Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 September 2026 | 18(9): 29729–29735
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.10249.18.9.29729-29735
#10249 | Received 11 November 2025 | Final received 09 June 2026| Finally
accepted 05 September 2026
DNA Barcode
based species delineation within Freshwater Garfish Xenentodon
cancila (Hamilton, 1822) (Beloniformes:
Belonidae) reveals the existence of multiple lineages
under single nominal taxon
Ranjit More 1 , Jiwan Sarwade
2 , A. Shabnam 3 & K.P. Dinesh 4
1 Department of Zoology, Pratapsinh Mohite-Patil Mahavidyala, Near Girdhardas Devi
Highschool, Pune Road Tal Karmala, Solapur District,
Maharashtra 413203, India.
2 Department of Zoology, Arts, Science and Commerce College Indapur, Pune-Solapur
Road (NH 65), Indapur, Pune, Maharashtra 413106,
India.
3,4 Zoological Survey of India,
Western Regional Centre, Vidya Nagar, Sector-29, P.C.N.T. (PO), Rawet Road, Akurdi, Pune, Maharashtra
411044, India.
1 zoologistranjit@gmail.com (corresponding
author),2 j.sarwade@rediffmail.com, 3 a.shabnam1312@gmail.com, 4 kpdinesh.zsi@gmail.com
Editor: Mandar Paingankar,
Government Science College, Gadchiroli, India. Date of publication: 26
September 2026 (online & print)
Citation: More,
R., Jiwan Sarwade, A.
Shabnam & K.P. Dinesh (2026). DNA Barcode based species delineation
within Freshwater Garfish Xenentodon cancila (Hamilton, 1822) (Beloniformes:
Belonidae) reveals the existence of multiple lineages
under single nominal taxon. Journal of
Threatened Taxa 18(9):
29729–29735. https://doi.org/10.11609/jott.10249.18.9.29729-29735
Copyright: © More et al. 2026. Creative Commons Attribution 4.0 International License.
JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: None.
Competing interests: The authors declare no competing interests.
Author details: Ranjit More is Assistant Professor and Head, Department of Zoology, Pratapsinh Mohite-Patil Mahavidyalaya, Karmala, Solapur (MS) 413203, India, working as a fisheries biologist specializing in taxonomy, ecology and conservation of freshwater ichthyofauna. Jiwan Sarwade is Principal, Arts, Science and Commerce College, Indapur, Pune, Maharashtra, India. A. Shabnam is a as Research Associate at Zoological Survey of India and works on cockroach taxonomy with expertise in DNA barcoding and molecular phylogenetics. K.P. Dinesh is an amphibian taxonomist with expertise in DNA barcoding and molecular phylogenetics and scientist at the Zoological Survey of India.
Author contributions: RM and JS designed the experiment. RM conducted the practical work. AS and KPD collected and analyzed the molecular data. RM, AS, and KPD contributed to manuscript writing. All authors have read and approved the final manuscript.
Acknowledgements: RM and JS thank the principal and head of the Department of Zoology, Modern College, Pune, and Arts, Science and Commerce College, Indapur. RM is grateful to SARTHI for financial support during the study period. The Maharashtra State Biodiversity Board for granting permission to access biological resources (No. MSBB/Research/CR
622/615/2019-2020) is duly acknowledged. AS and KPD thank the director, Zoological Survey of India, Kolkata, and the officer-in-charge, ZSI WRC, Pune, for their support. We acknowledge Sahil Shikalgar, research fellow, ZSI, WRC, Pune for help in photography.
Abstract: The present correspondence provides the first DNA barcode-based evidence
of multiple evolutionary lineages within the freshwater garfish Xenentodon cancila
across South Asia. Taxonomically validated specimens from the Ujani Reservoir, Maharashtra, India, were analyzed with 77
publicly available mt COI sequences. Phylogenetic and
species delimitation analyses revealed seven distinct clades within Xenentodon, substantially exceeding the currently recognized two species. The Ujani population formed a strongly supported
monophyletic cluster with sequences from India, Bangladesh, and Korea,
corresponding closely to the nominal X. cancila.
