Phylogenetic position and osteology of Pethiasetnai (Chhapgar & Sane, 1992), an endemic
barb (Teleostei: Cyprinidae)
of the Western Ghats, India, with notes on its distribution and threats
Unmesh Katwate 1, MandarS. Paingankar 2, Shrikant Jadhav 3 & Neelesh Dahanukar 4
1 Bombay Natural History Society (BNHS),
Hornbill House, Opposite Lion Gate, Shaheed Bhagat Singh Road, Mumbai, Maharashtra 400001, India
2,4 Systematics, Ecology & Conservation
Laboratory, Zoo Outreach Organization (ZOO), 96 KumudhamNagar, Villankurichi Road, Coimbatore, Tamil Nadu
641035, India
3 ZoologicalSurvey of India (ZSI), Western Regional Center, Akurdi, Pune, Maharashtra
411044, India
4Indian institute of Science Education and
Research (IISER), Dr. Homi Bhabha Road, Pashan, Pune,
Maharashtra 411008, India
1 theunmesh@gmail.com, 2 mandarpaingankar@gmail.com,3 shrikantj123@yahoo.com, 4 n.dahanukar@iiserpune.ac.in
(corresponding author)
Abstract: Pethia setnai is
an endemic and threatened freshwater fish of the Western Ghats of India. It has a restricted distribution in the
west flowing rivers in the states of Maharashtra, Goa and Karnataka. We clarify the phylogenetic position of Pethia setnai, provide osteological details of topotypic material, and morphometric data of specimens from
Maharashtra, Goa and Karnataka. We
also provide details on micro-level distribution, habitat and threats to the
species in its native range.
Keywords: Biometrics, Puntius setnai, threatened, topotype.
doi: http://dx.doi.org/10.11609/JoTT.o3857.5214-27 | ZooBank:urn:lsid:zoobank.org:pub:4F75DBB8-C599-402B-A9D1-C63D5CDD9DE7
Editor: Anonymity requested. Date
of publication: 26 December 2013 (online & print)
Manuscript details: Ms # o3857 | Received 21
November 2013 | Final received 17 December 2013 | Finally accepted 18 December
2013
Citation: Katwate, U., M.S. Paingankar, S. Jadhav& N. Dahanukar(2013). Phylogenetic position and osteology of Pethia setnai (Chhapgar & Sane, 1992), an endemic barb (Teleostei: Cyprinidae) of the
Western Ghats, India, with notes on its distribution and threats. Journal of Threatened
Taxa 5(17): 5214–5227; http://dx.doi.org/10.11609/JoTT.o3857.5214-27
Copyright: © Katwateet al. 2013. Creative Commons Attribution 3.0 UnportedLicense. JoTT allows unrestricted use of this
article in any medium, reproduction and distribution by providing adequate
credit to the authors and the source of publication.
Funding: Neelesh Dahanukar is a
DST-INSPIRE Faculty Fellow. This study was partially supported by DST-INSPIRE
Research Grant [IFA12-LSBM-21], Government of India to Neelesh Dahanukar and CEPF-ATREE Western Ghats Small Grant
(CEPF-ATREE-WGhats/SGP/WGSG186 – BNHS_FISHES)
to Unmesh Katwate.
Competing interests: Declared none. Funding source had no role in study design, data
collection, results interpretation and manuscript writing.
Author Contribution: UK, MP and ND collected fresh specimens. UK, SJ and ND studied museum
specimens. UK, MP, SJ and ND performed morphometry.
UK performed osteology. MP and ND performed genetic analysis. UK, MP, SJ and ND
wrote the manuscript.
