See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/327012928

Osteological characterization of the genus puntius (Teleostei: Cyprinidae)

recorded from six river systems of southern Western Ghats, India

Article · January 2018

DOI: 10.22034/iji.v5i2.252

CITATIONS READS

3 166

3 authors, including:

A A Arunkumar Anbu Manimekalan Arunachalam

Karpagam Academy of Higher Education Bharathiar University
6 PUBLICATIONS 6 CITATIONS 36 PUBLICATIONS 151 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Fish diversity View project

Effluent Treatment View project

All content following this page was uploaded by A A Arunkumar Anbu on 17 September 2020.

The user has requested enhancement of the downloaded file.

 Iran. J. Ichthyol. (June 2018), 5(2): 139-166 Received: November 2, 2017
© 2018 Iranian Society of Ichthyology Accepted: May 24, 2018
P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v5i2.252
http://www.ijichthyol.org

Research Article
Osteological characterization of the genus Puntius (Teleostei: Cyprinidae)
recorded from six river systems of southern Western Ghats, India
Anbu Aravazhi ARUNKUMAR, Nagaraj VIJAYA LAKSHMI, Arunachalam MANIMEKALAN*

Biodiversity and Molecular Lab, Department of Environmental Sciences, Bharathiar University, Coimbatore 641 046,
India.
*
Email: manimekalan@gmail.com
Abstract: This paper provides information on the osteological characterization of
the genus Puntius with respect to six river systems of Southern Western Ghats.
Fishes were collected using cast net, dip net, gill net and drag net from various
streams and rivers of Southern Western Ghats. Clearing and staining methods was
carried out for osteological study. After clearing and double staining, the specimens
were observed under a stereomicroscope and photographed using a digital camera.
Twenty-nine morphometric and meristic osteology characters were taken. Principal
component analysis and cluster analysis were performed to group the species and to
detect the similarity between the species. Comparing all the species it was observed
that the species were grouped into three groups. The first class has 10 species
P. mahecola, P. chola, P. bimaculatus, P. dorsalis, P. melanampyx, P. fasciatus,
P. ticto, P. denisonii, P. sophore, P. conchonius. The second class has 4 species;
P. filamentous, P. sarana spirulus, P.amphibious and P. ophicephalus. The third class
has only one species P. carnaticus well supporting the observations of Shantakumar
& Vishwanath (2006).
Keywords: Chalakudy, Meristics, Morphometrics, Preethmoids, Skeleton, Skull.
Citation: Arunkumar, A.A.; Vijaya Lakshmi, N.V. & Manimekalan, A. 2018. Osteological
characterization of the genus Puntius (Teleostei: Cyprinidae) recorded from six river systems
of southern Western Ghats, India. Iranian Journal of Ichthyology 5(2): 139-166.

Introduction validate or refute claims of species identity.

Osteological characters are considered to be
important diagnostic traits for interpreting
systematics and phylogenetic relationships within
Teleostei than other phenotypic tools (Nybelin 1973;
Keivany 2014a, b, c, d, 2017a, b). Around the world
the importance of cyprinid fishes in freshwater
communities and their fossil history greatly interests
many ichthyologists and other scientists. Moreover,
well-preserved fossils can increase knowledge
concerning the primitive or derived morphology of
osteological characters used in systematics. The
osteological characters clearly exhibit themselves
and also exhibit the variation within and between the
species of ambiguous genera, further helping to
139
Traditional morphology-based taxonomy is not
always correct in identification to the species level,
and therefore a multidisciplinary approach to
taxonomy which includes morphological,
osteological, molecular and distributional data is
essential (Krzywinski & Besansky 2003).
Ramaswami (1948, 1952a, b, c, d, 1953, 1955a, b)
has worked considerably on the osteology of Indian
cyprinid fishes. The record states that the teleostian
fishes have undergone marvelous and interesting
morphological changes in various aspects in
connection to the environment and he emphasized the
importance of the osteological study.
Many experimental attempts have been made by
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166
various workers to support with more appropriate
characters for the justification of the genus and
species on the basis of osteological study. Regan
(1911) has stated that the morphological characters
(external) were not sufficient enough for taxonomy
of cyprinids. Regan (1911) proposed a phylogenetic
key to different sub-families based on the skull,
Weberian apparatus and the pectoral girdle. Because
of this, osteological studies became an important tool
in the hands of taxonomists.
The oldest published cyprinid record (Hora 1938)
on 'cyprinid' scales from the lower Palaeogene
(Palaeocene) Intertrappean beds of the Central
Provinces. However, scales are poor diagnostic
fossils for the Cyprinidae because scale characters
overlap between Cyprinidae, Catostomidae and
Characoidea. The subfamily Cyprininae is one of the
most diverse groups within this family and order
Cypriniformes (Nelson 2006). The subfamily
Cyprininae generally split to four main groups:
barbins (i.e. Puntius, Barbus and Spinibarbus),
cyprinins (i.e. Cyprinus, Carassius, and
Carassioides), labeonins (i.e. Labeo, Garra, and
Osteochilus), and oreinins (i.e. snow trout) (Yang et
al. 2009). The systematic relationships of these
groups have been extremely chaotic, as various
numbers of groups have been recognized within the
subfamily (Chen et al. 2005, 2009; Fang 2003; Fang
et al. 2009, Yang et al. 2009).
Many ichthyologists like Gouan (1770), Weber
(1820), Agassiz (1833), Hallmann (1837), Muller
(1842), Owen (1848), Stannius (1854), Huxley
(1864, 1876), Parker (1873), Masse (1873), Stohr
(1884) and Wright (1884) have been attracted to
study on teleosts osteology. McClelland (1842) who
was a pioneer worker made use of osteological
studies of the family Cyprinidae in separating these
tribes. Sagemehl (1891), Regan (1911) and Sarbahi
(1933) were the significant contributors on the
endoskeleton studies.
In the middle of last century, Ramaswami (1948,
1952a, b, c, d, 1955a, b) and Harrington (1955) could
make use of many osteological characters, viz,
140
osteocranium apparatus in the taxonomy of family
Cyprinidae. Recently, workers like Vashist &Verma
(1968), Sorescu (1975, 1978) and Howes (1978) have
made good contributions by studying the
Osteocranium and associated structures. Hubbs
(1919, 1926), Gregory (1933), Eaton (1935), Matthes
(1963), Saxena & Khanna (1963), Saxena &Bakshi
(1964), Das & Daftri (1967a, b), Mc. Allister (1968),
Nelson (1969), Tiwari (1972), Mirza (1973), Gosline
(1973, 1975) and Takaya (1974) have restricted their
studies to the osteocranium or only parts of it Mehta
and Tandon (1984).
The first experimental investigation of species
groups within Puntius appears to have been by
Kortmulder (1972). Taki et al. (1978), from an
osteological study of 23 mostly Southeast Asian
species of Puntius s.l., proposed six groups, of which
two (their P. conconius group and the P. arulius
group) comprised South Asian species. Detrich et al.
(1999) stated that zebra fish, Danio rerio (Hamilton)
of the family Cyprinidae has become an important
model organism, especially for vertebrate
developmental studies. Fink & Fink (1981) have
stated that the caudal skeleton of Danio exemplifies
the otophysan synapomorphy of a compound
terminal centrum and the cypriniform synapomorphy
of having two or fewer epurals. Buhan (1972) stated
that the upper lobe of the caudal fin has four hypurals
is typical for Cyprinidae.
Roghan Pethiyagoda (2012) has found out through
an examination of external morphology and
osteology analysis, the genus Systomus possesses
maxillary and rostral barbels with last unbranched
dorsal-fin ray stiff (‘osseous’), serrated and lateral
line complete with 27-34 scales. Based on the above
characterization three new genera were proposed
Dawkinsia, Dravidia and Pethia. Also he has
separated Systomus and Eechalakanda as separate
genus from Puntius. According to Weitzman (1962),
the presence of a preethmoid is an evolutionary
advanced character of Cypriniforms. Presence of
preethmoid is a characteristic feature of all the
species of Puntius.
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
According to Weitzman (1962) and Ramaswami
(1956), the frontal bone of all Cyprinids except
Esomus are in contact with the pterotic and it is a
primitive feature. Fink & Fink (1981, 1996) have
stated that the caudal skeleton of Danio exemplifies
the Otophysan synapomorphy of a compound
terminal centrum and the Cypriniform
synapomorphy having two or fewer epurals. Buhan
(1972) stated that the upper lobe of the caudal fin has
four hypurals is typical for Cyprinidae.
Shantakumar & Vishwanath (2006) studied the
inter-relationship between 12 species of genus
Puntius based on the dorsal spine, pre-dorsal
vertebral counts, vertebral counts, colour, barbels and
caudal fin have been divided as three phyletic group
and stated that the number of vertebrae ranges from
29-36 inclusive of first four complex vertebrae and
last fused preural and ural centra.
Shantakumar & Vishwanath (2006) worked on the
interrelationships of 12 species of the genus Puntius
from Manipur and he has observed that among the 12
species 3 groups were formed viz. sarana, conconius
and sophore and placed P. sarana and P. sophore
under two different groups.
Moreover, Intra-specific variation in Puntius has
not been extensively reported yet, and therefore,
taxonomically important diagnostic variation in the
species is poorly understood. Many scientists get
confused with the ambiguous species for which so far
very little work has been carried out making use of
osteological characterization and comparison to
distinguish the individuality between these species.
Perusal of literature says not much work had been
carried out on the osteology of genus Puntius. Hence
an attempt is made to distinguish these species based
on the osteological study of genus Puntius from
Southern Western Ghats.
Materials and Methods
Study Area: Seventeen Puntius species were
collected from the long and meandering eastward
flowing river systems of Southern Western Ghats,
especially from Bhavani River, Moyar River,
141
Fig.1. Collection location of six river systems.
Chalakudy River, Periyar River, Cauvery River and
Nugu River. The study sites and its habitat
characteristics are presented in Table 1 and Figure 1.
Methodology: A total of 10 specimens from each
species were selected for osteological study, because
some species are coming under threatened category,
so to minimize the collection only 5 individuals were
collected. Osteological study was carried out by the
method described by Taylor & Van Dyke (1985).
After clearing and double staining, the specimens
were observed under a stereomicroscope and
specimens were photographed using a Cannon 6D
digital camera. Twenty-nine osteological
morphometric and meristic characters were measured
and counted based on the method described by
Rainboth (1996), Harrington (1955), Wilimovesky &
Weitzmann (1955) and Topp & Cole (1968). The
measurements are taken using a digital Vernier
caliper in mm. Principal component analysis were
performed for group the species and cluster analysis
was performed to know the similarity between the
closely related species using osteological
morphometric and meristic character data. To
remove the size component from the shape
measurements (Thorpe 1976), all the measurements
were standardized according to the formula:
Ms= Mo (Ls/Lo)b
Where Ms=standardized measurement; Mo=
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Table 1. Study sites and their habitat characteristics.