Other geographically structured lineages from southern, northeastern, and
central India, exhibit deep genetic divergence, indicating potential
undescribed taxa. These findings highlight persistent taxonomic gaps within Xenentodon and emphasize the need for
integrative revision combining molecular and morphological datasets.
Keywords: Cryptic diversity, freshwater
biodiversity, garfish, integrative taxonomy, morphological datasets, phylogeny,
polyphyletic, Ujani Reservoir.
INTRODUCTION
DNA barcoding
is a widely used molecular tool, enabling rapid species identification and
discovery of cryptic diversity in fishes (Hajibabaei
et al. 2007; Stein et al. 2014; Rahman et al. 2019). Although extensive DNA barcode datasets are
available through global repositories such as BOLD (Ratnasingham
& Hebert 2007) and GenBank, significant geographic gaps persist across
Asia, resulting in taxonomic uncertainties in many Indian fish species.
The family Belonidae
(order Beloniformes) comprises 47 species in 10
genera (Fricke et al. 2018). Among them, the southern and southeastern Asia
endemic genus Xenentodon includes two
recognized species, Xenentodon cancila (Hamilton, 1822) and X. canciloides
(Bleeker, 1854). X. cancila, originally described from the Ganga River,
India, is widely distributed in India, Bangladesh, Cambodia, Malaysia, Nepal,
Sri Lanka, Bhutan, Laos, Myanmar, Pakistan, Vietnam, Hawaii, Hong Kong,
Indonesia, and Thailand (Hossain et al. 2013), while X. canciloides
is restricted to Cambodia, Indonesia, Laos, Thailand, and Vietnam (Froese and Pauly 2026).
The freshwater garfish X. cancila is an economically important fish in southern
Asia and is widely traded in regional fisheries (Islam & Dutta 2018).
Despite its broad distribution range and commercial significance, molecular
studies have been limited and geographically fragmented (Thapliyal et al. 2013;
Khedkar et al. 2014; Patil
et al. 2018), resulting in its poorly resolved genetic identity. Against this
backdrop, the present study examines the genetic identity of morphologically
verified X. cancila specimens from Ujani Reservoir, Maharashtra, India, using mitochondrial
Cytochrome C Oxidase subunit 1 (mt COI) barcodes to
assess species monophyly of X. cancila through
single-gene-based phylogenetic analysis. The phylogenetic pattern recovered are
discussed and ascertained through ASAP species-delimitation analysis to
delineate the potential existence of cryptic species within Xenentodon.
MATERIALS AND METHODS
Specimens
were collected and studied from the Ujani Reservoir,
Maharashtra, India, during 2018–2021. The Identity of the specimens as Xenentodon cancila was
confirmed by a taxonomic expert at the Zoological Survey of India. The studied
specimens were deposited in the Zoology Museum, Modern College, Pune, and the
Arts, Science and Commerce College, Indapur
(ICASC/ZOO/105–106/2021–22). Fresh specimens were photographed (Image 1) before
DNA-barcoding studies.
Genomic DNA was extracted from
white muscle or fin clips using the DNeasy Blood
& Tissue Kit (Qiagen), following the manufacturer’s protocol. DNA quality
was assessed using a Qubit 2.0 fluorometer, and samples above 50 ng were
amplified with universal mitochondrial Cytochrome C Oxidase subunit 1 (mt COI) primers. PCR was performed in a 25 µL reaction
volume under the following thermal-cycling conditions: 95 ⁰C for three min; 35
cycles of 95 ⁰C for 30 s, 50 ⁰C for 30 s, 72 ⁰C for 45 s; final extension at 72
⁰C for three min (Patil et al. 2018). Amplified ~700 bp COI fragments were bidirectionally sequenced at Barcode
Biosciences, Bengaluru, India.
