Journal of Threatened Taxa | www.threatenedtaxa.org | 26
December 2019 | 11(15): 15061–15067
The windowpane oyster family Placunidae Rafinesque, 1815 with additional description of Placuna quadrangula (Philipsson, 1788) from India
Rocktim Ramen Das 1, Vijay
Kumar Deepak Samuel 2, Goutham Sambath 3, Pandian Krishnan 4, Purvaja Ramachandran 5 & Ramesh Ramachandran
6
1–6 National
Centre for Sustainable Coastal Management, Ministry of Environment, Forest
& Climate Change, Chennai, Tamil Nadu 600025, India.
1 Graduate
School of Engineering and Science, University of the Ryukyus, Nishihara,
Okinawa 903-0213, Japan.
3 King Fahd
University of Petroleum and Minerals, Dhahran, 31261, Kingdom of Saudi
Arabia.
1 k188604@cs.u-ryukyu.ac.jp,
2 deepakocean@gmail.com (corresponding author),3 taaz_goutham@yahoo.co.in,
4 krishnanars@yahoo.com,
5 purvaja.ramachandran@gmail.com, 6 rramesh_au@yahoo.com
doi: https://doi.org/10.11609/jott.5049.11.15.15061-15067 | ZooBank: urn:lsid:zoobank.org:pub:6E29E0AB-3A16-47F0-BC36-6C9D89F54568
Editor:
Anna Holmes, National
Museum Cardiff, Cathays Park, UK. Date of publication: 26
December 2019 (online & print)
Manuscript
details: #5049 | Received 07 May
2019 | Final received 18 November 2019 | Finally accepted 01 December 2019
Citation: Das, R.R., V.K.D. Samuel, G. Sambath, P. Krishnan, P. Ramachandran & R. Ramachandran (2019). The windowpane oyster family Placunidae Rafinesque,
1815 with additional description of Placuna
quadrangula (Philipsson,
1788) from India. Journal
of Threatened Taxa
11(15): 15061–15067. https://doi.org/10.11609/jott.5049.11.15.15061-15067
Copyright: © Das et al. 2019. Creative Commons Attribution 4.0 International
License. JoTT
allows unrestricted use, reproduction, and distribution of this article in any
medium by adequate credit to the author(s) and the source of publication.
Funding: National Centre for Sustainable Coastal Management, Ministry of Environment, Forests & Climate Change, Govt. of India.
Competing interests: The authors declare no competing interests.
Acknowledgements: The authors are grateful
to Dr. C. Stella (Department of Oceanography and
Coastal Area studies, Alagappa University, India) for
providing information regarding Placuna
ephippium. The comments provided by Dr. Graham Oliver (National Museum Wales, Cardiff, UK), Dr. Anna Holmes (National Museum Wales, Cardiff, UK) and Dr. Henk Dekker (Naturalis Biodiversity Center,
The Netherlands) greatly improved this manuscript. RRD is grateful to the curator of Ryukyu
museum, University of the Ryukyus and Kouri Shell
Museum, Okinawa for providing access to Placuna
samples. The Tamil abstract prepared by Dr. Lavanyaa Shri Goutham is
acknowledged.
Abstract: The Bivalvia family Placunidae
Rafinesque, 1815 in India is reviewed in this paper based on previous
literature and records. Additionally,
the species Placuna quadrangula
is described from the Indian sub-continent.
Being an economically important family in this geographic region, this
paper can be regarded as a baseline for further ecological, management and
policy-related studies pertaining to Placunidae
and other exploited species.
Keywords: India, Placuna quadrangula, Placunidae,
Taxonomy.
Abbreviations: ANI—Andaman & Nicobar Islands | GOM—Gulf of Mannar | GOK—Gulf of Kutch | MBRC—Marine Biological
Research Centre | NZC—National Zoological Collections | PB—Palk Bay | GBR—Great
Barrier Reef | QGIS—Quantum Geographic Information System | ZSI—Zoological
Survey of India | RUMF—Ryukyu University Museum, Fujukan.
