Training Manual
Training Manual
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Editors
S. Bijoy Nandan
P. Graham Oliver
Jayachandran P. R.
Asha C.V.
Published by
Directorate of Public Relations and Publications
Cochin University of Science & Technology
Kochi 682022, Kerala, India
English Language
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Training manual -1 International Training Workshop on Taxonomy of Bivalve Molluscs.
Editors
Bijoy Nandan
P. Graham Oliver
Jayachandran P.R.
Asha C.V.
Citation:
S. Bijoy Nandan, P. Graham Oliver, Jayachandran P.R. Asha C.V. (eds.). 2016. Training manual -
1st International Training Workshop on Taxonomy of Bivalve Molluscs. Directorate of Public
Relations and Publications, CUSAT, Kochi, India.
Cover Design
Mr. Jayachandran PR
Publisher
Director
Directorate of Public Relations and Publications
Cochin University of Science and Technology
Kochi- 682022, Kerala state, India.
Website: www.cusat.ac.in
Email: [email protected], bijoynandan @yahoo.co.in
Printed in India at
Indus offset printers
Kochi-682022, Kerala
No part of this publication may be reproduced, or transmitted in any form or by any means,
without prior written permission of the publisher or author. Plates/Images in the Manual is copy
righted to authors especially chapter 8 is the combination of power point slides with personal
collections of Dr. P. Graham Oliver and chapters 1-7, 9 are presentations of various authors.
These contents are strictly limited to personal use of participants in the workshop (ITW’01, 2016)
for learning purpose.
ISBN 978-93-80095-79-0
CONTENTS
About editors /i
Preface /ii
Acknowledgments /iii
CHAPTER ONE
Taxonomy and Faunistic Survey’s in India
Dr. Ramakrishna / 1
CHAPTER TWO
Annotated Classification and Diversity of Marine Bivalve Molluscs of India
Dr. N.V. Subba Rao /25
CHAPTER THREE
Overview of the bivalve fisheries of India
Dr. K. Sunilkumar Mohamed / 47
CHAPTER FOUR
Ashtamudi clam fishery - 1st MSC Certified fishery in India
Dr. K.K. Appukuttan / 54
CHAPTER FIVE
Taxonomy of Marine Molluscs of India: Status and Challenges Ahead
Dr. Biju Kumar A. / 65
CHAPTER SIX
Molecular approaches in taxonomy with special reference to bivalve mollusc
Dr. Hari Krishnan K./ 86
CHAPTER SEVEN
Sampling Techniques for molluscan fauna
S. Bijoy Nandan, Jayachandran P. R., Asha C.V. / 105
CHAPTER EIGHT
Taxonomy of Bivalve Molluscs
Dr. P. Graham Oliver / 115
CHAPTER NINE
Status and species diversity of Tridacna in Indian waters
Ecological determinants and stochastic fluctuations of Tridacna maxima survival rate in
Lakshadweep Archipelago, Monitoring densities of the giant clam Tridacna maxima in the
Lakshadweep Archipelago
Dr. Deepak Apte /326
Glossary
About the Editors
Dr. S. Bijoy Nandan, Professor, Department of Marine Biology, Microbiology &
Biochemistry, School of Marine Sciences, Cochin University of Science and Technology, is an
active researcher, teacher and administrator for over the last 2 decades. His area of interest is
marine ecology, marine ecotoxicology and biology of polar communities. He has received
several awards and fellowships of which the Jawaharlal Nehru Award of Indian Council of
Agricultural Research in 1993, Recognition Award of Zoological Society of India in 2008 and
U.S Fulbright Visiting Scholar in 2013-14 has been outstanding. He has about 130 peer
reviewed publications to his credit and implementing research projects of national and
international relevance, also been involved in U.N Compensation project in Arabian Gulf
coast. He has earlier served as the Head of Central Inland Fisheries Research Institute (ICAR),
Kerala Unit and Senior Officer in Central Institute of Fisheries, Nautical & Engineering
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Training, Government of India. Bijoy Nandan is the Organising Secretary of the 1
International Training Workshop on Taxonomy of Bivalve Molluscs
Dr. P. Graham Oliver, Specialist in systematics and functional morphology of marine and
freshwater Bivalvia (Mollusca). See Systematics of marine Bivalvia. Interests include:
Chemosymbiotic taxa (Thyasiroidea & Solemyoidea) of the deep-sea; world-wide Arcoidea;
provision of identification tools for British and western Indian Ocean bivalves. Author of
books on Red Sea and Arabian marine bivalves and chief scientist and co-editor of marine
biodiversity workshop in Rodrigues (West Indian Ocean). Current major project:- provision
of a web-based identification guide to the marine bivalves of the British Isles, inter-tidal to
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abyssal. Graham is the course director of the 1 International Training Workshop on Taxonomy of
Bivalve Molluscs.
