CHAPTER

Marine Molluscan Taxonomy and Biology - An

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Overview
V. VENKATESAN, K. S. MOHAMED, R. VIDYA, K. K. SAJIKUMAR, AND
P. S. ALLOYCIOUS
Molluscan Fisheries Division
ICAR-Central Marine Fisheries Research Institute

Introduction
Molluscs are a phylum of soft-bodied invertebrates which includes Gastropoda (snails, limpets
whelks and slugs), Bivalvia (oysters, mussels, clams, scallops, and cockles) and Cephalopoda (squids
cuttlefishes, and octopuses). Of these, the gastropods, with large and diverse group (about 80% of
all molluscs). The remaining groups such as Monoplacophora (cap-shaped neopilinids),
Polyplacophora (chitons), Scaphopoda (tuskshells), Solenogastres (crawling worm-molluscs), and
Caudofoveata (shell-less burrowing worm-molluscs) are known to a much lesser extent (Haszprunar
and Wanninger, 2012). Brief description about the major groups of molluscs is shown in Table 1.
Molluscs are originally originated in marine water, spread to freshwater and on to the land.
Freshwater and landforms are almost equal to the marine forms. Molluscs primarily inhabit in the
intertidal and littoral zones of the sea, occasionally descend to a greater depth upto 10,000 m.
Molluscs belonging to different taxonomic groups have been exploited for food, pearls, and shells.
The estimate number of species of molluscs vary from different parts of the world, however, estimates
number of existing species are about more than 1,00000 (Haszprunar and Wanninger, 2012).
About 5070 species have been reported from India belonging to 290 families and 784 genera which
are recorded from Gulf of Mannar (428 species), Lakshadweep (424 spp.), Gulf of Kutch (350 spp.),
Orissa coast (337 spp), West Bengal coast (425 spp.) and Andaman & Nicobar Islands (1434 spp).
Nearly 3,370 species of molluscs are recorded from marine habitat (Venkataraman and Wafar, 2005).
Among these, gastropods are the most diverse, followed by bivalves, cephalopods, polyplacophores
and scaphopods. At present over 1.5 lakh tonnes of cephalopods, over 1 lakh tonnes of bivalves
and nearly 20,000 t of gastropods are exploited from Indian waters. The large number of marine
gastropods (19 species)
Table 1. Major taxonomic groups of molluscs (Source: Haszprunar and Wanninger, 2012)
Gastropoda Comprises more than 1,00000 species that inhabit all marine, freshwater and terrestrial habitats
and size range from 0.5 mm to 100 cm in body length. All types of feeding habits (filter-
feeders, herbivores, predators, ecto- and endoparasites, and detritovores) and all mode of
reproduction are found in this group.
Bivalvia Includes more than 20,000 extant species (1mm to over 150 cm) that live in all kinds of marine
and freshwater habitats. They are not only filter feeder, but also include detritivorous and
carnivorous bivalves. Some of them also use symbiotic zooxanthellae for nourishment. Most
are epibenthic or burrow in soft bottoms, some burrow in limestone, wood (eg. shipworms).
Fertilization is mostly external. Trochophore, veliger and glochidia type larvae are known in
this group.
Cephalopoda Comprises only about 1,000 extant species that inhabit exclusively marine and range from 3
cm up to 7m in body length. Some members (Nautiloidea and Ammonitoidea) have external

