Cabdio crassus, a new species of cyprinid fish (Teleostei: Cyprinidae) from the Kaladan River of Mizoram, India

Abstract

Cabdio crassus, a new fish species, is described from the Kaladan River in Mizoram, India. The new species is distinguished from all its congeners by having a ventral keel extending from the middle of the chest, between the posterior base of the pectoral fin and along the abdomen up to the anus (vs. more or less keeled median scales from mid-point of abdomen between posterior base of pelvic fin up to anus in all other Cabdio) and 11½–12½ branched anal-fin rays (vs. 7 in C. jaya and 9 in both C. morar and C. ukhrulensis). It is further distinguished from C. morar and C. ukhrulensis by possessing more lateral-line scales (45–51 vs. 38–42 in C. morar and 35–37 in C. ukhrulensis), more predorsal scales (20–23 vs. 17–18 in C. morar and 14 in C. ukhrulensis) and more lateral transverse scales (½7/1/3½ vs. 5/1/2 in both C. morar and C. ukhrulensis). It also differs from C. jaya in having fewer lateral-line scales (45–51 vs. 52–60), more lateral transverse scales (½7/1/3½ vs. 5/1/3) and more pharyngeal tooth-rows (3 vs. 2). Furthermore, the cytochrome c oxidase sub unit I (coi) gene sequence separates Cabdio crassus from all other Cabdio species (interspecies distance ranges from 7.8–12.3%). The anomalies observed among the GenBank sequences of the genus Cabdio are discussed and resolved.

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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by R. Pethiyagoda: 19 Jul. 2019; published: 16 Aug. 2019 159
Zootaxa 4657 (1): 159–169
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4657.1.7
http://zoobank.org/urn:lsid:zoobank.org:pub:F3187B80-06D0-4025-A2AD-6598373AF89C
Cabdio crassus, a new species of cyprinid fish (Teleostei: Cyprinidae) from the
Kaladan River of Mizoram, India
LALRAMLIANA*1, SAMUEL LALRONUNGA2 & MAHENDER SINGH3
1Department of Zoology, Pachhunga University College, Aizawl-Mizoram, India.
Email: lrl_zoo@yahoo.co.in
2Department of Environmental Science, Pachhunga University College, Aizawl-Mizoram, India.
3ICAR-National Bureau of Fish Genetic Resources, Lucknow, India.
*Corresponding author
Abstract
Cabdio crassus, a new fish species, is described from the Kaladan River in Mizoram, India. The new species is distinguished
from all its congeners by having a ventral keel extending from the middle of the chest, between the posterior base of the
pectoral fin and along the abdomen up to the anus (vs. more or less keeled median scales from mid-point of abdomen
between posterior base of pelvic fin up to anus in all other Cabdio) and 11½–12½ branched anal-fin rays (vs. 7 in C. jaya
and 9 in both C. morar and C. ukhrulensis). It is further distinguished from C. morar and C. ukhrulensis by possessing
more lateral-line scales (45–51 vs. 38–42 in C. morar and 35–37 in C. ukhrulensis), more predorsal scales (20–23 vs.
17–18 in C. morar and 14 in C. ukhrulensis) and more lateral transverse scales (½7/1/3½ vs. 5/1/2 in both C. morar and
C. ukhrulensis). It also differs from C. jaya in having fewer lateral-line scales (45–51 vs. 52–60), more lateral transverse
scales (½7/1/3½ vs. 5/1/3) and more pharyngeal tooth-rows (3 vs. 2). Furthermore, the cytochrome c oxidase sub unit
I (coi) gene sequence separates Cabdio crassus from all other Cabdio species (interspecies distance ranges from 7.8–
12.3%). The anomalies observed among the GenBank sequences of the genus Cabdio are discussed and resolved.