Mr. Jayachandran P.R., Project Scientist, Department of Marine Biology, Microbiology &
Biochemistry, School of Marine Sciences, Cochin University of Science & Technology, is an
active researcher in marine benthic ecology and ecotoxicology. Jayachandran is the Associate
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Organising Secretary of the 1 International Training Workshop on Taxonomy of Bivalve Molluscs.
Mrs. Asha C.V., Senior Research Fellow, Department of Marine Biology, Microbiology &
Biochemistry, School of Marine Sciences, Cochin University of Science & Technology, is an
active researcher in marine benthic ecology.
PREFACE
Phylum Mollusca being the largest group of animals existing today are distributed in
large numbers in marine, estuarine and freshwater habitats. The class Bivalvia in Mollusca is
commonly referred to as clams, mussels, cockles, oysters, and scallops. More than 30,000
living species of bivalves have been described from different aquatic systems. Even though
records on the taxonomy, distribution and diversity of species are available from different
ecosystems, they lack clarity in specific identification characters and descriptions. Moreover
species identification especially in the Arabian Sea and associated regions has been inconsistent,
such that many ambiguities in bivalve taxonomy still remain. In view of this, as a pioneering
attempt the 1st International Training Workshop on Taxonomy of Bivalve Molluscs, a hands on training
using local species, was organized by the Department of Marine Biology, Microbiology &
Biochemistry, School of Marine Sciences, Cochin University of Science and Technology, Kochi
Kerala, India from 10-14 May, 2016, in collaboration with Department of Aquatic
Biology and Fisheries, University of Kerala, Kariavattom, Thiruvananthapuram, Kerala,
India and collaboration with Dr. P. Graham Oliver, National Museum of Wales, and School of
Ocean Sciences, Bangor University, North Wales, United Kingdom. This training manual is a
compilation of the 1st International Training Workshop on Taxonomy of Bivalve Molluscs, and elaborates
at depth on the taxonomy and fishery characteristics of molluscs especially the different species
of bivalves. Chapters 1-7, 9 are based on invited lectures delivered and chapter 8 is by
Dr. P. Graham Oliver the course Director of the training programme. Thus, the manual is
prepared for the participants of the international training workshop and can also serve as a
reference material for marine biologists and taxonomists working in molluscan studies.
The authors are indebted to the Head and faculty of the Department of Marine Biology,
Microbiology & Biochemistry, CUSAT, for the facilities and the National Fisheries Development
Board, Kerala State Council for Science, Technology & Environment, Kerala State Biodiversity
Board and University Grants Commission (UGC), for supporting to conduct the 1st International
Training Workshop on Taxonomy of Bivalve Molluscs and bring out this contribution. We are grateful
to researchers of the Ecology Division of the Department of Marine Biology for their unstinted
support in publishing this manual. We also sincerely thank the Director, Directorate of Public
Relations & Publications, CUSAT for his advice and support for the publication of this Training
Manual.
Taxonomy and Faunistic Survey’s in India
Ramakrishna 1
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species are still undescribed. The description of the new animal phyla
Loricifera in 1983, Cycliophora in 1995 and Mycrognathozoa in 2000, the
possibility that tropical arthropods alone could number over 10 million
species, and the fact that over 12,000 – 15,000 new animal species are
described yearly, exemplify how little is known regarding the magnitude of
global species richness (World Conservation Monitoring Centre, 1992 for an
overview; Simonetti, 1997). The smaller the organisms, more poorly known
the group to which it belongs (Wilson, 2003). About 69,000 species of fungi
have been named, but as many as 1.6 million are thought to exist. Of the
abundant nematodes, around 15,000 species are known but millions more
might await discovery. The bacteria and archaeans are the black hole of
systematics (Wilson, 2003); although only 6000 have been formally
recognized, approximately that many, almost all new to science, can be found
only a few grams of rich forest soil.
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The word systematics stem from the Latinised Greek word ‘Systema’
and can be defined as the classification of living organisms into hierarchical
series of groups emphasizing their phylogenetic interrelationships. It is the
science of arranging ‘species’ names into an order that reflects their
evolutionary relationships. As pointed out by Lincoln et al. (1998) the term
systematics has often been used as equivalent or synonymous with taxonomy,
but a lot of controversy exists about this, and several definitions of taxonomy
and systematics have been proposed by different authors to clarify the
situation. Mayr (1969) characterised taxonomy as ‘the theory and practice of
classifying organisms’ whereas he regarded systematics as ‘the science of the
diversity of organisms’. Wheeler (2008), believes that the systematics is a sub
discipline of taxonomy concerned with reconstructing phylogeny.
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Taxonomic Impediment:
Taxonomy provides scientifically based knowledge about species, the units of
biodiversity, and establishes a basic biological language about them that
enables meaningful communication between people.
