<br />
<strong>the</strong> Indian Region. Records <strong>of</strong> <strong>the</strong> Zoological Survey<br />
<strong>of</strong> India, Occasional Paper, 135: 1-178.<br />
— 1997. Nomenclatural and systematic status <strong>of</strong> Barbus<br />
mussallah Sykes, 1839. Journal <strong>of</strong> <strong>the</strong> Bombay Natural<br />
His<strong>to</strong>ry Society, 94: 48-55.<br />
— 2010. The freshwater <strong>fishes</strong> <strong>of</strong> <strong>the</strong> Indian region<br />
(2nd ed.). Narendra Publishing House, Delhi.<br />
xxxi + 616 pp., 39 pls.<br />
Jeanmougin, F., J. D. Thompson, M. Gouy, D. G. Higgins<br />
& T. J. Gibson. 1998. Multiple sequence alignment<br />
with Clustal X. Trends in Biochemical Sciences, 23:<br />
403-405.<br />
Jerdon, T. C. 1849. On <strong>the</strong> fresh-water <strong>fishes</strong> <strong>of</strong> sou<strong>the</strong>rn<br />
India. Madras Journal <strong>of</strong> Literature and Science, 15:<br />
302-346.<br />
Jordan, D. S. 1919. The genera <strong>of</strong> <strong>fishes</strong>, part II, from<br />
Agassiz <strong>to</strong> Bleeker, 1833-1858, twenty-six years,<br />
with <strong>the</strong> accepted type <strong>of</strong> each. A contribution <strong>to</strong><br />
<strong>the</strong> stability <strong>of</strong> scientific nomenclature. Leland<br />
Stanford Jr. University Publications, University<br />
Series, 36: i-ix + 163-284 + i-xiii.<br />
Jordan, D. S. & R. E. Richardson. 1908. Fishes from islands<br />
<strong>of</strong> <strong>the</strong> Philippine Archipelago. Bulletin <strong>of</strong> <strong>the</strong><br />
Bureau <strong>of</strong> Fisheries, 27: 233-287.<br />
Knight, J. D. M., K. Rema Devi & V. Atkore. 2011. Systematic<br />
status <strong>of</strong> Sys<strong>to</strong>mus rubrotinctus Jerdon<br />
(Teleostei: Cyprinidae) with notes on <strong>the</strong> <strong>Puntius</strong><br />
arulius group <strong>of</strong> <strong>fishes</strong>. Journal <strong>of</strong> Threatened Taxa,<br />
3: 1686-1693.<br />
Knight, J. D. M., K. Rema Devi, T.J. Indra & M. Arunachalam.<br />
2012. A new species <strong>of</strong> barb <strong>Puntius</strong> nigripinnis<br />
(Teleostei: Cyprinidae) from sou<strong>the</strong>rn<br />
Western Ghats, India. Journal <strong>of</strong> Threatened Taxa,<br />
4: 2409-2416.<br />
Kortmulder, K. 1972. A comparative study in colour<br />
patterns and behaviour in seven Asiatic Barbus<br />
species (Cyprinidae, Ostariophysi, Osteichthyes): a<br />
progress report. Behaviour, Supplement, 19: i-xiii,<br />
1-331.<br />
Kortmulder, K. 1986. Similar behaviour and colour<br />
patterns in 3 not closely related Barbus species. Is<br />
evolutionary convergence a likely explanation?<br />
(Barbus nigr<strong>of</strong>asciatus Gün<strong>the</strong>r, 1868, B. filamen<strong>to</strong>sus<br />
(Cuv. & Val. 1844), B. lateristriga Cuv. & Val. 1842;<br />
Pisces, Cyprinidae). Behaviour, 98: 180-212.<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
93<br />
Kortmulder, K. & R. J. van der Poll. 1981. The juvenile<br />
and adult pigment patterns <strong>of</strong> Barbus lateristriga<br />
Cuv. & Val. 1842, B. titteya (Deraniyagala 1929) and<br />
B. narayani Hora, 1927 (Pisces, Cyprinidae), and<br />
<strong>the</strong>ir taxonomic value. Ne<strong>the</strong>rlands Journal <strong>of</strong> Zoology,<br />
31: 453-465.<br />
Kottelat, M. 1982. A small collection <strong>of</strong> fresh-water<br />
