Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 April 2020 | 12(5): 15612–15618
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.5801.12.5.15612-15618
#5801 | Received 24 February 2020 | Finally
accepted 30 March 2020
Varying colour pattern, yet
genetically similar: Pebble Crab Seulocia vittata (Stimpson, 1858) (Brachyura:
Leucosiidae) from the southeastern
coast of India
Sanjeevi Prakash 1 & Amit Kumar 2
1,2 Centre for Climate Change
Studies, Sathyabama Institute of Science and
Technology, Jeppiaar Nagar, Rajiv Gandhi Salai, Chennai,
Tamil Nadu 600119, India.
1,2 Sathyabama Marine Research Station, 123 Sallimalai Street, Rameswaram, Tamil Nadu 623526, India.
1 prakash.cccs@sathyabama.ac.in
(corresponding author), 2 amit.kumar.szn@gmail.com
Editor: Mandar Paingankar, Government Science College Gadchiroli,
Maharashtra, India. Date
of publication: 26 April 2020 (online & print)
Citation:
Prakash, S. & A. Kumar (2020). Varying colour pattern, yet genetically similar:
Pebble Crab Seulocia vittata
(Stimpson, 1858) (Brachyura: Leucosiidae)
from the southeastern coast of India. Journal of Threatened Taxa 12(5): 15612–15618. https://doi.org/10.11609/jott.5801.12.5.15612-15618
Copyright: © Prakash & Kumar 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: Systematics
Research Fund (SRF) by the Linnaean Society and the Systematics Association, UK awarded to SP.
Competing interests: The authors declare no competing interests.
Author details: Sanjeevi Prakash is a marine biologist, specialized with the taxonomy
of coral-reef dwelling decapod crustaceans. Currently, he is focusing on the
integrative approach such as morphology and molecular taxonomy to understand
the cryptic species diversity in the coral reef associated caridean shrimps of
the world. Amit Kumar is a marine biologist. He has been using
molecular markers to understand marine biodiversity. He is also interested in
studying how marine organisms respond to predicted climate change stress
conditions.
Author contribution: SP and AK designed and performed the study. SP
identified the species using morphology. AK did the phylogeny. SP and AK
interpreted the results and wrote the manuscript.
Acknowledgements:
The authors would like to thank
the management, Sathyabama Institute of Science and
Technology for establishing Sathyabama Marine
Research Centre at Rameswaram and providing necessary facilities to carry out
research work. This work was partially
supported by the Systematics Research Fund (SRF) by the Linnean
Society and the Systematics Association, UK awarded to S.P. Authors thank P.M. Vatsala
for correcting the English language of the manuscript. We also thank the anonymous
reviewers and the editor for constructive criticism which improved the final
version of the manuscript.
Abstract: Five adult specimens of leucosiid crab Seulocia
vittata (Stimpson, 1858) were recently collected
off the coast of Palk Bay, southern India. Typical morphological examination revealed the
presence of two colour patterns: grey and red.
Interestingly, molecular analysis based on the barcoding gene cytochrome
oxidase sub unit I (COI) revealed that both grey and red colour patterns in S.
vittata showed 0% sequence divergence between the
specimens. This indicates a situation of
reverse cryptic behavior in this crab. Surprisingly, the evolutionary and ecological
processes leading to the absence of genetic divergence and variation in
morphology (colour pattern) in S. vittata
complex remain to be addressed.
Keywords: Colouration in crab, DNA
barcoding, leucosiid, Mandapam, molecular phylogeny,
Tamil Nadu.
Introduction
Pebble crab or leucosiid
crab belonging to the family Leucosiidae (Samouelle, 1819) is rich in diversity (Ng et al. 2008;
Galil & Ng 2015). It mostly inhabits
the sandy and silty areas adjacent to seagrass beds, coral reef flats, as well
as intertidal areas usually buried in the sand (Naderloo
& Apel 2012; Ng & Komatsu 2016). Leucosiid crabs of
India have a long history where the key to the Indian Leucosia
was first provided by Alcock (1896) during an
“Investigator” expedition. At present,
97 species belonging to 35 genera of the family Leucosiidae
have been reported from India (Trivedi et al. 2018). While revisiting the Leucosiidae
classification, Galil (2005) proposed a new genus Seulocia
which differs from other genera in the shared characters such as third to
sixth abdominal somites fused in males and the
straight shaft in the first pleopod of males twisted once on its axis. So far, 11 species have been described in
this genus (Ng et al. 2008; WoRMS 2019), of which six
species S. cristata (Galil, 2005), S. pubescens (Miers, 1884),
S. pulchra (Galil, 2005), S. rhomboidalis (De Haan, 1841),
S. truncata (Alcock,
1896), and S. vittata (Stimpson, 1858) have
been recorded in Indian waters (Trivedi & Vachhrajani
2017; Trivedi et al. 2018). All of these
above records were mainly based on the morphological characteristics and lack
of information on genetic relatedness among them.