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Bio NET International in its meeting held on 8th July, 2008 further said that in
order to overcome this crisis of taxonomic impediment we need to look into
the following viz.,
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the tree of life, makes baseline data available for conservation and ecology
studies, and affords humans the possibility to take advantage of the
underutilized resources offered by the earths’ biodiversity (Wilson 2004).
Taxonomic Keys
The main purpose of a key is to facilitate identification or to distinguish one
type of organism or object from another. A key may or may not reflect ideas
of evolutionary or phylogenetic relationship. A taxonomic key is a device for
identifying an object unknown but that someone else has already described.
The user chooses between alternative characteristics of the unknown object
and, by making correct choices, arrives at the name of the object. Keys that are
based on successive choices between two alternatives are known as
dichotomous keys. A dichotomous key is a tool that is the most widely used
form of classification in the biological sciences because it offers the user a
quick and easy way of identifying unknown organisms. Keys consist of a
series of choices that lead the user to the correct name of a given item.
"Dichotomous" means "divided into two parts." That is why dichotomous
keys always give two choices in each step. In each step, the user is presented
with two statements based on characteristics of the organism. If the user
makes the correct choice every time, the name of the organism will be
revealed at the end.
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or ratios; the terms large and small have no meaning in the absence of
quantitative values. If possible, use several easily seen and interpreted
morphological characters in each lead. There are two kinds of descriptions
that might be presented to the user of a dichotomous key: qualitative and
quantitative descriptions. Qualitative descriptions concern the physical
attributes, or qualities, of the item being classified. Quantitative descriptions
concern values that correspond with the item being classified. Examples of
quantitative descriptions are such phrases as "number of annulai as in
Annelida" or, "has 8 legs," or "weighs 5 grams". Knowing the difference
between these two types of descriptions can be immensely beneficial for
creators and users of dichotomous keys.
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In fact, one of the main objectives of this initiative, the discovery and
description of new taxa, cannot be accomplished with sequence data alone
(Ebach & Holdrege 2005b). Molecular evidence is mainly derived from
cytoplasmic organelles, whereas the taxonomic system is based on characters
that are derived mainly from chromosomal genome which is more
comprehensive. (Grant, 2003). As previously mentioned, the superposition of
intra- and inter-specific variation is a serious problem (Meyer & Paulay 2005,
Cognato 2006, Meier et al. 2006, Whitworth et al. 2007). This difficulty,
however, is not unique to molecular data, and is encountered with other sets
of data such as morphology, ecology and other sources (Will et al. 2005). The
problems with the sole use of morphology in taxonomy work are also well-
known (see Packer et al. 2009). Phenotypic plasticity, cryptic species and
identification of immature stages are good examples (Hebert et al. 2003). From
this perception that any character system used in taxonomy to the exclusion
of others will fall short of the task, the practice of an integrative taxonomy
that draws data from different sources is promising. This practice would
certainly be superior to the current chaos generated by single data sources
and illuminate taxonomic results with complementary sources of data (Will et
al. 2005).
Conclusion
In Linné’s time, biology was virtually identical with taxonomy. In the quarter
of a millennium elapsed since then, biology has undergone a huge amount of
evolution and diversification. Taxonomy is now only part of biology, and it is
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range of distribution (Alien and Invasive species). In the process, these non
native species alter ecosystem balance, impair economic stability and threaten
indigenous species.
The phase of field studies was initiated by the British Government for
the investigation of deep sea fauna, assigned to Surgeon Naturalists of the
Marine Survey of India. The deep sea explorations started in 1822, with Royal
Indian Marine Ship Investigator, laid a solid foundation for the systematic
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study of deep-sea fauna. Among the other earlier zoological explorer, the
contributions made by Dr. Jhon Anderson from his expeditions to Yunan
(China) in 1868 and 1875, collections form Persian Boundary Commission
(1870-1872), Second Yarkan Mission (1873 – 1874) and Dafla expedition (1874
– 1975) are significant in the history of the survey.
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Preston H B 1915. Mollusca. Freshwater Gastropoda & Pelecypoda. Taylor & Francis, London, 244
pp., 29 figs.
Fauna of Amphibia and Reptilia
Smith, M A (1931-1943) Reptilia and Amphibia. 3 Volumes. Volume 4 was to cover Amphibia.