Raw sequence files were manually
checked for quality to be used for further analysis. Additionally, 77 sequences
of Belonidae were downloaded from GenBank, cleaned,
and aligned in MEGA X (Kumar et al. 2018). Maximum--Likelihood (ML) analysis
was performed with 83 sequences (Table 1) in the IQ-TREE v1.6.12 web server (Trifinopoulos et al. 2016), with 1,000 ultrafast bootstraps
under the GTR+F+I+G4 model selected by Bayesian information criterion (BIC).
The obtained tree was visualized in FigTree v1.4. The
same dataset was subjected to species-delimitation analysis using the Assemble Species
by Automatic Partitioning (ASAP) method (Puillandre
et al. 2021). Sequences from this study were submitted to GenBank under
accessions OM865416–OM865419.
RESULTS AND DISCUSSION
The mt COI phylogenetic analysis of Belonidae
revealed multiple lineages within Xenentodon,
although with low bootstrap support indicating polyphyly of the genus (Image
2). However, X. cancila sequence from the Ujani Reservoir
formed monophyletic clade with
sequences from India, Bangladesh, and Korea. Overall, seven distinct clades
were recovered, far beyond the two currently recognized Xenentodon
species. Species delimitation using ASAP (Image 3) corroborated the
phylogenetic pattern, supporting the existence of multiple independently
evolving lineages.
Although mt COI inference has limited utility for resolving deeper
phylogenetic relationships, it remains valuable marker for detecting lineage
diversity and identifying taxonomic hypotheses for further taxonomic revisions
(Kalawate et al. 2022). Notably, most publicly
available Xenentodon sequences are from
India but lack associated voucher-specimen data, limiting taxonomic assignment
with confidence.
Type locality
of X. cancila is the Ganga River, India (Hamilton
1822), whereas that of X. canciloides is the
Kapuas River, Indonesia (Bleeker 1854) (Image 4). Considering the proximity of
sequences origin location and respective type localities tentatively Clade I
can be assigned to X. cancila (sequences from
India and Bangladesh) (Images 2 & 3) and Clade III to X. canciloides (sequences from Thailand) tentatively
(Images 2 & 3). The Remaining lineages corresponding to Clade II (from
Maharashtra, India), Clade IV (from Kerala and Tamil Nadu, India), Clade V
(from Manipur, India), Clade VI (from Uttarakhand, India), and Clade VII (from
Arunachal Pradesh, West Bengal, India) (Images 2 & 3) likely represent
undescribed species, warranting ‘Integrative Taxonomic Studies’ with broader
sampling from respective localities and taxonomic revision of the group.
CONCLUSION
The genetic
heterogeneity among populations suggests that current knowledge of species
distribution and diversity is far from complete. This brings into attention
multiple lineages and the need to revisit and re-evaluate riverine and
watershed-based biogeographic patterns of Xenentodon
species across southern Asia, primarily in India and Bangladesh, to better
understand the diversity, distribution, evolutionary relationships, and
ecology.
Data availability statement
All relevant
data supporting the findings of this study are included within the manuscript.
No additional datasets were generated or analyzed during the current
study.
Table 1. Sequences details of 600 bps fragment
of mt COI gene data used in maximum likelihood IQ tree construction.