No Study site Latitude and Altitude Forest type Stream Stream Stream Mean
longitude order Width Depth Velocity*
(m) (m) (m/sec)
Moyar River System
1 Gulithuraipatti 11°36’N, 76°47’E 312 Thorn forest 4 10 6 4
2 Kallampalayam 11°31’N, 77°00’E 300 Thorn forest 4 13 8 4
3 Belemeenthurai 11°36’N, 76°47’E 520 Dry deciduous 4 19 1.75 4
Chalakudy River System
4 Orukomban range 10°22’N, 76°39’E 450 Dry deciduous 4 6 0.5 3
5 Thenmudiparai 10°24’N, 76°36’E 510 Dry deciduous 5 25 1.5 3
6 Baghapallam 10°27’N, 76°43’E 748 Dry deciduous 5 8 0.5 3
7 Thellikal 10°27’N, 76°44’E 840 Dry deciduous 4 4 1 3
8 Puliyarkutti 8th 10°23’N, 76°40’E 527 Dry deciduous 4 19.2 1.2 3
9 bridge
Puliyarkutti 3rd 10°23’N, 76°41’E 512 Dry deciduous 4 37 1.5 3
10 bridge
Thunakadavu 10°25’N76°46’E 510 Dry deciduous 4 13.6 0.5 3
11 stream
Thunakadavu tunnel 10°20’N, 76°34’E 520 Dry deciduous 5 15 10 5
12 Urilikal 10°19’N, 76°53’E 3238 Dry deciduous 2 7 1.5 2
13 Athirappalli 10°18’N, 76°34’N 202 Semi evergreen 4 8 3 4
14 Pillapara 11°36’N, 76°47’E 267 Semi evergreen 4 5 2 4
Bhavani River System
15 Kovaikutralam falls 10°56’N, 76°41’E 560 Semi evergreen 2 5 1.2 4
16 Nellithurai 11°17’N, 76°53’E 380 Thorn forest 4 27 1.1 5
Periyar River System
17 Oorpannikaham 09° 28’N,77°16’E 884 Evergreen 4 12 2.1 2
18 Valukuparai 09° 28’N,77°17’E 869 Evergreen 4 7.5 0.3 3
19 Melaparai 09°26’N,77°18’E 965 Evergreen 4 11 4.2 3
20 Naduthotam 09°26’N, 77°19’E 950 Evergreen 4 7.5 0.3 3
21 Ummikuppamthodu 09°28’N, 77°14’E 943 Evergreen 4 5 3 4
22 Sorrakottaodai 09°28’N, 77°15’E 879 Evergreen 4 7 1.5 3
23 Mullaithodu 09°31’N, 77°16’E 869 Evergreen 4 10 0.6 3
24 Anjurily 11°36’N,76°47’E 912 Evergreen 4 20 5 2
25 Thenkasithodu 11°36’N, 76°47’E 872 Evergreen 4 11.3 0.5 2
Cauvery River System
26 Kadapilliyarthittu 12°07’N, 77°46’E 1137 Dry deciduous 4 75 1.5 2
27 Belikoondu 12°11’N, 77°43’E 267 Dry deciduous 4 80 10 5
28 Nadathittu 12°08'N, 77°44'E 262 Dry deciduous 4 70 6 3
29 Sinnaru 12°06’N, 77°46’E 225 Dry deciduous 4 55 0.5 3
30 Thonanthikla 12°07’N, 76°46’E 341 Dry deciduous 4 25 1 4
Nugu River System
31 Noolpuzha 11°41’N, 76°23’E 2810 Semi evergreen 3 25 4.1 4
*Velocity (m/sec): 1. Very slow (<.05); 2. Slow (0.05-0.2); 3. Moderate (0.2-0.5); 4. Fast (0.5-1.0); 5.Very fast (>1).

measured character length; Ls=overall mean standard
length for all fish samples; Lo=standard length of
specimen) Tigano et al. (2004, 2006). Parameter b
was estimated for each character from the observed
data by the allometric growth equation M=aLb, as the
slope of the regression of log Mo on log Lo using all
fish in all groups but allowing the intercept to differ
between groups (Elliot et al. 1995). Linear regression
determined the relationship between each character
and the species (N=15) was used to test whether
scaling coefficients were significantly different
142
between species (P≤0.05). XLSTAT and SPSS
software were used for statistical analysis.
Results
The Skull
Of the 15 Puntius species skull osteology studied,
only minor difference were observed in size, shape,
modifications, and the meristic characters between
the species. The skull is described to have two parts
the neurocranium and the branchiocranium. The
neurocranium is further classified into four regions
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
Fig.2. Dorsal view of neurocranium. (A) Puntius amphibious with fontanel. (B) Puntius dorsalis without fontanel. All
the bars indicate 1cm.
like olfactory region, orbital region, otic region and
the occipital region. Similarly the branchiocranium is
divided into four regions mandibular arch, palatine
arch, hyoid arch, opercula apparatus and branchial
arch.
The neurocranium of genus Puntius is elongated
and narrow. Dorsally, the pineal foramen in the mid
region, the latero-sensory canals at the lateral edges,
the semicircular ridge on parietals and the
supraoccipital crest and the two crests on epiotics arc
characteristic features of all the species studied.
Among the 15 species, P. chola, P. dorsalis,
P. sophore and P. amphibious have a prominent
dorso-median cranial fontanel. It extends from the
middle of the parietal to the middle of the frontal and
is characterized by its rectangular shape and
elongated structure (Fig. 2a). The rest of the 11
species did not show any changes in the
143
Neurocranium (Fig. 2b). The olfactory region in the
neurocranium expresses the variations between the
species in their structure, shape and position, which
are characterized to make out the variations among
the species.
Olfactory region: Olfactory region is found to be the
anterior most region of the skull, and it is composed
of three median bones (ethmoid, prevomer, rostral)
and four paired bones (lateral ethmoids, prefrontals,
nasals, preethmoids). In the olfactory region all the
three median bones and four paired bones looks
similar in respect to all the individuals (Fig. 3).
The ethmoid is considered as vertically oriented
median bone articulating with the anterior edges of
frontals and prevomer. The dorsal side of ethmoid is
pentagonal and its middle part is concaved while two
sides are convex. On either side of ethmoid have two
olfactory foramens. Each olfactory foramen is thus
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166
Fig.3. Dorsal view of neurocranium of Puntius species
in the olfactory region.
bordered by the posterior edge of ethmoid anteriorly
and by the lateral ethmoids dorsally and posteriorly
(Fig. 4). Among the 15 species four species namely,
P. chola, P. dorsalis, P. sophore and P. amphibious
the ethmoid appears to be slightly broader and
protected by a strong ossified prevormer above it
(Plate 1). The rest of the 11 species have a little
narrow ethmoid and non - ossified prevormer
comparing to the above said species (plate.1). The
paired preethmoids are articulated laterally with the
articular junction of ethmoid and prevormer in all the
species taken for study. The lateral ethmoid is an
angular bone with a broad base. Ventrally it
enunciates with the lateral side of prevomer while
antero-dorsally it enunciates with the ethmoid.
Latero-posteriorly it is inseparably fused to the
downward arching pre-frontals. The posterior end of
palatine normally rests against lateral ethmoid-
prefrontal complex bending downward (Fig. 3).
Among the 15 species the lateral ethmoid articulates
laterally with palatine, anterodorsally with ethmoid
and latero-posteriorly with prefrontal and moreover
no remarkable variations were noted in their shape or
size for species differentiation (Plate 1).
The Prefrontals are very thin curved paired
angular bones where one angle curves vertically
downward. The posterior part of this bone is super
144
imposed by the anterior end of two frontals. The
downward bending prefrontals form the orbits
anterior boundaries (Fig. 3). Among the 15 species
the prefrontals articulates laterally with prevomer,
anterodorsally with ethmoid and latero-posteriorly
with lateral ethmoid and moreover no remarkable
variations were noted in their shape and size for
species differentiation (Plate. 1).
Kinethmoid bone also termed as rostral is a
"question mark shaped" tiny bone which is easily
displaceable in handling of digested specimens. It fits
into the gap formed by the two premaxillaries and the
anterior groove of the ethmoid. Rostral helps to lever
the premaxillaries and the maxillaries in the edge of
the mouth in all the 15 species (Fig. 3). Moreover, no
remarkable variations were noted in their shape or
size for species differentiation (Plate 1).
Prevormer bone located the most antero-ventrally
of the cranium. It is a horizontal bone with a forked
head and a short pointed shaft. The pointed posterior
end of the shaft fits into the V-shaped anterior end of
parasphenoid to form the supporting axis of the skull.
The anterior end of prevormer split into two lobes,
which extend beyond the forward limits of dorsally
bending ethmoid. The branched end of prevormer
pronounces with the button shaped preethmoids (Fig.
3) and no remarkable variations were noted in their
shape and size in all the 15 species (Plate 1).
The preethmoids are exclusively supported by the
prevomer and it serve for the attachment of the
palatine. Preethmoids are tiny knob like bones firmly
fused to the anterior lateral ends of prevormer-
ethmoid joints. According to Weitzman (1962), the
presence of a preethmoid is an evolutionary advanced
character of Cypriniforms. Presence of preethmoid is
a characteristic feature genus Puntius (Plate 1; Fig.
3).
Nasals are small plate like bones located dorsally
and attached to the anterior most ends of frontals.
Unlike other bones of the skull they were not firmly
enunciated. Nasals are completely traversed by the
nasal section of the supraorbital latero-sensory canal.
In all the 15 species studied no remarkable variations
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
Fig.4. Dorsal view of the ethmoid of Puntius species. P. chola, P. dorsalis and P. sophore the ethmoid appears to be
slightly broader and protected by a strong ossified prevormer above it. P. bimaculatus, P. ticto and P. sarana having
little narrow ethmoid and non-ossified prevormer
were noted in their shape and size (Plate 1).
Orbital region: The orbital region is prominent by the
presence of eyes. Each orbit is surrounded anteriorly
by prefrontals, antero-dorsally by orbitosphenoids,
dorsally by frontals and pterosphenoids,
posterodorsally by sphenotics, posteriorly by
prootics and ventrally by the parasphenoid.
Anterodorsally and postero-ventrally the two
semicircular sclerotic bones surround the eye ball.
Externally the rim of the orbit is overlaid by a series
of ossicles called circumorbital bones. Dorsally the
entire orbital region is covered by the thin, elongate,
and paired frontals. Each frontal is about two third of
the entire length of the neurocranium. The narrow
anterior end of frontals overlay the posterior end of
ethmoid while anterior outer lateral part articulates
with the prefrontals. Behind the prefrontals the
supraorbital fuse with the outer lateral wall of the
frontals. The posterior edge of frontals articulates
with parietals and the sphenotics. Along the
middorsal line, the left frontal articulates with the
right frontal. The posterior end the two frontals are
separated from the elongated pineal of foramen bone.
Otic region: The otic region consists of parietals,
epiotics, prootics, and sphenotics. Inside the otic
145
capsules there are some bony structures called
otoliths. In regard with the otic region all the bones
looks similar in respect to all the species studied (Fig.
2a, b).
Rectangular parietals form the dorsal wall of the
otic region. Parietals articulate anteriorly with
frontals, posteriorly with supraoccipital and epiotics
and laterally with pterotics. The two paritals
articulate with each other at the middorsal line. The
anterior wall of left parietal form the posterior margin
of pineal foramen. The hind margins of parietals form
a semicircular ridge at the hind end of the skull which
includes a part of latero-sensory canal system called
supra-temporal canal. This is a characteristic feature
of all Puntius species. The parietals are smooth and
slightly convexed dorsally. In all the 15 species no
remarkable variations were noted in their shape and
size (Plate 1).
The two epiotics lie on either side of
supraoccipital bone and form the hindmost lateral
wall of the skull. The epiotics articulate anteriorly
with parietals, posteriorly with exoccipitals,
outerlaterally with pterotics and inner laterally with
the supraoccipital bone. Dorsally, the epiotics are
slightly convexed and pentagon in shape with a small
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Table 2. Osteological characters abbreviations.