Smith, M A (1931) Reptilia and Amphibia 1: Loricata and testudines pp. 101–103
Smith, M A (1935) Reptilia and Amphibia 2: Sauria
Smith, M A (1943) Reptilia and Amphibia 3: Serpentes
Boulenger, G. A. (1890) Reptilia and Batrachia
Fishes
Day, Francis (July 11, 1889) Fishes Volume I Chondropterygii, Teleoste (Physostomi;
Acanthopterygii: Percidae)
Day, Francis (September 21, 1889) Fishes Volume II Teleostei (Acanthopterygii excl. Percidae;
Anacanthini; Lophobranchii; Plectognathi), Leptocardii
Birds
Oates, EW (Blanford, W. T. ed.) (1889) Birds. 1 [582 p]
Blanford, W. T. (1890) Birds. 2 [424 p]
Blanford, W. T. (1895) Birds. 3 [450 p - 102 figs]
Blanford, W. T. (1898) Birds. 4 [500 p - 127 figs]
Mammilia
Blanford, WT (1888, 1891) Mammalia
Part 1 Primates, Carnivora, Insectivora
Part 2 Chiroptera, Rodentia, Ungulata, Cetacea, Sirenia, Edentata
Second edition
Pocock, R.I. (1939) Mammalia. I. Primates and Carnivora
Pocock, R.I. (1941) Mammalia. II. Carnivora: Aeluroidea, Arctoidea
The Government of India passed The Indian Museum Act in 1866 and
the museum of Asiatic Society was transferred in 1875 to the newly
constructed building. By this time the
zoological and anthropological collection
had expanded, a number of descriptive
catalogues were published and expeditions
were taken out to different parts of the
empire.
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however after his retirement, Alcock was given little support. Lord Curzon
decided to exhibit the collections of the Indian Museum as a memorial to
Queen Victoria in 1903 and Alcock was ordered to "to vacate the gallery of Fishes
at a moment's notice."
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The Zoological Survey of India (ZSI) commenced its work with four scientific
officers under the directorship of Dr. Thomas Nelson Annandale. The first
survey was undertaken at the request of the medical authorities of India.
During First World War, the government
became anxious that the disease
Schistosomiasis, hitherto unknown in India,
might be introduced by soldiers returning
from Middle East. A survey of Indian
freshwater Mollusca and their possible role in
acting as vectors of human Schistosomiasis
was under taken. Other significant
contribution from Dr Nelson Annandale
include expedition to Lake Galilee, Tiberas,
Palestine, to the Tale Sap in Thailand and to
Lake Biwa in Japan Dr. Nelson
Annandale
The seed of environmental research in ZSI were sown even in those early
years, by undertaking the study on the impact of engineering schemes, one
on Manchar Lake in Sind (Now in Pakistan) and the other at Vishakapatnam
to understand the changes seen in the construction of a harbour in the area.
The scientific contributions of Dr Sunder Lal Hora in the field of ichthyology,
for three and a half decades (1920-’55), dominated the scene of Indian
ichthyology in the 20th century.
During war years (1931-46) and great financial depression the activities
of ZSI was restricted to maintenance of rich collection in the survey. Even
during this period of natural crisis, extensive ecological and taxonomical
studies were carried out in and around Calcutta especially the wetlands of
north and south salt lakes of Sunderban and other swamps of the area.
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carried on either by men who knew the animals but were experts in science or
by those ‘rapid passengers’ who had no time to observe creatures, but worked
up their account afterwards. However, after the Independence, due
recognition was given to zoological science in their five year plans and got its
due share of importance in daily life. Establishment of research institutes such
as Forest Research Institute and Bombay Natural History Society and
University education and their contributions to Indian zoology need to be
taken into account for a thorough understanding of the status of Indian
zoology. A further boost to the scenario came recently after the Biodiversity
Convention of 1992 and establishment of Ministry of Environment and
Forests.
Suggested Readings
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Annotated Classification and Diversity of Marine
Bivalve Molluscs of India
N.V.Subba Rao 1
Introduction
The bivalves are laterally compressed and have two calcareous shell valves
joined together dorsally by a hinge. In general, the two valves enclose the
body or soft parts completely, except in few where the shell is reduced. They
have gills or ctenidia, which are used more for food collecting than
respiration. The hinge and ctenidia are two important structures on which the
earlier classifications were based. Bivalves exhibit remarkable ecological
diversities and adaptations to aquatic ecosystems, from freshwater to deep
sea. They have developed different life styles for their survival: filter feeding,
deposit feeding, photo autotrophic or photo symbiotic methods. They occur in
abundance in shallow coastal waters, estuaries and lagoons. They are
gregarious and settle permanently in a place and some, such as oysters, form
reefs. Some have developed byssal threads to stick to a particular place and to
overcome the force of waves. Many have resorted to burrowing habits and
settle in soft bottom or semisolid substratates. A few have adapted to
turbulent rocky intertidal zones of the sea and a few have specialized to live
as infauna in intertidal sandy beaches. Members of a few families such as
Lucinidae, Vesicomyidae etc have adapted to deep waters of oxygen
minimum zones of the sea. They have successfully overcome the difficulties in
such environments by developing endosymbiotic relationships with sulfur
oxidizing bacteria. Members of Lasaeidae and Montacutidae have specialized
in living as commensals with various infaunal or epifaunal invertebrates, like
crustaceans, polychaetes etc.