Author Details: Unmesh Katwate is a Zoology (Oceanography) postgraduate currently working as
Scientist-A in BNHS. He works on conservation, taxonomy and ecological aspects
of freshwater fishes and amphibians of the Western Ghats. Mandar S. Paingankar is a molecular
biologist and works on vector biology with an emphasis on host parasite
interactions. He works on molecular phylogeny and systematics of freshwater
fishes as a hobby. Shrikant Jadhav is a
scientist in Zoological Survey of India. He works on freshwater fish diversity
and taxonomy. Neelesh Dahanukar works in ecology and
evolution with an emphasis on mathematical and statistical analysis. He is also
interested in taxonomy, distribution patterns and molecular phylogeny of
freshwater fish and amphibians.
Acknowledgements: We
are grateful to Abhijeet Gharat,
Pratik, Keerthi Krutha, Shreya Mulherkar, Saahil Acharya and Nikhil Modak for helping with the collections. We thank Chetana Katwate for help in
osteology and morphometric study. We thank Dr. Siby Philip for help with the phylogenetic analysis and Dr. B.F. Chhapgar for helpful
discussions. We are grateful to Dr. Asad Rahmani, Director; Dr. Deepak Apte, COO; Rahul Khot, incharge Natural History
Collection department and Vithoba Hegde,
senior field assistant, for their help during study of the museum specimens and
registration of specimens in Bombay Natural History Society (BNHS), Mumbai. We
are also grateful to Dr. Sanjay Molur,
Founder Secretary; and Priyanka Iyer,
curator of fish collection, for holding our specimen vouchers in the museum
collection of Wildlife Information Liaison Development (WILD) Society,
Coimbatore. We thank the Director and Officer In-Charge (Collections),
Zoological Survey of India (ZSI), Kolkata for providing us the access to the
type material of the species. We are also thankful to Dr. Pankaj Bhatnagar,
Officer-in-charge, Zoological Survey of India, Western Regional Center, Pune, for encouragement and support.
This article forms part of a special
series on the Western Ghats of India, disseminating the results of work
supported by the Critical Ecosystem Partnership Fund (CEPF), a joint initiative
of l’Agence Française de Développement, Conservation International, the European
Commission, the Global Environment Facility, the Government of Japan, the
MacArthur Foundation and the World Bank. A fundamental goal of CEPF is to
ensure civil society is engaged in biodiversity conservation. Implementation of
the CEPF investment program in the Western Ghats is led and coordinated by the Ashoka Trust for Research in Ecology and the Environment
(ATREE).
For
figures, images, tables -- click here
Introduction
Two major hurdles in conservation of threatened and endemic species,
especially those that occur in biodiversity hotspots is the Wallaceanshortfall (arising from the incomplete information regarding the distribution
of species) and Linnean shortfall (several species
still not formally described) (Bini et al.
2006). The Western Ghats region,
part of the Western Ghats-Sri Lanka biodiversity hotspot is no exception to
this, as it has already been suggested that the freshwater fish fauna
(especially endemic and threatened species) of this region is poorly known with
regard to their taxonomy and distribution (Dahanukaret al. 2011; Raghavan et al. 2012). One reason for this gap in knowledge
regarding geographical distribution is the lack of detailed descriptions and
reliable genetic data, which can ultimately lead to misidentifications. As a result, genetic sequences from topotypic material and detailed morphometric data will not
only help in understanding the systematics of the species, but also provide
reliable identification criteria.
In a recent review of the South Asian barbs previously placed under the
genus Puntius, Pethiyagodaet al. (2012) tentatively placed Puntius setnai Chhapgar & Sane,
1992 in the genus Pethia based on the data
available in the original description of the species. The tentative placement was probably due
to the limited information on the morphology, osteology and/or genetic data of
the species. In the current study, we provide genetic information of Pethia setnai from topotypic material, osteologicaldetails to confirm the placement of the species under Pethia,
and reliable genetic and morphometric data for accurate identification of the
species. We also provide information on the distribution, habitat and threats
to the species.