The family Placunidae Rafinesque, 1815 is
comprised of the genera Placunanomia and Placuna, the latter with seven accepted living
species (Huber 2010). Distributed mostly
within the Indo-West Pacific region (Matsukuma 1987),
the genus Placuna Lightfoot, 1786 has a long
documentary history, as it is commercially exploited for pearl production,
food, lampshades and shell-craft items (Gallardo et al. 1995). From Indian waters, Placuna
placenta (Linnaeus, 1758) (Image 1) is the only species that has been
thoroughly studied due to its economic value (Laxmilatha
2015a). Placuna
sella (Gmelin, 1791) now P. ephippium (Philipsson, 1788) was first
reported from the Andaman & Nicobar Islands, India by Prashad (1932), followed by Gulf of Mannar,
southeastern coast of India (Rao & Dey 2000; Venkataraman et al. 2004) and was later revised
by Stella (2010) (Image 2) from the Mandapam coast (Adjacent to GOM). Placuna
ephippium is also exploited, especially in the Bay of Banate,
Philippines and its surrounding areas, and mainly known for its food value (Gallardo
1994). Currently, as per published
reports, two species of Placuna, P. ephippium
and P. placenta are reported from the Indian coast (Rao 2017)
(Image 3). In this study we confirm the
presence of a third species from India, P. quadrangula
(Philipsson, 1788).
This paper further attempts to distinguish all three species from the
Indian coast based on morphology.
Materials and Methods
Five valves of P. quadrangula were
collected from three different locations along the eastern coast of India
(Table 1). The collected dry specimens
were cleaned and preserved in zip-lock covers for further assessment. Identification to species level was made
based on the characters provided by Lynge (1909) and
Huber (2010). Specimens were deposited
in the National Zoological Collections (NZC) repository in the Marine
Biological Research Centre (MBRC), Zoological Survey of India (ZSI),
Chennai. A single specimen of P.
placenta deposited at (ZSI/MBRC) was assessed additionally. Measurements of specimens were recorded with
Yuri Digital Calliper 200x0.01mm. Study
area map was created using QGIS 3.6.3 Noosa.
Systematics
Order Pectinida Gray,
1854
Superfamily Anomioidea Rafinesque, 1815
Family Placunidae Rafinesque, 1815
Genus Placuna Lightfoot, 1786
Placuna is monomyarian, with low umbones, V-shaped crurae and pallial line, often obscured. Valves circular to sub-circular and laterally
compressed. All the characters are
common in the species observed from the Indian subcontinent, and a detailed
species wise description is given below.
Placuna placenta (Linnaeus, 1758)
(Image 1)
Description: Shell thin, very flat,
roughly circular or subcircular in shape,
inequivalve, periostracum absent. Inner
surface smooth, outer surface lamellate, growth lines present. Transparent when juvenile, turning opaque
with age. Lacking radial lines on the
external surface. Crurae
below the umbones, unequal in size, adductor muscle scar slightly anterior of
midline. Pallial line obscure and
non-sinuated. Specimen examined has a
damaged or broken outline.
Distribution: Extends from Gulf of Aden
in the west to Taiwan in the east (Matsukuma 1987;
Huber 2010).
Placuna ephippium (Philipsson, 1788)
Description: Shell saddle-shaped with
curved dorsal margin without periostracum.
Growth lines visible. External
colour purple brown/brownish with black shades to large red-purplish-blackish
spots on the interior surface. Lacking
radial lines on the external surface. Crurae prominent, equal in size and wideset. Single adductor muscle scar in the center of midline; purplish in colour.
Distribution: India to Australia (Matsukuma 1987; Huber 2010).
Remarks: See Discussion.