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shells, while all other (Coleoidea) have internal, reduced / lost shells. All bear 8 - 10 arms
(about 80 arms in Nautilus) for capturing prey. Fertilization is external.
Scaphopoda Includes about 800 marine species of 2mm to 20 cm body length. They burrow in sand or mud
and feed chiefly on foraminiferans. Fertilization is external and they have lecithotrophic
(trochophore-like) larvae.
Monoplacophora Comprises less than 30 extant species with a size range from 1-40 mm long. They inhabit from
about 200 m down to 7000 m depth. Dorsal surface is protected by a single cup-shaped shell
and the mode of feeding is more or less similar to those of chitons.
Polyplacophora Includes about 1000 extant marine species with range from 3mm to 30 cm body length. Dorsal
side is protected by eight serial plates. They are mostly either herbivorous or detritovorous.
They have strong rasping tongue for food uptake. Fertilization is external and they have
lecithotrophic (trochophore-like) larvae.
Solenogastres Includes small marine group (280 species) of 1mm to 30 cm body length covered with cuticle
with spicules or scales. They live interstitially and feed on cnidarians. The mode of reproduction
is through copulation and they have lecithotrophic (modified trochophore or pericalymma -
type) larvae.
Caudofoveata Comprises small marine group (180 species) of 2mm to 15 cm body length covered with cuticle
with spicules or scales. They burrow in sand or mud and lose their foot sole entirely. They have
lecithotrophic (modified trochophore type) larvae.

Table 2. List of scheduled marine molluscs from India

Endangered list of molluscs
Class: Gastropoda
1 Cassis cornuta (Linnaeus, 1758)
2 Charonia tritonis (Linnaeus, 1758)
3 Conus milneedwardsi Jousseaume,1894
4 Cypraecassis rufa (Linnaeus, 1758)
5 Tudicla spirillus (Linnaeus, 1767)
6 Staphylaea limacina (Lamarck, 1810) ( = Cypraea limacina)
7 Leporicypraea mappa (Linnaeus, 1758) ( = Cypraea mappa)
8 Talparia talpa (Linnaeus, 1758) ( = Cypraea talpa )
9 Pleuroploca trapezium (Linnaeus, 1758) (= Fasciolaria trapezium)
10 Harpulina arausiaca (Lightfoot, 1786)
11 Dolomena plicata sibbaldi (G.B. Sowerby II, 1842) (= Strombus plicatus sibbaldi)
12 Ophioglossolambis digitata (Perry, 1811) (= Lambis crocea)
13 Lambis millepeda (Linnaeus, 1758)
14 Lambis scorpius (Linnaeus, 1758)
15 Lambis truncata ([Lightfoot],1786)
16 Harpago chiragra (Linnaeus, 1758) (= Lambis chiragra)
17 Harpago arthriticus (Roding 1798) (= Lambis chiragra arthritica )
18 Rochia nilotica (Linnaeus,1767) (= Trochus niloticus)
19 Turbo marmoratus Linnaeus, 1758

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Class: Bivalvia
1 Hippopus hippopus (Linnaeus, 1758)
2 Tridacna maxima (Roding, 1798)
3 Tridacna squamosa Lamarck, 1819
4 Placuna placenta (Linnaeus, 1758)
Class:Cephalopoda
1 Nautilus pompilius Linnaeus, 1758

followed by bivalves (4 species) and cephalopod (1 species) has been placed in the endangered
list which is a major cause of concern (Table 2). The collection, possession and trading of these
scheduled molluscs (Table 2) or their products (live or dead) are prosecuted and will attract a
punishment of severe imprisonment upto 7 years along with heavy fine under section 50, 51 of
wildlife (Protection) Act 1972.
Commercially exploited molluscs of India
Cephalopods
Three groups of cephalopods viz., squids (order Teuthoidea), cuttlefishes (order Sepiiodea)
and octopuses (order Octopodidea), are exploited from Indian seas (Table 3). The main species
occurring in commercial catches are Uroteuthis (Photololigo) duvaucelii ( = Loligo duvauceli), Sepia
pharaonis, S. aculeata and Amphioctopus neglectus (= Octopus membranaceous).
Table 3. Commercially exploited cephalopods from Indian Seas
(Source: Mohamed and Venkatesan, 2017)
Species Common Name Distribution
Squids
Uroteuthis (P.) duvaucelii Indian squid All along Indian coast
Loliolus (N) uyii Little squid Chennai & Visakhapatnam
U (P) edulis Swordtip squid SW coast
U (P) singhalensis Long barrel squid SW & SE coast
Loliolus (L) hardwickei Little Indian squid All along Indian coast
Sepioteuthis lessoniana Palk Bay squid Palk Bay & Gulf of Mannar
Sthenoteuthis oualaniensis Purple-back Flying squid Oceanic Indian EEZ
Thysanoteuthis rhombus Diamond squid Oceanic Indian EEZ
Cuttlefishes
Sepia pharaonis Pharaoh cuttlefish All along Indian coast
S. aculeata Needle cuttlefish All along Indian coast
S. elliptica Golden cuttlefish Veraval & Kochi
S. prashadi Hooded cuttlefish SW & SE coast
S. brevimana Shortclub cuttlefish Chennai & Visakhapatnam
Sepiella inermis Spineless cuttlefish All along Indian coast

