Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 July 2020 | 12(10): 16262–16271
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.6040.12.10.16262-16271
#6040 | Received 24 April 2020 | Final
received 21 June 2020 | Finally accepted 19 July 2020
The identity and distribution of Bhavania annandalei
Hora, 1920 (Cypriniformes: Balitoridae),
a hillstream loach endemic to the Western Ghats of
India
Remya L. Sundar
1, V.K. Anoop 2, Arya Sidharthan
3, Neelesh Dahanukar
4 & Rajeev Raghavan 5
1,5 Center for Aquatic Resource Management
and Conservation (CARMAC), Kerala University of Fisheries and Ocean Studies
(KUFOS), Panangad P.O., Kochi, Kerala 682506, India.
2,3,5 School of Ocean Science and
Technology (SOST), Kerala University of Fisheries and Ocean Studies (KUFOS), Panangad P.O.,
Kochi, Kerala 682506, India.
4 Indian Institute of Science
Education and Research (IISER), Dr. Homi Bhabha Road, Pashan, Pune,
Maharashtra 411008, India.
4 Zoo Outreach Organization (ZOO),
No. 12 Thiruvannamalai Nagar, Saravanampatti,
Coimbatore, Tamil Nadu 641035, India.
5 Department of Fisheries Resource
Management, Kerala University of Fisheries and Ocean Studies (KUFOS), Panangad P.O.,
Kochi, Kerala 682506, India.
1 remyalsundar@gmail.com, 2 anoopanjukunnu@gmail.com,
3 aryaanandhu06@gmail.com, 4 n.dahanukar@iiserpune.ac.in,
5 rajeevraq@hotmail.com
(corresponding author)
Editor: Anonymity requested. Date of publication: 26 July 2020 (online & print)
Citation: Sundar, R.L., V.K. Anoop,
A. Sidharthan, N. Dahanukar
& R. Raghavan (2020).
The
identity and distribution of Bhavania annandalei Hora, 1920 (Cypriniformes:
Balitoridae), a hillstream
loach endemic to the Western Ghats of India. Journal of Threatened Taxa 12(10): 16262–16271. https://doi.org/10.11609/jott.6040.12.10.16262-16271
Copyright: © Sundar et al. 2020. 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: Kerala University of Fisheries and Ocean Studies.
Competing interests: The authors declare no competing interests.
Author details: Remya L. Sundar, is interested in taxonomy, diversity and distribution of freshwater and estuarine fishes of Kerala. V.K. Anoop is interested in the systematics and biogeography of loaches of the Western Ghats. Arya Sidharthan is interested in molecular ecology and phylogeography of balitorid loaches of the Western Ghats. Neelesh Dahanukar is interested in molecular phylogenetics and biogeography of freshwater fishes of the Western Ghats. RAJEEV RAGHAVAN is interested in conservation of aquatic biodiversity of Western Ghats with special reference to freshwater fishes.
Author contribution: RR, ND and AS conceived the study; RLS, VKA and AS carried out the field surveys and laboratory studies; all authors equally contributed to the writing of the manuscript.
Acknowledgements: VKA and AS thank the Kerala State
Biodiversity Board (KSBB) for PhD fellowship, and RLS and RR thank the Center for Aquatic Resource Management and Conservation
(CARMAC), Kerala University of Fisheries and Ocean Studies (KUFOS) for
funding. The authors are grateful to
M.R. Ramprasanth, Josin Tharian, Vishnu Raj, and Anvar
Ali for useful discussions and help in the field. Permits for collection inside forest areas of
Kerala were provided by the Kerala State Forest and Wildlife Department to VKA
and AS.
Abstract: Bhavania annandalei Hora 1920, is resurrected from
the synonymy of B. australis (Jerdon, 1849) based on examination of freshly collected topotypic specimens.