Keywords: Taxonomy, DNA barcode, freshwater fish, Indo-Burma Biodiversity hotspot
Introduction
Hamilton (1822) described Cyprinus morar and C. jaya under the divisions Chela and Cabdio respectively. Heckel
(1848) erected the genus Aspidoparia giving A. sardina as type species. This was subsequently acknowledged by
Günther (1868: 285–286), who tentatively included A. morar and A. jaya in the genus. Day (1878), however, con-
sidered Aspidoparia sardina to be a synonym of A. morar and included only two species under the genus, viz., A.
jaya and A. morar. Aspidoparia was regarded as a valid genus by subsequent authors (Talwar & Jhingran 1991;
Selim & Vishwanath 2001) until recent studies (Tang et al. 2010; Liao et al. 2011; Kottelat 2013) showed it to be
a junior synonym of Cabdio. There are currently three valid species of Cabdio, viz., C. jaya (Hamilton 1822), C.
morar (Hamilton 1822) and C. ukhrulensis (Selim & Vishwanath 2001).
The genus Cabdio morar (type locality ‘Yamuna and Tista Rivers, India’) has been reported from Iran, Paki-
stan, India, Nepal, Bangladesh, Myanmar and Thailand. However, the identity of the species reported as C. morar
from Iran, Myanmar, Pakistan and Thailand needs to be confirmed (Chaudhury 2010a). Cabdio jaya (type local-
ity ‘Northern Behar, India’) occurs over a narrow range along the Ganges-Brahmaputra basin in India, Nepal and
Bangladesh (Chaudhury 2010b); while C. ukhrulensis (type locality ‘Chatrickong River, Ukhrul District, Manipur,
India’) is restricted to the Chindwin-Ayeyarwaddy basin.
Recent collections of fishes in the Kaladan River included a species of Cabdio that differs from the three other
species of the genus already reported. A comparison of the morphology and cytochrome c oxidase sub unit I (coi)
sequences of the species with congeners revealed it to be an unnamed species, which is described herein as Cabdio
crassus, sp. nov.

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160 · Zootaxa 4657 (1) © 2019 Magnolia Press
Material and methods
Morphological analysis. Specimens were fixed in 10% formalin and subsequently preserved in 70% alcohol.
Counts and measurements were made on the left side of specimens following Kottelat & Freyhof (2007) (Table 1).
Measurements were made point to point with digital callipers to the nearest 0.1 mm. Measurements, except standard
length, are given as proportions of standard length (SL). Sub-units of the head are presented as proportions of head
length. Fin rays were counted under a stereo zoom microscope. For vertebral counts, two specimens were cleared
and stained in alizarin. Numbers in parentheses after a meristic value indicate the frequency of that value. Abbre-
viation of institutions: PUCMF for Pachhunga University College Museum of Fishes, Mizoram, India; MUMF for
Manipur University Museum of Fishes, Manipur, India; ZSI/ERS for Eastern Regional Station, Zoological Survey
of India, Shillong, Meghalaya, India; and ZSI for Zoological Survey of India, Kolkata. SD is standard deviation.
TABLE 1. Morphometric data for Cabdio crassus (n=15). (ZSI FF 8218, FF 8219; PUCMF 17017, 17018). Ranges
include values of holotype.
holotype Range Mean ± SD
Standard length (SL) in mm 88.1 85.8–111.4
% of SL
Body depth at dorsal 28.9 25.2–30.4 27.6±1.4
Body depth at anal 22.4 20.2–23.4 21.7±0.8
Maximum body depth 29.2 25.8–30.9 28.2±1.3
Predorsal length 60.3 56.8–61 59.3±1.2
Prepectoral length 25.7 23.3–26.1 25.0±0.7
Prepelvic length 54.1 50.9–54.2 52.9±0.8
Preanal length 77.8 73.8–78.7 76.3±1.2
Anal to pectoral 54.1 50.7–55.3 53.4±1.2
Anal to pelvic 25.5 22–26.8 25.1±1.2
Dorsal to hypural 45.3 42.8–46.1 44.3±0.9
Caudal-peduncle depth 10.3 10.3–11.3 10.6±0.3
Caudal-peduncle length 10 10–13.5 12.1±0.9
