J.Natn.Sci.Foundation Sri Lanka 2011 39 (2):121-128
RESEARCH ARTICLE
Osteological variation of the olive barb Puntius sarana (Cyprinidae)
in Sri Lanka
F. I. Irfan and K. B. Suneetha Gunawickrama*
Department of Zoology, Faculty of Science, University of Ruhuna, Matara.
Submitted: 17 December 2010 ; Accepted: 21 January 2011
Abstract: Osteological intra-specific variation was studied
in olive barb Puntius sarana Hamilton 1822, from six water
bodies in Sri Lanka. Discernible variation was reported in the
elements of the cranium, pharyngeal bones and caudal skeleton
of the fish. The curvature of the median suture between frontals
of the skull roof in fish from the rivers Walawe, Nilwala and
the Kirindi Oya was directed opposite to that in fish from the
rivers Menik, Gin and Kalu. Shape of the tip of dorsal limb
of the pharyngeal bones was either most commonly pointed
or truncated. Consistent distinctiveness in the rudimentary
neural arch of the compound centrum was observed in the
Walawe River population, while double neural spines were
present on preural centra (PU2 or PU3) of the caudal skeleton
in some populations. The size of the parhypural foramen and
epural distance in the caudal skeleton, and the total number
of vertebrae of the axial skeleton also showed variation. The
results are the first information on osteological variation of
P. sarana in Sri Lanka.
Keywords: Geographic variation, olive barb, osteological
comparison, Puntius sarana taxonomy.
INTRODUCTION
Puntius sarana Hamilton 1822 is a cyprinid fish widely
distributed across the southern part of Peninsular India
and in Sri Lanka (Jayaram, 1991, 1999; Talwar &
Jhingran, 1991) and is the largest species of the genus
in Sri Lanka (Pethiyagoda, 1991). External appearance
is unequivocal with olive coloured dorsal side, silvery
lateral sides with golden reflection and a conspicuous
black blotch on the caudal peduncle (Deraniyagala,
1952) (Figure 1A). Juveniles are reported to have an
additional black spot on the body just below the dorsal
fin, and fins vary in colour ranging from dusky brown to
orange (Deraniyagala, 1952). Valued as an edible fish in
Sri Lanka, its distribution ranges from lowland waters up
to an elevation of about 500 m, particularly in clear, slow
flowing rivers, streams and lakes with sandy, pebble and
occasionally muddy bottoms (Pethiyagoda, 1991). It can
tolerate brackish water conditions and strong currents
(Kortmulder et al., 1990).
Studies on intra-specific variation are important in
problems of species delineation, as it is acknowledged
that inadequate information on intra-specific geographic
variation can lead to erroneous species descriptions
(Ishihara, 1987). Among other phenotypic tools,
osteological characters are considered to be important
diagnostic traits for interpreting systematics and
phylogenetic relationships within Teleostei (Nybelin,
Figure 1: P. sarana (A) Illustration of a specimen collected from Walawe River (scale bar = 1 cm), and (B) an
osteological preparation (52.4 mm SL; from Nilwala River at Godapitiya)
*
Corresponding author (suneetha@zoo.ruh.ac.lk)
F. I. Irfan & K. B. Suneetha Gunawickrama
122
1973; Arratia, 1983, 1997, 1999; Schultze & Arratia,
1988). Dunn (1983, 1984) reviewed the use of skeletal
structures and the utility of developmental osteology
in taxonomic studies. Osteological characters may also
provide a major tool in examining variability within a
species (Eastman, 1980) since different populations of
the same species, which share external appearance, may
vary in skeletal structures (Hilton & Bemis, 1999). Some
studies (Weisel, 1960; Eastman, 1980; Arratia, 1983;
Sanger & McCune, 2002) have reported intra-specific
variation in caudal and cranial osteology of fishes. Intraspecific variation in P. sarana in Sri Lanka has not been
extensively reported yet, and therefore, taxonomically
important diagnostic variation in the species is poorly
understood. A recent study by Shantakumar & Vishwanath
(2006) has reported some osteological information of
P. sarana group in India, but such studies on P. sarana in
Sri Lanka are not available. The present study aimed to
reveal osteological variation in P. sarana among selected
geographic populations in Sri Lanka. Information on
osteology of P. sarana in Sri Lanka would facilitate
comparative studies with data from other locations of the
distribution range within Asia, and thereby contribute to
the taxonomic clarification of the species in Sri Lanka.