Placuna quadrangula (Philipsson, 1788)
(Image 3. A–J)
Description: Shell thin, brittle,
papery, laterally compressed but slightly concave (Image 3. A-J). Outline quadrangular, periostracum
inconspicuous. Surface smooth,
lamellate, growth lines fine with closely arranged radial threads. Externally pinkish to whitish with
non-uniform white radial rays originating from umbones; internally pinkish to
whitish. Prominent crurae
of equal size. Adductor muscle scar
centrally situated, rounded. Pallial
line obscure and non-sinuated. Internal
margins smooth. Internal and external
surface of the specimens with attached fouling organisms.
Distribution: Present study – Tamil
Nadu, Puducherry (eastern coast of India), Mergui Archipelago, Thailand,
Indonesia, Philippines and Australia (Matsukuma 1987;
Sanpanich 2011).
Remarks: The five examples collected from three different
localities possess the typical radially rayed pattern.
Discussion
Comparative analysis among the Placuna
species from India with some notes on other Indo-pacific species
The radial colour patterns originating from the umbones (Image 3) a
typical character for P. quadrangula is
emphasized in Huber 2010. Although
confused with P. ephippium which is larger, the specimen described from
Mandapam has no mention of purple muscle scar nor the color
predominance in the shell as seen in Huber 2010. Rather it is described as “almost
transparent” along with light brownish nature with black patches (Image 3)
(see Stella 2010). The transparent
nature fits well with P. placenta juvenile as observed in the
collections of the Ryukyu Museum (RUMF-ZM-03693) and in Kouri
Shell Museum, Okinawa (Rocktim Ramen Das, pers. obs.)
but is known to turn white and shiny after maturity. The presence of brown radial rays in the
latter from southeastern Asian specimens (see Matsukuma 1987) might be misleading and needs
reassessment. P. ephippium lacks
such brown radial rays and possesses large red, purplish-black spots on the
interior region (Henk Dekker pers. comm.).
Life history of genus Placuna
A review of literature revealed that there is a limited amount of
research based on the life history of genus Placuna
with exclusive information available only for P. placenta. Adam Young (1980), who did an extensive study
on the larval growth and development of P. placenta, revealed that the
shells remain inequivalve and transparent from a very early stage. The author also revealed that from fertilized
egg to the formation of spat, it takes about 10–11 days and the final sedentary
phase is reached when around 600µm in size is reached during which several key
morphological changes take place, viz., active foot appearance during larval
metamorphosis. Narasimham
(1984), who later studied the biology of P. placenta from the eastern
coast of India (Kakinada Bay) (Figure 1) mentioned about the biannual spawning
strategy of the species and based on the gonadal appearance and morphology
divided the maturity into four stages, viz.: active, ripe, partially spawned
and spent/resting. Interestingly, a
recent observation from the coast of Sonmiani (Balochistan) indicated that P. placenta spawns
all-round the year (Parveen et al. 2018) which contradicts the findings of
Hornell (1909) and Moses (1939) whose studies were from a not so distant area
of Okha, Gujarat, India (~400 km). This probably indicates the local
environmental parameters like temperature, salinity and monsoonal
characteristics can play an important role at regional scales (Ladja 2002).
Status of genus Placuna in India
Along the coast of India, the windowpane oyster (P. placenta) was
initially reported by Hornell (1909a,b).
In the 1970s, Narasimham documented its
utilization due to its high economic value while Laxmilatha
(2015) reviewed the economic value.
Though the species is reported from various places of the Indian
sub-continent (Table 3), it is commercially exploited only from specific areas
along the coastline (Table 3).
Exploitation of the species in areas of Gulf of Kutch was mainly for
pharmaceutical purposes (Alagaswami & Narasimham 1973; Narasimham et
al. 1993). Presently the exploitation
levels are low in the Gulf of Kutch. In
Kakinada Bay, P. placenta is regarded as one of the most important
bivalve resources, but the stock is under threat due to overexploitation (Rao
& Somayajulu 1996; Laxmilatha
2015b). Nauxim
Bay in Goa had a minor fishery where the meat was locally consumed (Narasimham et al. 1993).