Morphometric Characters Abbreviations Meristic Characters Abbreviations
Distance rudimentary neural arch & DRNA&E Length of hypural 6 LH6
Epural jaw Length
Upper UJL Length of neural spine LNS
Lower Jaw Length LJL Length of haemal spine LHS
Dorsal fin spine length DFSL Eye ball diameter EBD
Dorsal fin ray length DFRL Last dorsal spine LDS
Pectoral fin spine length PFSL Number of dorsal fin spines DFS
Pectoral fin ray length PFRL Number of anal fin spines AFS
Pelvic fin spine length PVFSL Number of pectoral fin spines PFS
Pelvic fin ray length PVFRL Number of pelvic fin spines PVFS
Anal fin spine length AFSL Number of dorsal fin rays DFR
Anal fin ray length AFRL Number of anal fin rays AFR
Length of epural LE Number of pectoral fin rays PFR
Length of rudimentary neural arch LRNA Number of pelvic fin rays PVFR
Length of Compound centrum LCC Vertebral counts VC
Width of compound centrum WCC No. of supraneurals S
Vertebral column length VCL No. of predorsal vertebra PDV
Vertebral column width VCW Parahypurals P
Length of parahypural LP Hypurals H
Length of hypural 1 LH1 Fontanel F
Length of hypural 2 LH2 Dorsal proximal pterygiophores DPP
Length of hypural 3 LH3 Anal proximal pterygiophores APP
Length of hypural 4 LH4 Operculum length OL
Length of hypural 5 LH5 Operculum width OW

crest on the middle. These crests arc typical in
Puntius species. Just near the semicircular ridge, the
dorsolateral part of the epiotic articulates with the
post-temporal which act as a suspensor of the
pectoral girdle. In all the 15 species no remarkable
variations were noted in their shape and size (Plate
1).
Pterotics are postero-laterally located elongate
bones which lie between sphenotic and epiotic. They
form sutures with sphenotic and frontal anteriorly,
with parietals laterally and with epiotics and
exoccipitals posteriorly. The pterotic articulates
ventrally with sphenotic, prootic and opisthootic. The
posterior most end of the pterotic is ventro-laterally
directed and elongated as a fine process. The
dorsolateral surface of pterotic articulates with
lamellar portion of supra temporal bone which helps
to suspend the pectoral girdle to the skull. In all the
15 species considered for study no remarkable
variations were noted in their shape or size for
species differentiation (Plate 1).
146
Sphenotic is a small irregular bone located at the
lateral margin of the cranium. It articulates dorsally
with frontals, posteriorly with pterosphenoids and
ventrally with prootics. Sphenotic is drawn out
antero-laterally as a prominent process, against
which the super mesial surface of the fifth suborbital
rests. An elongated concavity is enclosed between
sphenotics, prootics and pterotic bones for the
condyle of hyomandibular. In all the 15 species
considered for study no remarkable variations were
noted in their shape or size for species differentiation
also in splanchnocranium/gills, hyoids, jaws (Plate
1).
Axial skeleton
The first four vertebrae are fused and reduced to form
the Weberian apparatus, which is a characteristic
feature of all Cypriniformes (Kotalawala 1992). The
total number of vertebrae in the axial skeleton varies
between the species of Puntius studied and can be
used in taxonomical studies (Plate 4). The number of
vertebral count ranges from 29-36 in different
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

Table 3. Vertebra, supraneurals and pre-dorsal vertebra count of Puntius species.

Species N Vertebral counts Supraneurals Pre-dorsal
30 31 32 33 34 35 36 37 38 39 40 41 42 4 5 6 8 vertebrae
9 10
1. 5 5 5 5
2. 3 3 3 3
3. 5 5 5 5
4. 2 2 2 2
5. 5 5 5 5
6. 5 5 5 5
7. 5 5 5 5
8. 5 5 5 5
9. 2 2 2 2
10. 5 5 5 5
11. 2 2 2 2
12. 3 3 3 3
13. 5 5 5 5
14. 5 5 5 5
15. 3 3 3 3

Puntius species. The number is inclusive of the first
four vertebrae of the complex vertebrae and the last
fused pre-ural and ural centrum. The total number of
vertebrae of Puntius ranges from 27 to 41.
Vertebral column count in the species was noted
as follows: P. filamentosus (31), P. melanampyx
(30), P. fasciatus (32), P. sarana spirulus (34),
P. chola (33), P. amphibious (30), P. mahecola (35),
and P. denisonii (30). Rather there are few
exceptional cases in the genus Puntius which has
vertebral counts more that the value stated by
Shanthakumar (2006) in P. ophicephalus (38) and
P. carnaticus (42). The total vertebral count ranges
from 35-36 in P. sarana, 29-33 in P. conchonius and
P. sophore groups (Shanthakumar 2006).
The predorsal vertebral count shows two features
i.e., eight and ten with nine as the intermediate
between the two counts. Here, P. bimaculatus and
P. denisonii has 9 and P. carnaticus has 10 predorsal
vertebral counts (Table 3). A series of midsaggital
ossification are found dorsal of the vertebral column
in the region between the rear of the neurocranium
and the anterior most proximal radial pterygiophore
of the dorsal fin. These bones, the supraneurals along
with proximal radial pterygiophores of the dorsal fin
interdigitate basally between the distal portions of the
neural spines of the proximate vertebrae. There are
four to six supraneurals in Puntius (Shanthakumar,
2006). The 1st supraneural is located anterior to
neural spine of the 5th vertebras. Among the 15
species P. carnaticus and P. sophore have 6
supraneurals in the rest of the species only 4 were
noted. Species like P. sophore, P. chola, P. terio,
P. conconius, P. stoliczkanus, P. ornatus and
P. bizonatus have only 4 supraneurals and 8 predorsal
vertebral counts, rather P. manipuriensis have 5
supraneurals and 9 predorsal vertebral counts
(Shanthakumar 2006).
Appendicular Skeleton
Dorsal fin: The dorsal fin constitutes of three-way
pterygiophores, rays and spines. There are 9-11
proximal, intermediate and distal pterygiophores, 2-
4 dermal spines and 7-9 dermal rays in the dorsal fin
of Puntius species studied. All pterygiophores are
three-way sagittal bony elements supporting the
spines and rays of dorsal fin. The proximal
pterygiophores are long and dowel-like, interlinking
with the neural spines of the precaudal vertebrae. The
intermediate pterygiophores are very short and rod
shaped and articulate with the dorsal end of proximal
pterygiophores. The distal pterygiophores are small

147
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Table 4. Dorsal and anal spines, dorsal and anal rays and dorsal and anal proximal pterygiophores count of Puntius
species.
Dorsal Anal
Species N Proximal Nature of Spines Rays Proximal
Spines Rays pterygiophores spines pterygiophores
1. 5 4 8 10 smooth 3 5 6
2. 3 4 8 9 smooth 3 6 6
3. 5 4 8 9 smooth 3 6 6
4. 2 4 8 9 serrated 3 5 6
5. 5 4 7 9 smooth 4 5 6
6. 5 3 7 8 smooth 3 6 6
7. 5 4 8 9 serrated 3 5 6
8. 5 3 8 9 serrated 3 5 6
9. 2 3 8 9 smooth 2 5 6
10. 5 3 8 9 smooth 3 5 6
11. 2 4 8 7 smooth 3 5 6
12. 3 4 8 10 smooth 3 5 6
13. 5 3 7 8 smooth 3 5 6
14. 5 3 8 9 smooth 2 5 6
15. 3 4 8 10 smooth 3 5 6
1. P. filamentosus, 2. P. melanampyx, 3. P. fasciatus, 4. P. sarana spirulus, 5. P. ophicephalus, 6. P. chola, 7. P. ticto,
8. P. conconius, 9. P. dorsalis, 10. P. mahecola, 11. P. denisonii, 12. P. carnaticus, 13. P. bimaculatus, 14. P. sophore,
15. P. amphibious.

proximal, intermediate and distal pterygiophores.