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Octopuses
Amphioctopus neglectus Webfoot octopus SW & SE coast and islands
A. marginatus Veined octopus SW & SE coast and islands
A. aegina Marbled Octopus SW & SE coast and islands
O. lobensis Lobed octopus SW & SE coast and islands
O. vulgaris Common octopus SW & SE coast and islands
Cistopus indicus Old woman octopus SW & SE coast and islands

Bivalves
Various groups of bivalves such as clams, oysters, mussels, and windowpane oysters are
exploited along the Indian coast for food and shells (Table 4).
Table 4. Commercial important bivalves of India
Resource Common name
Clams and cockles
Villorita cyprinoides Black clam
Paphia malabarica, Paphia sp Short neck clam, textile clam
Meretrix casta, Meretrix meretrix Yellow clam
Mercia opima Baby clam
Mesodesma glabaratum
Sunetta scripta Marine clam
Donax sp Surf clam
Geloina bengalensis Big black clam
Tegillarca granosa (= Anadara granosa) Cockle
Placuna placenta Window pane oyster
Tridacna sp, Hippopus hippopus Giant clam
Mussel
Perna viridis Green mussel
Perna indica Brown mussel
Pearl oyster
Pinctada fucata Indian pearl oyster
Pinctada margaritifera Blacklip pearl oyster
Edible oyster
Crassostrea madrasensis Indian backwater oyster
Saccostrea cucullata Rock oyster

Molluscan Fisheries in India

Cephalopods are the most important group of molluscs with estimated all India production of
about 2, 61,663 tonnes in 2017 which was 11.6 % more compared to the previous year. They are
landed either as by-catch or as a targeted fishery. Targeted fishery is mostly carried out in mechanized
trawlers operating upto 200 m depth, and beyond in some areas.

