Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 March 2023 | 15(3): 22834–22840
ISSN 0974-7907
(Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.6894.15.3.22834-22840
#6894 | Received 16
November 2020 | Final received 02 December 2022 | Finally accepted 01 February
2023
Genetic evidence on the
occurrence of Channa harcourtbutleri
(Annandale, 1918) in Eastern Ghats, India: first report from mainland India
Boni Amin Laskar
1 , Harikumar Adimalla
2, Shantanu Kundu 3, Deepa Jaiswal 4 & Kailash Chandra 5
1 High Altitude Regional Centre of
Zoological Survey of India, Solan, Himachal Pradesh
173211, India.
1,4 Freshwater Biology Regional
Centre, Zoological Survey of India, Attapur,
Hyderabad, Telangana 500032, India.
2 House No. 2-60, Village Turkapalle, Nalgonda, Telangana 508266, India.
3,5 Zoological Survey of India, New
Alipore, Kolkata, West Bengal 700052, India.
1 boniamin.laskar@gmail.com
(corresponding author), 2 harikumaradimalla92@gmail.com, 3 shantanu1984@gmail.com,
4 deepajzsi@gmail.com, 5 kailash616@gmail.com
Abstract: Channa harcourtbutleri (Annandale) was described from Inle Lake (Southern Shan State) in Myanmar, and is
currently considered as a valid species in the Channa
gachua species-group. Notwithstanding several
detailed studies on Channa from India
in the recent, none has mentioned the occurrence of C. harcourtbutleri in the Indian mainland. In
continuation to the faunal diversity exploration in Eastern Ghats, India, a few
specimens in the C. gachua species-group were
collected from the river Sabri sub-basin of the river Godavari basin in the
East Godavari District of Andhra Pradesh which was identified as C. harcourtbutleri through DNA barcoding. This is a first
report on occurrence of the species in the wild in the Eastern Ghats, India.
Keywords: Channidae, DNA barcoding, Godavari basin, Inle Lake, phylogeny, taxonomy.
Editor: Mandar Paingankar, Government Science College Gadchiroli,
Maharashtra, India. Date of
publication: 26 March 2023 (online & print)
Citation: Laskar, B.A., H. Adimalla, S. Kundu, D. Jaiswal & K. Chandra (2023). Genetic evidence on the occurrence of
Channa harcourtbutleri
(Annandale, 1918) in Eastern Ghats, India: first report from mainland India. Journal of Threatened Taxa 15(3): 22834–22840. https://doi.org/10.11609/jott.6894.15.3.22834-22840
Copyright: © Laskar et al. 2023. 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: The research is funded by the Core Funding of Zoological Survey of India (ZSI), Kolkata, Ministry of
Environment, Forest and Climate Change (MoEF&CC), New Delhi.
Competing interests: The authors declare no competing interests.
Author details: Boni Amin Laskar is currently working as scientist-E in High Altitude Regional Centre of Zoological Survey of India, Solan. His field of research is molecular studies and taxonomy of freshwater fishes. He has over 20 years of research experience including field surveys throughout various biogeographic zones in India. Harikumar Adimalla is a
budding researcher in the field of molecular studies of freshwater fishes from Deccan peninsular biogeographic zone.
Shantanu Kundu is a molecular biologist with over 15 years of experience including field surveys and molecular studies of Indian fauna, especially the Himalayan and northeastern Indian region. He is currently working as a post-doctoral fellow in the Department of Marine Biology, Pukyong National University, Busan, South Korea.
Deepa Jaiswal is working as scientist-E in Freshwater Biology Regional Centre of Zoological Survey of India, Hyderabad. Her field of specialization is taxonomy of aquatic Insects. Kailash Chandra is the former director of Zoological Survey of India. He is a renowned taxonomist in India and is a recipient of E.K. Janaki Ammal National Award on taxonomy.
Author contributions: BAL & HA did field surveys and collected the specimens. BAL studied the morphology and meristics of the specimens for taxonomic identification. HA generated the DNA data. BAL & SK did the molecular analysis. BAL, SK & DJ wrote the article. BAL & KC reviewed the article.