The two species can be distinguished by a combination of morphological
characters including low, dense, and sparsely distributed tubercles on dorsal
surface of head and operculum, rostral barbels
reaching anterior border of upper lip, rostral flaps between the rostral barbels fleshier, 11–12 scale rows above the lateral line,
and caudal peduncle stout with its depth to width ratio less than 2.5. The two species formed significantly distinct
clusters in multivariate space. Further,
the two species have a raw genetic distance of 6.4% in the mitochondrial cytochrome
oxidase subunit 1 gene. The distribution
of B. annandalei is restricted to the river
systems draining the Agasthyamalai Hills, below the Shencottah Gap in southern Western Ghats, while B. australis occurs in rivers north of the Shencottah Gap.
Keywords: Agasthyamalai,
Cobitoidea, Kerala, mountain loach, synonymy.
Introduction
The hillstream
loach Bhavania annandalei
was described by Hora (1920; p203) from Tenmalai,
erstwhile Travancore State (= current day southern Kerala), and suggested that
the species occurs throughout the southern Western Ghats in the Nilgiris, Malabar, and Travancore. Hora (1920) diagnosed B. annandalei from its only known congener, B. australis (Jerdon, 1849)
(type locality: Walliar Jungle = Walayar),
by a combination of characters; the most prominent of which included a broad
snout (vs. pointed), interrupted lower lip (vs. continuous), caudal-lobes equal
(vs. lower lobe longer), and presence of a pair of papillae on the lower lip
(vs. absence).
Hora’s (1920) description of B.
annandalei was however, based on a single adult
female specimen collected by Dr. Annandale from
Travancore, Kerala. Though, Hora (1920)
seemed to have access to additional juvenile specimens collected by Captain
Sewell from the Nilgiris (Cherambadi)
and Wayanad (Nellimunda, Mananthavady,
and near Vythiri), he did not examine them or provide
other details. Subsequently, Hora (1937;
p8) extended the distribution of the species to Mysore, based on four specimens
collected by M.S. Bhimachar from a stream between Kottigehar and Balehonnur
(erstwhile Mysore State = current day Tunga River
System in Karnataka). No details of the
specimens were provided.
In his review on ‘Homalopterid fishes from Peninsular India’, Hora (1941)
synonymized B. annandalei with B. australis, after examining specimens from throughout
its distribution range including Kallar/South
Travancore (current day Vamanapuram River, Kerala); Pampadumpara/North Travancore (current day Periyar River, Kerala); Sethumadai
Hills/ Mysore (current day Anamalai hills near
Pollachi, Tamil Nadu); and Kottigehar/Mysore (current
day Tunga River, Karnataka), and realizing that his
description of B. annandalei was based mainly
on immature specimens. This synonymy was
subsequently adopted by Menon (1987) in his review of the homalopterid
loaches of India, but without examining the type (or fresh topotypes) of B. annandalei, or the topotypes of B. australis. Later
workers followed this synonymy and considered Bhavania
to be monotypic (Talwar & Jhingran 1991;
Menon 1999; Kottelat 2012). ‘Bhavania
arunachalensis’, described by Nath et al. (2007)
from Naodhing drainage in Arunachal Pradesh, is
considered to be a ‘species inquirenda et incertae sedis’ (i.e., doubtful
identity and uncertain placement) (Kottelat 2012),
and is most likely a species of the genus Balitora
(see Fricke et al. 2020).
Given their hill-stream
adaptations (widespread paired fins, flattened ventral surfaces with body
suckers and rasping mouths on their ventral surface allowing them to firmly
grasp rock or gravel surfaces necessary in the mountain torrents) (Chen 1980; Kottelat 2012), and the fact that the type locality of B.
annandalei (Tenmalai)
and B. australis (Walayar)
are at least 300km apart and separated by two significant biogeographic
barriers - the Palghat Gap and the Shencottah Gap
(see Anoop et al. 2018), it is highly unlikely that the two are
conspecific. Collection of fresh topotypic specimens of both B. australis
and B. annandalei and detailed
examination and comparison of their biometrics, and genetic distance analysis
based on the mitochondrial cox1 gene, revealed that the two species are clearly
distinct. We, therefore, resurrect Bhavania annandalei
Hora, 1920, from the synonymy of B. australis
(Jerdon, 1849) and provide notes on the distribution
range of this species.