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Bivalve fishery is the next in importance and fishing is practiced in limited extent mostly at a
subsistence level in various estuaries and coastal seas. Clams and cockles contribute 73.8%, followed
by oysters (12.5%), mussels (7.5%) and windowpane oysters (6.2%) (Mohamed and Venkatesan,
2017). The annual average clam production is about 57,000 t, oysters about 18,800 t, and marine
mussels about 14,900 t (Mohamed and Venkatesan, 2017). At present, there was no fishery for
marine pearl oysters, but it was the major fisheries before 1962 in the Gulf of Mannar area. Scallops
occur in certain area in stray numbers and do not contribute in fishery, whereas the windowpane
oyster formed considerable fishery till a few year back (Mohamed and Venkatesan, 2017).
Gastropods in India are exploited for both as food and as curios. Among gastropods, the sacred
chank is most important with annual production of over 1,000 t till a few years back (Mohamed and
Venkatesan, 2017). The fishing of top shell viz., Rochia nilotica and Turbo marmoratus has been
banned as they have been declared as endangered. One species of Abalone viz., Haliotis varia
occur in stray numbers and are not fished. Mining for subsoil shell deposits was carried out from
time immemorial especially in the Ashtamudi and Pulicat Lakes for industrial purposes.
Mollusc biology
Molluscs are extremely large group and diverse in all phases of life. They occur in all marine
habitats of the world including deep-sea hydrothermal vents, freshwater environments upto 40° C,
land (gastropod alone) and permanent ice (Haszprunar and Wanninger, 2012). They range in size
from 0.4 mm (omalogyrid gastropods) to more than 15 m (Architeuthis squids) (Haszprunar and
Wanninger, 2012). Their longevity can range from a few months to up to more than 150 years
(Deep sea giant bivalves) (Haszprunar and Wanninger, 2012). They mostly crawl or glide through
cilia or muscle waves with mucous (Haszprunar and Wanninger, 2012). Some animals can
permanently cement to the substrate, such as giant clam and edible oyster while some can attach
to the substrate through byssus thread such as mussels. Modes of feeding are also diverse including
filterfeeders , omnivores, predators, grazers, detritovores , ecto- and endoparasites, and various
kind of symbioses with bacteria, plankton (Zooxanthellae), and algae.
The body of theoretical molluscs comprises five fundamental parts – the foot, the head, the
visceral mass, the mantle and the shell. The alimentary tract or system of theoretical molluscs
consists of ingestion, digestion, absorbtion and assimilation of food. The system starts with mouth
which leads to the buccal cavity having pair of jaws in each side. Pharynx, located at the anterior of
the buccal cavity, is occupied by the odontophore which supports the tongue like structure called
radula. Ducts from one or two pairs of salivary glands are present at the anterior of pharynx which
in some species (Conus sp) are modified into organs to secret venom used to paralyze or kill the
prey. The tract goes on with the esophagus and then enlarges in a stomach where the food has
been partially digested as threads of particles linked together by mucus. Food is mostly digested in
the ducts of two large digestive glands by tiny cilia (whiplike structure). These digestive glands
occupy almost all the space within the visceral mass. Digestion of molluscs takes place both
extracellularly and intracellularly. Extracellular digestion occurs especially in the stomach while,
intracellular digestion takes place especially in the hepatopancreas. These organs (stomach /
hepatopancreas) do the dual functions – secretion of digestive enzymes and absorption of food

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particles. The structure of posterior portion of the stomach is conical in many molluscs and a
translucent rod shape in bivalves. This structure is known as crystalline style which secrets enzymes
to digest certain carbohydrates. After the stomach comes the intestine which opens at the anus
into the pallial cavity.
Circulatory system in molluscs is open except cephalopods. Heart, made up of two dorsal
auricles/atria and a single ventricle, gets only oxygenated blood from gills and send it to different
regions of the body through posterior aorta. Blood/hemolymph transport through blood vessels
directly to the openings or spaces between the organs. Respiratory pigments in molluscs are of two
main types viz., red hemoglobin and blue, copper containing hemocyanin.
Excretory system removes the waste materials that are formed from the breakdown of
assimilated food chiefly nitrogenous waste such as ammonia and urea. This function is carried out
by one or more kidneys which are diverse in the various groups of molluscs. In primitive group,
these organs are linked to the pericardial cavity and at least one of the excretory passages is modified
to form a gonoduct for transfer of gametes. Excretory system opens into the pallial chamber. Pallial
chamber is also an important structure which mediates between the animal and its external
environment.
Respiratory system in molluscs is generally formed by the pair of gills in the pallial chamber.
However, most of the gastropods have single gill. Gills are the site of gas exchange and look like a
feather, with a central axis. Gills are of different forms in different group of the molluscs depending
on their environment and feeding habits. Land snails do not possess gills instead they have primitive
form of lung.
Molluscs show various mode of reproduction. Most of them are either gonochoristic or
hermaphroditic. Percentage of gonochoristic and hermaphroditic species are more or less equal
(Haszprunar and Wanninger, 2012). Few of them occasionally show parthenogenesis. Majority of
the molluscs, especially gastropods and cephalopods transfer sperm by means of copulatory organs,
whereas, many species, especially gastropods, scaphopods and chitons shed their gametes liberally
into the water. Their egg sizes range from about 80 µm (many bivalves and gastropods) to 2 cm
(Nautilus spp) (Haszprunar and Wanninger, 2012).
Larvae of them are either intracapsular or direct development into miniature form or
planktotrophic or lecithotrophic. Larvae may look different from adult form. Typical molluscan larvae
are veligers which are usually more or less modified form of Trochophore larvae. Example of special
type of larvae is glochidium of freshwater unionoids which is well known as parasite on fish gills.
Biology of commercially important cephalopods
All cephalopods are active predators that feed on live prey, mainly fishes and crustaceans. Fish
always occurs in the diet of squid U.(P.) duvaucelii of all sizes (Mohamed and Joseph, 2005). The
fondness of crustacean diet diminishes with increase in size and there is indication of cannibalism
above 80 mm DML (Oommen, 1977). Cephalopods are one of the major preys for a variety of
marine fishes including tunas, billfishes, cetaceans, and whales (Silas, 1985). Many researchers
have observed the high proportion of empty stomachs in samples and fatigue in feeding during
spawning (Oommen, 1977).