METHODS AND MATERIALS
Sample collection: Samples of P. sarana were collected
from six freshwater bodies, namely, the Menik River
(M) at Kataragama, the Kirindi Oya (K) at Tissa, the
Nilwala River (N) at Godapitiya, the Gin River (G) at
Wakwella, the Kalu River (KR) at Athwelthota, and the
Walawe River (W) at Pattiyapola (tank hydrologically
connected) in Southern and Western Provinces of Sri
Lanka (Figure 2). Sample collection was carried out from
January to December 2008. A total of 3 – 6 specimens
of sub-adults from each location (standard length range
46.4 – 69.5 mm) were used for osteological preparations.
Fish were collected from multi-meshed gill nets and gape
nets, or collected from catches of fishermen. Fish were
identified using taxonomic descriptions given in Munro
(1955).
Osteological analysis: Osteological preparation of the
whole fish specimens (Figure 1B) (initially stored in
5% formalin and subsequently fixed in 70% alcohol)
followed the method described by Taylor & Vandyke
(1985) using Alizarin Red-S. Line drawings of the
separated bones of 3-6 specimens from each location
were made using a Camera Lucida fixed to a Wild M5A
stereomicroscope. The nomenclature of the caudal and
cranial elements follows published osteological work
(Weisel, 1960; Arratia, 1983; Sanger & McCune, 2002).
In addition, the distance between the distal end of the
rudimentary neural arch of compound centrum and
the proximal end of the epural is referred to as epural
distance.
RESULTS
Cranial portion of P. sarana skull-roof was more or less
asymmetrical in the dorsal view (Figure 3). Dorso-median
cranial fontanel is absent in all studied specimens, and the
adjacent bones are abutted firmly. The bony elements of
the neurocranium from the dorsal view are the frontals,
parietals, pterotic, and supraoccipital. Frontals, the largest
of the skull bones covering a considerable part of the
skull on the dorsal side, are flat and elongated anteriorly
while broad and thick posteriorly. The frontal bones are
single ossified plates, which overlap medially and their
dorsal view in some skulls reveal an underlying ridge,
which gives a virtual appearance of each frontal being
divided into two parts. The paired parietals are small and
relatively thin bones situated in between frontals and
supraoccipital in the hindmost region of the skull. They
are attached to the supraorbital margin of the frontal bone
antero-laterally, and to the pterotic laterally. The pterotics
occupy the postero-lateral corners of the skull and are
conspicuously irregular in shape.
Figure 2: Collection localities (triangles) of P. sarana
(Codes for rivers; 1: Kalu River, 2: Gin River, 3: Nilwala
River, 4: Walawe River, 5: Kirindi Oya, 6: Menik River)
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Although the composition of cranial bones did not
show any variation, the shapes and the arrangement
Journal of the National Science Foundation of Sri Lanka 39 (2)
123
Osteology of the olive barb
of cranial bones, particularly the direction/ path of
suture between left and right halves of the cranium,
showed variation among studied specimens of P. sarana
(Figure 4). The specimens of the rivers Menik (M), Gin
(G), and Kalu (KR) had oppositely directed curvature at
the suture between left and right frontals compared to the
specimens from the rivers Walawe (W), Nilwala (N), and
the Kirindi Oya (K).
Figure 3: The cranial skeleton of P. sarana, 53.8 mm SL.
(FR- frontal bones; PR- parietal bone; PTO- pterotic bone;
scale bar = 1 mm).