Six specimens of putative topotypic Bhavania annandalei were collected from Palaruvi
falls at Tenmala (Kallada
River), Kerala, and six specimens of putative topotypic
B. australis were collected from near
the Kavarakund falls, upstream of Malampuzha
Reservoir, Kerala, India (Fig. 1).
Samples were collected using a hand net/scoop net during early morning
hours, fixed in 10% formalin and transferred to 70% ethanol for permanent
voucher storage in the museum collections of the Kerala University of Fisheries
and Ocean Studies (KUFOS), Kochi, India. Gill tissues were obtained from fresh
specimens and preserved in absolute ethanol.
Morphometric measurements were taken for 37 characters (measured to the
nearest 0.1mm using digital calliper) and meristic values were determined for
10 characters using a stereo-zoom microscope.
For meristic counts, values in parenthesis after the count respresent its frequency.
For fin ray counts, unbranched fin rays are expressed as small roman
numerals. For pectoral fin, fin rays are
provided as padded fin rays + branched fin rays + unbranched fin rays. For statistical analysis of morphometric
data, subunits of body were taken as percentage of standard length and subunits
of head were taken as percentage of head length. Principal component analysis (PCA) was
performed to check whether the two species formed distinct clusters in
multivariate space using correlation matrix.
Null hypothesis that the clusters are not significantly different from
each other was tested using analysis of similarities (ANOSIM) employing
Euclidian distances and 9999 permutations.
Statistical analysis was performed in PAST 4.02 (Hammer et al.
2001). Genetic sequences of
mitochondrial partial cytochrome oxidase subunit 1 (cox1) of topotypic Bhavania annandalei and B. australis
were obtained from our ongoing study (Sidharthan et
al. Unpublished). Additional sequences
were downloaded from GenBank database.
Gene sequences were aligned using MUSCLE (Edgar 2004) and raw genetic
distance was estimated using MEGA 7 (Kumar et al. 2016). Data were partitioned into three codon
positions of cox1 gene. Partition
analysis (Chernomor et al. 2016) and ModelFinder (Kalyaanamoorthy et
al. 2017) were used to find the right partitioning scheme and nucleotide
substitution model for the partition scheme employing minimum Bayesian
information criterion (BIC). Maximum
likelihood (ML) analysis was performed in IQ Tree (Nguyen et al. 2015) with
best partition scheme and ultrafast bootstrap support for 1,000 iterations
(Hoang et al. 2018). Phylogenetic tree
was edited in FigTree v1.4.2 (Rambaut
2009).
Bhavania annandalei
Hora, 1920
(Images 1-–3)
Materials examined: KUFOS.19.AS.BH.02.1–6, 6
ex., 07.ii.2019, 8.945N & 77.158E, 32.7–37.6 mm SL, Palaruvi
falls, Tenmala, Kallada
River, Kerala, India, coll. Arya Sidharthan, E.S. Abhijith, & George Joseph.
Diagnosis. Bhavania
annandalei is distinguished from its only known
congener B. australis by a combination of
characters: low density and sparsely distributed tubercles on dorsal surface of
head, especially on operculum, (vs. high density of tubercles on dorsal surface
of head and operculum) (Image 3); gape of mouth comparatively farther from
snout tip, as a result the rostral barbels reaching
anterior border of upper lip, (vs. gape of mouth closer to snout tip, and
rostral barbels reaching posterior border of upper
lip) (Image 3); rostral flaps between the rostral barbels
fleshier (vs. less fleshier) (Image 3); fewer post-dorsal scales (34–36 vs.
38–41); fewer scales above the lateral line (11–12 vs. 14–15); and caudal
peduncle stout with its depth to width ratio 1.8–2.3 (vs. laterally compressed
caudal peduncle with depth to width ratio 2.8–3.6). This species has a fin formula of D. ii+7+i;
P. 6+10+i; V. ii+7–8; A. ii+5, and scale counts of Ll. 65–67 and L.tr.
11–12/9–10.
Description: Morphometric and meristic data
of Bhavania annandalei
are provided in Table 1 and Table 2, respectively. General body form as per Image 1a and Image
2a. Head details as in Image 3a, c.