1
(Rtd:) Chief Scientist, Zoological Survey of India
E-mail: [email protected]
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Depending on the nature of their life habits bivalves can be grouped broadly
as follows:
Some Bivalves as per their Life Habits
Epifaunal
Free living/ Swimming: shell equilateral but not eqivalved. One valve,
usually lower valve (left valve) slightly larger than the other valve.
Monomyarian, anterior adductor muscle absents, only posterior
adductor muscle present. Ex. Pectinidae.
Byssally attached (epibyssate): Equivalved or inequivaled, with reduced
anterior region. Some have a byssal notch and a well-developed auricle
and some such as ark shells have ventral gape Ex. some ark shells,
mytilids, living in exposed habitats, Pteriids, hammer- oysters, Some
epifaunal nestlers ex. Barabatia and Mytilopsis.
Dorsally attached: Byssus absent, some may have in the juvenile stage,
ex.Tridacna. It remains attached on its dorsal end.
Cemented to the substratum: Commonly equivalved, attaches by the
lower left valve.It assumes the shape of the object to which it gets
attached. Shell exhibits various shapes. Ex. Oysters.
Semi-infaunal
Byssally attached (Endobyssate): Remains partly buried in the
substratum. Ex. Pen shells. Lives in fissures or crevices ex. some ark
shells and a few mussels, like Modiolus. Limopsids occur epi- or
endobyssally in soft sediments.
Infaunal
Burrowing: Shell eqivalved and isomyarian (with identical muscles) or
anisomyarian (with two dissimilar muscles), with a distinct pallial line
without a pallial sinus (nonsiphonata), ex nuculids, Yoldiidae. Deep
burrowing forms have well developed siphons that is seen as a pallial
sinus in an empty shell. exs. lucinids, tellinids, semelids and venerids.
Boring: Shell usually equivalved, with ridges or spines on the exterior
ex. Coral borers, rock borers- Lithophaginae, Petricolinae,
Gastrochaenidae.
Woodborers: Shell with supplementary plates, body modified into a
tubular form like a worm and the shell reduced, covering only a part of
the anterior end. Ex. shipworms (Teredinidae).
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The first classification that had found general acceptance was proposed by
Thiele (1935; English translation, 1998). He based his classification on the
nature of hinge teeth, form of adductor muscles and the structure of ctenidia.
He recognized three orders; Taxodonta, with taxodont teeth, Anisomyaria,
with two types of muscles and Eulammellibranchiata with ctenidial gills. The
last mentioned was again divided into four suborders; Schizodonta,
Heterodonta, Adapedonta and Anomalodesmata. This classification was in
use till the publication of Treatise on Invertebrate Paleontology in which Newell
(1965, 1969), based on shell structure, hinge teeth and anatomy proposed six
subclasses;Palaeotaxodonta, Cryptodonta, Pteriomorphia, Palaeoheterodonta,
Heterodonta and Anomalodesmata. Till recently this classification was
adopted by many incorporating a few minor changes (Boss, 1985; Vaught,
1989). A new classification of Bivalvia was proposed in 2010 (Bieler, Carter
and Coan, 2010). This system of classification is adopted by WoRMS (World
Register of Marine Species). In their subsequent studies Bieler et al. (2014),
proposed a revised classification and classified Bivalvia into Protobranchia
and Autobranchia. The latter again divided into Pteriomorphia and
Heteroconchia. They recognized six major monophyletic lineages:
Protobranchia, Pteriomorphia, Palaeoheterodonta, Archiheterodonta,
Anionomalodesmata, and Imparidentia. But Autobranchia and Heteroconchia
are not used to avoid additional Linnaean ranks in the Classification (Gofas,
2015). According to this classification there are two superorders and fifteen
orders to accommodate 110 families.
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very complicated, so also the classification. There are still some unresolved
issues. We may expect some stabilization in the classification of Bivalvia with
the publication of Bivalvia Treatise.
As per the latest classification mainly by Bieler et al. (2010 and 2014) Indian
bivalves, up to families, are arranged as follows:
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Order Arcida Stoliczka, 1871, Ark clams, False ark clams, bitter sweet clams
Shell thick, quadrate, trapezoidal or ovate. Shell covered with thick
periostracum. Sculpture consists of radial ribs. Anterior adductor muscle
smaller than the posterior adductor or very much reduced. Hinge almost
straight and bears transverse, rather perpendicular plications or denticles.
Mantle not fused ventrally. Outer fold of mantle bears ocelli. Sedentary forms
with free or byssate adults. Shallow burrowers with epifaunal or semi-
infaunal nestling habits.