Figure 4: Variation in the cranial skeleton of P. sarana from six
locations (the arrows indicate the curvature direction along
mid cranial region for each row; scale bar = 1mm).
Pharyngeal bones of P. sarana are relatively broad
and stout (Figure 5). The tip of the dorsal limb (dorsal
lip) is usually pointed, and the upper angle along the
outer edge is unrecognizable. Congregated teeth on the
pharyngeal bones are moderate in size and arranged
in three rows. The common dental formula on the two
bones is 2, 3, 5 - 5, 3, 2, where the outermost or the first
row has two slender teeth. The thickness and the size of
the teeth increase inwards, thus the last or the inner main
row has five conspicuously larger broad teeth. However,
two populations (the Walawe and the Menik rivers) had
slightly different shapes of pharyngeal bones, where its
dorsal lip is truncated and can be clearly distinguished
from those in other specimens (Figure 5). Dental formula
was not different among the populations.
The vertebral column of P. sarana generally consists
of 36 vertebrae, but varied from 34 to 36, where the
specimens from the rivers Gin, Kalu and Menik had 36
vertebrae. Specimens from the Nilwala River and the
Walawe River mostly had 34 vertebrae, while those from
the Kirindi Oya commonly had 35. The last three caudal
vertebrae form the caudal skeleton (Figure 6) supporting
the caudal fin with 18 principal rays (asymmetrical with
10 and 8 in upper and lower lobes, respectively). The last
caudal vertebra is commonly interpreted as a compound
Figure 5: (A) Fifth ceretobranchial of P. sarana, 68.3 mm SL.
(DLB- dorsal limb; DLP- dorsal lip; ExA- external angle; VLB- ventral limb; VLP- ventral lip;
I- First tooth row; II- Second tooth row; III- Main row; 5- Fifth tooth in the main row).
(B). Variation in the shape of the dorsal lip of pharyngeal bones of P. sarana: (i) truncated, in
Walawe River and Menik River; (ii) pointed, in other five locations (scale bar = 1mm).
Journal of the National Science Foundation of Sri Lanka 39 (2)
June 2011
F. I. Irfan & K. B. Suneetha Gunawickrama
124
Figure 6: Caudal skeleton and preural region of P. sarana, 68.3 mm
SL.
(EP-epural; HF-hypural foramen; HS-haemal spines;
H (1-6)- six hypurals; HYP-hypurapophysis; NA-neural
arch; NS-neural spines; PF-parhypural foramen;
PH-parhypural; PU-preural centrum; CC-compound
centrum; RNA-rudimentary neural arch; PL-pleurostyle
(scale bar = 1mm).
Figure 7: Caudal skeleton of P. sarana from six locations showing
the variation in various elements (W – Walawe River; G –
Gin River; N – Nilwala river; KR – Kalu River; M – Menik
River; K – Kirindi Oya) (scale bar = 1mm).
centrum formed by an assumed fusion of the first preural
centrum with one or more ural centra (PU1 + U1), and
three other elements fused with it, i.e. parhypural in the
lower lobe, and pleurostyle and rudimentary neural arch
in the upper lobe. The parhypural bears a hypurapophysis
at the base, which is a tiny and bony process. Other
caudal skeleton elements include six hypurals (four and
two in the upper and lower lobes respectively), and a
June 2011
detached tiny bone called epural located antero-dorsal to
the pleurostyle. In some specimens, hypurals are abutting
to one another but not fused. In addition to the hypural
foramen (HF) between proximal regions of the first
and second hypurals, the proximal regions between the
parhypural and the first hypural creates the parhypural
foramen (PF). The size and the shape of PF seem to vary
with the presence or absence of an invagination along the
lateral margin of the proximal end of parhypural. One
uroneural remains as a thin bone slip along the side of
the pleurostyle, and the distal end of it can be seen in
all specimens (not shown in Figure 6). The rudimentary
neural arch on the dorsal facet of the compound centrum
is commonly narrow, elongated and curved dorsoanteriorly near its tip. The distance between the distal
end of the rudimentary neural arch and the proximal end
of the epural (epural distance) seems to vary. Each of the
two anterior vertebrae of the caudal element, i.e. preural
centrum 2 and 3, give rise to a dorsal neural spine and a
ventral haemal spine.