Materials and Methods
Study site
Topotypes of Pethia setnai were
collected from Salaulim River, a tributary of Zuari River in Sanguem(15.234⁰N & 74.182⁰E, 19m), Goa. Additional material was also collected
from Terekhol River at MadkholVillage (15.935⁰N & 73.910⁰E, 43m), Maharashtra (Fig. 1). Collections were done responsibly
following the guidelines set by IUCN (2008). Material collected in the present study
is deposited in the museum of Bombay Natural History Society (BNHS), Mumbai; Wildlife Information Liaison Development (WILD),
Coimbatore, and the Zoological Survey of India, Western Regional Center (ZSI-WRC), Pune.
Morphometry
Measurements were taken point to point using
dial calipers to the nearest 0.01mm and then rounded
to 0.1mm. Subunits of the body are
presented as percent of standard length (SL), and
subunits of the head are presented as percent of head
length (HL). All pored scales were
counted when reporting the lateral line scales. Methods for taking counts and
measurements follow Kullander (2008) and Pethiyagoda et al. (2012).
Materials examined
Pethia setnai: 9 exs., BNHS FWF 53, 63 to 70,
10.viii.2013, collected from Sanguem, Goa, by U. Katwate, M. Paingankar and N. Dahanukar; 3 exs., WILD-13-PIS-043
to 045, 10.viii.2013, collected from Sanguem, Goa, by
U. Katwate, M. Paingankarand N. Dahanukar; 2 exs.,
ZSI-WRC P/3567, 10.viii.2013, collected from Sanguem,
Goa, by U. Katwate, M. Paingankarand N. Dahanukar; 9 exs.,
BNHS FWF 54 to 62, 12.vi.2013, collected from TerekholRiver at Madkhol, Maharashtra, by U. Katwate and N. Dahanukar; 3 exs., WILD-13-PIS-046 to 48, 12.vi.2013, collected from Terekhol River at Madkhol,
Maharashtra, by Unmesh Katwateand Neelesh Dahanukar; 2 exs., ZSI-WRC P/3568, 12.vi.2013, collected from Terekhol River at Madkhol,
Maharashtra, by U. Katwate and N. Dahanukar;
1 ex., ZSI-WRC P/3572, 5.ii.1971,
collected from Kaneri River at Gund,
District Karwar (Mysore), by B.S. Lambaand Party; 3 exs., ZSI-WRC P/3571, 17.ii.1971, collected
from Ramanguli, District Karwar(Mysore), by B. S. Lamba and Party; 26 exs. BNHS FWF 17-42, collected from Sanguem,
Goa by S.R. Sane.
Photographs examined
Pethia setnai: Holotype, ZSI Kolkata FF2766, 01.iii.1985, collected from Sanguem, Goa, by S.R. Sane; Paratypes,
6 exs., ZSI Kolkata FF2767,
01.iii.1985, collected from Sanguem, Goa, by S.R.
Sane.
Pethia narayani: Syntypes, 2 exs.,ZSI Kolkata F12180/1, collected from Cauvery River, Coorg,
by C.R.N. Rao.
Osteology
Two specimens (BNHS FWF 55 and BNHS FWF 70) were cleared and stained
using the procedure described by Potthoff(1984). Osteological nomenclature follow Conway (2011) and descriptions of
osteology follow Pethiyagoda et al. (2012) so as to
allow easy comparison with other related taxa. Osteologicalillustrations were made from images captured by a digital camera fitted on
stereo-zoom light microscope (Leica S8 APO, USA).