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Order Ostreida Ferussac, 1822. True oysters, Pen shells, Pearl oysters
Synonym: Pterioidea Newell, 1965
Shell of variable shapes and sizes. Resilium of fibrous material and either
continuous or separated by lamellae. Ventral margin of mantle not fused and
bears tentacular processes. Byssal notch absent in the adult. Mainly cemented
to the substratum by one valve.
Order Pectinida Gray, 1854. Scallops, Jingle shells, Kitten paws, Thorny
oysters, Window pane oyster
Shell often inequivalve, sculptured with radial ribs. Hinge margin moderately
long and without true hinge teeth. Ligament in a median cartilage in a pit, the
lamellar area either reduced or absent. Byssal notch completetly in the right
valve. Mantle open ventrally. Gill filaments connected by interlocking cilia
projecting from ciliated discs. Many members with two auricles dorsally on
either side of the shell. Many have pallial tentacles, short or long, on the
ventral margin of the mantle.
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Carter et al. (2011) included all the above families in order Cardiida and
recognized suborder Cardiina Ferussac, 1822 to include superfamilies
Cardioidea and Tellinoidea as given below. For the rest of the families
Hyporder Veneroidei J.Gray, 1854 and Minorder Veneroitei J.Gray, 1854 were
proposed. The super family Gastrochaenoidea was placed in a separate
suborder Gastrochaenidina Morretes, 1949. A suborder Leptonidina Dall,
1889 was proposed to include the superfamily Galeommatoidea Gray, 1840.
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Superfamily Hiatelloidea
Family 49. Hiatellidae
Superfamily Solenoidea. Carter et al. (2011) placed this superfamily in a
separate order Solenida Dall, 1889.
Family 50. Pharidae
Family 51. Solenidae
Superfamily Cardioidea
Family 52. Cardiidae
Superfamily Tellinoidea
Family 53. Donacidae
Family 54. Psammobiidae
Family 55. Semelidae
Family 56. Solecurtidae
Family 57. Tellinidae
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Superfamily Lucinoidea
Family 58. Lucinidae
Superfamily Thyasiroidea
Family 59. Thyasiridae
Superfamily Dreissenoidea
Family 60. Dreissenidae (introduced into India).
Superfamily Myoidea
Family 61. Corbulidae
Family 62. Myidae
Superfamily Pholadoidea
Family 63. Pholadidae
Family 64. Teredinidae
Family 65. Xylophagidae
Superfamily Arcticoidea
Family 66. Trapezidae
Superfamily Cyrenoidea (partly freshwater)
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The first mentioned families are comparatively more speciose than all
others. Of the total 298 genera and 760 species, these families account for 196
genera 520 species. Veneridae is the most conspicuous with 29 genera and 95
species followed by Tellinidae with 34 genera and 71 species.
Coral reef ecosystem is the most preferred habitat for bivalves and
their species diversity in the four important ecosystems of India are given in
Table .1. Of these bivalves, twenty-four species are known to bore into corals
(Appukuttan, 1974 and Subba Rao & Surya Rao, 1981).
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Gastropoda 60 55 16 33 9 16
Species
Source: Das & Dev Roy, 1989; Subba Rao et al. (1995); Bhoominathan et al.
(2012)
Majority of the 760 species reported from India are from the shallow
waters. But from the records it is seen a total of about 47 species belonging to
23 families were dredged from the deep sea, beyond 100 meters’ depth. There
are 20 species from Bay of Bengal, 16 species from Lakshadweep Sea, seven
each from Arabian Sea and Andaman Sea. All the species of Propeamussiidae
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(11 species) and Cuspidariidae are from the deep sea. A species of the first
mentioned was hauled from a maximum depth of 3297 meters and a species
of the latter was from 1165 meters. Limatula subtilis (Limidae) was dredged
from 1987 meters’ depth. The families Malletiidae (2 species), Vesicomyidae
(3 species), Myochamidae (one species), and Euciroidae (one species) are
exclusively from the deep Sea. Excluding these six families (from out of 23),
all other families have majority of the species in shallow waters with a few
species extending their distribution into deep sea.
All the deep sea collections were made by Marine Survey, RIMS (Royal
Indian Marine Survey Steamer) Investigator from 1884 to 1911. RIMS
Investigator carried out biological research from 1884 to 1914 and 1914 to
1926. It collected samples by establishing 233 deep water stations between
1884 and 1911. All the collections made by the Investigator were studied by
E.A.Smith (1894-!906) and published under the title “Natural History Notes
from Indian Marine Survey Steamer Investigator’. Smith dealt with about 400
species based on Investigator Collections and of which 176 new species (of all
classes of Mollusca) were described. Some of the species were illustrated by
Annandale & Stewart (1897-1909) in six parts and 23 plates. There are no
further significant collections or additions to our knowledge of deep sea
molluscs of Indian Seas. After a lapse of more than hundred years recently
Raman et al. (2002), on a cruise of FORV Sagar Sampada, dredged a few
bivalves from the oxygen minimum zone at a depth of 300-700 meters in the
Northeast Bay of Bengal.