Acknowledgements: We are grateful to the
director, Zoological Survey of India, Kolkata, for their continued support and
enthusiasm, and for providing necessary facilities throughout the study. We
further express our gratitude to the officer-in-charge, Freshwater Biology Regional
Centre, Zoological Survey of India, Hyderabad, for providing necessary
facilities to carry out this study. The third author (SK) acknowledges the
fellowship grant received from the Council of Scientific and Industrial
Research (CSIR) Senior Research Associateship (Scientists’ Pool Scheme) Pool
No. 9072-A.
Introduction
Snakehead fishes belong to the
family Channidae, are most popular in ornamental fish
trade. The species in the genus Channa Scopoli, 1777 are currently distributed in southern,
eastern, and southeastern Asia while their allied species in the genus Parachanna Teugels
& Daget, 1984 are endemic in Africa. Snakehead
fishes are broadly categorized into two major groups based on the presence of
gular scales. The first group having gular scales comprises of all the African
species of the genus Parachanna and
four Asian species of the genus Channa while
the second group lacking gular scales comprises of all the rest of the channid
species (Musikasinthorn & Taki 2001; Zhang et al.
2002). Further, the Channa gachua species group (sensu
Britz 2008) was characterized by a varying number
of dark and light semi-circular bands on the pectoral-fin, which has been
subsequently followed by other ichthyologists (Britz
et al. 2019; Praveenraj et al. 2019). The presence of
gular scales is regarded as a plesiomorphic state within the Channidae (Li et al. 2006). As mentioned in Li et al.
(2006), the African taxa of Parachanna differ
from the Asian taxa of Channa by the absence
of supporting ‘lamellas’ or process of the first epibranchial and hyomandibular
in the suprabranchial organs (Senna 1924; Bonou & Teugels 1985).
Further, the study has shown that the species lacking pelvic fins clade with
the species having the pelvic fins, and has explained that the loss of the
pelvic fins occurred at least three times independently during the evolution of
the taxa in Channidae (Bonou
& Teugels 1985). The gular region with scales or
without scale is one of the key characters in channid taxonomy (Talwar & Jhingran 1991; Li et al. 2006), and the shape of isthmus
was also shown to be a key feature in differentiating marulius
and gachua species-groups (Vishwanath & Geetakumari 2009). Among the currently reported 24 channid
species in India, 18 species (15 with pelvic fins and 3 without pelvic fins)
are included in the C. gachua
species-group (sensu Britz
2008). Britz et al. (2019) mentioned that snakehead
fishes have a centre of diversity in the eastern part
of the Himalaya Biodiversity Hotspot (Conte-Grand et al. 2017; Rüber et al. 2019). In the recent two decades, quite a good
number of new species, mostly in the C. gachua
species-group, have been described. However, a few of the recent
descriptions have been retained in synonymy by Britz
et al. (2019).
Channa harcourtbutleri (Annandale) was described from Inle Lake (southern Shan State) in Myanmar (Annandale,
1918), but it was placed in synonymy with C. gachua
(Hamilton) by Hora & Mukerji (1934). However, since the latter species is
younger than the former, this synonymy at its first instance appears
incorrect. Ng et al. (1999) resurrected
the species and discussed the differences between them. The taxonomy of C.
gachua has been a complex problem (Ng et al.
1999), but the recent phylogenetic study suggested two distinct lineages within
the C. gachua species-complex
(Conte-Grand et al. 2017). The true C. gachua
as referred in Conte-Grand et al. (2017) is restricted to the area west of the
Indo-Burman ranges (i.e., Rakhine Yoma and Chin
Hills) and covers Sri Lanka, India, Nepal, Bangladesh, and the Rakhine area of
Myanmar. The taxon previously recorded as C. gachua
from Sri Lanka has been revalidated as C. kelaartii
(Gunther), and has its population also distributed in southern peninsular India
(Sudasinghe et al. 2020). The eastern lineage of the C.
gachua species-complex, nominally referred to
as C. limbata, is distributed to the
east of the Indo-Burman ranges from Myanmar reaching east to Vietnam and
southern China and south to Indonesia and Malaysia (Conte-Grand et al. 2017).