Genetic analysis
Gills were taken from four fresh specimens (BNHS FWF 53, BNHS FWF 54,
WILD-13-PIS-043 and WILD-13-PIS-046) and preserved in absolute ethanol. The
tissue was digested at 600C for two hours using the STE buffer (0.1MNaCl, 0.05M Tris-HCl, 0.01M
EDTA, 1%SDS) with 15µl Proteinase K (20mg/ml) per 500ml of STE buffer. DNA was extracted using conventional
phenol-chloroform method and re-suspended in TE (10mM Tris,
1mM EDTA, pH 8) buffer. Polymerase
chain reaction was performed to amplify cytochrome b (cytb)
gene using primer pair L14724 (5’-GACTTGAAAAACCACCGTTG-3’) and H15915
(5’-CTCCGATCTCCGGATTACAAGAC-3’) (Chen et al. 2007). PCR reaction was performed in a 25µl
reaction volume containing 5µl of template DNA (~200ng), 5µl of 10X reaction buffer
(100 mM Tris pH 9.0, 500 mM KCl, 15 mMMgCl2, 0.1% Gelatin), 3µl of 25mM MgCl2,
1µl of 10mM dNTPs, 1µl of each primer, 0.5µl Taq polymerase (2.5 units) and 8.5µl nuclease free water.
The thermal profile was 10 minutes at 950C, and 35 cycles of 1
minute at 940C, 1 minute at 520C and 2 min at 720C,
followed by extension of 10 min at 720C. Amplified DNA fragments were purified
using the Wizard Gel and PCR clean up system (Promega,
USA). The purified PCR products
were sequenced using BigDye Terminator v3.1 cycle
sequencing kit (Applied Biosystems, USA) and ABI
prism 3730 sequencer (Applied Biosystems,
USA). Sequences were analyzed by BLAST tool (Altschul et al. 1990). All sequences used in the paper have been deposited in GenBank.
We used the cytb gene dataset of South Asian
cyprinid fishes by Pethiyagoda et al. (2012), Raghavan et al. (2013a) and additional sequences of Pethia, Dawkinsiaand Haludaria available from NCBI GenBank (http://www.ncbi.nlm.nih.gov/). Accession numbers for the sequences used
for the analysis are provided in Fig. 1. Gene sequences were aligned using
MUSCLE (Edgar 2004). Phylogeny was
performed using the freeware MEGA 5 (Tamura et al. 2011). Best
fit model for nucleotide substitution was selected from 24 models
available in MEGA 5 based on minimum AkaikeInformation Criterion (AIC) value (Posada & Crandall 2001). Tamura & Nei(1993) nucleotide substitution model including invariant sites and a Gamma
parameter was obtained as a best fit model (TN93+G+I,
AIC = 12894.38, lnL = -6328.70). This model was then used for constructing
a phylogenetic tree using the maximum likelihood method. Reliability of the phylogenetic tree was
estimated using bootstrap values run for 1000 iterations.
Results and discussion
Before it was formally described, Pethia setnai collected from Ponda,
Goa, was first reported as Pethia nigrofasciata (Günther, 1868)
by Yazdani(1977). Chhapgar& Sane (1980) criticized this report, after collecting this species from Sanguem, Goa, as well as examining the specimens studied byYazdani (1977), and suggested that the species was Pethia narayani (Hora, 1937). However, subsequently, Chhapgar & Sane
(1992) described the species as Puntius setnai. Subsequently, Yazdani & Ghate (1994) provided further information on this species
based on the specimens collected from Hosangadi in
Karnataka.
Phylogenetic position
Pethia setnaiforms a monophyletic group (Fig. 1) with the genus Pethia sensu stricto as defined byPethiyagoda et al. (2012). The genetic divergence in
the cytb gene between the topotypicmaterial collected from Sanguem, Goa and additional
material collected from Terekhol River at Madkhol, Maharashtra, was 0.325%, while the divergence
between the two Terekhol specimens as well as two Sanguem specimens was 0.2%. Because of the very low genetic divergence,
the two populations have to be considered as genetically similar. We could not
collect fresh specimens of the species for genetic analysis from Karnataka
State, and as a result, future studies are essential to establish the extent of
genetic variation in different known populations of the species. Nevertheless, as we are providing the
genetic information of the topotypic material, future
studies can have a more reliable comparative material.