Dorsal-fin height 21 19.1–22.1 20.5±0.9
Anal-fin length 14.2 12.6–15.6 13.8±0.8
Pectoral-fin length 25 23.2–26.5 24.7±1.0
Pelvic-fin length 15.8 15.4–17.2 16.0±0.5
Head length 23.2 21.1–23.6 22.4±0.7
% of HL
Snout length 31.9 28.5–31.9 30.1±1.1
Eye diameter 25.5 23.7–28.6 25.5±1.1
Internarial 20.1 19–22.2 20.8±0.8
Interorbital width 36.8 34.5–37.6 36.3±1.0
Head width at eye 46.6 40.4–46.9 44.0±2.3
Head width at opercle 51 47–53.1 49.8±1.7
Head depth at eye 57.8 55.8–62.1 59.8±1.7
Head depth posterior to eye 69.6 64.6–71.6 69.6±1.7
Head depth at occiput 80.4 76.5–83.7 81.2±1.9
DNA extraction, PCR amplification and DNA sequencing. Approximately 40 mg of muscle tissue samples (5
paratype specimens for Cabdio crassus, 7 for C. morar and 3 for C. ukhrulensis) were extracted from the voucher
specimens and preserved in 100% ethanol prior to fixation of the specimens in formalin. DNA extraction follows
Sambrook & Russel (2001). Amplification of the coi gene were performed using the primers Fish-F1 and Fish-R1
(Ward et al. 2005) in Veriti 96 fast thermal cycler (Applied Biosystems, Inc., USA). A total of 25 µl PCR volume

NEW SPECIES OF CABDIO FROM MIZORAM, INDIA Zootaxa 4657 (1) © 2019 Magnolia Press · 161
containing 1X buffer, 100 µM dNTPs, 2 mM MgCl2, 5 pmol of each primer, 2U Taq DNA polymerase and 100 ng
template DNA were prepared. The PCR conditions are: initial denaturation of 3 min at 94°C, followed by 35 cycles
of denaturation at 94°C for 50 sec, annealing at 54°C for 30 sec., extension at 72°C for 80 sec. with final extension
of 10 min at 72°C. Sequencing was performed in forward direction only using an automated ABI 3500 Genetic
Analyzer (Applied Biosystems, Inc, USA).
TABLE 2. List of sequences (n=39) analysed in this study.
Sl No. Species NCBI Accn. No. Country Reference
1. Cabdio crassus MF145C MN065162 Mizoram, India This study
2. Cabdio crassus MF145D MN065163 Mizoram, India This study
3. Cabdio crassus MF3 MN065164 Mizoram, India This study
4. Cabdio crassus MF4 MN065165 Mizoram, India This study
5. Cabdio crassus MF5 MN065166 Mizoram, India This study
6. Cabdio morar 8076U MN065167 Lucknow, India This study
7. Cabdio morar 8076V MN065168 Lucknow, India This study
8. Cabdio morar 8076W MN065169 Lucknow, India This study
9. Cabdio morar MF31A MN065170 Mizoram, India This study
10. Cabdio morar MF31B MN065171 Mizoram, India This study
11. Cabdio morar MF31C MN065172 Mizoram, India This study
12. Cabdio morar MF31E MN065173 Mizoram, India This study
13. Cabdio ukhrulensis MF114A MN065174 Myanmar (Fish Market) This study
14. Cabdio ukhrulensis MF114B MN065175 Myanmar (Fish Market) This study
15. Cabdio ukhrulensis MF114C MN065176 Myanmar (Fish Market) This study
16. Salmostoma bacaila EU417785 India GenBank
17. Salmostoma bacaila EU417789 India GenBank
18. Salmophasia bacaila KJ936881 Meghalaya, India GenBank
19. Salmostoma bacaila EU417786 India GenBank
20. Salmophasia bacaila KT364775 Tanguar Haor, Bangladesh GenBank
21. Salmophasia bacaila KX266823 South India GenBank
22. Salmophasia bacaila KT896724 Manipur, India GenBank
23. Salmophasia bacaila JN815302 Assam, India GenBank
24. Salmophasia bacaila KT896723 Manipur, India GenBank
25. Salmostoma bacaila JX260969 Andhra Pradesh, India GenBank
26. Salmostoma bacaila JX260968 Andhra Pradesh, India GenBank
27. Salmostoma sp. KY290070 Assam, India GenBank
28. Salmostoma sp. KY290071 Assam, India GenBank
29. Cabdio morar KJ909343 Manipur, India GenBank
30. Cabdio morar KJ909344 Manipur, India GenBank
31. Cabdio morar AP011335 Aquarium trade (unknown source) Tang et al. 2010
32. Aspidoparia jaya FJ459446 India GenBank
33. Aspidoparia morar FJ459447 Assam, India GenBank