Body elongate, dorso-ventrally depressed anteriorly, laterally compressed
posteriorly; dorsal profile convex, deepest at dorsal-fin origin. Body wider than its depth at dorsal-fin
origin, deeper than wide at anus. Head
small, rounded, less than one-fourth of standard length; depressed, longer than
broad, with minute sparsely distributed indistinct tubercles on dorsal surface
of head. Eyes small, dorso-laterally
positioned, not visible from underside of head. Snout pointed in lateral view,
round in dorsal view. Nostrils
positioned dorsally, closer to anterior border of eye than to snout tip,
anterior nostril situated inside a skin flap covering the posterior
nostrils. Mouth inferior. Lips fleshy.
Gape of mouth less than three times maximum head width. Barbels three
pairs, two rostral: outer rostral barbels shorter
than inner ones; one pair of maxillary barbels,
situated slightly anterior to the angle of mouth. Three fleshy rostral flaps interspaced
between rostral barbels. Gill opening small, restricted above the base
of the pectoral fin.
Body with scales except chest and
belly. Lateral line complete, with 68–72
small scales. Caudal peduncle slender,
its length almost three times its depth.
Dorsal-fin originating slightly behind the pelvic-fin origin, closer to
tip of snout than to caudal-fin base; with two unbranched, followed by seven
branched and a simple ray. Pectoral fin
elongated, longer than head, with six unbranched, followed by 10 branched and a
simple ray. Pelvic-fin length almost
equal to head length; fin origin closer to snout tip than to end of caudal
peduncle, its posterior end not reaching anus, with two unbranched and eight
branched rays. Anal fin with two
unbranched and five branched rays.
Caudal fin forked, with 19 principal rays.
Colouration:
In life (Image 1a), body is chestnut brown on dorsal and lateral sides, creamish-white on chest and belly; 3–4 prominent broad dark
brown ventral bands; two broad ventral bands on the dorsal-fin base. There are three black-coloured bands on the
dorsal fin, 6–7 bands on the pectoral, three bands on the pelvic, 1–2 bands on
the anal, and four bands across the caudal fin.
Morphometric analysis: In the morphometric analysis,
using all the size-adjusted characters (Table 1), the two species clustered
separately on the first two PCA axes (Fig. 2a).
The clusters were significantly different from each other (ANOSIM, 9999
permutations, R = 0.2315, P = 0.0271) indicating that the species formed
distinct clusters in multivariate space.
While length-length relationships for most characters showed similar
trends for both the species, there were two relationships that showed marked differences. Length-length relationship between caudal
peduncle depth and width (Fig. 2b) suggested that width increased rapidly with
increasing depth in the case of B. annandalei
compared to B. australis. Similarly, length-length relationship between
head length and head depth at nape (Fig. 2c) suggested that head depth
increased rapidly with increasing head length in the case of B. annandalei compared to B. australis.
Genetic analysis: Partition analysis and
model selection identified separate nucleotide substitution models for all
three codon positions, TNe+I for first codon, F81+F
for second codon, TN+F+G4 for third codon position of cox1 gene (BIC =
5570.419, lnL = -2633.96, df = 47). Maximum likelihood phylogenetic tree based on
best partition scheme and model selection (Fig. 3) recovered Bhavania annandalei
and B. australis as a clade sister to
Southeast Asian congeners of Balitoridae. Topotypic B.
annandalei (MT002520) differed from topotypic B. australis
(MT002518) with a raw genetic distance of 6.4% in the cox1 gene.
Distribution: Bhavania
annandalei is known with certainty from the Kallada, Vamanapuram, and Neyyar river systems in southern Kerala, India. These river systems drain the western slopes
of the Agasthyamalai Hill ranges, south of the Shencottah Gap. It
is highly likely that the species also occurs on the eastern slopes of the Agasthyamalai Hills particularly in the Tambaraparini
River system in Tamil Nadu, but detailed surveys and voucher specimens are
required to confirm this. In this
context, we believe that previous records of B. australis
from several tributaries of the Tambaraparini, Manimuthar, and Chittar draining
the eastern slopes of the Agasthyamalai (Johnson
& Arunachalam 2009), could most likely represent B. annandalei.