<strong>fishes</strong> from Kalimantan, Borneo, with descriptions<br />
<strong>of</strong> one new genus and three new species <strong>of</strong> Cyprinidae.<br />
Revue Suisse de Zoologie, 89: 419-437.<br />
— 1992. The identity <strong>of</strong> Barbus johorensis Duncker, 1904<br />
(Teleostei: Cyprinidae). Raffles Bulletin <strong>of</strong> Zoology,<br />
40: 187-192.<br />
— 1996. The identity <strong>of</strong> <strong>Puntius</strong> eugrammus and diagnoses<br />
<strong>of</strong> two new species <strong>of</strong> striped barbs (Teleostei:<br />
Cyprinidae) from Sou<strong>the</strong>ast Asia. Raffles Bulletin<br />
<strong>of</strong> Zoology, 44: 301-316.<br />
— 1999. Nomenclature <strong>of</strong> <strong>the</strong> genera Barbodes, Cyclocheilichthys,<br />
Rasbora and Chonerhinos (Teleostei:<br />
Cyprinidae and Tetraodontidae), with comments<br />
on <strong>the</strong> definition <strong>of</strong> <strong>the</strong> first reviser. Raffles Bulletin<br />
<strong>of</strong> Zoology, 47: 591-600.<br />
— 2000. Diagnoses <strong>of</strong> a new genus and 64 new species<br />
<strong>of</strong> <strong>fishes</strong> from Laos (Teleostei: Cyprinidae, Bali<strong>to</strong>ridae,<br />
Bagridae, Syngnathidae, Chaudhuriidae and<br />
Tetraodontidae). Journal <strong>of</strong> <strong>South</strong> <strong>Asian</strong> Natural<br />
His<strong>to</strong>ry, 5: 37-82.<br />
— 2001. Fishes <strong>of</strong> Laos. WHT Publications, Colombo,<br />
198 pp., 48 pls.<br />
Kottelat, M. & K. K. P. Lim. 1996. Freshwater <strong>fishes</strong> <strong>of</strong><br />
Sarawak and Brunei Darussalam: a preliminary<br />
annotated check-list. Sarawak Museum Journal, 48<br />
(1995 [1996]): 227-256.<br />
Kottelat, M. & H. H. Tan. 2011. Sys<strong>to</strong>mus xouthos, a new<br />
cyprinid fish from Borneo, and revalidation <strong>of</strong><br />
<strong>Puntius</strong> pulcher (Teleostei: Cyprinidae). Ichthyological<br />
Exploration <strong>of</strong> Freshwaters, 22: 209-214.<br />
Kottelat, M. & R. Pethiyagoda. 1989. Eine neue Barbenart<br />
von Sri Lanka: <strong>Puntius</strong> asoka spec. nov. Die Aquarium<br />
und Terrarium Zeitschrift, 42: 472-476.<br />
Kottelat, M. & R. Pethiyagoda. 1991. Descriptions <strong>of</strong><br />
three new species <strong>of</strong> cyprinid <strong>fishes</strong> from Sri Lanka.<br />
Pp. 298-313 in: R. Pethiyagoda, Freshwater <strong>fishes</strong><br />
<strong>of</strong> Sri Lanka. Wildlife Heritage Trust <strong>of</strong> Sri Lanka,<br />
Colombo.<br />
Kottelat, M., A. J., Whitten, S. N. Kartikasari & S. Wirjoatmodjo.<br />
1993. Freshwater <strong>fishes</strong> <strong>of</strong> Western Indonesia<br />
and Sulawesi. Periplus, Hong Kong, xxxviii<br />
+ 259 pp, 84 pls.<br />
Kottelat, M. & E. Widjanarti. 2005. The <strong>fishes</strong> <strong>of</strong> Danau<br />
Sentarum National Park and <strong>the</strong> Kapuas Lakes area,<br />
Kalimantan Barat, Indonesia. Raffles Bulletin <strong>of</strong><br />
Zoology, Supplement, 13: 139-173.<br />
Koumans, F. P. 1940. On a new species <strong>of</strong> <strong>Puntius</strong> from<br />
Borneo. Temminckia, 5: 189-190.<br />
Kumar, K. K., F. G. B. Pereira & K. V. Radhakrishnan.<br />
2011. <strong>Puntius</strong> madhusoodani (Teleostei: Cyprinidae),<br />
a new species <strong>of</strong> barb from Manimala River, Kerala,<br />
<strong>South</strong> India. Biosystematica, 5: 31-37.