Notwithstanding several detailed studies on channid taxa from India recently, none
has mentioned the occurrence of C. harcourtbutleri
in mainland India (Conte-grand et al. 2017; Britz
et al. 2019; Sudasinghe et al. 2020).
In continuation to the faunal
diversity exploration in Eastern Ghats, India, several specimens of the genus Channa were collected from various localities. Among
the examined specimens, a few specimens collected from the river Sabri
sub-basin of the river Godavari basin in the East Godavari District of Andhra
Pradesh, were morphologically identified as similar to C. harcourtbutleri and were confirmed through DNA
barcoding. C. harcourtbutleri is a Burmese
species, hitherto not recorded from mainland India.
Materials
and Methods
The study incorporates several
specimens of the genus Channa from various
localities within India. However, this study is specifically aimed to resolve
the identity of the specimens in the Channa
gachua species-group collected from the northern
Eastern Ghats, within a range of around 30 km to the north-east of Papikonda National Park.
DNA isolation, PCR and DNA
sequence
DNA isolation followed basic
methods after partial modification (Sambrook & Russell 2001; Laskar et al. 2018). Partial segment of the
mitochondrial cytochrome oxidase C subunit I (COI) gene was amplified using the
primer pairs FishF1-FishR1 (Ward et al. 2005). A total of 28 COI sequences for
nine channid species from India were generated in this study. All the examined
specimens were registered in the National Zoological Collections of Zoological
Survey of India (Freshwater Biology Regional Centre), Hyderabad, and the COI
sequences generated in this study were submitted to NCBI GenBank and BOLD. The
accession numbers are given in the material examined section as well as in the
phylogenetic tree and are marked by orange stars. We also retrieved COI
sequences from NCBI, and BOLD. Representative sequences for 22 clearly defined
channid taxa from India as referred in Conte-Grand et al. (2017) were
retrieved from databases. Further, the sequences of a few recently described
species whose accessions are referred by their original authors were retrieved
from NCBI. The dataset of 157 COI sequences included an outgroup Parachanna obscura (MK074551). Genetic
divergence analyses and the neighbor-joining phylogenetic tree visualization
were performed in MEGA7.0 (Kumar et al. 2016). Bayesian inferences were drawn
in Mr. Bayes (Ronquist & Huelsenbeck
2003) and the tree topology was developed in iTOL (Letunic & Bork 2007). We used the Kimura 2-parameter
model, mostly applied in DNA barcoding studies (www.bold.org), to calculate the
mean genetic distance between the groups. The study is limited by the lack of
COI sequence of a few of the recent species, like, C. brahmacharyi, C. pomanensis.
Further, we limit to discuss only the conspecific status of the COI sequences
generated in the study, C. harcourtbutleri
in particular.
Materials examined
C. gachua
(Hamilton, 1822): FBRC/ZSI/F1979, 1, Himayat Sagar, Telangana,
17.35N & 78.42E, GenBank accession:
KM272635; FBRC/ZSI/F/3317, 1, Small stream at Basavagu
village, Andhra Pradesh, 17.70N 81.02E, GenBank accession: MT118102;
FBRC/ZSI/F/3450, 6, Stream at Mothugudem, Andhra
Pradesh, 17.80N & 81.64E, GenBank accessions: MW002473, MW002474, MW002475;
FBRC/ZSI/F/2662, 1, Manjeera Dam, Telangana, 17.692N
& 78.171E, GenBank accession: MH795975; FBRC/ZSI/F/3628, 1, Gubbagurthi near Wyara Lake,
Telangana, 17.27N & 80.37E, GenBank accession: MW002494.
C. harcourtbutleri
(Annandale, 1918): FBRC/ZSI/F/3393, 3, Papikonda
National Park at G. M. Valasa Road, Andhra Pradesh,
17.59N & 81.68E, GenBank accession: MW002468; FBRC/ZSI/F/3615, 4, 62.0-83.0
mm SL, Stream at Mothugudem-Donkarayi Road, E.