34. Salmostoma bacalia HM224244 Bangladesh Tang et al. 2010
35. Salmostoma bacalia MG954366 West Bengal, India GenBank
36. Salmostoma bacalia NC015549 West Bengal, India Tang et al. 2010
37. Salmostoma bacalia KY290087 Assam, India GenBank
38. Salmostoma bacalia KY290072 Assam, India GenBank
39. Barilius vagra FJ459413 Uttarakhand, India GenBank

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162 · Zootaxa 4657 (1) © 2019 Magnolia Press
DNA sequence analysis. The partial coi genes of fifteen individuals of three species of the genus Cabdio
(5 individuals of C. crassus, 7 of C. morar and 3 of C. ukhrulensis) were sequenced. In addition, 23 sequenc-
es comprising Cabdio and Salmostoma bacaila species along with one outgroup, Barilius vagra, were included
from GenBank for comparative analysis (Table 2). The sequences were aligned using CLUSTAL_W, blasted in
NCBI (http://www.ncbi.nlm.nih.gov) for the nearest matches and submitted to NCBI GenBank (Accession Nos.
MN065162–MN065176). The maximum likelihood (ML) and Bayesian phylogenetic tree (using the coi dataset),
implemented in MEGA7 (Kumar et al. 2016) and BEAST (Bouckaert et al. 2014) respectively, resulted into well-
supported trees with similar topologies. Since both the resultant trees exhibited similar topology, only the ML tree
is presented herein (Fig. 1). The genetic distance was calculated by averaging pairwise comparisons of sequences
across close relatives of Cabdio by the Kimura 2 parameter in MEGA7. Based on the lowest BIC (Bayesian Infor-
mation Criterion), the best fit nucleotide substitution model (out of 24 models) for present coi data set was K2+G+I
given by Nei & Kumar (2000).
FIGURE 1. Maximum Likelihood (ML) tree of coi dataset of 39 sequences using K2+G+I nucleotide substitution model with
1000 bootstraps
Results
Cabdio crassus, sp. nov.
(Fig. 2)

NEW SPECIES OF CABDIO FROM MIZORAM, INDIA Zootaxa 4657 (1) © 2019 Magnolia Press · 163
Holotype: ZSI FF 8218, 88.1 mm SL; India; Mizoram, Kaladan River, in the vicinity of Kawlchaw village;
23°28’29”N; 92°23’27”E; Lalramliana et al., 24 Nov. 2015.
Paratypes: ZSI FF 8219, (2), 91.2–94.4 mm SL; PUCMF 17017 (5), 85.8–111.4 mm SL; PUCMF 17018 (3),
90.0–95.5 mm SL; (dissected and preserved in glycerin with thymol) data as holotype.
Diagnosis. Cabdio crassus markedly differs from all other valid species of Cabdio (Fig. 3) by having the ven-
ter keeled from mid-point of chest, between the posterior base of the pectoral fin and along the abdomen up to the
anus, and possessing 11½–12½ (vs. 7–9) branched anal-fin rays. It is further distinguished from C. morar and C.
ukhrulensis by possessing more lateral-line scales (45–51 vs. 38–42 in C. morar and 35–37 in C. ukhrulensis), more
predorsal scales (20–23 vs. 17–18 in C. morar and 14 in C. ukhrulensis) and more lateral transverse scales (½7/1/3½
vs. 5/1/2 in both C. morar and C. ukhrulensis). It also differs from C. jaya in having fewer lateral-line scales (45–51
vs. 52–60), more lateral transverse scales (½7/1/3½ vs. 5/1/3) and more pharyngeal tooth-rows (3 vs. 2).
FIGURE 2. Cabdio crassus ZSI FF 8218, holotype, 88.1 mm SL; India, Mizoram, Kaladan River
Description. See Table 1 for morphometric data and Figure 2 for general appearance. Body elongate, laterally
compressed, body depth greatest at dorsal-fin origin. Dorsal profile rising from tip of snout to dorsal-fin origin, then
gradually sloping towards caudal peduncle. Ventral profile convex up to caudal peduncle; dorsal and ventral profile
of caudal peduncle slightly concave. Chest and abdomen keeled from mid-point of chest between posterior base of
pectoral fin through abdomen till anus. Head compressed along entire length; depth greater than width.