Fig.5. Skeletal structure of the dorsal fin of Puntius.
and cone shaped and articulate with the dorsal end of
intermediate pterygiophores. However, all the
pterygiophores of the dorsal fin of Puntius do not
contain this tripartite structure. The first and last
pterygiophores of all the species studied do not
contain intermediate or distal pterygiophores and
distal, Medial and proximal part did not fused
together. The three or four pterygiophores following
the first pterygiophores do not contain intermediate
pterygiophores. In these, the distal pterygiophores
directly articulates with the proximal pterygiophores.
The other five or six pterygiophores contain
148
The last pterygiophore is a vestige of a full structural
element. The number of dorsal spine and how they
articulate with the pterygiophores varies in all the 15
species (Table 4, Fig. 5, Plate 2). It is clearly noted
that in all the species spines of the dorsal fin always
articulates with the proximal pterygiophores of the
first two pterygiophores and not with intermediate
and distal pterygiophores. Among the 15 species
P. melanampyx, P. fasciatus, P. ticto, P. carnaticus,
P. denisonii, P. filamentosus, P. amphibious,
P. sarana spirulus and P. ophicephalus have 4 dorsal
spines and rest of the species have only 3 dorsal
spines.
All the 15 species of the genus Puntius have a
well-developed ossified dorsal fin spine. It was well
noted that two different types of dorsal spines viz. (1)
strong and serrated and (2) weak and smooth spine
occurred in the inner side of the dorsal spine. The
serrations are two types one is serrations in the upper
half of the spine and other is serrations in the lower
half of the spine (Fig. 5). In P. chola, P. terio and
P. sophore weak and smooth spine was noted and
species like P. conconius, P. toliczkanus, P. ornatus,
P. bizonatusi, P. ticto, P. manipuriensis, P. sarana,
P. orphoides and P. jayarami have serrated last dorsal
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
Fig.6. (a) Dorsal spine with serrations in the lower
part of P. sarana, (b) dorsal spine with serrations in
the upper part of P. conconius and P. ticto and (c)
speices with weak and smooth dorsal spine.
spine (Shanthakumar 2006). In species like
P. conconius, P. ticto and P. sarana the spine is
articulated with the second pterygiophore and
serrated, while the others are smooth in nature.
Anal fin: Anal fin resembles the pattern of dorsal fin,
with spines and rays supported by tripartite
pterygiophores. The number of spines, rays and
pterygiophores of anal fin are less than dorsal fin. The
anal fin contained 7 proximal pterygiophores, 3
spines and 5 rays. The pterygiophores of anal fin
inter-connected with the haemal spines of anterior
caudal vertebrae. The first and last pterygiophores of
anal fin do not contain intermediate pterygiophores
or distal pterygiophores and the last one is a vestige
because it appears in the larval stages. Out of 3 anal
spines first two directly comprehensible with the first
proximal pterygiophores while the other one
articulates with the 2nd proximal pterygiophores. The
second and third pterygiophores are without
intermediate pterygiophores or distal pterygiophores
while the others are with proximal pterygiophores,
intermediate pterygiophores and distal
pterygiophores. The rays of anal fin articulate with
the cone shaped distal pterygiophores of 2nd to 6th
pterygiophores. The distal pterygiophores of anal fin
bend over to meet the succeeding proximal
pterygiophores and articulate with the rays.
Therefore each ray is associated with a
149
Fig.7. Skeletal structure of the anal fin of Puntius.
pterygiophores but is displaced posteriorly so that it
receives greater support from the pterygiophores of
the succeeding ray. Typically all rays of anal fin are
branched, segmented and bilaterally paired in all the
species of Puntius studied. The numbers of
pterygiophores, spines and rays of anal fin of Puntius
studied are more or less constant in regard to all the
species studied at present. Among the 15 species
taken for study P. ophicephalus have 4 anal spines
and P. sophore, P. dorsalis have 2 and in rest of the
species have 3 spines (Table 4, Fig. 7).
Caudal fin: The caudal fin supported by the last 3
caudal vertebrae of the axial skeleton (Jinadasa &
Kotalawala 1991). Basically there are six hypurals
and one parypural. The 5th and 6th hypurals are very
small in size. There is an epural above the specialized
neural process and anterior of the urostyle. Caudal fin
of Puntius has 10+9 principal rays. A small pair of
uroneurals are noted where one of its end is
connected to distal end of the urostyle and another
end extends little beyond urostyle between the first
and second principle rays. In species of Puntius there
are no inter-muscular bones attached to pleural ribs,
but to heamal arch heamal spine, neural arch and
neural spines. Among the 15 species
P. melanamplyx, P. ticto, P. chola, P. dorsalis and
P. conconius have 7 hypurals and in rest of the
species have only 6 hypurals (Table 5, Fig. 8).
Comparison among the species: Length of lower Jaw
in P. filamentosus ranges between 2.09 to 2.24 (2.18)
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Fig.8. Caudal fin.

Table 5. Comparative meristic characters of Puntius species.

Species
Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
LDS 1 1 1 2 1 1 2 2 1 1 1 1 1 1 1
DFS 4 4 4 4 4 3 4 3 3 4 4 3 4 3 3
AFS 3 3 3 3 4 3 3 3 3 3 3 2 3 2 3
PFS 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
PVFS 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2
DFR 8 8 8 8 7 7 8 8 8 8 8 8 8 8 7
AFR 5 6 6 5 5 5 5 5 5 5 5 5 5 5 6
PFR 12 13 14 16 14 12 12 12 12 12 15 13 13 12 11
PVFR 8 7 9 8 9 8 8 8 8 8 8 8 8 8 7
VC 31 30 32 34 38 32 33 32 35 30 42 30 30 33 33
S 4 4 4 4 4 4 4 4 4 4 6 6 4 4 4
PDV 8 8 8 8 8 9 8 8 8 9 10 8 8 8 8
P 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
H 6 7 6 6 6 6 7 7 6 6 6 6 6 7 7
F 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2
DPP 10 9 9 9 9 8 9 9 9 7 10 9 10 9 8
APP 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6
1. Puntius filamentosus, 2. P. melanampyx, 3. P. fasciatus, 4. P. sarana spirulus, 5. P. ophicephalus, 6. P. bimaculatus, 7.
P. ticto, 8. P. conconius, 9. P. mahecola, 10. P. denisonii, 11. P. carnaticus, 12. P. Sophore, 13. P. amphibious, 14. P.
dorsalis, 15. P. chola

and in P. fasciatus 1.19 to 1.69 (1.40) which is greater
than that in P. melanampyx 1.14 to 1.72 (1.34).
Distance between rudimentary neural arch and epural
of P. filamentosus is 3.30 to 3.40 (3.36) and in
P. fasciatus is 2.90 to 2.95 (2.92) which is greater
than that in P. melanampyx 2.30 to 3.30 (2.63).
Vertebral column length of P. filamentosus 32.26 to
40.34 (35.32) and P. melanampyx 24.56 to 26.23
(25.67) is greater than P. fasciatus 24.04 to 27.18
150
(25.39). Vertebral column width of P. filamentosus
0.17 to 0.22 (0.20) and P. fasciatus 0.16 to 0.50 (0.36)
is greater than P. melanampyx 0.16 to 0.17 (0.17).
Eye ball diameter of P. filamentosus 3.56 to 3.78
(3.70) and P. fasciatus 1.38 to 2.21 (1.88) is greater
than P. melanampyx 1.40 to 1.66 (1.58). Operculum
length of P. filamentosus 6.30 to 6.92 (6.67) and
P. fasciatus 5.47 to 6.02 (5.80) is greater than
P. melanampyx 5.33 to 5.49 (5.39). Operculum width
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

Table 6. Comparative osteological characters of Puntius species.

Characters P. filamentosus(n=5) P. melanampyx(n=3) P. fasciatus(n=5)

Range Mean SD Range Mean SD Range Mean SD
DRNA&E 3.30-3.40 3.36 0.05 2.30-3.30 2.63 0.57 2.90-2.95 2.92 0.03
UJL 2.58-2.71 2.66 0.06 1.60-1.80 1.67 0.12 1.50-1.80 1.62 0.16
LJL 2.09-2.24 2.18 0.06 1.14-1.72 1.34 0.33 1.19-1.69 1.4 0.27
DFSL 11.75-12.58 12.25 0.31 7.17-7.46 7.27 0.17 7.32-7.90 7.64 0.3
DFRL 11.71-12.54 12.21 0.31 8.13-8.43 8.23 0.17 7.76-8.26 8.01 0.23
PFSL 9.28-9.94 9.68 0.26 6.50-6.52 6.51 0.01 4.68-6.40 5.39 0.93
PFRL 9.18-9.73 9.51 0.21 7.19-7.41 7.34 0.13 4.68-7.09 5.66 1.3
PVFSL 9.45-10.10 9.84 0.25 4.72-4.78 4.74 0.03 4.71-5.48 5.16 0.41
PVFRL 8.84-9.43 9.2 0.23 5.09-5.39 5.19 0.17 5.31-5.48 5.4 0.09
AFSL 6.77-7.25 7.06 0.18 4.99-5.06 5.01 0.04 4.96-5.12 5.04 0.08
AFRL 6.31-6.97 6.7 0.27 5.50-5.85 5.62 0.2 5.05-5.74 5.35 0.35
LE 1.01-1.06 1.04 0.02 0.03-0.31 0.12 0.16 0.31-0.52 0.44 0.12
LRNA 0.32-0.41 0.37 0.05 0.16-0.32 0.27 0.09 0.11-0.16 0.13 0.03
LCC 0.06-0.09 0.08 0.02 0.06-0.32 0.15 0.15 0.32-0.32 0.32 0
WCC 0.02-0.04 0.03 0.01 0.02-0.20 0.08 0.1 0.20-0.20 0.2 0
VCL 32.26-40.34 35.32 3.02 24.56-26.23 25.67 0.96 24.04-27.18 25.39 1.35
VCW 0.17-0.22 0.2 0.02 0.16-0.17 0.17 0.01 0.16-0.50 0.36 0.18
LP 2.22-2.61 2.46 0.17 1.31-1.57 1.4 0.15 1.50-1.54 1.53 0.02
LH1 2.27-2.63 2.48 0.16 1.34-1.39 1.36 0.03 1.36-1.43 1.4 0.03
LH2 1.80-2.02 1.93 0.1 0.53-0.85 0.74 0.18 0.52-1.41 1.05 0.49
LH3 1.72-1.94 1.85 0.1 1.45-1.76 1.66 0.18 1.35-1.42 1.39 0.03
LH4 0.67-0.94 0.83 0.15 0.62-0.67 0.65 0.03 0.62-0.74 0.69 0.07
LH5 0.58-0.62 0.6 0.02 0.53-0.58 0.57 0.03 0.50-0.54 0.52 0.02
LH6 0.43-0.52 0.48 0.04 0.32-0.43 0.39 0.07 0.25-0.32 0.28 0.04
LNS 3.34-3.93 3.69 0.28 1.44-1.48 1.45 0.02 1.45-1.82 1.67 0.2
LHS 3.27-3.54 3.36 0.11 2.51-2.72 2.58 0.12 2.47-2.64 2.56 0.08
EBD 3.56-3.78 3.7 0.08 1.40-1.66 1.58 0.15 1.38-2.21 1.88 0.45
OL 6.30-6.92 6.67 0.32 5.33-5.49 5.39 0.09 5.47-6.02 5.8 0.3
OW 4.12-4.96 4.46 0.43 2.48-2.91 2.76 0.25 2.49-2.84 2.69 0.18

of P. filamentosus 4.12 to 4.96 (4.46) and
P. melanampyx 2.48 to 2.91 (2.76) is greater than
P. fasciatus 2.49 to 2.84 (2.69) in standard length.
Based on meristic analysis it shows similarity in
counts of dorsal fin, anal fin, pectoral fin, pelvic fin
and vertebral column count of P. filamentous (31),
P. melanampyx (30) and P. fasciatus (32) shows
variations.
Similarly with P. dorsalis, P. mahecola and
P. amphibious are deep bodied. Upper jaw length of
P. dorsalis is 3.50 to 3.60 (3.55) and P. mahecola 2.79
to 2.92 (2.87) is greater than P. amphibious 1.90 to
3.57 (2.72). Lower jaw length of P. mahecola 3.06 to
3.17 (3.10) and P. dorsalis 2.81 to 2.89 (2.85) is
151
greater than P. amphibious 1.80 to 2.61 (2.33).
Distance between rudimentary neural arch and
Epural of P. mahecola 0.87 to 3.40 (1.71) and
P. dorsalis 0.43 to 0.44 (0.43) is greater than
P. amphibious 0.02 to 0.08 (0.04). Vertebral column
length of P. amphibious 48.99 to 51.59 (50.17) and
P. mahecola 21.93 to 25.69 (24.34) is greater than
P. dorsalis 19.98 to 21.91(20.94). Vertebral column
width of P. amphibious 1.96 to 2.37 (2.13) and
P. mahecola 0.42 to 0.44 (0.43) is greater than
P. dorsalis 0.02 to 0.04 (0.03). Eye ball diameter of
P. amphibious 4.04 to 4.26 (4.19) and P. mahecola
2.84 to 2.93 (2.87) is greater than P. dorsalis 1.55 to
2.51 (2.03). Operculum length of P. amphibious 9.24
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Table 6. Continued.