Godavari, Andhra Pradesh, 17.84N & 81.67E, GenBank accession: MW002479;
FBRC/ZSI/F/3630, 2, 72.0–78.0 mm SL, River Pamuleru at
Egavalasa village, Andhra Pradesh, 17.7N &
81.78E, GenBank accession: MW002470 (Image 1).
C. kelaartii
Gunther,1868: FBRC/ZSI/F/3124, 13, Nilavoor Lake,
Tamil Nadu, 12.56N & 78.64E, GenBank accessions: MT720842, MT720843,
MT720844, MT720845, MT720846, MT720847; FBRC/ZSI/UN9604/DNA582, 1, Small pond
at Valvanthinadu village, Namakkal
District, Tamil Nadu, 11.280N & 78.364E, GenBank accession: MN685707.
C. punctata
(Bloch, 1973): FBRC/ZSI/F/2405, 1, Kaddem Dam,
Telangana, 9.026N & 76.385E, GenBank accession: MF601323; FBRC/ZSI/F/2717,
1, Maharashtra, 20.450N & 74.403E,
GenBank accession: MH795988; FBRC/ZSI/F/3627, 1, Gubbagurthi
near Wyara Lake, Telangana, 17.27N & 80.37E,
GenBank accession: MW002493; FBRC/ZSI/F/3421, 1, Small stream near Basavagu Village, Andhra Pradesh, 17.70N & 81.02E,
GenBank accession: MT654658.
C. striata
(Bloch,1793): FBRC/ZSI/DNA357, 1, Namsai, Assam,
27.57N & 95.39E, GenBank accession: MK681748.
C. marulius
(Hamilton, 1822): FBRC/ZSI/F2337, 1, Singur Dam,
Telangana, 17.802N & 77.892E, GenBank accession: KY694512; FBRC/ZSI/F/2715,
1, Maharashtra, 20.450N &
74.403E, GenBank accession: MH795986; FBRC/ZSI/DNA267, 1, Tungabhadra River,
Andhra Pradesh, 16.169N & 77.934E, GenBank accession: MK336898.
C. bleheri
Vierke, 1991: FBRC ZSI DNA354, 1, Tinsukia,
Assam, 27.57N & 95.39E, GenBank accession: MK632315.
C. aurantimaculata
Musikasinthorn, 2000: FBRC/ZSI/DNA359, 1,
Tinsukia, Assam, 27.57N &
95.39E, GenBank accession: MK632318.
C. stewartii
(Playfair, 1867): FBRC/ZSI/DNA356, 1, Tinsukia, Assam, 27.57N & 95.39E, GenBank accession:
MK632316.
Results
The phylogenetic tree shows
distinctive cladding of the Asian channid taxa with reference to the African Parachanna used herein to root as out-group. The
generated sequences of the C. gachua
species-group from the Yelagiri Hills in Tamil Nadu
form a distinct clade that comprises of a few database sequences that were
referred in a recent study as C. kelaartii.
Thus, this study contributes further specimens of C. kelaartii
from the southern Eastern Ghats. The sequences of the C. gachua like specimens collected from near to
the type locality of Ophicephalus marginatus and various other localities in the Godavari
River basin are nested as a subclade to the topotypic
C. gachua. Our generated sequences of
other congeners like C. marulius, C.
punctata, C. stewartii,
C. bleheri, C. aurantimaculata, and C. striata
show distinct cladding and each comprises of conspecific database sequences
wherever available. The sequence of C. pomanensis
(referred in Praveenraj et al. 2019) was found to be
cohesively claded with a sequence of C. melanostigma. Hence, in the absence of further
specimens for both the taxa, the identity of the two sequences is not confirmed
and remained beyond the limit of this study. Our sequences of C. stewartii are nested in the 65th clade of
C. stewartii of Conte-Grand et al.
(2017) (BIN:AAF3764) while status of the taxonomic assignment of the 64th
clade of C. stewartii of Conte-Grand et
al. (2017) (BIN:AAF3772) is also beyond the limit of this study. However, the
generated sequences of the specimens in C. gachua
species-group from near the Papikonda National Park,
East Godavari District, in the Eastern Ghats, are cohesively claded with the two database sequences from southern India
of Conte-Grand et al. (2017) (MF462283 BOLD:ADL6569, MF462269) and formed a
sub-clade to the species C. harcourtbutleri
from Myanmar (BIN:AAC3926) suggesting their close genetic similarity with the
latter, hence referred hereafter as C. harcourtbutleri
India.