Eye ovoid, located in anterior half of head, visible in both dorsal and ventral view. Snout rounded with cluster of
tubercles; its length equal to eye diameter. Mouth subterminal, rictus not reaching vertical through anterior margin
of orbit. Barbel absent.
Dorsal fin with 2 (15) unbranched and 6½ (1) or 7½ (14) branched rays, its origin at vertical through posterior
base of pelvic fin, posterior margin slightly concave. Pectoral fin with 1 unbranched and 12 (3), 13 (9) or 14 (3)
branched rays, adpressed fin tip reaching two-thirds distance between its origin and pelvic-fin origin, axillary scale
present at its base. Pelvic fin with 1 unbranched and 7 (15) branched rays, adpressed fin tip reaching midway be-
tween its origin and anal-fin origin, axillary scale present at its base. Anus closer to anal-fin origin than to base of
last pelvic-fin ray. Anal fin with 3 unbranched and 11½ (2) or 12½ (13) branched rays. Caudal fin forked, with 10 +
9 (15) principal caudal-fin rays, its lobes more or less equal.
Lateral line complete, prominent, curved, with 45 (2), 46 (6), 47 (3), 49 (1), 50 (2), 51 (1) on body plus 1 or 2
scales on caudal-fin base; ½7/1/3 (4) or ½7/1/3½ (11) scales in transverse line from dorsal-fin origin to pelvic-fin
origin. Circumpeduncular scales 16 (15), predorsal scales 20 (3), 21 (3), 22 (7) or 23 (2), pre-pelvic and pre-anal
region scaled. Five infraorbital bones (IO1–5); IO3 largest, deep, partly overlapping preoperculum; post cleithrum
absent; 3 rows of pharyngeal teeth. Vertebrae (abdominal + caudal): 24 + 16 = 40 (1) or 25 + 15 = 40 (1).

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FIGURE 3. Image of other Cabdio spp. A) Cabdio jaya, ZSI FF 5607, 74.3 mm SL; India, Arunachal Pradesh. B) C. morar, ZSI
FF 4968, 70.6 mm SL; India, West Bengal, Ganga River. C) C. ukhrulensis, PUCMF 15054, 72.7 mm SL.
Colouration. (Fig. 2) Dorsum brownish, lighter towards flank. Lower half of body and belly silvery or pale
white. Caudal fin hyaline with black pigment distally. Other fins hyaline without markings.
Geographical distribution. Known from the main channel of the Kaladan River drainage of Mizoram, India
(Fig. 5).
Etymology. The specific name crassus is a Latin adjective meaning stout or thick, referring to the stout or thick
body in comparison with other species.
Molecular analysis. In the coi tree, Cabdio crassus is nested within Cabdio (interspecies distance ranges from
7.8 to 12.3%) (Fig. 1). The genetic distance between Cabdio crassus and C. morar is 11.7–12.3%, and between it
and C. ukhrulensis is 7.8–8.0%. The inability to collect fresh specimens results in non-availability of tissue sample
in the case of C. jaya.
Identity of Cabdio reported in GenBank. Analysis of the generated Cabdio sequences with close species
available in the GenBank revealed several potentially misidentified specimens (Table 3). Our 7 developed sequenc-
es of Cabdio morar (morphologically identified through a thorough comparison of specimens from ZSI, Kolkata
and ZSI, Shillong: see Comparative Material, below) clustered in the ML tree with GenBank sequences labelled
Salmostoma bacaila (downloaded from the NCBI KT364775, KX266823, EU417785, EU417786, EU417789,
KJ936881, KJ896723, KJ896724 and JN815302) with 0.0–1.2 % K2P distance, indicating that these GenBank
sequences in fact represent Cabdio morar. Further, the calculated genetic distance between Cabdio morar and the
species labelled as Salmostoma sp. (JX 260968, JX 260969), which is 2.0–3.0%, and between C. morar and Salmo-
stoma sp. (KY 290071), which is 4.4%, suggested that the latter two may represent species of Cabdio, rather than
to Salmostoma. Our sequences of Cabdio ukhrulensis show a very low genetic distance (0.0–0.2%) and cluster to-
gether in the ML tree with GenBank sequences labelled C. morar (AP011335, KJ909343 and KJ909344) indicating
that they too, are conspecific. Furthermore, the GenBank sequences labelled as Aspidoparia morar (FJ459447) and
A. jaya (FJ459446) clustered together with Salmostoma bacaila (HM224244, MG954366, NC015549, KY290072,
KY290087) with a low genetic distance of 0.3–0.7 % indicating that they represent the latter species.