DNA barcoding along with morphological examination has
been considered as a useful tool for the validation of species (Madhavan et al. 2020).
This method can effectively identify cryptic species due to differences
in their genetic character (Bucklin et al. 2007). No such study has been reported for the genus
Seulocia.
Hence, in the present study along with the detailed morphological
examination, we used mitochondrial cytochrome oxidase subunit I (COI) gene to
validate the taxonomy of S. vittata from the southeastern coast of India.
Materials and Methods
The fish landing centre at Mandapam in Ramnad District of Tamil Nadu, India is one of the major
landing sites in the southeastern coast of India
(9.2860N & 79.1530E).
A total of five crab specimens were hand-picked from the freshly
discarded by-catch of commercial trawlers at the fish landing during June–July
2019. Specimens were quickly cleaned to
remove sediments and photographed (Cannon Powershot
G16) in the field to record fresh colouration.
The specimens were preserved in 95% ethanol and brought to Sathyabama Marine Research Station, Rameswaram for further
detailed examination.
Morphological examination
The specimens were examined by comparing key
morphological features and photographs described by Galil (2005). Four specimens (3 male, 1 female) were in red
colouration (LR) and 1 male was in typical bluish-grey colour (LG) as described
by Galil (2005). The carapace length (cl
in mm) was measured from the tip of the rostrum in the anterior region to the
posterior border of the carapace. The
carapace width (cw in mm) was measured from the
lateral margins of the carapace. The
specimens were then deposited in the national zoological collection of the
Marine Biological Regional Centre (MBRC), Zoological Survey of India (ZSI),
Chennai, Tamil Nadu, India.
Molecular identification
One representative each of red and grey coloured
specimens was subjected to molecular identification. Total genomic DNA from the
propodus/meri region of the
major cheliped of the crab was extracted using OMEGA BIO-TEK E.Z.N.A. Blood & Tissue DNA Kit, USA following the
manufacturer’s protocol. PCR
amplification was done for the mitochondrial cytochrome oxidase subunit I (COI)
gene using LCO-1490 (5′-GGTCAACAAATCATAAAGATATTGG-3′) and HCO-2198
(5′-TAAACTTCAGGGTGACCAAAAAATCA-3′) primers (Folmer et
al. 1994). Each PCR contained 12.5μL 2X
PCR master mix (Ampliqon, Denmark), 2.5μL each of the
two primers (10nM), and 2.5μL of template DNA (10–20 ng) and water to make a
final volume of 25μL. PCR conditions
were as follows: initial denaturation at 95°C for 10 min, 35 cycles of 95°C for
45 sec, 50°C for 45 sec, and 72°C for 45 min and final extension at 72°C for 10
min. PCR products were then visualized on 1% agarose and products with the high
intensity band were sequenced with ABI Prism 3730 Genetic Analyzer based on BigDye Terminator Chemistry.
Chromatograms were visualized, edited, and contigs
were prepared using consensus sequences from both the strands in the BioEdit (Hall 1999).
Sequences obtained in the present study were deposited in NCBI
GenBank. Sequences of COI from the
present study were then compared with published COI sequences of related taxa
from NCBI GenBank using BLASTn tool (https://blast.ncbi.nlm.nih.gov/Blast.cgi). COI sequences of species of the genus Seulocia and other related genus belonging to
the family Leucosiidae were downloaded and aligned in
the web version of Clustal Omega (https://www.ebi.ac.uk/Tools/msa/clustalo/). The alignment consisted of two sequences
obtained in the present study (MN786514, MN786515), two published sequences of
the genus Seulocia: S. vittata
(MH675982), S. latirostrata (MH675981) as
ingroup terminals, as well as four sequences from other genera as outgroup
terminal: Leucosia rubripalma
(MH675986), L. craniolaris (MH675985), Euclosiana scitula (MH675980),
and E. crosnieri (MH675978). The pairwise genetic distance between the
species was determined by the Kimura 2-parameter method (Kimura 1980) using
MEGA 7 (Kumar et al. 2016). The
phylogenetic tree was constructed using maximum likelihood method based on the
Tamura-Nei model in MEGA 7 (Kumar et al. 2016). Bootstrap test was conducted using 1,000
replications to get the best topology from a 75% majority rule consensus tree (Felsenstein 1985).
Results
Systematic accounts
Order Decapoda (Latreille, 1802)
Infraorder Brachyura (Latreille, 1802)
Family Leucosiidae (Samouelle, 1819)
Genus Seulocia
(Galil, 2005)
Seulocia vittata (Stimpson, 1858), Image 1 & 2.