Osteological preparations from six locations
revealed consistent variation in various components of
the caudal element as described herein (Figure 7). The
shape, length and the stoutness of the rudimentary neural
arch in fish from the Walawe River (W) apparently differ
from the common pattern, where it is distinctively shorter
and stouter than those in other rivers. The size of the
parhypural foramen is also a distinctive feature where the
specimens from the Walawe River (W) and the Gin River
(G) showed relatively larger parhypural foramen than in
other specimens. No intra-specific variation was detected
in the shape or the position of the hypurapophysis of
P. sarana. The epural distance considerably varies among
locations where the largest gap is found in fish from the
Walawe River, while it is smaller and varies slightly
in fish from other locations. In contrast to the common
pattern, double neural spines on preural centrum 2 (PU2)
is seen in five specimens from the Walawe River (W)
and in three (out of 6) specimens from the Kirindi Oya.
However, the length of the doubled neural spines on PU2
showed variation between these two locations, where the
spines are of similar length in fish from the Walawe River
but contrastingly of different length (one is very short) in
fish from the Kirindi Oya. Doubling of the neural spine
of the preural centrum 3 is seen in four specimens (out of
6) from the Nilwala River (N).
DISCUSSION
The present study revealed certain differences in
osteology among the six geographic populations of
P. sarana in Sri Lanka. There is no variation in the bones
forming the skull roof, but the bone arrangement seems to
Journal of the National Science Foundation of Sri Lanka 39 (2)
Osteology of the olive barb
be a character for population differentiation. The median
inter-frontal suture showed discernible variation, which
categorized the six studied populations into one of two
broad patterns, having leftward or rightward curvature.
However, the examination of more specimens is required
to assess whether that feature can be considered as a
population specific phenotypic trait.
The pharyngeal bones of adult cyprinoids represent
the fifth (V) ceratobranchials (Eastman, 1971). The
shape of the pharyngeal bones has a functional role in
muscle attachment and thus facilitating mastication of
food (Eastman, 1971). Pharyngeal bones and teeth are
important in identification of some fish species (Eastman,
1977) when supplemented with other taxonomically
useful phenotypic characters. Eastman (1977) reviewed
the use of bones and teeth of the pharyngeal apparatus of
22 species of catostomid fishes (family Catostomidae),
and showed that the differences in pharyngeal bones
may exist even if the species share similarities in diets.
The shape of the dorsal tip is a diagnostically variable
character in many cyprinoids focused in Eastman
(1971). Differences in the shape of the pharyngeal bone
of some populations of P. sarana may hence represent
a taxonomically informative character. According to
Shantakumar & Vishwanath (2006), the dorsal limb in
Indian P. sarana has a blunt tip. Apparently a similar
dorsal limb shape was found in fish from all Sri Lankan
locations, except the Walawe River and the Menik River.
This raises a question as to whether the latter populations
in Sri Lanka are unique, but requires more taxonomical
information from both Indian and other Sri Lankan
populations. Similar to that in Indian P. sarana
(Shantakumar and Vishwanath, 2006), and Cyprinus
carpio (Eastman, 1971), three rows of pharyngeal teeth
occur on each ceratobranchial of P. sarana studied.
The number of teeth on the pharyngeal bone may be
variable as they may be replaced through the life of
individuals, thus it is likely to show considerable interspecific variation (Eastman, 1977) apparently connected
to trophic adaptations. However, the present study did
not reveal any variation in the arrangement of teeth of
P. sarana specimens in Sri Lanka.
The caudal skeleton is an important system for
interpreting systematic and phylogenetic relationships of
actinopterygian fishes (Gosline, 1961; Nybelin, 1972).