94<br />
Kullander, S. O. 2008. Five new species <strong>of</strong> <strong>Puntius</strong> from<br />
Myanmar (Teleostei: Cyprinidae). Ichthyological<br />
Exploration <strong>of</strong> Freshwaters, 19: 59-84.<br />
Kullander, S. O. & R. Britz. 2008. <strong>Puntius</strong> padamya, a<br />
new species <strong>of</strong> cyprinid fish from Myanmar (Teleostei:<br />
Cyprinidae). Electronic Journal <strong>of</strong> Ichthyology,<br />
2: 56-66.<br />
Kullander, S. O. & F. Fang. 2005. Two new species <strong>of</strong><br />
<strong>Puntius</strong> from nor<strong>the</strong>rn Myanmar (Teleostei: Cyprinidae).<br />
Copeia, 2005: 290-302.<br />
Linthoingambi, I. & W. Vishwanath. 2007. Two new<br />
fish species <strong>of</strong> <strong>the</strong> genus <strong>Puntius</strong> Hamil<strong>to</strong>n (Cyprinidae)<br />
from Manipur, India, with notes on P. tic<strong>to</strong><br />
(Hamil<strong>to</strong>n) and P. s<strong>to</strong>liczkanus (Day). Zootaxa, 1450:<br />
45-56.<br />
Marcus Knight, J. D., K. Rema Devi & V. Atkore 2011.<br />
Systematic status <strong>of</strong> Sys<strong>to</strong>mus rubrotinctus Jerdon<br />
(Teleostei: Cyprinidae) with notes on <strong>the</strong> <strong>Puntius</strong><br />
arulius group <strong>of</strong> <strong>fishes</strong>. Journal <strong>of</strong> Threatened Taxa,<br />
3: 1686-1693.<br />
M’Clelland, J. 1839. Indian Cyprinidae. Asiatic Researches,<br />
19: 217-471, pls. 37-61.<br />
McClelland, J. 1838. Observations on six new species <strong>of</strong><br />
Cyprinidae, with an outline <strong>of</strong> a new classification<br />
<strong>of</strong> <strong>the</strong> family. Journal <strong>of</strong> <strong>the</strong> Asiatic Society <strong>of</strong> Bengal,<br />
7: 941-948, pls. 55-56.<br />
— 1844. Description <strong>of</strong> four species <strong>of</strong> <strong>fishes</strong> from <strong>the</strong><br />
rivers at <strong>the</strong> foot <strong>of</strong> <strong>the</strong> Boutan Mountains. Calcutta<br />
Journal <strong>of</strong> Natural His<strong>to</strong>ry, 5: 274-282.<br />
Meegaskumbura, M., A. Silva, K. Maduwage & R.<br />
Pethiyagoda. 2008. <strong>Puntius</strong> reval, a new barb from<br />
Sri Lanka (Teleostei: Cyprinidae). Ichthyological<br />
Exploration <strong>of</strong> Freshwaters, 19: 141-152.<br />
Menon, A. G. K. 1999. Check list – fresh water <strong>fishes</strong> <strong>of</strong><br />
India. Records <strong>of</strong> <strong>the</strong> Zoological Survey <strong>of</strong> India,<br />
Miscellaneous Publication, Occasional Paper, 175:<br />
i-xxviii + 1-366.<br />
Menon, A. G. K. & K. Rema Devi. 1992. <strong>Puntius</strong> mudumalaiensis,<br />
a new cyprinid fish from Mudumalai,<br />
Tamil Nadu. Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry<br />
Society, 89: 229-231.<br />
Menon, A. G. K. & K. Rema Devi. 1993. <strong>Puntius</strong> sharmai,<br />
a new cyprinid fish from Madras. Journal <strong>of</strong> <strong>the</strong><br />
Bombay Natural His<strong>to</strong>ry Society, 89: 353-354.<br />
Menon, A. G. K., K. Rema Devi & M. P. Thobias.1999.<br />