NEW SPECIES OF CABDIO FROM MIZORAM, INDIA Zootaxa 4657 (1) © 2019 Magnolia Press · 165
FIGURE 4. Ventral view of A) Cabdio crassus, ZSI FF 8218, holotype, 88.1 mm SL (arrow showing keeled chest); B) C. ukh-
rulensis, PUCMF 15054, 72.7 mm SL; C) C. morar, PUCMF 2094, 64.3 mm SL; D) C. jaya, ZSI FF 5607, 74.3 mm SL.
Discussion
Cabdio crassus conforms to the conception of the genus (as Aspidoparia) by Heckel (1848), Day (1878), Talwar &
Jhingran (1991) and Vishwanath et al. (2007) except that it possesses a keeled ventral surface from the mid-point of
the chest between the posterior base of the pectoral fin and along the abdomen, up to the anus, and a higher number
of branched anal-fin rays (12–13 vs. 7–9 in all other Cabdio species).
A keeled chest and abdomen as in Cabdio crassus is present also in species of the cyprinid genera Securicula
and Salmostoma; this apparently apomorphic character has never yet been reported in other species of Cabdio.
However, C. crassus differs from both Securicula and Salmostoma in having the lower jaw shorter than the upper

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166 · Zootaxa 4657 (1) © 2019 Magnolia Press
jaw (i.e. a sub-terminal mouth) and lacking a symphyseal process (vs. lower jaw longer or jaws subequal, a sym-
physeal process present). Moreover, the species is clearly nested within the clade comprising species of Cabdio in
the coi inferred ML tree (Fig. 1). It is perhaps worth mentioning that, although it has not been reported by previous
authors, C. ukhrulensis, C. morar and C. jaya exhibit an abdomen in which the median scales more or less form a
keel (see Fig 4).
DNA barcoding represents a rapid, accurate and automatable species-identification method using short and
standardized gene regions (Hebert & Gregory 2005). However, inaccurate identification of the barcoded species
in the GenBank register retards the utility of identification using barcodes and leads to confusion (Conte-Grand et
al. 2017; Lalramliana et al. 2018; Lalramliana et al. 2019). For instance, the genetic distance between the seven
sequences of Cabdio morar developed here and species labelled as Salmostoma bacaila (KT364775, KX266823,
EU417785, EU417786, EU417789, KJ936881, KJ896723, KJ896724 and JN815302) in GenBank is very low (99%
identity with 0.0–0.8 % K2P distance) indicating that they belong to the same species.
FIGURE 5. Drainage Map showing the locality of Cabdio crassus sp. nov. (▲) and the distribution of the genus Cabdio ex-
amined in this study.
▲ Cabdio crassus ■ Cabdio jaya ● Cabdio morar ▼Cabdio ukhrulensis (▽ holotype)
The examples of Cabdio morar studied here were collected from the Barak River Drainage (Brahmaputra ba-
sin) of Mizoram and Ganges River near Lucknow, India, and identified morphologically through comparison with
specimens from the Ganga basin present in ZSI, Kolkata, India. Cabdio morar is characterized by having 2 simple
and 7½ branched dorsal-fin rays, 1 simple and 7½ branched pelvic-fin rays, 1 simple and 14 branched pectoral-fin
rays, 3 simple and 9½ branched anal-fin rays, a complete lateral line with 38–42 + 1–2 scales, ½4/1/2½–3 scales in
transverse line from dorsal-fin origin to pelvic-fin origin and 3 pharyngeal teeth rows (Hamilton 1822; Day 1878;
Selim & Vishwanath 2001; ZSI FF 4968; ZSI/V/ERS 8354; PUCMF 15051–53). It therefore appears that the spe-
cies identified as Salmostoma bacaila (mentioned above) in the NCBI Genbank in fact represents Cabdio morar.