It may also provide evidence for intra-specific variation
in fishes (Eastman, 1980). Caudal fins of cyprinids
in general, and of P. sarana fishes typically have six
hypurals including four in the upper lobe (Buhan, 1972;
Shantakumar & Vishwanath, 2006). P. sarana specimens
examined in the present study followed the same pattern.
All specimens of P. sarana bear a parhypural foramen
Journal of the National Science Foundation of Sri Lanka 39 (2)
125
that may have a functional significance linked to caudal
fin movements, similar to the function described for the
hypural foramen (the space between the first and second
hypurals) in fishes (Kampmeier, 1969). These inter-bone
spaces are known to be involved in facilitating the lymph
communication (Kampmeier, 1969). Variation in the size
of the parhypural foramen, which is not related to fish size
was seen where the specimens from the rivers Walawe
and Gin bear relatively large parhypural foramen. The
presence and structure of the hypurapophysis seem to have
some functional significance linked to fast swimming
capability as it has been linked to the caudal musculature
(Eastman, 1980), particularly as an attachment site for
the hypochordal longitudinal muscle (Nursall, 1963).
The absence of variation in hypurapophysis of P. sarana
may indicate its significance in all fish, as this may be
a required characteristic of any fast swimmer. However,
the caudal musculature of cyprinids has not been studied
well (Eastman, 1980), and therefore, making any
peculiar functional interpretations is difficult (Gosline,
1961). Nevertheless, according to Nursall (1963), there
is no apparent change in the degree of development of
the muscle with or without hypurapophysis. Uroneurals
are modified ural neural arches (Arratia & Schultze,
1992; Sanger & McCune, 2002), and the presence of
one thin uroneural along the side of the pleurostyle was
a homogenous feature in all studied specimens with no
intra-specific variation. First of the paired uroneurals
is usually not seen as it has fused to the compound
centrum, but the second one usually remains as seen in
Danio species as a thin slip of bone along the side of the
pleurostyle (Sanger & McCune, 2002). They probably
provide functions in protecting the terminal part of the
spinal cord, as well as in ensuring that the tail functions
hormocercally (Patterson, 1968).
The axial skeleton of P. sarana commonly consists
of 36 vertebrae (Shanthakumar & Vishwanath, 2006),
and the variation in vertebral number seems to be linked
to the doubling of the neural spines in the preural caudal
vertebrae. For instance, specimens from the rivers
Nilwala, Walawe and Kirindi, which had less than 36
vertebrae, all showed doubling of the neural spines on
preural centra. It is generally believed that the doubling
process is a result of vertebral fusion or more correctly,
non-separation during development (Kändler, 1932).
The number of vertebrae in fishes is fixed early in
development, usually by the time of hatching (Gwyne,
1940; Garside, 1966), and believed to be influenced by
physical, chemical and biological conditions of the water
where eggs and larvae are laid and develop (Arratia,
1992). Vertebral fusion is known to be a common feature
in other cyprinids as well (Eastman, 1980). The observed
results are in agreement with the above interpretation,
June 2011
F. I. Irfan & K. B. Suneetha Gunawickrama
126
and indicate that the vertebral number seems to be a
powerful adjunct in studies of intra-specific variation
in P. sarana fish. Similar complex vertebrae have been
encountered in the preural region of variety of fishes
(Kändler, 1932; Barrington, 1937; Ford, 1937; Buhan,
1972; Patterson and Rosen, 1977). The preural vertebrae
and caudal skeleton comprise a significant component
of the morphological evidences presented in papers on
fish inter-relationships (Patterson, 1968; Greenwood &
Rosen, 1971; Patterson & Rosen, 1977; Arratia, 1997,
1999), thus the knowledge of intra-specific variation in
the preural vertebrae is greatly important in fish anatomy
and taxonomy.