Apart from the above-mentioned locations, the collection of windowpane
oyster from the coastal waters of Tamil Nadu mainly for the pearl and shell
craft industries. Vellapatti
fishing hamlet near Tuticorin is the hub for the utilization of P. placenta
in the cosmetic and paint industry.
Rameswaram is famous for the windowpane oyster lampshades and mirrors. It is important to highlight that due to such
activities, previous densely populated areas of P. placenta are
lost (Tripathy & Mukhopadhyay 2015). The saddle-shaped oyster P. ephippium collected
in Mandapam, Gulf of Mannar is also used in the
production of a variety of curios/souvenirs, viz., trays, lampshades (Stella et
al. 2010). The third species P. quadrangula, reported here is either invasive or has
been overlooked over the past decade. It
is important to highlight that Iredale in the
scientific reports published related to the GBR expedition refers to a
publication from the Bolten Museum dating back to
1798 which mentions Ephippium anomia (synonym P. quadrangula) from Tranquebar (now Tharangambadi, ~230km from Chennai). Further in-depth analysis revealed the
information is related to its morphological characters, leaving the information
regarding its geography being rather vague (see Iredale
1939; Röding 1906).
Conclusion
Placuna placenta is the only species under
the genus that is listed (as Placenta placenta) under Schedule IV of the
Indian Wildlife Protection Act, 1972. As
mentioned in the act, “No person shall hunt any wild animal specified in
Schedules I, II, III, and IV except as provided under section 11 and section 12”. Furthermore, Section 11 and 12 allows for
hunting only in special cases and with proper documents and permissions from
the concerned government authority; however, surreptitious fishing of this
species continues even in protected areas, e.g., Kakinada Bay (Coringa Wildlife Sanctuary) (Laxmilatha
2015a,b) apart from other areas as mentioned above. Moreover, the recent news of P. placenta
being smuggled to western Asian and South American countries in alternate forms
(Ravinesh et al. 2018) further highlights the urgent
need to assess the genus. As our study
highlights the morphological aspects of this genera, a thorough comparative
assessment of the internal organs and the application of molecular methods
should provide essential insights. On
the other hand, biogeographical assessment of P. quadrangula,
its ecology and its implications on P. placenta distributions is
urgent. As the Indian Ocean is home to
large and unknown malacofauna (see Das et al. 2017), continuous surveys to
discover these understudied resources remains imperative.
Table 1. Materials examined.
Locality |
Coordinates (decimal) |
No. of valves |
Dimensions Lx WxH (mm) |
Date of collection |
Collected by |
Deposition no. |
Kottivakkam (Chennai) |
12.9660N, 80.2650E |
2 |
80.38 x 78.77 x 8.23; 57.83 x 49.48 x 4.88 |
28.ii.2015 |
GS |
ZSI/MBRC/M.2004; ZSI/MBRC/M.2005 |
Serenity Beach, Kottakuppam
(Puducherry) |
11.9760N, 79.8450E |
1 |
92.09 x 91.30 x 9.59 |
19.x.2015 |
RRD |
ZSI/MBRC/M.2006 |
Kasimedu, Royapuram (Chennai) |
13.1230N, 80.2970E |
2 |
71.94 x 61.37 x 7.32; 50.55 x 47.71 x 4.91 |
13.iii.2016 |
RRD; GS |
ZSI/MBRC/M.2007; ZSI/MBRC/M.2008 |
Table 2. Summary of important morphological features
among the Indian Placuna species.