The present dataset is comprised
of representative sequences from the clearly defined taxa. However, in order to
estimate the range of intra-species genetic divergence, the sequences are
grouped based on the clustering inferred from neighbor-joining phylogeny
(Figure 1). Sequences in the cluster with topotypic C.
gachua (referred in Britz
et al. 2019) are named as C. gachua Topotypic. The sequences of C. gachua like specimens from near to the type locality of
O. marginatus are named as C. gachua Godavari. Similarly, the sequences of C. royi from Andaman Islands (Praveenraj et al. 2019) are tentatively named as C. royi, sequences in the cluster with C. harcourtbutleri BIN:AAC3926 from Myanmar
(Conte-Grand et al. 2017) are named as C.
harcourtbutleri Myanmar, and the
sequences of C. gachua species-group
from near the Papikonda National Park, East Godavari
District, along with other database sequences in the same cluster are named as
C. harcourtbutleri India. The
overall genetic divergence between groups in the dataset is lying in the range
from 1.4–25.8 % (Table 1). The divergence matrix revealed that the sequences of
C. gachua Godavari are
genetically diverged by 3.2% K2P distance from C. gachua
Topotypic, and maintained 9.4–22.8 % K2P distance
from all the congeners. C. pseudomarulius
is diverged by 4.3% K2P from C. marulius,
C. pardalis is diverged by 5.8% from C.
bipuli, C. melanostigma
by 6.3% from C. aurantimaculata, and so
on. Similarly, the divergence matrix revealed that the sequences of C. harcourtbutleri India are genetically
diverged by 1.4% K2P distance from C. harcourtbutleri
Myanmar (BIN: AAC3926), 2.3% K2P distance from C. royi
Andaman Islands, and maintained 13.4–24.3 % K2P distance from all the
congeners.
Discussion
Conte-Grand et al. (2017)
recovered a total number of 90 BINs in their dataset having a total number of
38 valid species at time, and inferred higher species diversity in snakeheads.
However, they neither included any COI sequence data of C. limbata in the phylogenetic analysis nor
assigned any BIN for the species. As of now, a search for BINs with the name ‘Channa’ in BOLD yielded a record of 93 BINs.
Conte-Grand et al. (2017) had an extensive dataset covering various
geographical areas and populations. In fact, the representative sequences for
almost all the species described or validated after 2017 were present either as
an unnamed clade (potential new BIN in Conte-Grand et al. 2017) or has been
assigned with BOLD BIN.
The taxonomic history of the
snakehead fishes finds two descriptions, O. marginatus
and O. coramota, with their type
locality in Vizagapattam (=Visakhapatnam, Andhra
Pradesh), part of the Eastern Ghats region. However, both these species have
been synonymized with C. gachua
(Roberts 1993; Ng et al. 1999; Courtenay & Williams 2004; Kottelat 2013). In fact, Britz et
al. (2019) examined specimens of topotypic O. marginatus, and found a very similar colour pattern as well as a very little genetic difference
with topotypic C. gachua
(2.4% uncorrected p-distance). Britz et al. (2019)
pointed out several flaws in the description of C. shingon
by Endruweit (2017) and provided various valid
reasons to consider C. shingon as a
junior synonym of C. harcourtbutleri. Britz et al. (2019) was also not convinced enough by the
morphological descriptions to consider C. royi
as a distinct species from C. harcourtbutleri,
and a very low genetic distance of 2.4–2.8 % uncorrected p-distance was stated
to be in the range of intra-species variation, and therefore considered Andaman
C. royi as a junior synonym of C.
harcourtbutleri. However, the distribution
limit of C. harcourtbutleri and C.
limbata is not yet clear. Conte-grand et al.