Characters P. sarana spirulus (n=2) P. ophicephalus (n=5) P. chola (n=5)

Range Mean SD Range Mean SD Range Mean SD
DRNA&E 0.34-0.40 0.37 0.04 0.07-0.54 0.26 0.26 0.43-0.54 0.47 0.06
UJL 3.25-3.51 3.38 0.18 5.26-5.64 5.44 0.13 3.59-3.98 3.74 0.21
LJL 4.55-4.69 4.62 0.1 3.38-4.34 3.79 0.35 2.85-3.20 2.98 0.16
DFSL 9.55-9.62 9.59 0.05 9.72-12.79 10.91 1.21 7.22-8.56 7.73 0.62
DFRL 12.35-12.64 12.49 0.21 12.02-15.18 13.21 1.45 6.09-6.86 6.37 0.34
PFSL 12.30-12.31 12.31 0.01 9.90-13.23 11.37 1.19 4.90-5.40 5.08 0.22
PFRL 12.29-12.32 12.31 0.01 9.90-13.21 11.36 1.19 4.50-5.29 4.8 0.37
PVFSL 10.31-10.48 10.4 0.12 9.09-11.13 9.86 0.85 4.92-5.51 5.14 0.26
PVFRL 10.31-10.48 10.4 0.12 9.09-11.12 9.86 0.85 4.91-5.49 5.13 0.26
AFSL 10.26-10.32 10.29 0.04 9.51-11.73 10.32 1.02 5.44-5.93 5.72 0.19
AFRL 10.26-10.32 10.29 0.04 9.48-11.74 10.29 1.03 5.78-6.14 5.9 0.14
LE 0.74-0.82 0.78 0.05 1.59-1.83 1.69 0.09 1.54-1.96 1.79 0.22
LRNA 0.21-0.34 0.27 0.09 0.53-0.67 0.61 0.07 0.35-0.43 0.38 0.04
LCC 0.12-0.19 0.15 0.05 0.32-0.53 0.45 0.11 0.02-0.05 0.04 0.02
WCC 0.01-0.05 0.03 0.03 0.22-0.35 0.29 0.07 0.01-0.04 0.03 0.02
VCL 40.81-43.44 42.13 1.86 24.22-53.35 40.35 13.49 21.98-27.13 25.29 2.06
VCW 0.77-0.90 0.84 0.09 0.54-0.74 0.65 0.08 0.22-0.29 0.25 0.03
LP 2.12-2.18 2.15 0.04 2.28-2.93 2.58 0.31 1.59-2.55 1.96 0.49
LH1 2.23-2.29 2.26 0.05 2.28-2.77 2.51 0.24 0.56-0.82 0.71 0.13
LH2 1.85-1.93 1.89 0.05 2.01-2.47 2.22 0.2 0.43-0.85 0.68 0.22
LH3 1.56-1.58 1.57 0.01 1.80-2.19 1.95 0.16 0.32-0.73 0.56 0.22
LH4 1.42-1.45 1.44 0.02 1.02-1.05 1.03 0.01 0.22-0.55 0.42 0.18
LH5 1.19-1.26 1.23 0.05 0.58-0.77 0.68 0.07 0.18-0.50 0.37 0.17
LH6 1.01-1.08 1.04 0.05 0.48-0.65 0.56 0.07 0.15-0.24 0.2 0.05
LNS 2.34-2.40 2.37 0.05 2.19-2.81 2.44 0.23 2.05-2.59 2.26 0.27
LHS 2.34-2.42 2.38 0.06 2.69-3.83 3.11 0.52 2.48-3.10 2.71 0.28
EBD 4.03-4.04 4.04 0 3.35-4.23 3.71 0.33 1.80-2.17 1.94 0.18
OL 9.22-9.24 9.23 0.01 7.26-8.03 7.64 0.29 3.42-4.44 4.03 0.55
OW 6.13-6.24 6.18 0.07 4.09-4.75 4.46 0.28 2.65-2.96 2.77 0.15

to 9.46 (9.38) and P. dorsalis 4.51 to 5.49 (5.00) is
greater than P. mahecola 4.66 to 4.69 (4.67).
Operculum width of P. amphibious 7.23 to 8.19
(7.87) and P. dorsalis 3.13 to 3.59 (3.36) is greater
than P. mahecola 2.97 to 3.02 (3.00) in standard
length. Based on meristic analysis it shows similarity
in counts of dorsal fin, anal fin, pectoral fin, pelvic
fin and vertebral column count of P. dorsalis (33),
P. mahecola (35) and P. amphibious (30) shows
variations.
Likewise comparing P. sarana spirulus with
P. ophicephalus are deep bodied, having two pair of
barbels (maxillary and rostral) both species. Upper
jaw length of P. ophicephalus 5.26 to 5.64 (5.44) is
greater than P. sarana spirulus 3.25 to 3.51 (3.38).
Lower jaw length of P. sarana spirulus 4.55 to 4.69
(4.62) is greater than P. ophicephalus 3.38 to 4.34
152
(3.79). Distance between rudimentary neural arch
and epural of P. sarana spirulus 0.34 to 0.40 (0.37) is
greater than P. ophicephalus 0.07 to 0.54 (0.26).
Vertebral column length of P. sarana spirulus 40.81
to 43.44 (42.13) is greater than P. ophicephalus 24.22
to 53.35 (40.35). Vertebral column width of P. sarana
spirulus 0.77 to 0.90 (0.84) is greater than
P. ophicephalus 0.54 to 0.74 (0.65). Eye ball
diameter of P. sarana spirulus 4.03 to 4.04 (4.04) is
greater than P. ophicephalus 3.35 to 4.23 (3.71).
Operculum length of P. sarana spirulus 9.22 to 9.24
(9.23) is greater than P. ophicephalus 7.26 to 8.03
(7.64). Operculum width of P. sarana spirulus 6.13 to
6.24 (6.18) is greater than P. ophicephalus 4.09 to
4.75 (4.46) in standard length. Based on meristics
analysis it shows similarity in counts of dorsal fin,
pectoral fin, and pelvic fin, and only vertebral
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

Table 6. Continued.
Characters P. ticto (n=5) P. conconius (n=5) P. dorsalis (n=2)
Range Mean SD Range Mean SD Range Mean SD
DRNA&E 0.34-0.40 0.37 0.04 0.07-0.54 0.26 0.26 0.43-0.54 0.47 0.06
UJL 3.25-3.51 3.38 0.18 5.26-5.64 5.44 0.13 3.59-3.98 3.74 0.21
LJL 4.55-4.69 4.62 0.1 3.38-4.34 3.79 0.35 2.85-3.20 2.98 0.16
DFSL 9.55-9.62 9.59 0.05 9.72-12.79 10.91 1.21 7.22-8.56 7.73 0.62
DFRL 12.35-12.64 12.49 0.21 12.02-15.18 13.21 1.45 6.09-6.86 6.37 0.34
PFSL 12.30-12.31 12.31 0.01 9.90-13.23 11.37 1.19 4.90-5.40 5.08 0.22
PFRL 12.29-12.32 12.31 0.01 9.90-13.21 11.36 1.19 4.50-5.29 4.8 0.37
PVFSL 10.31-10.48 10.4 0.12 9.09-11.13 9.86 0.85 4.92-5.51 5.14 0.26
PVFRL 10.31-10.48 10.4 0.12 9.09-11.12 9.86 0.85 4.91-5.49 5.13 0.26
AFSL 10.26-10.32 10.29 0.04 9.51-11.73 10.32 1.02 5.44-5.93 5.72 0.19
AFRL 10.26-10.32 10.29 0.04 9.48-11.74 10.29 1.03 5.78-6.14 5.9 0.14
LE 0.74-0.82 0.78 0.05 1.59-1.83 1.69 0.09 1.54-1.96 1.79 0.22
LRNA 0.21-0.34 0.27 0.09 0.53-0.67 0.61 0.07 0.35-0.43 0.38 0.04
LCC 0.12-0.19 0.15 0.05 0.32-0.53 0.45 0.11 0.02-0.05 0.04 0.02
WCC 0.01-0.05 0.03 0.03 0.22-0.35 0.29 0.07 0.01-0.04 0.03 0.02
VCL 40.81-43.44 42.13 1.86 24.22-53.35 40.35 13.49 21.98-27.13 25.29 2.06
VCW 0.77-0.90 0.84 0.09 0.54-0.74 0.65 0.08 0.22-0.29 0.25 0.03
LP 2.12-2.18 2.15 0.04 2.28-2.93 2.58 0.31 1.59-2.55 1.96 0.49
LH1 2.23-2.29 2.26 0.05 2.28-2.77 2.51 0.24 0.56-0.82 0.71 0.13
LH2 1.85-1.93 1.89 0.05 2.01-2.47 2.22 0.2 0.43-0.85 0.68 0.22
LH3 1.56-1.58 1.57 0.01 1.80-2.19 1.95 0.16 0.32-0.73 0.56 0.22
LH4 1.42-1.45 1.44 0.02 1.02-1.05 1.03 0.01 0.22-0.55 0.42 0.18
LH5 1.19-1.26 1.23 0.05 0.58-0.77 0.68 0.07 0.18-0.50 0.37 0.17
LH6 1.01-1.08 1.04 0.05 0.48-0.65 0.56 0.07 0.15-0.24 0.2 0.05
LNS 2.34-2.40 2.37 0.05 2.19-2.81 2.44 0.23 2.05-2.59 2.26 0.27
LHS 2.34-2.42 2.38 0.06 2.69-3.83 3.11 0.52 2.48-3.10 2.71 0.28
EBD 4.03-4.04 4.04 0 3.35-4.23 3.71 0.33 1.80-2.17 1.94 0.18
OL 9.22-9.24 9.23 0.01 7.26-8.03 7.64 0.29 3.42-4.44 4.03 0.55
OW 6.13-6.24 6.18 0.07 4.09-4.75 4.46 0.28 2.65-2.96 2.77 0.15

column count of P. sarana spirulus (34), and
P. ophicephalus (38) shows variations.
In regard with P. denisonii and P. bimaculatus
both are deep bodied and has one pair of barbels
(maxillary) whereas rostral and mandibular barbels
are absent. Upper jaw length of P. denisonii 1.01 to
1.07 (1.04) is greater than P. bimaculatus 0.10 to 0.11
(0.10). Lower jaw length of P. bimaculatus 1.31 to
1.69 (1.54) is greater than P. denisonii 0.96 to 1.06
(1.01). Distance between rudimentary neural arch
and epural of P. bimaculatus 0.16 to 0.18 (0.17) is
greater than P. denisonii 0.02 to 0.04 (0.03).
Vertebral column length of P. denisonii 23.48 to
29.10 (26.29) is greater than P. bimaculatus 24.18 to
26.48 (25.40). Vertebral column width of P. denisonii
0.37 to 0.54 (0.46) is greater than P. bimaculatus 0.10
to 0.20 (0.16). Eye ball diameter of P. bimaculatus
1.45 to 2.32 (1.84) is greater than P. denisonii 0.26 to
0.44 (0.35). Operculum length of P. bimaculatus 3.59
to 4.82 (4.40) is greater than P. denisonii 2.56 to 2.64
(2.60). Operculum width of P. bimaculatus 1.58 to
1.83 (1.74) is greater than P. denisonii 1.21 to 1.61
(1.41) in standard length. Based on meristics analysis
it shows similarity in counts of dorsal fin, pectoral
fin, and pelvic fin, and only lateral line scale, anal fin
and caudal fin of P. bimaculatus (32) and P. denisonii
(30) show variations.
Similarly, with P. conchonius, P. chola and P. ticto
are deep bodied. Upper jaw length of P. conconius
4.48 to 5.51 (4.70) and P. chola 3.59 to 3.98 (3.74) is
greater than P. ticto 2.95 to 3.95 (3.45). Lower jaw
length of P. ticto 3.10 to 3.70 (3.46) and P. conconius
2.85 to 3.94 (3.10) is greater than P. chola 2.85 to
3.20 (2.98). Distance between rudimentary neural

153
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Table 6. Continued.