|
Placuna placenta |
Placuna ephippium |
Placuna quadrangula |
Morphology (Figure 2) |
Roughly circular |
Saddle shape with curved dorsal margin |
Quadrangular, thin, papery |
Colour |
Semi-transparent (juvenile); opaque and white
(adult). |
Purplish, large spots in the interior region (Red,
Purple or Black) |
Pinkish to whitish, with prominent white radial
lines. |
Crurae (Figure 2) |
V-shaped, narrow and unequal |
V-shaped, wide apart and equal |
V-shaped, wide apart and equal |
Radial lines |
Absent |
Absent |
Irregular radial lines originating from umbones |
Pallial Line |
Obscure |
Obscure |
Obscure |
Posterior adductor muscle |
Monomyarian |
Monomyarian |
Monomyarian |
Adductor muscle scar |
Slightly anterior of midline |
Center of midline, purplish |
Center of midline, rounded |
Umbones |
Low |
Low |
Low |
Table 3. Reports of genus Placuna
from various parts of the Indian subcontinent*
Source |
Locality/Region |
Species |
Hornell
1909b |
Balapur
Harbour, Beyt Island and Okha
(Gujarat); Ennore (Tamil Nadu); Buckingham canal
(Tamil Nadu); Pulicat lake (Andhra Pradesh) |
Placuna
placenta |
Prashad
1932 |
Andaman
Islands |
Placuna sella+ |
Rai 1933 |
Bombay
coast (Maharashtra) |
Placuna
placenta |
Alagaswami
& Narasimham 1973 |
Gulf of
Kutch (Gujarat); Malabar coast, Vembanad lake (Kerela); Tuticorin**, Nagapattinam**,
Kakinada bay** (Andhra Pradesh) |
Placenta
placenta++ |
Narasimham et
al. 1993 |
Nauxim
Bay** (Goa) |
Placenta
placenta++ |
Hameed
& Somasundaram 1998 |
Gulf of Mannar (Tamil Nadu) |
Placenta
placenta++ |
Rao & Dey 2000 |
West Bengal
|
Placuna
placenta |
Venkataraman
et al. 2004 |
Andaman
& Nicobar Islands |
Placuna
placenta |
Samuel et
al. 2005 |
Dhanuskodi
(Tamil Nadu) |
Placenta
placenta++ |
Stella et
al. 2010 |
Mandapam**
(Tamil Nadu) |
Placuna
ephippium |
Boominathan et
al. 2012, 2014 |
Kali River;
Uttara Kannada district (Karnataka) |
Placuna
placenta, Placenta placenta++ |
Murugesan
et al. 2013 |
Parangipettai
(Tamil Nadu) |
Placenta
placenta++ |
Prabhu et
al. 2013 |
Mallipattinam
(Tamil Nadu) |
Placenta
placenta++ |
Thilagavathi et
al. 2013 |
Muthupettai
(Tamil Nadu) |
Placenta
placenta++ |
Bijukumar et
al. 2015 |
Kavaratti
Island (Lakshadweep) |
Placuna
placenta |
Tripathy
& Mukhopadhyay 2015 |
Murud-Jinjira**
(Maharashtra); Pouchitra**, Raida**,
Goomara** (GOK, Gujarat), Chennai** |
Placuna
placenta |
Mahapatro et
al. 2016 |
Chilika
Lake (Odisha) |
Placuna
placenta |
Rao 2017 |
Pindara
Bay (Gujarat); Baitkal cove and Pavin
halla (Karnataka); Pambam,
Kundugal point (Tamil Nadu); Eatimukkala
and Kalingapatnam (Andhra Pradesh); Jharkali and Jambu Island (West
Bengal) |
Placuna
placenta |
Ravinesh et
al. 2018 |
Navi Mumbai
(Maharashtra) |
Placuna
placenta |
Present
study |
Chennai
(Tamil Nadu); Kottakuppam (Puducherry) |
Placuna quadrangula |
*The information may be non-exhaustive +Placuna sella is the synonym of Placuna
ephippium ++Placenta placenta is a group
under which Placuna placenta was
assigned by Gray 1849, thus can be regarded as a
synonym in this context, **Exploited Areas
For
figures & images - - click here
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