Restricted Synonymy
Cancer craniolaris; Herbst, 1783: 90, pl. 2, fig. 17.
Leucosia craniolaris; Fabricius, 1798: 350
(part); K. Sakai, 1999: 19, pl. 7E.
Leucosia vittata Stimpson, 1858: 159; Shen & Dai, 1964: 28, fig.; Chhapgar, 1968: 609; Chen & Sun, 2002: 436, fig. 197,
pl. 16.8.
Leucosides craniolaris; Rathbun, 1910: 310 (part).
Leucosia sinica Shen & Chen, 1978: 80, pl. 2, figs 12, 13,
text-fig. 5; Huang, 1994: 580; Chen & Sun, 2002: 440, fig. 199.
Materials examined: Grey colouration: MBRC/ZSI D1-609, 11.vii.2019, 1 male, (cl 22mm; cw 20mm), India, Tamil Nadu, Mandapam fish landing site
(Palk Bay), 9.2860N & 79.1530E , depth 10–15 m, col.
Prakash & Amit Kumar.
Red Colouration: MBRC/ZSI D1-610, 11.vii.2019, 1 male
and 1 female, (cl 22 each; cw 19 and 20), same
collection data as above.
MBRC/ZSI D1-610, 11.vii.2019, 2 males, (cl 19 and 22; cw 18 and 20), same collection data as above.
Short description
Carapace sparsely punctate anteriorly, anterior margin
tridenticulate with median denticle slightly larger
than the adjacent ones. Anterolateral
margin with minute beaded lines. Margin
of epibranchial angle of carapace finely milled, epimeral
margin evenly milled throughout.
Posterior margin of carapace slightly rounded in male specimens (Both
Grey and Red) and rounded in female specimens (Red). Thoracic sinus deep, pterygostomian
region anteriorly defined by scalloped, overhanging and oblique margin. Fused abdominal segment bearing granule
medially in male, smooth without granules in females. Merus of the major
cheliped perliform, tubercles on the lateral margins,
few tubercles (3–5) on the dorsal as well as ventral region. The upper margin of carpus and palm smooth
and lower margins perliform, movable finger with
upper margin carinate (both grey and red).
Meri of the remaining pereiopods bearing beaded lines on the dorsal and
ventral margin, carpi prominently carinate dorsally. Propodi of the
pereiopods carinate in dorsal and ventral margins, dactyli flat and
non-carinate on the lateral margins.
Colouration – Grey (Male)
Carapace bluish-grey, with median dorsal reddish-brown
band becoming broader posteriorly (Image 1A–C).
Presence of two oblique bands of reddish-brown on each side diverging
from the front. Major cheliped meri, carpi and pal with a combination of reddish-brown and
bluish-grey band, distal region reddish, fixed and movable finger whitish
anteriorly and reddish posteriorly.
Meri, carpi, propodi, and dactyli of the
remaining pereiopods with a combination of reddish and white bands, tip of the
dactyli brown to black. Abdominal region
almost whitish (Image 1B). Terminal end
of the maxillipeds dark bluish-grey (Image 1C).
Colouration – Red (Male and Female)
Anterior region of carapace bluish-grey, latter half
of the carapace brick red to reddish-brown in colour in females (Image
2A–C). In males, carapace with
bluish-grey extended to latter half, posterior region of carapace brick-red
(Image 2D–F). Oblique bands are not
visible on carapace (Image 2D). Dorsal
and ventral region of the meri, carpi, and propodi of the major chelipeds brick-red to reddish colour,
dactyli of the movable dark red with a whitish tip (Image 2A,B). Meri of the remaining pereiopods brick-red in
colour, carpi, propodi, and dactyli dark brown to
black in colour. Abdomen brick red with
big black patch at the centre in both females and males (Image 2B, 2E). Propodi and carpi
of the major cheliped were black ventrally (Image 2B, 2E). Terminal end of the maxillipeds dark
bluish-grey in both females and males (Image 2C, 2F).
DNA barcode and phylogenetic relationship
BLAST analysis revealed that the sequences for LG and
LR exhibited 99.67% and 99.83% similarity with existing COI sequence of Seulocia vittata sequence
in the NCBI GenBank. The phylogenetic
analysis based on ML tree constructed using single mitochondrial gene fragment
(COI – 653bp) resulted in tree topology that S. vittata
(grey and red colour patterns) are closely related to S. vittata
available in the GenBank (Figure 1).
Out of 653 sites, 148 were parsimony informative sites. In the ML tree, all the S. vittata clustered together to form a monophyletic
clade. In addition, S. vittata is sister to its congener comprising Seulocia latirostrata and
other related genera Leucosia and Euclosiana (Figure 1).