<strong>Puntius</strong> chalakkudiensis, a new colourful species <strong>of</strong><br />
<strong>Puntius</strong> (family: Cyprinidae) fish from Kerala, south<br />
India. Records <strong>of</strong> <strong>the</strong> Zoological Survey <strong>of</strong> India,<br />
97 (4): 61-63.<br />
Menon, A. G. K., K. Rema Devi & W. Vishwanath. 2000.<br />
A new species <strong>of</strong> <strong>Puntius</strong> (Cyprinidae: Cyprininae)<br />
from Manipur, India. Journal <strong>of</strong> <strong>the</strong> Bombay Natural<br />
His<strong>to</strong>ry Society, 97: 263-268.<br />
Mercy, T. V. A. & E. Jacob. 2007. A new species <strong>of</strong><br />
Teleostei: Putius pookodensis (Cyprinidae) from<br />
Wayanad, Kerala, India. Journal <strong>of</strong> <strong>the</strong> Bombay<br />
Natural His<strong>to</strong>ry Society, 104: 76-78.<br />
Ng, H. H. & H. H. Tan. 1999. The <strong>fishes</strong> <strong>of</strong> <strong>the</strong> Endau<br />
drainage, Peninsular Malaysia with descriptions <strong>of</strong><br />
two new species <strong>of</strong> cat<strong>fishes</strong> (Teleostei: Akysidae,<br />
Bagridae). Zoological Studies, 38: 350-366.<br />
Palumbi, S. R. 1996. Nucleic acids II: The polymerase<br />
chain reaction. Pp. 205-248 in: D. M. Hillis, C. Moritz,<br />
and B. K. Mable (Eds.), Molecular systematics.<br />
Sinauer Associates, Sunderland.<br />
Parenti, L. R. & K. K. P. Lim. 2005. Fishes <strong>of</strong> <strong>the</strong> Rajang<br />
Basin, Sarawak, Malaysia. Raffles Bulletin <strong>of</strong> Zoology,<br />
Supplement, 13: 175-208.<br />
Pethiyagoda, R. & M. Kottelat. 2005a. A review <strong>of</strong> <strong>the</strong><br />
barbs <strong>of</strong> <strong>the</strong> <strong>Puntius</strong> filamen<strong>to</strong>sus group (Teleostei:<br />
Cyprinidae) <strong>of</strong> sou<strong>the</strong>rn India and Sri Lanka. Raffles<br />
Bulletin <strong>of</strong> Zoology, Supplement, 12: 127-144.<br />
Pethiyagoda, R. & M. Kottelat. 2005b.The identity <strong>of</strong><br />
<strong>the</strong> south Indian barb <strong>Puntius</strong> mahecola (Teleostei:<br />
Cyprinidae). Raffles Bulletin <strong>of</strong> Zoology, Supplement,<br />
12: 145-152.<br />
Pethiyagoda, R., A. Silva, K. Maduwage & M. Meegaskumbura.<br />
2008. <strong>Puntius</strong> kelumi, a new species <strong>of</strong><br />
cyprinid fish from Sri Lanka (Teleostei: Cyprinidae).<br />
Ichthyological Exploration <strong>of</strong> Freshwaters, 19: 201-<br />
214.<br />
Pillay, T. V. R. 1951. A morphometric and biometric<br />
study <strong>of</strong> <strong>the</strong> systematics <strong>of</strong> certain allied species <strong>of</strong><br />
<strong>the</strong> genus Barbus Cuv. & Val. Proceedings <strong>of</strong> <strong>the</strong><br />
National Institute <strong>of</strong> Science, India, 17: 331-348.<br />
Rainboth, W. J. 1985. Neolissochilus, a new genus <strong>of</strong> <strong>South</strong><br />
<strong>Asian</strong> cyprinid <strong>fishes</strong>. Beaufortia, 35: 25-35.<br />
— 1996. FAO species identification field guide for<br />
fishery purposes. Fishes <strong>of</strong> <strong>the</strong> Cambodian Mekong.<br />