The environment plays an important role in
establishing phenotypes (Arratia & Schultze, 1992). It
has been demonstrated that the degree of constancy is
often affected and characters may even vary considerably
when a species comes under the influence of different
environmental conditions (Tatarko, 1968). Most of the
ossified structures are subjected to broad environmentally
induced variation (Taning, 1952; Tatarko, 1968; Fowler,
1970) and temperature in particular is known to affect
the developmental rate of ossified meristic characters,
coupled to slowing or accelerating growth and
differentiation (Eastman, 1980). However, in this study,
temperature variations between locations are insignificant
and therefore temperature would not be a strong factor
that creates differences in osteological features in
studied specimens. Morphological (and /or osteological)
structures and their development are strongly correlated
with functional requirements (Weisel, 1960; Strauss,
1990; Mabee, 2000; Maglia et al., 2001). For instance,
in Barbus barbus, bony elements of the opercular region
and the cranial bones involved in feeding ossify earlier
than other bones (Vandewalle et al., 1992), hence, it is
assumed that functional needs lead to the developmental
sequences where bones implicated in early functional
needs probably respond by earlier ossification.
The taxonomic significance of the variation
observed in the present study has to be assessed in
relation to the available taxonomic information on the
species. The species described by Hamilton in 1822
from the Ganges was named as Cyprinus sarana, and
later it was synonymized with Puntius sarana. However,
its first description from Sri Lanka appeared as Puntius
pinnauratus Day 1878 (Day 1865). Deraniyagala (1930)
synonymized it with P. chrysopoma, while a related
species with superficial appearance to it was later named
P. timbiri (Deraniyagala, 1963). The identification
of this species in Sri Lanka appears to have certain
ambiguities, because of these synonyms and the lack of
information on whether there is only a single species of
June 2011
P. sarana–like fish (Pethiyagoda, 1991). The taxonomic
complexity of this species has further been indicated
by Talwar & Jhingran (1991) reporting intra-specific
variants that have been designated as four sub species,
namely P. sarana orphoides, P. sarana subnasutus and
P. sarana sarana from India, and P. sarana spilurus
from Sri Lanka (Ceylon; type locality unknown). In
the absence of systematic studies, there is an apparent
taxonomic uncertainty of the Sri Lankan P. sarana, while
all fish having gross external similarities to Deraniyagala
(1952) description are currently named Puntius sarana.
Until an extensive taxonomic study is undertaken on the
species by examining all the relevant type specimens,
Pethiyagoda (1991) has suggested retaining the name
Puntius sarana for all related varieties that have close
superficial resemblance.
According to Shantakumar & Vishwanath (2006),
there are specific osteological characters, which belong
to the P. sarana group in Manipur, India. However, the
same study has not revealed which sub-species they have
focused on. Therefore, a useful comparison between the
Sri Lankan specimens and those from other subspecies
originally described from India using many osteological
features was not possible. Such comparisons may
contribute to the taxonomic clarification of P. sarana
fishes in the region. Observed osteological variations
provide good evidences for intra-specific heterogeneity
among P. sarana populations, where the Walawe River
population of P. sarana can be differentiated from other
studied populations based on the shape of the rudimentary
neural arch, the size of the parhypural foramen, the
epural distance and the shape of the dorsal lip of fifth
ceretobranchials. Accordingly, the results of the present
study reveal a convincing phenotypic heterogeneity of
the species among populations in rivers in Sri Lanka.
Examination of more specimens representing other
geographic samples from its distribution range will
contribute to a wider taxonomic perception of this
species.
Acknowledgement
Assistance in fieldwork by D. A. M. Dilrukshana, C.
H. Priyantha, and P. H. Dinesh Dilhan, funding by
National Research Council (grant NRC 05-60) and
help from Dr Rohan Pethiyagoda and the research crew
previously at WHT biodiversity center, Agarapathana
are acknowledged. The comments of two anonymous
referees on the initial manuscript are appreciated.
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