Remarks: The density of chromatophores
in Bhavania is likely to be dependent on the
micro-habitat as well as the colour and type of substratum it inhabits. Other ecological factors that may influence
body colour are forest/canopy cover, intensity of light, turbidity, water flow
and water temperature (V.K. Anoop pers. obs. 2018 and 2019). This is reflected in the different body
colours shown by the two species in different habitats and locations (see Image
1), an observation which was also made by Hora (1941).
Comparative material: Bhavania
australis, KUFOS.19.AS.BH.01.1–6, 6ex.,
13.iv.2019, 10.8636N & 76.6904E, 46.4–58.8 mm SL, near Kavarakund
falls, upstream of Malampuzha Reservoir, Kerala,
India, coll. M.R. Ramprasanth.
Table 1. Morphometric data of Bhavania annandalei
(KUFOS.19.AS.BH.02.1-6, n=6) and B. australis
(KUFOS.19.AS.BH.01.1-6, n=6) putative topotypes.
|
Characters |
Bhavania annandalei |
Bhavania australis |
||
|
Mean (sd) |
Range |
Mean (sd) |
Range |
|
|
Total length (mm) |
62.3 (18.2) |
40.2–85.8 |
76.1 (10.6) |
62.4–90.2 |
|
Standard length (SL, mm) |
50.8 (14.5) |
33.2–70.2 |
62.4 (8.8) |
51.6–74.2 |
|
Head length (HL, mm) |
11.1 (2.6) |
8.2–14.7 |
13.0 (1.7) |
11.3–15.2 |
|
% SL |
|
|
|
|
|
Head length |
22.2 (1.6) |
20.9–24.7 |
20.9 (0.9) |
19.8–21.9 |
|
Pre-pectoral length |
18.0 (0.9) |
17.2–19.7 |
18.2 (0.9) |
17.1–19.4 |
|
Pre-dorsal length |
49.8 (1.9) |
46.7–52.1 |
47.3 (1.8) |
44.5–49.4 |
|
Pre-pelvic length |
44.0 (1.1) |
42.4–45.1 |
44.3 (1.5) |
42.0–46.6 |
|
Pre-vent length |
70.3 (2.5) |
67.4–73.2 |
69.6 (1.6) |
67.7–71.6 |
|
Pre-anal fin length |
79.1 (1.8) |
77.1–81.6 |
78.7 (1.6) |
77.3–81.8 |
|
Origin of pelvic fin to anus
distance |
29.3 (1.5) |
26.6–30.7 |
28.7 (3.1) |
26.2–34.4 |
|
Anal fin to anus distance |
9.2 (1.3) |
7.6–11.0 |
9.9 (0.6) |
9.1–10.4 |
|
Post dorsal length |
44.3 (1.2) |
42.9–46.1 |
44.7 (1.0) |
43.6–46.2 |
|
Body depth at dorsal fin origin |
14.3 (0.8) |
12.9–15.2 |
13.2 (0.6) |
12.0–13.8 |
|
Body width dorsal fin origin |
17.7 (1.2) |
15.5–18.8 |
18.5 (0.7) |
17.4–19.3 |
|
Height of dorsal fin |
19.2 (1.2) |
16.9–20.0 |
20.1 (0.9) |
19.1–21.7 |
|
Dorsal-fin base length |
11.9 (0.7) |
11.1–12.7 |
12.0 (0.4) |
11.3–12.4 |
|
Body depth at anal fin origin |
11.4 (0.5) |
10.6–11.9 |
11.0 (0.5) |
10.4–11.9 |
|
Body width at anal fin origin |
7.0 (0.7) |
6.2–7.9 |
6.9 (0.6) |
5.9–7.6 |
|
Length of upper caudal lobe |
20.7 (1.4) |
18.4–22.0 |
20.6 (1.9) |
19.0–24.1 |
|
Length of lower caudal lobe |
24.3 (1.5) |
22.3–26.1 |
22.2 (1.4) |
20.8–24.7 |
|
Length of median caudal rays |
17.6 (1.4) |
15.2–19.2 |
16.2 (0.7) |
15.3–16.7 |
|
Anal fin length |
14.1 (0.9) |
12.8–15.5 |
15.5 (0.5) |
14.9–16.3 |
|