Furthermore, the calculated pairwise genetic distance
of COI gene fragment using Kimura-2 parameter revealed that LR, LG, and
published S. vittata in the GenBank have no
genetic divergence. However, the genetic distance of 0.177 was calculated
between S. vittata and Seulocia
latirostrata, which is comparable to the genetic
distance of more than 0.200 with other genera such as Leucosia
and Euclosiana in the family Leucosiidae (Table 1).
Discussion
A nomenclatural and taxonomical validation of the
brachyuran crabs of the world includes over 6793 species belonging to 1,271
genera and 93 families (Ng et al. 2008).
Interestingly, recent advances in the molecular techniques have received
greater attention in understanding the evolutionary perspectives of marine
brachyuran crabs (Hultgren & Stachowicz 2008; Lai
et al. 2013; Fratini et al. 2018; Mantelatto
et al. 2018; Chen et al. 2019). The
integrative approach of both morphological and molecular analyses offers robust
information not only on the taxonomic ambiguity of the species, including
cryptic species (Baeza & Prakash 2019) but also
in the monitoring of commercial crabs for seafood safety (Rath
et al. 2018). In India, the molecular
based study in brachyuran crabs was very limited (Vartak
et al. 2015; Apreshgi et al. 2016; Ravichandran et
al. 2017; Rath et al. 2018; Madhavan
et al. 2020). In the present study, we
performed the molecular phylogeny as well as identified the pairwise sequence
divergence study of Seulocia vittata and its congeners based on the COI gene
fragment. The ML tree suggests that
S. vittata was sister to the S. latirostrata and other related species that are
supported with high bootstrap values.
Seulocia vittata has a wide geographic distribution in the
Indo-Pacific from Mauritius, India, Singapore, Malaysia, Thailand, Indonesia,
China, and Philippines (Galil 2005). The
present study represents a rediscovery of S. vittata
in the south east coast of India after the original report of Alcock (1896) during the “investigator” expedition. Moreover, no information exists on its
varying colour pattern. We observed that
though the individuals of S. vittata differed
in their colour pattern (grey and red variants), their genetic distance showed
no variation between these two-colour forms.
This indicates a situation of reverse cryptic behaviour in this crab.
Surprisingly, the evolutionary and ecological processes leading to the absence
of genetic divergence and variation in morphology (colour pattern) in S. vittata complex remain to be addressed. There could be several possible explanations:
a) S. vittata possess the capacity to change
colour and camouflage in nature as anti-predatory mechanisms (Stevens et al.
2014); b) the colour variation could be due to ecological adaptation to
different depths and habitats such as reefs and open sand flats (Darnell 2012);
c) morphological colour variation with low genetic structuring may indicate
high dispersal capacities of S. vittata throughout
the evolutionary history of this species. However, to validate the above hypotheses,
extensive sampling efforts and detailed examinations at larger geographical
scales are required.
Based on the outcome of this study, we recommend
integrative taxonomic and phylogeographic approaches to demonstrate the extent
and magnitude of species complexity in the leucosiid
crabs. This goal needs to be prioritized
as there is a recent increase in the trawl net operations in the south eastern
coast of India that could lead to decline in the benthic biodiversity (Purohit
2017). This could cause profound
implications in the conservation planning, stock assessment, biogeography,
evolutionary as well as the natural history of leucosiids. Lastly, the species complexity in S. vittata will provide the opportunity to understand the
important mechanisms of speciation among the leucosiid
crabs.
Table 1. Pairwise genetic distance calculated using
Kimura 2-parameter based on COI gene fragment of S. vittata
and other closely related taxa.
Species name |
1 |
2 |
3 |
4 |
5 |
6 |
7 |
Seulocia vittata (MH675982) |
|
|
|
|
|
|
|
Seulocia latirostrata (MH675981) |
0.177 |
|
|
|
|
|
|
Seulocia vittata - LG (Present study) |
0 |
0.177 |
|
|
|
|
|
Seulocia vittata - LR (Present study) |
0 |
0.177 |
0 |
|
|
|
|
Leucosia rubripalma (MH675986) |
0.214 |
0.212 |
0.214 |
0.214 |
|
|
|
Leucosia craniolaris (MH675985) |
0.202 |
0.196 |
0.202 |
0.202 |
0.026 |
|
|
Euclosiana scitula (MH675980) |
0.215 |
0.204 |
0.215 |
0.215 |
0.188 |
0.182 |
|
Euclosiana crosnieri (MH675978) |
0.215 |
0.204 |
0.215 |
0.215 |
0.188 |
0.182 |
0 |
For
figure & images - - click here
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