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The characteristic of length weight relationship of Indian cephalopods has been reported to be
hypoallometric with the ‘b’ value is lower than 3 (Meiyappan et al., 1993). This relationship is also
significantly different for males and females (Mohamed, 1996).
Cephalopods along the Indian coast are reported to spawn almost throughout the year. The
earlier work on the reproductive biology of the Palk Bay squid Sepioteuthis lessoniana has been
carried out by Rao (1954). Later on, the maturity of three species of squids and six species of
cuttlefishes has been reported by Silas et al. (1985ab). Size at first maturity (Lm) and peak spawning
seasons of some of the studied species is given in Table 5. Maturity stages for biological studies of
squids and cuttlefishes have been standardized (Silas, 1985) and described as four-point ( Immature,
Maturing, Mature, and Spent) maturity scale. This maturity scale has been used by all workers on
Indian cephalopods.
Mature and partially spawned individuals of U. (P) duvaucelii are found throughout the year
along both the coasts, but along the west coast, peak spawning has been observed during post
monsoon i.e. Sep-Nov. (Silas et al., 1985a; Mohamed, 1993). This species forms large congregation
during this season and becomes vulnerable to the purse seine fleet operating along Karnataka
coast (Mohamed, 1993) and also to cast netters along coastal water of Alleppey (Meiyappan and
Srinath, 1989). This squids congregate for spawning in near shore waters after which the female
migrate to the shallow subtidal regions with hard substratum for laying the fertilized eggs (Mohamed,
1993). Fertilized eggs from the subtidal regions of Karwar seas have been collected for rearing
(Asokan and Kakati, 1991). Based on sex ratio (M 80:F20) of such squid schools, it would be easy to
conclude that female was semelparous. However the evidence such as relatively low GSI levels and
the occurrence of mature females over a wide range of size classes, suggests that this species is
multiple spawner and not a semelparous species (Mohamed, 1993). Similar studies in other
commercial cephalopods are not availableTable 5.Biology of the commercially important
cephalopods (Source: Silas et al, 1985ab; Abdussamad et al., 2004; Abdussamad & Somayajulu,
2004; John Chembian, 2013, Sajikumar (unpublished))