Characters P. mahecola (n=3) P. denisonii (n=2) P. carnaticus (n=3)

Range Mean SD Range Mean SD Range Mean SD
DRNA&E 0.87-3.40 1.71 1.46 0.02-0.04 0.03 0.01 0.05-0.08 0.07 0.02
UJL 2.79-2.92 2.87 0.07 1.01-1.07 1.04 0.04 3.52-3.57 3.55 0.03
LJL 3.06-3.17 3.1 0.06 0.96-1.06 1.01 0.07 2.57-4.71 3.29 1.24
DFSL 12.22-12.78 12.42 0.32 7.53-8.14 7.84 0.43 9.61-15.56 13.55 3.41
DFRL 8.89-9.32 9.05 0.24 7.32-7.98 7.65 0.46 15.72-18.29 17.4 1.45
PFSL 4.39-4.54 4.44 0.08 2.48-2.95 2.71 0.33 13.37-18.35 16.67 2.85
PFRL 3.90-4.04 3.95 0.07 2.46-2.83 2.65 0.26 14.37-17.53 16.45 1.8
PVFSL 8.29-8.60 8.4 0.17 2.75-3.07 2.91 0.23 15.36-16.10 15.83 0.41
PVFRL 7.76-8.05 7.86 0.16 2.67-2.73 2.7 0.04 15.36-16.10 15.83 0.41
AFSL 5.69-5.93 5.77 0.14 3.84-3.89 3.86 0.03 15.31-16.99 16.4 0.94
AFRL 5.69-5.93 5.77 0.14 3.65-3.67 3.66 0.01 15.31-16.99 16.4 0.94
LE 1.38-1.41 1.39 0.02 0.34-0.68 0.51 0.24 0.82-3.11 2.35 1.32
LRNA 0.25-0.25 0.25 0 0.22-0.55 0.39 0.24 0.03-0.34 0.13 0.18
LCC 0.20-0.20 0.2 0 0.13-0.20 0.16 0.05 0.19-0.45 0.36 0.15
WCC 0.08-0.08 0.08 0 0.05-0.09 0.07 0.03 0.05-0.23 0.17 0.1
VCL 21.93-25.69 24.34 2.1 23.48-29.10 26.29 3.98 62.61-68.72 66.26 3.22
VCW 0.42-0.44 0.43 0.01 0.37-0.54 0.46 0.12 0.89-2.37 1.88 0.85
LP 1.53-1.61 1.56 0.04 0.24-0.33 0.29 0.07 2.20-3.09 2.79 0.51
LH1 1.60-1.66 1.62 0.04 0.10-0.19 0.15 0.06 2.31-3.48 3.08 0.67
LH2 1.50-1.55 1.52 0.03 0.13-0.15 0.14 0.01 1.94-2.79 2.5 0.49
LH3 0.61-0.63 0.62 0.01 0.11-0.93 0.52 0.58 1.59-2.14 1.95 0.31
LH4 0.55-0.55 0.55 0 0.10-0.80 0.45 0.5 1.42-1.63 1.56 0.12
LH5 0.22-0.22 0.22 0 0.49-0.92 0.7 0.31 1.03-1.25 1.11 0.12
LH6 0.12-0.12 0.12 0 0.29-0.51 0.4 0.15 0.64-1.07 0.78 0.25
LNS 2.49-2.58 2.52 0.05 0.45-0.97 0.71 0.37 2.42-4.78 3.99 1.36
LHS 2.11-2.26 2.16 0.09 0.98-1.40 1.19 0.3 2.44-4.53 3.82 1.19
EBD 2.84-2.93 2.87 0.05 0.26-0.44 0.35 0.13 4.05-4.26 4.19 0.11
OL 4.66-4.69 4.67 0.02 2.56-2.64 2.6 0.06 9.25-9.46 9.38 0.12
OW 2.97-3.02 3 0.03 1.21-1.61 1.41 0.28 7.22-8.20 7.87 0.56

arch and epural of P. conconius 1.24 to 1.42 (1.32)
and P. ticto 0.54 to 0.67 (0.62) is greater than P. chola
0.43 to 0.54 (0.47). Vertebral column length of
P. conconius 27.98 to 32.72 (29.94) and P. ticto 25.48
to 26.09 (25.60) is greater than P. chola 21.98 to
27.13 (25.29). Vertebral column width of P. ticto
0.29 to 0.73 (0.55) and P. conconius 0.34 to 0.54
(0.42) is greater than P. chola 0.22 to 0.29 (0.25). Eye
ball diameter of P. conconius 1.68 to 2.74 (2.11) and
P. ticto 0.65 to 2.10 (1.52) is greater than P. chola
1.80 to 2.17 (1.94). Operculum length of
P. conconius 5.87 to 6.91 (6.29) and P. chola 3.42 to
4.44 (4.03) is greater than P. ticto 3.29 to 4.44 (3.57).
Operculum width of P. conconius 3.57 to 4.62 (3.99)
and P. chola 2.65 to 2.96 (2.77) is greater than P. ticto
1.94 to 2.96 (2.55) in standard length. Based on
meristic analysis it shows similarity in counts of
154
dorsal fin, anal fin, pectoral fin, pelvic fin and
vertebral column count of P. conconius (32), P. chola
(33) and P. ticto (33) shows variations.
Osteological morphometric adds more as a
quantitative element to descriptions of the individual,
allowing more clarified comparisons. It enables to
describe complex body shapes in exact and permits
numerical comparison between different species and
also when combined with multivariate statistical
methods (e.g. Principal Component Analysis, Cluster
Analysis etc.) they offer powerful tool for testing and
displaying differences between the individuals. PCA
on morphometric measurements normally results in a
first principal component characterized by
consistently positive loadings, where size is the
primary source of variance in the data. However,
because we analyzed standardized measurements to
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

Table 6. Continued.

Characters P. sophore (n=5) P. amphibious (n=3) P. bimaculatus (n=5)

Range Mean SD Range Mean SD Range Mean SD
DRNA&E 0.10-0.54 0.36 0.24 0.02-0.08 0.04 0.03 0.16-0.18 0.17 0.01
UJL 0.15-2.63 1.59 1.31 1.90-3.57 2.72 0.84 0.10-0.11 0.1 0.01
LJL 1.31-3.83 2.53 1.16 1.80-2.61 2.33 0.46 1.31-1.69 1.54 0.2
DFSL 6.12-8.13 6.56 0.87 14.30-15.53 14.85 0.62 6.00-6.28 6.12 0.11
DFRL 4.23-6.16 4.78 0.8 14.21-18.26 15.65 2.26 6.03-6.34 6.16 0.13
PFSL 3.29-4.32 3.86 0.51 10.63-18.33 13.39 4.29 4.32-4.48 4.42 0.07
PFRL 3.49-4.30 3.87 0.35 9.97-17.51 12.83 4.09 4.01-4.54 4.33 0.27
PVFSL 3.05-4.49 3.79 0.71 9.64-16.08 11.85 3.66 3.44-3.53 3.48 0.04
PVFRL 3.05-4.35 3.73 0.64 9.56-16.08 11.78 3.72 3.35-3.49 3.43 0.06
AFSL 3.00-3.95 3.21 0.41 8.78-16.96 11.59 4.66 3.01-4.51 3.91 0.8
AFRL 3.06-4.00 3.31 0.39 8.25-16.96 11.42 4.81 3.19-4.53 3.99 0.71
LE 0.18-0.33 0.25 0.07 1.23-3.11 2.17 0.94 0.28-0.31 0.3 0.02
LRNA 0.04-0.45 0.28 0.22 0.03-1.01 0.51 0.49 0.04-0.11 0.08 0.04
LCC 0.02-0.16 0.08 0.08 0.42-0.65 0.51 0.12 0.08-0.16 0.11 0.04
WCC 0.01-0.11 0.05 0.05 0.23-0.57 0.38 0.17 0.11-0.20 0.16 0.05
VCL 26.40-29.06 27.97 0.98 48.99-51.59 50.17 1.32 24.18-26.48 25.4 0.83
VCW 0.01-0.29 0.16 0.14 1.96-2.37 2.13 0.21 0.10-0.20 0.16 0.06
LP 0.10-0.65 0.33 0.29 2.19-3.09 2.79 0.52 0.64-0.84 0.7 0.08
LH1 0.20-0.41 0.29 0.11 2.30-3.48 3.09 0.68 0.22-0.41 0.34 0.08
LH2 0.30-0.37 0.33 0.03 1.93-2.79 2.5 0.49 0.14-0.35 0.23 0.11
LH3 0.30-0.37 0.32 0.03 1.58-2.14 1.95 0.32 0.12-0.31 0.2 0.11
LH4 0.21-0.26 0.23 0.03 1.42-1.63 1.56 0.12 0.10-0.26 0.17 0.09
LH5 0.16-0.21 0.19 0.02 1.06-1.25 1.12 0.11 0.08-0.21 0.13 0.07
LH6 0.08-0.12 0.1 0.02 0.64-1.07 0.78 0.25 0.05-0.12 0.08 0.04
LNS 1.24-1.61 1.45 0.19 2.41-4.78 3.99 1.37 1.52-1.63 1.58 0.05
LHS 0.75-1.48 1.12 0.36 2.43-4.52 3.82 1.21 1.20-1.51 1.33 0.16
EBD 0.65-2.30 1.34 0.88 4.04-4.26 4.19 0.12 1.45-2.32 1.84 0.44
OL 3.29-4.81 3.93 0.8 9.24-9.46 9.38 0.12 3.59-4.82 4.4 0.52
OW 1.77-3.84 2.66 1.07 7.23-8.19 7.87 0.55 1.58-1.83 1.74 0.11

Table 7. Distances between the central objects.