Taxonomy
Pethia setnai (Chhapgar & Sane, 1992)
Puntius nigrofasciata (nonGünther, 1868): Yazdani(1977)
Puntius narayani (non Hora, 1937): Chhapgar& Sane (1980)
Puntius setnai Chhapgar & Sane, 1992: Chhapgar & Sane (1992), Yazdani & Ghate (1994), Knight et al. (2012)
Pethia setnai (Chhapgar & Sane, 1992): Pethiyagoda et al. (2012)
Type material information
Holotype and
six paratypes of Pethia setnai collected from Sanguem,
Goa, by S.R. Sane on 1.iii.1985 are available in good condition in the museum
collection of ZSI Kolkata under the accession numbers FF2766 (Image 1) and
FF2767 (Image 2). Chhapgar & Sane (1992) described the species based on
25 specimens (the holotype and 24 paratypes)
with the note that “…holotype and some of the paratypes will be deposited with the Zoological Survey of
India, Calcutta”. The whereabouts
of remaining 18 paratypes is not known.
Interestingly, 26 specimens of P. setnai are
in the museum collection of BHNS under the accession numbers BNHS FWF 17-42
(see Materials examined). However,
whether some of the specimens from BNHS constitute the missing paratypes could not be deciphered.
Morphometric and meristic data
Morphometric characters and meristics of the
specimens collected for the present study from the type locality Sanguem, Goa; a newly discovered population from Terekhol River near MadkholVillage, Maharashtra and from Karwar, Karnataka are
provided in Table 1 and Table 2, respectively. Photographs specimens collected for the
present study are provided in Images 3 and 4.
Osteology
Osteology of the specimen from the type locality is provided in Image
5. Gill rakerssimple, acuminate (not branched or laminate); post-epiphysial fontanelle absent (Image 5b); infraorbital3 deep, partially overlapping the cheek and preoperculum(Image 5c); last unbranched dorsal-fin ray stiff,
serrated (Image 5d); free uroneural absent (Image
5e); 4 supraneurals; 13 precaudaland 13 caudal vertebrae. Osteological details of P.setnai matches with the diagnosis of the genus Pethia provided by Pethiyagoda et al. (2012).
Color variation
In life Pethia setnaihas silvery-grey or dull golden color with three
black transverse bands; anterior one situated between the dorsal profile and
below the lateral line encompassing 3rd and 4th lateral line scales; middle
band between the posterior half of the dorsal fin base and one scale below
lateral line encompassing 9th to 11th lateral line scale; posterior band
between the middle of end of dorsal fin base and caudal fin and the posterior
half of the anal fin base (Image 4). Posterior most and anterior most bands are darker than the middle band
(Image 4a). Dorsal, ventral and
anal fins are bright red in certain specimens in their natural habitat (Image
4a) but the fin colors fade rapidly in captivity
(Image 4b). Some specimens have
indigo blue tinge on the dorsolateral area and therefore the species is also
known as indigo barb among hobbyists. While, specimens with yellow fins are
also known from this species we have not come across such specimens in our
study.
Distribution
Based on the current collections as well as locality information
deciphered from literature (Table 3), Pethia setnai is known to be
distributed in the west flowing rivers of the Western Ghats in the
states of Karnataka, Goa and Maharashtra between 12.66⁰N to 15.94⁰N
latitude (Fig. 2). The species is currently known from seven fragmented populations, most of them reported from lower altitudes
(Table 3) except the one from Agumbe, Karnataka
located at 669m (Knight et al. 2012).
To our knowledge, there are no published and reliable records of Pethia setnai from
the Cauvery River system, or any other east flowing rivers. Anecdotal reference to the presence of
this species in the Cauvery, especially among the aquarium fish hobbyists, is
as a result of the misidentification of Pethia setnai as Pethia narayani by Chhapgar & Sane(1980). While criticizing the report of Pethia nigrofasciata from Goa by Yazdani (1977), Chhapgar & Sane (1980) suggested that the species was Pethia narayani (Image
6), a species originally described from Cauvery River system in Coorg by Hora (1937). Mr. S.R. Sane
who was active in the aquarium fish trade marketed the species occurring in Goa
as Narayan’s Barb until Chhapgar & Sane (1992)
realized that they had overlooked the presence of osseous and serrated last unbranched dorsal fin ray in the specimens from Goa (as
opposed to non-osseous and non-serrated last unbrancheddorsal fin ray in Pethia narayani), and described Pethia setnai (B.F. Chhapgar,
pers. comm. July 2013). Apart from
having strongly osseous and serrated (vs. smooth articulated) last unbranched dorsal fin ray, P. setnai differs from P. narayani based on 8 (vs.