Rome, FAO, 265 pp, 27 pls.<br />
Rambaut, A. 1996. Se-Al: Sequence Alignment Edi<strong>to</strong>r<br />
(Accessed 25 May 2005). Available: http://evolve.<br />
zoo.ox.ac.uk/<br />
Rema Devi, K., T. J. Indra & J. D. Marcus Knight. 2010.<br />
<strong>Puntius</strong> rohani (Teleostei: Cyprinidae), a new species<br />
<strong>of</strong> barb in <strong>the</strong> <strong>Puntius</strong> filamen<strong>to</strong>sus group from <strong>the</strong><br />
sou<strong>the</strong>rn Western Ghats <strong>of</strong> India. Journal <strong>of</strong> Threatened<br />
Taxa, 2: 1121-1129.<br />
Rendahl, H. 1922. Fische, gesammelt von Herrn Carl<br />
Lumholtz in Bulungan, Nordost-Borneo, 1914. Nytt<br />
Magasin for Naturvidenskapene, 60: 199-204.<br />
Roberts, T. R. 1998. Review <strong>of</strong> <strong>the</strong> tropical <strong>Asian</strong> cyprinid<br />
fish genus Poropuntius, with descriptions <strong>of</strong> new<br />
species and trophic morphs. Natural His<strong>to</strong>ry Bulletin<br />
<strong>of</strong> <strong>the</strong> Siam Society, 46: 105-135.<br />
— 1989. The freshwater <strong>fishes</strong> <strong>of</strong> western Borneo<br />
(Kalimantan Barat, Indonesia). Memoirs <strong>of</strong> <strong>the</strong><br />
California Academy <strong>of</strong> Sciences, 14: i-xii + 1-210.<br />
Roberts, T. R. & D. Catania. 2007. Designation <strong>of</strong> lec<strong>to</strong>types<br />
and neotypes, systematic status, and biological<br />
remarks on Red River Cyprinidae described<br />
by Nguyen and Doan, 1969. Natural His<strong>to</strong>ry Bulletin<br />
<strong>of</strong> <strong>the</strong> Siam Society, 55: 85-97.<br />
Schäfer, F. 2009. Oreichthys crenuchoides, a new cyprinid<br />
fish from West Bengal, India. Ichthyological Exploration<br />
<strong>of</strong> Freshwaters, 20: 201-211.<br />
Schut, J., S. S. De Silva & K. Kortmulder. 1984. Habitat<br />
associations and competition <strong>of</strong> eight Barbus (= Pun<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>
tius) species (Pisces, Cyprinidae) indigenous <strong>to</strong> Sri<br />
Lanka. Ne<strong>the</strong>rlands Journal <strong>of</strong> Zoology, 34: 159-181.<br />
Senanayake, F. R. 1985 (1982). Barbus srilankensis, a new<br />
species <strong>of</strong> cyprinid fish from Sri Lanka. Ceylon<br />
Journal <strong>of</strong> Science (Biological Science), 15: 165-172.<br />
Shantakumar, M. & W. Vishwanath. 2006. Interrelationship<br />
<strong>of</strong> <strong>Puntius</strong> Hamil<strong>to</strong>n-Buchanan (Cyprinidae:<br />
Cyprininae) found in Manipur, India. Zoo’s Print<br />
Journal, 21: 2279-2283.<br />
Silas, E. G. 1953. Notes on <strong>fishes</strong> from Mahableshwar<br />
and Wai (Satara district, Bombay state). Journal <strong>of</strong><br />
<strong>the</strong> Bombay Natural His<strong>to</strong>ry Society, 51: 579-589,<br />
Pl. 1.<br />
— 1954. New <strong>fishes</strong> from <strong>the</strong> Western Ghats, with<br />
notes on <strong>Puntius</strong> arulius (Jerdon). Records <strong>of</strong> <strong>the</strong><br />
Indian Museum (Calcutta), 51: 27-37, pl. 5.<br />