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Table 3. Bivalve molluscs recorded from two important sea grass ecosystems
Conclusion
An effort has been made in this paper to give a broad picture of diversity of
bivalves and to draw the attention of Indian malacologists to the latest
classification on the subject. During the last three decades interesting data has
been added on the phylogeny of Mollusca. Molecular and palaeontological
studies are providing new tools for understanding the relationships of
different groups of molluscs. This has led to many changes in the
classification of Mollusca and it is still in the process of stabilization as new
molecular sequence data is being generated and new hypotheses developed
on superfamilial ranks. Studies in India, as reflected in many recent studies,
are not touched by any of these latest changes in the phylogeny of Mollusca.
Many lists and ‘Check Lists’ are published, often including species not
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References
Appukuttan, K.K. 1974. Distribution of coral boring bivalves along the Indian
coasts. Journal of the Marine Biological Association of India 15(1): 427-430
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Boss, K.J. 1982. Mollusca. In: Synopsis and classification of living organisms (Ed.
S.P.Parker), vol.1: 945-1166, New York, McGraw Hill.
Carter, J.G., C.R. Altaba, L.C. Anderson, R.Araujo, A.S.Biakov, A.E. Bogan,
D.C.Campbell, M. Campbell, C. Jin-hua, J. C. W. Cope, G.Delvene,
H.H. Dijkstra, F. Zong-jie, R. L. Gardner, V.A.Gavrilova, I.A.
Goncharova, P.J. Harries, J.H.Hartman, M. hautmann, W.R. Hoeh, J.
Hylleberg, J.Bao-yu, P. Johnston, L. Kirkendale, K. Kleemann, J.
Koppka, J. Kriz, D. Machado, N. Malchus, A.Marquez- Aliaga, J.P.
Masse, C.A. McRoberts, P.U. Middlefart, S.Mitchell, L.A. Nevesskaja, S.
Ozer, J. Pojeta, jr, I.V. Polubotko, J.M. Pons, S.Popov, T. Sanchez,
A.F.Sartori, R.W.Scott, I.I. Sey, J.H. Signorelli, V.V. Silantiev,
P.W.Skeleton, T. Steuber, J.B.Waterhouse, G.L. Wingard and T. Yancey
2011. A Synoptical Classification of the Bivalvia (Mollusca).
Paleontological Contributions Number 4: 1-47. Paleontological Institute,
The University of Kansas,Lawrence, USA.
Daniel, A. & A.S. Rajagopal 1974. Molluscs of economic value from Great
Nicobar Island. Journal of the Bombay Natural History Society 70 (2):394-
398.
Das, A.K. & M.K.Dev Roy 1989. A general account of the mangrove fauna of
Andaman and Nicobar Islands. Pp.1-173, Fauna of Conservation Areas
4, Zoological Survey of India
Giribet, G. 2008. Bivalvia, pp. 105-141, in: Phylogeny and Evolution of the
Mollusca (Eds. W.F. Ponder & D.R. Lindberg), Berkeley, University of
California Press
Giribet, G. & D. L. Distel 2003. Bivalve Phylogeny and Molecular Data, pp. 45-
90, in: Molecular Systematics and Phylogeography of Mollusks (Eds. C.
Lydeard & D.R. Lindberg), Smithsonian Books, Washington.
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Subba Rao, N.V. (In Press). Indian Seashells, Part 2-Bivalvia. Zoological
Survey of India, Kolkata.
Subba Rao, N.V. & K.V.Surya Rao 1981. Occurrence of a coral boring bivalve
Gregariella coarctata (Carpenter) (Bivalvia: Mytiliidae) in the Indian
waters. Bulletin of the Zoological Survey of India 4(2): 213-215
Subba Rao, N.V., A.Dey & S. Barua 1995. Mollusca. In: Estuarine Ecosystem
Series, Part 2: Hugli Matla Estuary: 41-91.
Susan, D., N.G.K. Pillai & P. Satheeskumar 2012. Checklist and Spatial
Distribution of: Molluscan Fauna in Minicoy Island, Lakshadweep,
India. World Journal of Fish and Marine Sciences 4: 449-453.
Vaught, K.C. 1989. A Classification of the living Mollusca. Xii+ 195 pp., (Eds.
R.T. Abbott & K.J.Boss), Ameican Malacologists, Melbourne, U.S.A.