Anal fin base length |
7.0 (0.7) |
6.4–8.3 |
7.2 (0.5) |
6.6–7.9 |
|
Pelvic fin length |
22.5 (1.3) |
21.3–24.8 |
22.8 (0.8) |
21.6–23.9 |
|
Pectoral fin length |
26.8 (1.7) |
24.4–29.7 |
26.4 (1.2) |
24.2–27.3 |
|
Length of caudal peduncle |
13.3 (1.8) |
11.7–15.9 |
14.2 (0.7) |
13.1–15.1 |
|
Caudal peduncle depth |
9.3 (0.3) |
8.9–9.7 |
9.3 (0.8) |
8.4–10.7 |
|
Caudal peduncle width |
4.5 (0.5) |
3.9–5.1 |
3.0 (0.1) |
2.8–3.2 |
|
% HL |
|
|
|
|
|
Snout-supra-occipital distance |
93.3 (5.6) |
86.7–101.0 |
100.6 (5.4) |
94.2–107.5 |
|
Gape of mouth |
23.8 (3.3) |
19.7–26.9 |
29.6 (3.2) |
25.5–35.3 |
|
Head depth at eye |
41.2 (2.1) |
39.3–44.9 |
42.6 (3.2) |
37.9–45.9 |
|
Head width at eye |
75.3 (5.5) |
68.3–80.9 |
83.4 (6.5) |
75.7–93.0 |
|
Head depth at nape |
52.9 (5.3) |
47.3–60.3 |
41.8 (9.8) |
30.7–51.6 |
|
Snout length |
57.6 (5.3) |
51.4–64.9 |
58.6 (2.8) |
56.2–63.3 |
|
Maximum head width |
83.0 (9.0) |
71.1–95.8 |
88.9 (4.3) |
84.1–94.3 |
|
Eye diameter |
20.3 (2.8) |
16.5–23.7 |
17.5 (1.2) |
15.3–18.8 |
|
Interorbital width |
35.7 (4.8) |
30.6–42.9 |
39.0 (4.1) |
33.2–45.0 |
|
Internarial width |
27.3 (2.2) |
24.2–30.3 |
29.9 (2.6) |
26.1–33.7 |
Table 2. Meristic data of Bhavania australis
(KUFOS.19.AS.BH.01.1-6, n=6), and B. annandalei
(KUFOS.19.AS.BH.01.1-6, n=6) putative topotypes. Numbers in parenthesis indicate frequency of
character state in the materials examined.
|
Characters |
Bhavania annandalei |
Bhavania australis |
|
Dorsal-fin rays |
ii+7+i (6) |
ii+7 (3), ii+7+i (3) |
|
Pectoral-fin rays |
6+10+i (6) |
6+9+i (1), 6+10 (1), 6+10+i (4) |
|
Pelvic-fin rays |
ii+7 (2), ii+8 (4) |
ii+7 (4), ii+7+i (2) |
|
Anal-fin rays |
ii+5 (6) |
ii+5 (4), ii+5+i (1); ii+6 (1) |
|
Caudal-fin rays |
19 (6) |
19 (6) |
|
Lateral line scales |
65+4 (2), 66+3 (1), 67+3 (2),
67+3 (1) |
65+3 (2), 65+4 (1), 66+3 (1),
68+3 (1), 69+3 (1) |
|
Pre dorsal scales |
29 (1), 30 (2), 31 (3) |
28 (3), 29 (2), 30 (1) |
|
Post dorsal scales |
34 (3), 35 (2), 36 (1) |
38 (1), 39 (2), 40 (2), 41 (1) |
|
Scales above lateral line |
11 (2), 12 (4) |
14 (4), 15 (2) |
|
Scales below lateral line |
9 (2), 10 (4) |
10 (3), 11 (3) |
For
figures & images - - click here
References
Anoop, V.K., N. Dahanukar., S. Philip., L. Thomas & R. Raghavan (2018). Phylogeny of the hillstream loach genus Mesonoemacheilus
reveals widespread diversification through ancient drainage connections in the
Western Ghats Biodiversity Hotspot. Molecular Phylogenetics and Evolution
129: 77–84. https://doi.org/10.11646/zootaxa.4683.1.8
Chen, Y.Y. (1980). Systematic studies on the fishes
of the family Homalopteridae of China III. Phyletic
studies of the Homalopterid fishes. Acta Zootaxonomica Sinica 5:
200–211.