— 1956. The systematic position <strong>of</strong> <strong>the</strong> Indian cyprinid<br />
fish Cirrhinus fasciatus Jerdon (1849), with a new<br />
name for Barbus fasciatus Bleeker (1853). Copeia,<br />
1956: 194.<br />
Silva, A., K. Maduwage & R. Pethiyagoda. 2008. <strong>Puntius</strong><br />
kamalika, a new species <strong>of</strong> barb from Sri Lanka<br />
(Teleostei: Cyprinidae). Zootaxa, 1824: 55-64.<br />
Silva, A., K. Maduwage & R. Pethiyagoda. 2011. A review<br />
<strong>of</strong> <strong>the</strong> genus Rasbora in Sri Lanka, with description<br />
<strong>of</strong> two new species (Teleostei: Cyprinidae).<br />
Ichthyological Exploration <strong>of</strong> Freshwaters, 21: 27-<br />
50.<br />
Smith, H. M. 1945. The fresh-water <strong>fishes</strong> <strong>of</strong> Siam, or<br />
Thailand. Bulletin <strong>of</strong> <strong>the</strong> United States National<br />
Museum, 188: i-xi + 1-622, pls. 1-9.<br />
Srivastava, G. J., K. P. Verma & R. B. Sharma. 1977. A<br />
new species <strong>of</strong> <strong>the</strong> genus <strong>Puntius</strong> from Tirhut division,<br />
Bihar. Matsya, 2: 72-73.<br />
Taki, Y., A. Katsuyama & T. Urushido. 1978. Comparative<br />
morphology and interspecific relationships <strong>of</strong><br />
<strong>the</strong> cyprinid genus <strong>Puntius</strong>. Japanese Journal <strong>of</strong><br />
Ichthyology, 25:1-8.<br />
Talwar, P. K. & A. G. Jhingran. 1991. Inland <strong>fishes</strong>, Vol. 1.<br />
Oxford and IBH Publishing, New Delhi, 541 pp.<br />
Tan, H. H. & M. Kottelat. 2008. Revision <strong>of</strong> <strong>the</strong> cyprinid<br />
fish genus Eirmotus, with description <strong>of</strong> three new<br />
species from Sumatra and Borneo. Raffles Bulletin<br />
<strong>of</strong> Zoology, 56: 423-433.<br />
Taylor, W. R. & G. C. Van Dyke. 1985. Revised procedures<br />
for staining and clearing small <strong>fishes</strong> and<br />
o<strong>the</strong>r vertebrates for bone and cartilage study.<br />
Cybium, 9: 107-119.<br />
Tilak, R. 1973. A study <strong>of</strong> <strong>the</strong> freshwater and estuarine<br />
<strong>fishes</strong> <strong>of</strong> Goa, 2: Notes on <strong>the</strong> <strong>fishes</strong> found within<br />
<strong>the</strong> terri<strong>to</strong>ry <strong>of</strong> Goa. Records <strong>of</strong> <strong>the</strong> Zoological<br />
Survey <strong>of</strong> India, 67: 87-120.<br />
Ichthyol. Explor. Freshwaters, Vol. 23, No. 1<br />
95<br />
Tirant, G. 1885. Notes sur les poissons de la Basse-<br />
Cochinchine et du Cambodge. Excursions et reconnaissances,<br />
9: 413-438, 10: 91-198.<br />
Vaillant, L. L. 1902. Résultats zoologiques de l’expédition<br />
scientifique Néerlandaise au Bornéo central. Poissons.<br />
Notes from <strong>the</strong> Leyden Museum, 24: 1-166,<br />
pl. 1-2.<br />
Vishwanath, W. & J. Laisram. 2004. Two new species<br />
<strong>of</strong> <strong>Puntius</strong> Hamil<strong>to</strong>n-Buchanan (Cypriniformes:<br />
Cyprinidae) from Manipur, India, with an account<br />
<strong>of</strong> <strong>Puntius</strong> species from <strong>the</strong> state. Journal <strong>of</strong> <strong>the</strong><br />