(2017) mentioned an unexpected placement of two specimens from southern
peninsular India in the middle of the eastern lineage of the C. gachua species-complex. The same statement was
repeated in Ruber et al. (2019). Conte-grand et al.
(2017) showed that the two specimens from southern peninsular India (one from Chunchi falls, Cauvery River, 12.351N & 77.443E; and
the other from Kali River, 15.381N & 74.403E) were included in a Putative
BIN:ACM5826 new that claded away from the BIN of C.
harcourtbutleri (AAC3926).
Following the previous studies,
it may be figured out that the nominal species in the genus Channa
with fewer or no morphological differences can have intra-species genetic
divergence as high as 2.2–2.4 %, and the nominal taxa falling within such range
of genetic divergence could be considered as a single species. Therefore, a
clear understanding of the range of intra-species genetic divergence would be
helpful in taxonomic assignment of the channid taxa. Based on the analysis of
COI barcode sequences, we confirm that the specimens of C. gachua species-group from East Godavari District,
Eastern Ghats, India, along with the sequences from southern India, are
actually a single species which may be named as C. harcourtbutleri
because of low genetic divergence with the conspecific sequences from Myanmar.
Thus, this study claims the presence of C. harcourtbutleri
in the wild in Eastern Ghats region, in mainland India. Nonetheless, C. harcourtbutleri has already been recorded to
be distributed in Andaman Islands through the synonymization
of C. royi with C. harcourtbutleri. Hence, this study reports for the
first time the occurrence of C. harcourtbutleri
in mainland India (Image 2). Unexpectedly, no specimens from northeastern India
are available to place in the clade of C. harcourtbutleri.
In this background, the distribution of C. harcourtbutleri
is appearing disjunct.
Table 1. Estimated genetic divergence (% K2P) between
the studied groups in the genus Channa. The nalysis reveals a low K2P genetic divergence among the
specimens of C. royi Andaman Islands, C. harcourtbutleri Myanmar and C. harcourtbutleri
India, suggesting their conspecific status.
|
Grouped taxa |
Within- group (K2P %) |
Between-group (K2P %) |
|||||||||||||||||||||||
|
|
|
1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
12 |
13 |
14 |
15 |
16 |
17 |
18 |
19 |
20 |
21 |
22 |
23 |
24 |
|
C. harcourtbutleri
Myanmar |
0.013 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. harcourtbutleri
India |
0.002 |
2.3 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. royi
Andaman Islands |
0.003 |
2.3 |
1.4 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. gachua
Godavari |
0.002 |
17.3 |
17.4 |
16.7 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. gachua
Topotypic |
0.006 |
19.3 |
19.5 |
19.3 |
3.2 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. andrao |
0.000 |
16.0 |
14.7 |
15.7 |
18.6 |
19.5 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. aurantimaculata |