|
|
Accessions |
Name in GenBank |
Locality as per GenBank |
References |
|
1 |
OM865416.1 |
Xenentodon
cancila |
Ujani reservoir, Maharashtra, India |
This study |
|
2 |
OM865417.1 |
X. cancila |
Ujani reservoir, Maharashtra, India |
This study |
|
3 |
OM865418.1 |
X. cancila |
Ujani reservoir, Maharashtra, India |
This study |
|
4 |
OM865419.1 |
X. cancila |
Ujani reservoir, Maharashtra, India |
This study |
|
5 |
MH087053.1 |
X. cancila |
Bangladesh |
Unpublished |
|
6 |
MK359874.1 |
X. cancila |
Korea |
Unpublished |
|
7 |
KF742433.1 |
X. cancila |
Maharashtra, India |
Unpublished |
|
8 |
MK359936.1 |
X. cancila |
Korea |
Unpublished |
|
9 |
MK359971.1 |
X. cancila |
Korea |
Unpublished |
|
10 |
MK572631.1 |
X. cancila |
Moulvibazar, Sylhet,
Bangladesh |
Rahman et al. 2019 |
|
11 |
MK814506.1 |
X. cancila |
Rajasthan, India |
Unpublished |
|
12 |
MK814492.1 |
X. cancila |
Rajasthan, India |
Unpublished |
|
13 |
JX983511.1 |
X. cancila |
Narmada river, Dindori, MP, India |
Khedkar
et al. 2014 |
|
14 |
JX983513.1 |
X. cancila |
Narmada river, Dindori, MP, India |
Khedkar
et al. 2014 |
|
15 |
KX399218.1 |
X. cancila |
Tripura, India |
Unpublished |
|
16 |
KX399217.1 |
X. cancila |
Tripura, India |
Unpublished |
|
17 |
KX399216.1 |
X. cancila |
Tripura, India |
Unpublished |
|
18 |
JX983512.1 |
X. cancila |
Narmada river, Dindori, MP, India |
Khedkar
et al. 2014 |
|
19 |
KU685520.1 |
X. cancila |
Tripura, India |
Unpublished |
|
20 |
MK814522.1 |
X. cancila |
Rajasthan, India |
Unpublished |
|
21 |
KX399222.1 |
X. cancila |
Tripura, India |
Unpublished |
|
22 |
KU685522.1 |
X. cancila |
Tripura, India |
Unpublished |
|
23 |
KX399223.1 |
X. cancila |
Tripura, India |
Unpublished |
|
24 |
KX399221.1 |
X. cancila |
Tripura, India |
Unpublished |
|
25 |
KU685524.1 |
X. cancila |
Tripura, India |
Unpublished |
|
26 |
KX399220.1 |
X. cancila |
Tripura, India |
Unpublished |
|
27 |
KX399219.1 |
X. cancila |
Tripura, India |
Unpublished |
|
28 |
KU685521.1 |
X. cancila |
Tripura, India |
Unpublished |
|
29 |
KU685523.1 |
X. cancila |
Tripura, India |
Unpublished |
|
30 |
JX260995.1 |
X. cancila |
Maharashtra, India |
Unpublished |
|
31 |
MK572630.1 |
X. cancila |
Bangladesh |
Rahman et al. 2019 |
|
32 |
MK572629.1 |
X. cancila |
Bangladesh |
Rahman et al. 2019 |
|
33 |
KX946829.1 |
X. cancila |
Maharashtra, India |
Patil et al.
2018 |
|
34 |
KX946828.1 |
X. cancila |
Maharashtra, India |
Patil et al.
2018 |
|
35 |
MK049515.1 |
X. cancila |
Thailand |
Panprommin
et al.
2019 |
|
36 |
MK049514.1 |
X. cancila |
Thailand |
Panprommin
et al.
2019 |
|
37 |
MK049516.1 |
X. cancila |
Thailand |
Panprommin
et al.