Bombay Natural His<strong>to</strong>ry Society, 101: 130-137.<br />
Vishwanath, W. & H. Tombi Singh. 1986. A new species<br />
<strong>of</strong> <strong>the</strong> genus <strong>Puntius</strong> Hamil<strong>to</strong>n from Manipur.<br />
Records <strong>of</strong> <strong>the</strong> Zoological Survey <strong>of</strong> India, 83: 129-<br />
133.<br />
Weber, M. & L. F. de Beaufort. 1912a. Fische. Pp. 522-<br />
541, pls. 11-12 in: A. Maass, Durch Zentral-Suma tra.<br />
Behr, Berlin & Leipzig, vol. 2, fasc. 3.<br />
Weber, M. & L. F. de Beaufort. 1916. The <strong>fishes</strong> <strong>of</strong> <strong>the</strong><br />
Indo-Australian archipelago. III. Ostariophysi: II<br />
Cyprinoidea, Apodes, Synbranchi. Brill, Leiden,<br />
xv + 455 pp.<br />
Whitehead, P. J. P. & P. K. Talwar. 1976. Francis Day<br />
(1829-1889) and his collections <strong>of</strong> Indian <strong>fishes</strong>.<br />
Bulletin <strong>of</strong> <strong>the</strong> British Museum (Natural His<strong>to</strong>ry),<br />
His<strong>to</strong>rical Series, 5: 1-189, pls. 1-4.<br />
Wood, C. E. 1968. Two species <strong>of</strong> Cyprinidae from north<br />
central Mindanao. Philippine Journal <strong>of</strong> Science, 95:<br />
411-423.<br />
Yang, L., R. L. Mayden, T. Sado, S. He, K. Sai<strong>to</strong>h & M.<br />
Miya. 2010. Molecular phylogeny <strong>of</strong> <strong>the</strong> <strong>fishes</strong><br />
traditionally <strong>referred</strong> <strong>to</strong> Cyprinini sensu stric<strong>to</strong><br />
(Teleostei: Cypriniformes). Zoologica Scripta, 39:<br />
527-550.<br />
Yazdani, G. M. & M. Babu Rao. 1976. A new species <strong>of</strong><br />
<strong>the</strong> genus <strong>Puntius</strong> (Hamil<strong>to</strong>n) (Pisces: Cypriniformes:<br />
Cyprinidae) from western India. Journal <strong>of</strong><br />
<strong>the</strong> Bombay Natural His<strong>to</strong>ry Society, 73: 171-175.<br />
Yazdani, G. M. & D. F. Singh. 1994. <strong>Puntius</strong> crescentus,<br />
a new cyprinid fish from south India with observations<br />
on <strong>the</strong> taxonomic status <strong>of</strong> some related species.<br />
Journal <strong>of</strong> <strong>the</strong> Bombay Natural His<strong>to</strong>ry Society,<br />
91: 107-109.<br />
Yazdani, G. M. & S. Talukdar. 1975. A new species <strong>of</strong><br />
<strong>Puntius</strong> (Cypriniformes: Cyprinidae) from Khasi<br />
and Jaintia Hills (Meghalaya), India. Journal <strong>of</strong> <strong>the</strong><br />
Bombay Natural His<strong>to</strong>ry Society, 72: 218-221.<br />
Zheng, L.-P., J.-X. Yang, X.-Y. Chen & W.-Y. Wang. 2010.<br />
Phylogenetic relationships <strong>of</strong> <strong>the</strong> Chinese Labeoninae<br />
(Teleostei, Cypriniformes) derived from two<br />
nuclear and three mi<strong>to</strong>chondrial genes. Zoologica<br />
Scripta, 39: 559-571.<br />
Received 14 April 2011<br />
Revised 4 March 2012<br />
Accepted 6 May 2012
96<br />
Pethiyagoda et al.: Synopsis <strong>of</strong> <strong>South</strong> <strong>Asian</strong> <strong>Puntius</strong>