9) branched dorsal fin rays, 5 (vs. 6) branched anal fin rays and 19–20
(vs. 22) lateral line scales. Because of the misleading common name ‘Narayan’s Barb’ for P. setnai, it is likely that hobbyists have treated P. narayani from Cauvery River as P. setnai.
We failed to come across P. setnaiduring our repeated surveys in Mysore, Coorg and
several other areas from Cauvery river system. As a result, unless a reliable report ofP. setnai is available from Cauvery River
system, backed up with genetic data, the species is considered as restricted to
west flowing rivers of the Western Ghats based on Yadav(2003) and information presented in this paper.
Habitat
Chhapgar& Sane (1992) described Pethia setnai from clear streams of Sanguemand Ponda in Goa. However, they did not provide any
information on the habitat in which the species was found. As part of the
present study, fresh collection of P. setnai was
made from the Salaulim River, a tributary of Zuari River near Sanguem, Goa
(Image 7). The habitat consisted of
a slow flowing, clear water stream with maximum substratum composed of sand and mud. Riparian cover was rich in vegetation, and mostly dominated by Pundanus plant species. Pethia setnai was mostly found in small shoals
with 7–8 individuals in each shoal. During each collection attempt, maximum catch was
dominated by females with one to two males in each shoal. Juveniles were
observed to be confined at shallow depth (0.1–0.5 m), whereas
adults were confined to greater depths (1–2 m). We also discovered a new population of P.setnai in the main river channel of Terekhol River at Madkhol Village
(Image 8). In contrast with type
locality, at Terekhol, the individuals of P. setnai were found in deep pools and fast flowing rapids
(Image 8). Co-occurring species
included Dawkinsia cf. filamentosa,Haludaria pradhani,Devario aequipinnatus,Rasbora sp. and Salmophasia sp.
Threats
Based on IUCN Categories and Criteria (IUCN 2001), Dahanukar(2011) assessed Pethia setnai as a ‘Vulnerable’ species owing to its restricted distribution and on-going
threats to habitat because of tourism, urbanization and agricultural
pollution. The Goa populations are
subjected to heavy mining operations in the headwaters, apart from the pressure
from increasing tourism (Image 9). Habitats of the species in Terekhol River are
severely threatened by organic wastes and sewage. Further, the species is also known to
occur in aquarium trade under the common name Indigo Barb (Chhapgar& Sane 1992). Although no
statistics is available on the extent of wild caught Pethia setnai in the trade, it has already been
suggested that the unregulated aquarium trade is a plausible threat to the
endemic and threatened species of freshwater fishes in India (Raghavan et al. 2013b). Further studies on the
extent of collection and trade in P. setnai istherefore essential.
Conclusions
A comprehensive assessment of the status of freshwater fishes of Western
Ghats for the IUCN Red List of Threatened Species revealed that little to no
information exists regarding distribution, population status and life history
of several threatened and endemic species (Dahanukaret al. 2011). This has led to
renewed research on various aspects including taxonomic issues and those
explaining detailed distribution of several threatened freshwater fishes of the
Western Ghats (Ali et al. 2013a,b; Emmanuel et al. 2013; Katwateet al. 2012; Knight et al. 2013). We hope that the detailed biometric and molecular data of topotypic material of Pethia setnai will aid in the reliable identification of
this threatened taxon, and the information on its distribution and threats will
be helpful in directing future conservation efforts.
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