Chernomor, O., A. von Haeseler & B.Q. Minh (2016). Terrace aware data structure for
phylogenomic inference from supermatrices. Systematic
Biology 65: 997–1008. https://doi.org/10.1093/sysbio/syw037
Edgar, R.C. (2004). MUSCLE: multiple sequence
alignment with high accuracy and high throughput. Nucleic Acid Research
32: 1792–1797. https://doi.org/10.1093/nar/gkh340
Fricke, R., W.N. Eschmeyer & R. Van der Laan (eds) (2020). Eschmeyer’s Catalog of Fishes: Genera, Species, References. Electronic
version accessed 10 June 2020. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp
Hammer, Ø., D.A.T. Harper &
P.D. Ryan (2001). PAST: Paleontological statistics software package for education and
data analysis. Palaeontologia Electronica
4(1): 1–9.
Hoang, D.T., O. Chernomor, A. Von Haeseler, B.Q.
Minh & L.S. Vinh (2018). UFBoot2: improving the ultrafast bootstrap approximation. Molecular
Biology and Evolution 35: 518–522. https://doi.org/10.1093/molbev/msx281
Hora, S.L. (1920). Revision of the Indian Homalopteridae and of the genus Psilorhynchus
(Cyprinidae). Records of the Indian Museum 19(5):
195–215.
Hora, S.L. (1937). Notes on fishes in the Indian
Museum. XXVIII. On three collections of fish from Mysore and Coorg, south India.
Records of the Indian Museum 39(1): 5–28.
Hora, S.L. (1941). Homalopterid
fishes from peninsular India. Records of the Indian Museum 43(2):
221–232.
Johnson, J.A. & M.
Arunachalam (2009). Diversity, distribution and assemblage structure of fishes in streams of
southern Western Ghats, India. Journal of Threatened Taxa 1(10):
507–513. https://doi.org/10.11609/JoTT.o2146.507-13
Kalyaanamoorthy S., B.Q. Minh, T.K.F. Wong, A. von Haeseler & L.S. Jermiin
(2017). ModelFinder: fast model selection for accurate phylogenetic
estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285
Kottelat, M. (2012). Conspectus Cobitidum: an inventory of the
loaches of the world (Teleostei: Cypriniformes:
Cobitoidei). Raffles Bulletin of Zoology 26:
1–199.
Kumar, S., G. Stecher
& K. Tamura (2016). MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger
datasets. Molecular Biology and Evolution 33: 1870–1874. https://doi.org/10.1093/molbev/msw054
Menon, A.G.K. (1987). The fauna of India and the adjacent
countries. Pisces. Vol. IV. Teleostei - Cobitoidea. Part 1. Homalopteridae.
Zoological Survey of India, Calcutta, 259pp.
Menon, A.G.K. (1999). Check list - Fresh water fishes
of India. Records of the Zoological Survey of India, Occasional Paper
No. 175: 1–366.
Nath, P., D. Dam., P.T. Bhutia., S.C. Dey & D.N. Das
(2007) A new fish
species of the genus Bhavania (Homalopteridae: Homalopterinae)
from river Noadhing drainage, Arunachal Pradesh,
India. Records of the Zoological Survey of India 107(3): 71–78.
Nguyen, L.T., H.A. Schmidt, A.
von Haeseler & B.Q. Minh (2015). IQ-TREE: A fast and effective
stochastic algorithm for estimating maximum likelihood phylogenies. Molecular
Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300
Rambaut, A. (2009). FigTree, ver
1.4.3. Available online at: http://tree.bio.ed.ac.uk/software/figtree
Talwar, P.K. & A.G. Jhingran (1991). Inland Fishes of India and
Adjacent Countries. In 2 vols. Oxford & IBH Publishing Co., New Delhi, Bombay,
Calcutta, 1158pp.