P. sarana spirulus P. ticto P. carnaticus

P. sarana spirulus 0
P. ticto 23.391 0
P. carnaticus 33.271 54.568 0

eliminate the size effect, no such effect is expected.
The Dendrogram below expresses that the species
has their individuality between the species but still
the genus Puntius seems to have ambiguities among
themselves. The Cluster Analysis (CA), shows that
the species were grouped into three groups (Table. 7).
The first class has 10 species P. mahecola, P. chola,
P. bimaculatus, P. dorsalis, P. melanampyx,
P. fasciatus, P. ticto, P. denisonii, P. sophore,
P. conchonius. The second class has 4 species;
P. filamentous, P. sarana spirulus, P. amphibious and
155
P. ophicephalus. The third class has only one species
P. carnaticus. In regard to all the 15 species from
southern Western Ghats it is clear that the variation
within the species is 24.88% and between the species
is 75.12%. It is observed that there is no overlapping
of osteological bone measurements and meristic
characters between the species (Fig. 9).
The species mainly vary with contribution of the
observation (%) to factor I in vertebral column length
(33.943), vertebral column count (40.081), (Figs. 10,
11). All other character plays a supportive role to
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166

Fig.9. Dendrogram showing the similarity between the species.

express the variations between the species. The Discussion

above plots clearly explain that based on the Osteo –
morphometric measurements and meristic characters
the species are clubbed together which strongly
expresses that all species have a common ancestral
phylogeny.
156
As per Jayaram (2010), a total of 73 valid Puntius
species have been described worldwide, out of this
39 species is distributed in India. Twenty-nine
species are distributed to Southern Western Ghats
and the rest of 10 species are widely distributed in
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

Fig.10. The scores of PCA plotted against component I for pooled osteo-morphometric measurements representing the
characters.

Fig.11. The scores of PCA plotted against component I for pooled osteo-morphometric measurements representing the
characters and species.

157
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166
India. Among the 15 species P. chola, P. dorsalis,
P. sophore and P. amphibious the ethmoid appears to
be slightly broader and protected by a strong ossified
prevormer above it. While the rest of the 11 species
have a little narrow ethmoid and non-ossified
prevormer comparing to the above said species.
The paired preethmoids are articulated laterally
with the articular junction of ethmoid and prevomer
in all the species studied. According to Weitzman
(1962) the presence of a preethmoid is an
evolutionary advanced character of Cypriniformes.
While comparing the other orders, it was also noted
in Notropis bifrenatus (Harrington, 1955), Cyprinus
carpio (Wilimovsky and Weitzman, 1955) and
Schizothoraichthys niger (Yousuf et al. 1988).
However it was not found in Brycon meeki which
belong to the family Characidae of Cypriniformes
(Weitzman 1962).
The rostral bone or kinethmoid is a specialization
of cyprinids (Weitzman 1962). As in other cyprinids,
the rostral bone is also present in Puntius
(kinethmoid; Harrington 1955; Yousuf et al. 1988). It
helps to lever the premaxilla in the protrusion of
mouth. The lateral ethmoid articulates laterally with
prevomer, anterodorsally with ethmoid and latero-
posteriorly with prefrontals. The anterior margin of
prevomer is little ahead of the ethmoid.
In Cyprinids, the position of the anterior margin of
the prevomer of Puntius is always extends more
forward than that of preethmoid. Among the 15
species, four species namely, P. chola, P. dorsalis,
P. sophore and P. amphibious have a prominent
dorso-median cranial fontanel. The above character
supports the statement given by Weitzman (1962)
that there is a long cranial fontanel in the dorsal side
of the skull between the two frontals and parietals.
There is a pineal foramen on the dorsal side of the
skull surrounded by the frontals and parietals.
However, in Puntius, there is fontanel in certain
species and in the rest, a small pineal foramen
surrounded by frontals and parietals.
The pineal foramen is found also in Notropis
bifrenatus (Harrington 1955), but not reported in
158
S. niger (Yousuf et al. 1988). According to Weitzman
(1962) and Ramaswami (1956), the frontal bones of
all cyprinids except Esomus are in contact with the
pterotic and it is a primitive feature of the Cyprinids.
In all the species of Puntius studied, the frontal is in
contact with pterotic. Therefore, genus Puntius is
characterized as more primitive than Esomus.
The rostral bone is well-developed. The two orbits
are completely separated by interorbital septum. The
basisphenoid is absent in Puntius species. Mouth is
terminal and protrusible. There are five infraorbitals
and one supraorbital. The supraorbital firmly
articulates with the frontals.
In the species of Puntius studied, there seems to be
a relationship between the number of rays and the
proximal pterygiophores of the dorsal fin. The
number of proximal pterygiophores is equal to the
number of rays plus two. According to Weitzman
(1962) in evolutionarily advanced fishes the tripartite
character of pterygiophores of median fins greatly
reduced and form one piece of bone. The
intermediate pterygiophores are absent only on first
four or five pterygiophores of the dorsal fin. In other
pterygiophore the tripartite character is easily visible.
Therefore, based on the structure of pterygiophores
of the dorsal fin, the species of Puntius studied are
not evolutionarily advanced. There are three spines
and 5 rays in the anal fin and one spine and 8 or 9
rays in the pelvic fin. The number of rays of the
pectoral fin vary from 10-17. The number of
vertebrae vary from 27-41.
In the present study, the caudal fin has 6 hypurals,
1 parahypural, 2 pairs of uroneurals and 2 epiurals on
the caudal fin were noted in most of the species
analyzed. Based on Jinadasa & Kotalawala (1991)
caudal skeleton, all the species of Puntius are
primitive fishes and also the caudal fin has 6
hypurals, 1 parahypural, 2 pairs of uroneurals and 2
epiurals on the caudal fin. Caudal fin rays Celestial
Pearl Danio of were supported by the neural and
hemal spines of the 2nd and 3rd preural caudal centra,
the pleurostyle, a single epural, 5 hypural elements
and the parahypural (Conway et al. 2008).
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
Among the 15 species P. carnaticus and P. sophore
have 6 supraneurals whereas in rest of the species
only 4 was noted. Moreover, P. bimaculatus and
P. denisonii has 9 and P. carnaticus has 10 predorsal
vertebral counts supports the data given by
Shantakumar & Vishwanath (2006). Conway et al.
(2008) worked on The “Celestial Pearl Danio” a
miniature species and stated that there were five
small plate-like supraneurals, situated between the
neural spines. Based on the number of predorsals and
the number of spines articulated to the first
pterygiophores and the others, the dorsal fin formulae
can be formulated for the species of Puntius. Among
the 15 species P. melanamplyx, P. fasciatus, P. ticto,
P. carnaticus, P. denisonii, P. filamentosus,
P. amphibious, P. sarana spirulus and
P. ophicephalus 4 dorsal spines were noted whereas
in rest of the species only 3 were noted. In evolution,
the changes of these formulae occur due to the loss
of anterior spines, reduction of predorsals and
backward shifting of anterior spines. Therefore, the
differences of dorsal fin formulae of Puntius are also
may be due to the backward shift of spines. Moreover
species of Puntius contains only one postcleithrum.
The reduction of the number of postcleithra is
probably an evolutionarily advanced character in the
pectoral girdle of B. meeki (Cypriniforms:
Characidae) there are three post cleithra (Weitzman
1962). In Bonitos and Sciaenids there are two post
cleithra (Collettle & Chao 1975; Topp & Cole 1968).
These osteological features of Puntius may be
common to other species of the genus and also to the
other genera of the subfamily Cyprininae.
In regard to the vertebral column count
P. filamentosus (31), P. melanampyx (30),
P. fasciatus (32), P. sarana spirulus (34), P. chola
(33), P. amphibius (30), P. mahecola (35), and
P. denisonii (30). Rather there are few exceptional
cases in the genus Puntius which has vertebral counts
more than the value stated by Shanthakumar (2006).
P. ophicephalus (38) and P. carnaticus (42). The total
vertebral count ranges from 35- 36 in P. sarana group,
29-33 in P. conchonius and P. sophore groups
159
(Shanthakumar 2006).
Hence comparing all of the above results it was
observed that the species were grouped into three
groups. The first class has 10 species i.e. P. mahecola,
P. chola, P. bimaculatus, P. dorsalis, P. melanampyx,
P. fasciatus, P. ticto, P. denisonii, P. sophore and
P. conchonius. The second class has 4 species;
P. filamentous, P. sarana spirulus, P. amphibious and
P. ophicephalus. The third class has only one species
P. carnaticus. The result supports that the
observations of Shantakumar & Vishwanath (2006)
in regard to the placement of P. sarana and P. sophore
under two groups. Among the three groups obtained
the larger Puntius species group is distinct from the
smaller species. Our result well supports earlier
classification report of Jayaram (1991) stating that
Puntius spp. were classified under 10 groups where
the larger Puntius species group is distinct from the
smaller brightly and beautiful coloured species.
Acknowledgments
We sincerely thank department of Science and
Engineering Research Board, New Delhi
(SERB/F/2284/2013-14. DT. 10.07.2013) funding
for this research, Tamil Nadu, Kerala and Karnataka
Forest department for the permission and logistic
support, our lab research scholars J. Kumar and M.
Kumar for assisting during the field collection.
References
Britz, R. & Conway, K.W. 2009. Osteology of
Paedocypris, a miniature and highly developmentally
truncated fish (Teleostei: Ostariophysi: Cyprinidae).
Journal of Morphology 270: 389-412.
Buhan, P.J. 1972. The comparative osteology of the
caudal skeleton of some North American minnows
(Cyprinidae). American Midland Naturalist 88: 484-
490.
Chen, G.; Fang, F. & Chang, M.M. 2005. A new cyprinid
closely related Tcultrins xenocyprinins from the mid-
tertiary of South China. Journal of Vertebrate
Paleontology 25: 492-501.
Chen, W.J. & Mayden, R.L. 2009. Molecular
systematics of the Cyprinoidea (Teleostei:
 Iran. J. Ichthyol. (June 2018), 5(2): 139-166
Cypriniformes), the world's largest clade of
freshwater fishes: further evidence from six nuclear
genes. Molecular Phylogenetics and Evolution 52:
544-549.
Conway, K.W.; Chen, W.J. & Mayden, R.L. 2008. The
“celestial Pearl Danio” is a miniature Danio (S.S)
(Ostariophysi: Cyprinidae): evidence from
morphology and molecules. Zootaxa 1686: 1-28.
Conway, K.W.; Chen, W. & Mayden, R.L., 2008. The
“Celestial Pearl danio” is a miniature Danio (s.s)
(Ostariophysi: Cyprinidae): evidence from
morphology and molecules. Zootaxa 1686: 1-28.
Dahanukar, N.; Raut, R. & Bhat, A. 2004. Distribution,
endemism and threat status of freshwater fishes in the
Western Ghats of India. Journal of Biogeography
31(1): 123-136.
Detrich, W.H.; Westerfield, M. & Zon, L. 1999. The
Zebra fish, Gentics and Genomics, Vol. 60. San
Diego: Academic Press.
Eastman, J.T. 1980. The caudal skeletons of Catostomid
fishes. American Midland Naturalist 103(1): 133-
148.
Elliott, N.G.; Haskard, K. & Koslow, J.A. 1995.
Morphometric analysis of orange roughy
(Haplostethus atlanticus) off the continental slope of
southern Australia. Journal of Fish Biology 46: 202 -
220.
Fang F. 2003. Phylogenetic analysis of the Asian
cyprinid genus Danio (Teleostei, Cyprinidae). Copeia
714-728.
Fink, S.V. & Fink, W.L. 1981. Interrelationships of the
Ostariophysan fishes (Teleostei). Zoological Journal
of the Linnean Society 72: 297-353.
Fujita. K. 1989. Nomenclature of Cartilaginous elements
in the caudal skeleton of Teleostean fishes. Japanese
Journal of Ichthyology 36(1): 22-29.
Harrington, R.W. 1955. The Osteocranium of the
American cyprinoid fish, Notropis bifrenatus, with an
annotated synonymy of teleost skull bones. Copeia 4:
267-290.
Hora, S.L. & Misra, K.S. 1938. Fish of Deolali, Part: III.
Journal of Bombay Natural History Society 40: 20-
38.
Ishihara, H. 1987. Revision of the western North Pacific
species of the genus Raja. Japan Journal Ichthyol.
34(3): 241-285.
160
Jayaram, K.C. 2010. The Freshwater Fishes of Indian
Region, 2nd edition. Narendra Publications, New
Delhi.
Jinadasa, J. & Kotalawala, A.B. 1991. Interrelationships
among the species of the genus Puntius (Teleostei,
Cyprinidae) as indicated by the caudal skeletons.
Vidyodaya Journal of Science 3(1): 99-107.
Keivany, Y. 2014a. Comparative osteology of the
suspensorial and opercular series in representatives of
the eurypterygian fishes. Iranian Journal of
Ichthyology 1(2): 73-90.
Keivany, Y. 2014b. Comparative osteology of the jaws
in representatives of the eurypterygian fishes.
Research in Zoology 4(2): 29-42.
Keivany, Y. 2014c. Pectoral girdle bones in
eurypterygian fishes. International Journal of Aquatic
Biology 2(5): 253-274.
Keivany, Y. 2014d. Osteology of hyobranchial arches in
eurypterygian fishes. Iranian Journal of Ichthyology
1(3): 129-151.
Keivany, Y. 2017a. Eurypterygii caudal skeleton. Iranian
Journal of Ichthyology 4(1): 11-30.
Keivany, Y. 2017b. Osteological features of
eurypterygian pelvic girdles. Iranian Journal of
Science & Technology 41: 989-1002.
Kortmulder, K. 1972. A comparative study in colour
patterns and behavior in seven Asiatic Barbus
species: a progress report. Behaviour, Supplement 19:
1-331.
Krzywinski, J. & Besansky, N.J. 2003 Molecular
systematics of Anopheles: from subgenera to
subpopulations. Annual Review of Entomology 48:
111-139.
Kullander, S. & Fang, F. 2005. Two new species of
Puntius from Northern Myanmar. American society
of Icthyologists 2: 290-302.
Mehta, R. & Tandon, K.K., 1984. The comparative
morphology of the Osteocranium, the Weberian
apparatus, the girdles and the caudal skeleton of
Indian Cyprinid fishes with their value in systematics.
Records of the Zoological Survey of India.
Occasional Paper no. 58: 1-167.
Nelson, J.S. 2006. Fishes of the World, 4th Edition.
Wiley, New York.
Nybelin O. 1973. Comments on the caudal skeleton of
actinopterygians. In: Interrelationships of Fishes.
 Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India
369-372.
Rainboth, J. 1996. The Taxonomy, Systematics and
Zoogeography of Hypsibarbus a new genus of large
barbs from rivers of south eastern Asia. University of
California press, vol- 129.
Ramaswami, L.S. 1948. The homalopteroid skull. Proc.
Zool. Soc. Lond. 188: 515-530.
Ramaswami, L.S. 1952a. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. The systematic
position of Gyrinochelius Vaillant. Proceedings of the
National Academy of Sciences, India 18: 125-40.
Ramaswami, L.S. 1952b. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 2. The systematic
position of Psilorhynchus Mc Celland. Proceedings of
the National Academy of Sciences, India 18: 141-
150.
Ramaswami, L.S. 1952c. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 3. The skull and
other skeleton structures of homalopteroid fishes.
Proceedings of the National Academy of Sciences,
India 18: 495-519.
Ramaswami, L.S. 1952d. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 4. The skull and
other skeleton structures of gastromyzonid fishes.
Proceedings of the National Academy of Sciences,
India 18: 520-537.
Ramaswami, L.S. 1953. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 5. The skull and the
gas bladder capsule of Cobitidae. Proceedings of the
National Academy of Sciences, India 19: 323-347.
Ramaswami, L.S. 1955a. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 6. The skull and the
weberian apparatus in the subfamily Gobioninae
(Cyprinidae). Acta zool. Stockh 36: 127-158.
Ramaswami, L.S. 1955b. Skeleton of cyprinoid fishes in
relation to phylogenetic studies. 6. The kull and the
weberian apparatus in the subfamily Gobioninae
(Cyprinidae). Acta zool. Stockh 36: 127-158.
Regan, C.T. 1911. The classification of teleostean fishes
of the order Ostariophysi. 1. Cyprinoidei. Annals and
Magazine of Natural History 8(8): 13-39.
Pethiyagoda, R., Meegaskumbura, M. & Maduwage, T.
2012. A synopsis of the south Asian fishes referred to
Puntius (Pisces: Cyprinidae). Ichthyological
Exploration of Freshwaters 23(1): 69-95.
Sanger, J. 2002. Comparitive osteology of the Danio
161
axial skeleton with comments on Danio relationships
based on molecules and morphology. Zoological
Journal of the Linnean society 135: 529-546.
Shantakumar, M. & Viswanath, W. 2006. Inter-
relationship of Puntius Hamilton - Buchanan
(cyprinidae: Cyprininae) found in Manipur, India.
Zoo's Print 21(6): 2279-2283.
Taki, K., Katsuyama, A. & Urushido, T. 1978.
Comparitative morphology and interspecific
relationships of the cyprinid genus Puntius. Japanese
Journal of Ichthyology 25: 1-8.
Talwar, P.K. & Jhingran, A.G. 1991. Inland fishes of
India and adjacent countries. Vol. I, Oxford & IBH
Publishing Co. Pvt. Ltd, New Delhi.
Taylor, W.R. & Van Dyke, G.C. 1985. Revised
procedures for staining and clearing small fishes and
other vertebrates for bone and cartilage study.
Cybium 9: 107-119.
Thorpe RS (1976) Biometric analysis of geographic
variation and racial affnities. Biol Rev 51: 407–452
Tigano, C.; Canapa, A.; Ferrito, V.; Barucca, M.;
Arcidiacono, I.; Deidun, A.; Schembri, P.J. & Olmo,
E. 2006. A study of osteological and molecular
differences in populations of Aphanius fasciatus
Nardo 1827, from the central Mediterranean
(Teleostei, Cyprinodontidae); Marine Biology 194-
224.
Weitzman, S.H. 1962. The osteology of Brycon meeki, a
generalized characid fish, with an osteological
definition of the family. Stanford Ichthyology
Bulletin 8: 1–77.
Yang, C.; Cao, L.; Wang, W.; Yang, Y.; Abbas, K.; Yan,
B.; Wang, H.; Su, L. & Sun, Y. 2009. Comparative
and evolutionary analysis in natural diploid and
tetraploid weather loach Misgurnus anguillicaudatus
based on cytochrome b sequence data in central
China. Environmental Biology of Fishes 86: 145-153.
Iran. J. Ichthyol. (June 2018), 5(2): 139-166