Similarly, the same anomaly is observed between the three developed sequences of Cabdio ukhrulensis and species
labelled as C. morar (AP011335, KJ909343 and KJ909344). The Cabdio ukhrulensis analysed in this study were
collected from the local fish market in Tahan, Myanmar, presumably from the nearby Myittha River, a tributary of
Chindwin River (as per communication with the fishermen). Examination of, and comparison with the type speci-
mens revealed these specimens to be C. ukhrulensis (type locality: Chatrickong River, Chindwin drainage, Mani-
pur). Surprisingly, the 3 developed sequences of Cabdio ukhrulensis shared 99% identity (0.2% K2P distance) with
the three GenBank sequences of C. morar (AP011335, KJ909343 and KJ909344).

NEW SPECIES OF CABDIO FROM MIZORAM, INDIA Zootaxa 4657 (1) © 2019 Magnolia Press · 167
TABLE 3. Summary of synonymized species/probable misidentification and resolution of the genus Cabdio and
Salmostoma bacaila in the NCBI GenBank, accession numbers and habitat locality
S l
No.
Specimen ID in NCBI Specimen ID corrected NCBI Accn.
No.
Country Reference
1. Salmostoma bacaila Cabdio morar EU417785 India GenBank
2. Salmostoma bacaila Cabdio morar EU417789 India GenBank
3. Salmophasia bacaila Cabdio morar KJ936881 Meghalaya, India GenBank
4. Salmostoma bacaila Cabdio morar EU417786 India GenBank
5. Salmophasia bacaila Cabdio morar KT364775 Tanguar Haor, Bangladesh GenBank
6. Salmophasia bacaila Cabdio morar KX266823 South India GenBank
7. Salmophasia bacaila Cabdio morar KT896724 Manipur, India GenBank
8. Salmophasia bacaila Cabdio morar JN815302 Assam, India GenBank
9. Salmophasia bacaila Cabdio morar KT896723 Manipur, India GenBank
10. Salmostoma bacaila Cabdio morar (?) JX260969 Andhra Pradesh, India GenBank
11. Salmostoma bacaila Cabdio morar (?) JX260968 Andhra Pradesh, India GenBank
12. Salmostoma sp. Cabdio morar KY290070 Assam, India GenBank
13. Salmostoma sp. Cabdio sp.KY290071 Assam, India GenBank
14. Cabdio morar Cabdio ukhrulensis KJ909343 Manipur, India GenBank
15. Cabdio morar Cabdio ukhrulensis KJ909344 Manipur, India GenBank
16. Cabdio morar Cabdio ukhrulensis AP011335 Aquarium trade (unknown
source)
Tang et al.
2010
17. Aspidoparia jaya Salmostoma bacalia FJ459446 India GenBank
18. Aspidoparia morar Salmostoma bacalia FJ459447 Assam, India GenBank
It is interesting that Cabdio morar and C. ukhrulensis are superficially similar. Cabdio morar differs from the
latter, however, in having more lateral line scales (38–42 vs. 35–37), more predorsal scales (17–18 vs. 14) and more
pharyngeal tooth-rows (3 vs. 2). Further, analysis of the three C. morar sequences in NCBI revealed the provenance
of the two specimens (KJ909343 and KJ909344) as the Chindwin drainage of Manipur (as per data from NCBI),
while AP011335 is from aquarium trade, with unknown locality (Tang et al. 2010). It appears therefore that the
sequences of Cabdio morar previously submitted in NCBI have been misidentified: they belong to C. ukhrulensis
rather than to C. morar. Furthermore, the available sequences of Cabdio jaya (as Aspidoparia jaya, FJ459446) and
C. morar (as Aspidoparia morar, FJ459447) also appear to cases of misidentification. These two sequences appear
to be conspecific and represent Salmostoma, based on the low genetic distance and further consensus with Salmos-
toma bacaila in the ML tree.