2019 |
|
38 |
KU569073.1 |
Xenentodon
sp. |
South Africa |
Unpublished |
|
39 |
MK348178.1 |
X. cancila |
Ashtamudi lake, Kerala,
India |
Unpublished |
|
40 |
MH377838.1 |
X. cancila |
Kerala, India |
Unpublished |
|
41 |
MH377837.1 |
X. cancila |
Kerala, India |
Unpublished |
|
42 |
MH377836.1 |
X. cancila |
Kerala, India |
Unpublished |
|
43 |
MK348194.1 |
X. cancila |
Ashtamudi lake, Kerala,
India |
Unpublished |
|
44 |
MK348176.1 |
X. cancila |
Ashtamudi lake, Kerala,
India |
Unpublished |
|
45 |
KX826913.1 |
X. cancila |
Tamil Nadu, India |
Unpublished |
|
46 |
MG923408.1 |
X. cancila |
Vembanad
Lake, Kerala, India |
Unpublished |
|
47 |
MG923407.1 |
X. cancila |
Vembanad Lake, Kerala, India |
Unpublished |
|
48 |
MK348177.1 |
X. cancila |
Ashtamudi lake, Kerala,
India |
Unpublished |
|
49 |
MG923406.1 |
X. cancila |
Vembanad Lake, Kerala, India |
Unpublished |
|
50 |
EF607569.1 |
S. strongylura |
China |
Unpublished |
|
51 |
KF715035.1 |
Tylosurus gavialoides |
Philippines |
Unpublished |
|
52 |
EF607566.1 |
Strongylura
leiura |
China |
Zhang 2011 |
|
53 |
FJ237566.1 |
Strongylura
leiura |
India |
Lakra et al. 2011 |
|
54 |
GU674374.1 |
Tylosurus
crocodilus |
Indonesia |
Unpublished |
|
55 |
GU674427.1 |
Ablennes sp. |
Indonesia |
Unpublished |
|
56 |
MT323766.1 |
Ablennes hians |
Gulf of Mexico |
Unpublished |
|
57 |
KC970513.1 |
Tylosurus
acus |
Philippines |
Unpublished |
|
58 |
MH377844.1 |
Tylosurus acus melanotus |
India |
Unpublished |
|
59 |
GU702391.1 |
Tylosurus acus acus |
Brazil |
Unpublished |
|
60 |
KY176687.1 |
Tylosurus acus imperialis |
Turkey |
Unpublished |
|
61 |
GU225490.1 |
Strongylura
timucu |
Mexico |
Unpublished |
|
62 |
HQ937019.1 |
Strongylura marina |
USA |
April et al. 2011 |
|
63 |
KX245130.1 |
X. cancila |
India |
Unpublished |
|
64 |
KX245129.1 |
X. cancila |
India |
Unpublished |
|
65 |
JN965212.1 |
X. cancila |
Dehradun, India |
Thapliyal et al. 2013 |
|
66 |
FJ459540.1 |
X. cancila |
India |
Unpublished |
|
67 |
FJ459539.1 |
X. cancila |
India |
Unpublished |
|
68 |
FJ459538.1 |
X. cancila |
India |
Unpublished |
|
69 |
FJ459541.1 |
X. cancila |
India |
Unpublished |
|
70 |
MT812048.1 |
X. cancila |
Arunachal Pradesh, India |
Unpublished |
|
71 |
MN096221.1 |
X. cancila |
West Bengal, India |
Unpublished |
|
72 |
MT812049.1 |
X. cancila |
Arunachal Pradesh, India |
Unpublished |
|
73 |
MT812050.1 |
X. cancila |
Arunachal Pradesh, India |
Unpublished |
|
74 |
MT812052.1 |
X. cancila |
Arunachal Pradesh, India |
Unpublished |
|
75 |
MT812051.1 |
X. cancila |
Arunachal Pradesh, India |
Unpublished |
|
76 |
GU225484.1 |
Strongylura notata |
Mexico |
Unpublished |
|
77 |
JQ843010.1 |
Platybelone
argala |
Trinidad and Tobago |
Weigt et al.
2012 |
|
78 |
MF124076.1 |
Tylosurus
choram |
Israel |
Unpublished |
|
79 |
JF494139.1 |
Petalichthys
capensis |
South Africa |
Unpublished |
|
80 |
KJ768216.1 |
Belone belone |
Portugal |
Landi
et al. 2011 |
|
81 |
KC500332.1 |
Belone belone |
Turkey |
Keskin & Atar 2014 |
|
82 |
MK572415.1 |
Oryzias dancena |
Bangladesh |
Rahman et al. 2019 |
|
83 |
MK572416.1 |
Oryzias dancena |
Bangladesh |
Rahman et al. 2019 |
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