162
Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

163
Iran. J. Ichthyol. (June 2018), 5(2): 139-166

164
Arunkumar et al.- Osteological characterization of the genus Puntius from six river systems of southern Western Ghats, India

165
 ‫‪Iran. J. Ichthyol. (June 2018), 5(2): 139–166‬‬ ‫‪Received: November 2, 2017‬‬
‫‪© 2018 Iranian Society of Ichthyology‬‬ ‫‪Accepted: May 24, 2018‬‬
‫‪P-ISSN: 2383-1561; E-ISSN: 2383-0964‬‬ ‫‪doi: 10.22034/iji.v5i2.25202‬‬
‫‪http://www.ijichthyol.org‬‬

‫مقاله پژوهشی‬
‫ویژگیهای استخوانشناسی جنس ‪( Puntius‬ماهیان استخوانی حقیقی‪ :‬کپورماهیان) از شش‬
‫سیستم رودخانهای جنوب غربی قاتس‪ ،‬هندوستان‬

‫انبو آراواژی آرونکومار‪ ،‬ان ویجایا الکشیمی‪ ،‬آرونچاالم مانیمکاالن‬

‫آزمایشگاه تنوع زیستی و مولکولی‪ ،‬گروه علوم محیطی‪ ،‬دانشگاه بهاراتیار‪ ،‬کویمباتور ‪ ،641046‬هندوستان‪.‬‬

‫چکیده‪ :‬این مقاله اطالعاتی را در مورد ویژگیهای استخوانشناسی جنس ‪ Puntius‬در شش سیستم رودخانهای در جنوب غربی قاتس فراهم‬
‫میآورد‪ .‬ماهیان بااستفاده از تور پرتابی‪ ،‬ساچوک‪ ،‬تور گوشگیر و تور کشش از نهرها و رودخانههای مختلف جنوبغربی قاتس جمعآوری شدند‪.‬‬
‫روشهای مختلف شفافسازی و رنگآمیزی برای مطالعات استخوانشناسی مورد استفاده قرار گرفت‪ .‬پس از شفافسازی و رنگآمیزی دوگانه‪،‬‬
‫نمونهها زیر لوپ مشاهده و با استفاده از دوربین دیجیتال از آنها عکسبرداری گردید‪ .‬در مجموع ‪ 29‬ویژگی ریختشناسی‪ ،‬شمارشی و صفات‬
‫استخوانی اندازهگیری شد‪ .‬آنالیزهای تجزیه به مولفههای اصلی و خوشهای برای گروهبندی گونهها و تشخیص شباهت بین گونهای استفاده‬
‫شد‪ .‬مقایسه همه گونهها نشان داد که گونههای مورد بررسی به سه گروه تقسیم شدند‪ .‬گروه اول شامل ده گونه ‪P. mahecola, P. chola,‬‬
‫‪ ،P. bimaculatus, P. dorsalis, P. melanampyx, P. fasciatus, P. ticto, P. denisonii, P. sophore, P. conchonius‬گروه دوم‬
‫شامل ‪ 4‬گونه ‪ P. filamentous, P. sarana spirulus, P.amphibious and P. ophicephalus‬و گروه سوم تک گونه ‪P. carnaticus‬‬
‫را شامل میشدکه بهخوبی نتایج ‪ Shantakumar‬و ‪ )2006( Vishwanath‬را تایید میکرد‪.‬‬
‫کلماتکلیدی‪ :‬چاالکوری‪ ،‬صفات شمارشی‪ ،‬اندازشی‪ ،‬پیشاتموئید‪ ،‬اسکلت‪ ،‬جمجمه‪.‬‬

‫‪166‬‬

‫‪View publication stats‬‬