Cabdio jaya is a species described by Hamilton (1822) from the rivers of northern Bihar (River Ganges) without
any extant type material. Its original description did not include the lateral line scales count. However, the species
was characterized by subsequent workers (Day 1878; Talwar & Jhingran 1991; Selim & Vishwanath 2001; Vish-
wanath et al. 2007), including the lateral-line scales count, and we have relied on these data to identify this species.
Our many attempts to obtain fresh material of C. jaya from the type locality, unfortunately, failed. The examined
specimens of C. jaya (labelled as Aspidoparia jaya) from ZSI Kolkata are from Brahmaputra River of Arunachal
Pradesh, India. Though, the examined specimens are not from the exact type locality of C. jaya, data from literature
suggest that this identification is correct.
Kar & Sen (2007) listed Cabdio morar (as Aspidoparia morar) from the Kaladan River of Mizoram without
providing a description or illustration, and not mentioning where their specimens are deposited. It is likely that the
specimens they reported as C. morar were in fact C. crassus.
In conclusion, the discovery of Cabdio crassus from the Kaladan raises the number of Cabdio species to four.
Despite the stated weakness in using GenBank sequences for comparison, the present study demonstrates that the
coi gene fragment is an effective tool for identification of species of Cabdio.

LALRAMLIANA ET AL.
168 · Zootaxa 4657 (1) © 2019 Magnolia Press
Comparative material
Cabdio jaya: ZSI FF 5607, 11 ex., 56.0–74.5 mm SL; India: Arunachal Pradesh: Namdapha National Park & Tiger Reserve.—
ZSI/V/F/ERS 222; 11 ex., 46.4–64.2 mm SL; India: Arunachal Pradesh: D. Ering Sanctuary, E. Siang.
Cabdio morar: ZSI FF 4968, 2 ex., 67.4–70.6 mm SL; India: West Bengal: Ganga River in the vicinity of Palaghat, Panchanan-
dapur.—ZSI/V/ERS 8354; 9 ex., 46.4–81.1 mm SL; India: Assam: River at Goalpara, Goalpara District.—PUCMF 2094,
1, 64.3 mm SL; India: Mizoram: Terei River, a tributary of the Tlawng river (Barak River drainage) in the vicinity of
Saikhawthlir village.—PUCMF 15051, 2, 52.7–63.2 mm SL; India: Mizoram: Terei River, a tributary of the Tlawng river
(Barak River drainage) in the vicinity of Chuhvel village. —PUCMF 15052–53, 2, 61.5–64.3 mm SL; India: Mizoram: Mar
River, a tributary of the Karnaphuli River in the vicinity of Phuldungsei village.
Cabdio ukhrulensis: MUMF 1025, 1, holotype, 75.1 mm SL; India: Manipur: Ukhrul District, Chatrickong River (Chindwin
drainage).—PUCMF 15054, 2, 49.0–72.7 mm SL; Myanmar: Tahan, Tahan Market (presumably from the nearby Myittha
River, a tributary of Chindwin River).—PUCMF 15055, 2, 45.3–54.2 mm SL; data same as PUCMF 15054 (cleared and
stained).
Salmostoma bacaila: ZSI/V/F/ERS 794, 1 ex., 72.3 mm SL; India: Assam: Banawh River, 4 km East of Bijni, Bongaigaon
District.
Salmostoma phulo: ZSI/V/F/ERS 942, 2 ex., 48.7–50.3 mm SL; India: Assam: Bortari, Kamrup District.—ZOONEHU 49, 2 ex.,
51.6–53.9 mm SL; India: Meghalaya: Bugi River.
Acknowledgements
We gratefully acknowledge the financial support from Department of Biotechnology, Government of India (No.
BT/388/NE/TBP/2012). We are also grateful to Director, Zoological Survey of India, Kolkata, Laishram Kosygin
(ZSI, Kolkata), Dimos Khynriam (ZSI, Shillong) and Waikhom Vishwanath (MUMF) for permission to access ma-
terials under their care; Henry Lalmawizuala and K. Lalchhandama for language editing; Vanlalmalsawma (Sawm-
tea) for field assistance; Tawnenga, Principal, Pachhunga University College and Director, ICAR-National Bureau
of Fish Genetic Resources, Lucknow for providing laboratory facilities.
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