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 Table 5.Biology of the commercially important cephalopods (Source: Silas et al, 1985ab; Abdussamad et al., 2004; Abdussamad & Somayajulu,
2004; John Chembian, 2013, Sajikumar (unpublished))
Species Lm (mm) Spawning Peak Region/coast
period Spawning
U.(P) duvaucelii 76 (M);86 (F) Throughout Feb & Jun - Sep (Chennai) East coast
the year Jan, Jul & Sep (Waltair)
113 (M);118 (F) Feb-Mar, May-Jul, Sep-Oct (Kochi) West coast
S. lessoniana 102 (M);98 (F) Jan-Jun East coast
U.(P) singhalensis 126(M);135(F) Jan-Mar; Oct&Dec West coast
S. aculeata 102-118 (F) Throughout the year (Mandapam) Aug-Sep(Mandapam) East coast
Mar-Jun &Nov (Portonovo)
Feb-Dec (Chennai) Feb, Jun-Aug &Oct-Dec(Chennai)
Nov-Jul (Waltair) Apr, Jul, & Nov-Dec (Waltair)
Aug-Mar(Kakinada) Nov-Dec (Kakinada)
130-132(F) Nov-Dec (Kochi) West coast
May, Aug, Sep,& Dec (Mumbai)
S.pharaonis 120 (F)(Waltair) Jan-Feb, Apr & Sep-Dec(Waltair) Sep-Dec, Feb, & Apr-Jun East coast
Aug-Mar(Kakinada) Nov-Dec (Kakinada)
138 (F) (Chennai) Throughout the year (Chennai)
160 (F) (Vizhinjam) Jan, Mar-Apr, Sep-Oct, Dec (Vizhinjam) Oct-Dec, Mar-Apr West coast
157(F) (Kochi) Feb, Apr, Jun, Oct & Dec (Kochi)

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S. brevimana 63(F) (Chennai) Jan-Feb & Jul-Dec East coast
59(F) (Waltair)
S. elliptica 93-96 (M&F) Aug-April & Jun (Kochi) West coast
S. inermis 52(Waltair) Apr,Jun-Sep & Nov-Dec (Waltair) East coast
61 (Chennai) Feb-Mar, & Jul-Dec (Chennai) Sep, Dec & Mar (Chennai)
50 (Portonovo) Mar-Oct (Portonovo)
Apr-Nov (Kakinada)
83(Kochi) Apr, Sep-Dec (Kochi) Sep & Oct (Kochi) West coast
S. oualaniensis 128 (M);170 (F) Throughout the year Oct-Dec & Mar-May West coast
T. rhombus Oct -Feb West coast
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Fecundity studies on Indian cephalopods are few. Individuals of U.(P) duvaucelii produced 5300
eggs on an average (Rao, 1988) while, fecundity estimate of S. inermis ( 69 - 71 mm DML) was
ranged between 470 - 850 eggs (Unnithan, 1982). There was good correlation between length,
ovary weight, and fecundity in U.(P) duvaucelii. In S. lessoniana (120 - 196 mm DML), total number
of ripe eggs was from 180 to 1054 egg (average 497 eggs) (Venkatesan and Rajagopal, 2013).
Fecundity, ovary weight and nidamental gland weight showed strong correlation, while fecundity,
DML, and body weight showed relatively weak correlation.
Biology of commercially important bivalves
The biology of commercially important species of bivalves from India is given in Table 6. Physical
factors such as temperature and salinity are the important factors for influencing the reproductive
cycles and spawning in bivalves (Sastry, 1979). In addition to temperature, food
Table 4. The biological details of the commercially important bivalves (Source: Kripa and Appukuttan, 2003)
Species Lm Spawning L max Length (mm) in Distribution
period I yr II yr III yr
V.cyprinoides 20 - 25 May-June 52 30 41 - West coast
& Nov
P. malabarica 20 Sep-Feb 55 43.1 - - West coast
P. viridis 15.5-28 Dec-Jan 91.5 117 129 East coast
Jul-Nov 96 117 129 West coast
C. madrasensis 12-14(M) Nov-Feb 128 86 112 East coast
24-26(F) Jul-Sep 70-80 90-110 120-130 West coast
Feb-Apr
M. casta 11-17. Throughout 55 42.6 East &West coast
the year
M. meretrix 21-26 May-June 91 47 61.5 East coast
Feb - Sep
M. opima 11-20. Dec 53.8 30 43.5 East coast
May-Aug 22 31 43 West coast
T. granosa 20-24 Throughout 73.4 41.1 55.3 66.3 East coast
the year