0.001 |
12.7 |
13.4 |
13.1 |
15.4 |
16.9 |
15.7 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. aurantipectoralis |
0.001 |
16.1 |
16.0 |
16.9 |
17.2 |
16.9 |
18.3 |
14.9 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. barca |
0.000 |
16.9 |
16.3 |
17.5 |
15.2 |
16.5 |
15.4 |
8.8 |
15.4 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. bipuli |
n/c |
17.7 |
17.8 |
18.8 |
15.3 |
18.2 |
16.1 |
12.3 |
18.2 |
12.2 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. bleheri |
0.001 |
17.4 |
16.8 |
16.8 |
15.4 |
16.9 |
16.7 |
14.8 |
17.0 |
14.0 |
12.4 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. brunnea |
0.000 |
16.1 |
16.1 |
16.4 |
16.4 |
17.2 |
17.2 |
14.5 |
17.5 |
14.9 |
12.3 |
9.7 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
C. diplogramma |
0.004 |
22.8 |
23.3 |
24.1 |
22.7 |
22.0 |
22.3 |
22.0 |
23.0 |
23.1 |
25.8 |
23.9 |
25.0 |
|
|
|
|
|
|
|
|
|
|
|
|
|
C. kelaartii |
0.007 |
16.3 |
15.5 |
15.8 |
10.5 |
9.9 |
19.6 |
16.3 |
16.4 |
17.7 |
16.5 |
16.2 |
16.5 |
22.1 |
|
|
|
|
|
|
|
|
|
|
|
|
C. lipor |
n/c |
15.1 |
15.6 |
15.6 |
16.0 |
14.4 |
17.2 |
14.5 |
11.9 |
15.7 |
21.4 |
16.7 |
17.7 |
22.6 |
15.5 |
|
|
|
|
|
|
|
|
|
|
|
C. marulius |
0.004 |
24.6 |
24.3 |
24.7 |
21.0 |
24.7 |
19.5 |
21.1 |
21.1 |
20.3 |
18.6 |
18.4 |
21.7 |
19.1 |
21.2 |
23.3 |
|
|
|
|
|
|
|
|
|
|
C. melanostigma |
0.008 |
14.5 |
15.5 |
15.4 |
16.3 |
17.8 |
14.6 |
6.3 |
18.8 |
10.3 |
11.7 |
14.7 |
13.4 |
23.6 |
15.5 |
15.7 |
21.4 |
|
|
|
|
|
|
|
|
|
C. pardalis |
0.002 |
16.2 |
16.5 |
17.2 |
14.8 |
16.2 |
14.7 |
11.2 |
16.4 |
10.9 |
5.8 |
12.0 |
12.2 |
23.1 |
13.8 |
17.2 |
18.2 |
12.9 |
|
|
|
|
|
|
|
|
C. pseudomarulius |
0.000 |
23.3 |
22.2 |
23.1 |
22.8 |
25.0 |
18.7 |
19.8 |
20.5 |
19.8 |
19.2 |
19.2 |
23.1 |
16.9 |
20.7 |
24.2 |
4.3 |
21.0 |
18.0 |
|
|
|
|
|
|
|
C. punctata |
0.015 |
21.4 |
20.6 |
19.9 |
19.3 |
19.9 |
19.1 |
19.0 |
20.1 |
20.0 |
17.6 |
18.2 |
20.3 |
19.6 |
17.9 |
21.5 |
16.1 |
16.8 |
19.5 |
15.4 |
|
|
|
|
|
|
C. quinquefasciata |
0.008 |
14.9 |
14.3 |
15.3 |
16.8 |
17.2 |
15.9 |
12.6 |
16.0 |
13.3 |
15.0 |
15.7 |
14.4 |
22.2 |
15.7 |
15.4 |
19.2 |
12.3 |
13.7 |
19.1 |
19.3 |
|
|
|
|
|
C. rara |
0.002 |
16.1 |
16.6 |
15.7 |
9.4 |
10.8 |
19.2 |
16.3 |
15.6 |
18.0 |
18.8 |
17.3 |
18.3 |
22.1 |
8.8 |
14.5 |
25.4 |
17.1 |
15.2 |
25.0 |
19.7 |
18.4 |
|
|
|
|
C. stewartii |
0.005 |
17.1 |
16.6 |
17.7 |
17.9 |
19.0 |
16.6 |
12.6 |
15.8 |
13.8 |
15.8 |
16.4 |
12.8 |
21.6 |
15.7 |
16.3 |
20.7 |
12.3 |
12.2 |
19.3 |
19.1 |
8.3 |
19.1 |
|
|
|
C. stiktos |
0.013 |
23.6 |
22.3 |
22.2 |
21.3 |
21.2 |
22.8 |
21.3 |
20.2 |
19.6 |
18.7 |
17.7 |
19.9 |
24.2 |
20.1 |
20.7 |
21.4 |
20.6 |
18.6 |
22.5 |
23.0 |
21.3 |
22.1 |
18.4 |
|
|
C. striata |
0.001 |
21.1 |
19.8 |
18.4 |
22.0 |
23.2 |
19.3 |
21.0 |
24.0 |
21.2 |
17.0 |
22.2 |
23.8 |
20.3 |
20.6 |
23.0 |
18.1 |
18.7 |
19.4 |
17.5 |
16.7 |
21.7 |
22.1 |
22.0 |
23.7 |
For
figure & images - - click here for full PDF
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