supply and latitudinal distribution effects the reproductive cycle of bivalves (Newell et al.,
1982). The number of spawning events and duration of spawning period can also differ greatly with
respect to species, geographic area and environmental conditions (Gosling, 2003). In general, an
environment play an important role to influence the growth, reproduction and recruitment of
bivalves and same species shows different growth rates and spawning periods in different areas
(Kripa and Appukuttan, 2003). They found that the combination of different hydrographic parameters
like salinity, availability of settlement substrate and current pattern are responsible for controlling
the spat fall, population growth, zonation and species dominance. Although, most of bivalves are
gonochoristic, in certain bivalves like oysters hermaphroditism has been observed.

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Literature cited:
Abdussamad, E.M., Meiyappan, M.M., and Somayajulu, K.R., 2004. Fishery, population characteristics and stock
assessment of cuttlefishes, Sepia aculeata and Sepia pharaonis at Kakinada along the east coast of India.
Bangladesh J. Fish. Res., 8(2):143-150.
Abdussamad, E.M. and Somayajulu, K.R., 2004. Cephalopod fishery at Kakinada along the east coast of India: Resource
characteristics and stock assessment of Loligo duvauceli. Bangladesh J. Fish. Res., 8(1):61-69.
Asokan, P.K. and Kakati, V.S. 1991. Embryonic development and hatching of Loligo duvaucelii Orbigny (Loliginidae,
Cephalopoda) in the laboratory. Indian J. Fish., 38 (4): 201-206.
Gosling E. 2003. Bivalve molluscs—biology, ecology and culture. Fishing News Books, Oxford, 443 pp.
Haszprunar, G. and Wanninger, A., 2012. “Molluscs,” Current Biology, vol. 22, no. 13, pp. R510-R514.
John Chembian, A. 2013. Studies on the biology, morphometrics and biochemical composition of the ommastrephid
squid, Sthenoteuthis oualaniensis (Lesson, 1830) of the south west coast of India. Ph. D. Thesis, Cochin University
of Science and Technology, Kochi, India.
Kripa, V and Appukuttan, K.K., 2003. Marine Bivalves. In: Mohan Joseph, M., and Jayaprakash, A. A., (eds.), Status
of Exploited Marine Fishery Resources of India. ICAR-CMFRI, Cochin, p. 211-220.
Meiyappan, M.M., Srinath, M., Prabhakaran Nair, K., Satyanarayana Rao, K., Sarvesan, R., Syda Rao, G., Mohamed,
K.S., Kuber Vidhyasagar, Sundharam, K.S., Lipton, A. P., Natarajan, P., Radhakrishnan, G., Narasimham, K.A.,
Balan, K., Kripa, V. and Sathianandan, T.V. 1993. Stock assessment of the Indian squid Loligo duvauceli
Orbigny. Indian J. Fish., 40 (1&2): 74-84.
Meiyappan, M.M and Srinath, M., 1989. Growth and mortality of the Indian squid (Loligo duvauceli) off Cochin,
India. In: Contributions to tropical fish stock assessment in India: FAO/DANIDA/ICAR National Follow-up
Training Course on Fish Stock Assessment, 2 - 28 November 1987, Cochin.
Mohamed, K.S. 1993. Spawning congregations of Indian squid Loligo duvauceli (Cephalopoda Loliginidae) in the
Arabian Sea off Mangalore and Malpe. Indian J. Mar. Sci., 22:172-175.
Mohamed, K.S. 1996. Estimates of growth, mortality and stock of the Indian squid Loligo duvauceli orbigny, exploited
off Mangalore Southwest coast of India. Bull. Mar. Sci., 58 (2): 393-403.
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