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Overview of Bivalve fisheries of India
K. Sunil Mohamed 1 & Geetha Sasikumar
Introduction
Bivalves are commercially important molluscs belonging to the Class Bivalvia
(Lamellibranchia or Pelecypoda), which is the second largest Class under the
Phylum Mollusca. They are bilaterally symmetrical, laterally compressed
molluscs, with extensive mantle lobes which secrete a single shell composed
of two valves. Bivalves are reported to have originated in the euryhaline
warm shallow coastal waters prior to their gradual invasion to estuarine,
brackish, fresh and all the reaches of marine, ecosystems. Although, none
have invaded the land, the bivalves are more successful in marine and a few
species are found in freshwater habitats. Nearly 652 species of marine
bivalves are reported from India, of which 88 species are endemic to Indian
waters.
Habitat
The adult bivalves are benthic or bottom dwelling, with varying levels of
evolutionary adaptations to the benthic habitat. This can be generally
classified as 1) buried in soft sediments within burrows, 2) cemented or
attached by byssal threads to hard substratum and 3) semi-mobile as part of
the epibenthos. Thousands of square kilometers of the shallow coastal waters
encompassing the estuaries as well as the backwaters are habitats for the
bivalves, catering to the regional fishery in India. In estuarine areas, clear
zonation in bivalve resources occur in relation to the salinity gradient, with
the stenohaline species inhabiting the areas near the bar-mouths.
Exploitation
The commercially important bivalves in India are the clams, mussel and
oysters. Clams are exploited from the soft substratum by hand-picking or by
using manually operated dredges. In shallow estuaries the clams are located
1
Molluscan Fisheries Division
Central Marine Fisheries Research Institute [CMFRI]
PO Box 1603, Ernakulam North PO|Kochi 682018 | Kerala | India
Tel: +91 484 2394867 | +91 484 2394794 (Per) |Cell: +91 9447056559
E-mail: [email protected], [email protected] |www.cmfri.org.in
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Species composition
Clams are the most important group among bivalves forming 85.8%, followed
by mussels 9.6% and oysters 4.6%. Commercially exploited clams are the
Villorita cyprinoides, Paphia malabarica, Meretrix casta, Sunetta scripta, Anadara
granosa, Meretrix meretrix, Marcia opima, Cardium sp., Anadara rhombea, Geloina
bengalensis, Gafrarium diverticulum, Gafrarium tumidum. Nearly 93.3% of the
contribution to the average annual clam production during 2009-2015 was by
three species, namely the V. cyprinoides, P. malabarica, M. casta. The Indian
backwater oyster, Crassostrea madrasensis is the most important edible oyster
exploited (90.1%) followed by the rock oyster, Saccostrea cucullata (5.9%) and
windowpane oyster Placuna placuna (3.6%) along the Indian Coast.
Commercial fishery of mussels along the Indian coast is mainly for the green
mussels, Perna viridis, contributing 83.7% on an average, and the remaining by
the brown mussel Perna indica which is limited to the fishery along southern
tip of Indian peninsula.
Production trends
The average edible bivalve annual production from 2009 to 2015 from the
coastal states of Kerala, Karnataka, Andhra Pradesh, Tamil Nadu and
Maharashtra, was estimated at 1.03 lakh tonnes. The estimated landings
decreased by 47% when compared to the period 1996-2000, which was
estimated at 1.52 lakh tonnes. Major share of the bivalve production of the
country is from the State of Kerala (85.8%) where, the clams form nearly 89%
of bivalve production in the State followed by mussels and edible oysters.
Vembanad and Ashtamudi Lakes in Kerala are the two main estuaries, which
have well organized clam fishery. The short-neck clam (P. malabarica) fishery
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for M. opima, M. meretrix, A. granosa and Donax sp. M. meretrix occurs for
industrial purposes.
Utilization
Clams and mussels are generally marketed as shell-on, whereas oysters are
transported to the important domestic markets and sold shell-on and as wet-
shucked oyster meat. India has been exporting bivalves, especially clam and
mussel meat to other nations. Bivalves fished along the west coast are utilized
for human consumption. Some bivalve products like smoked and canned
oysters have good market in Indian metro-cities. In Kerala, Tamil Nadu and
Andhra Pradesh, part of the clam landings is used as a major ingredient of
shrimp feed. The extensive shrimp farms also use dried and boiled clam meat
as shrimp feed. Apart from these, the shells of bivalves are used in the
manufacture of cement, calcium carbide, sand-lime bricks and lime. The lime
shell is used as manure in coffee plantations, as mortat in building
construction, in the treatment of effluents, as a pesticide by mixing with
copper sulphate and in glass, rayon, polyfibre, paper and sugar industries.
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Bivalve shells with attractive sculpture are used by the ornamental shell craft
industry.
Management
Bivalves offer one of the important examples of marine resource management
along the Indian coast. However, apart from the restriction on the pearl oyster
fishery by the Government of Tamil Nadu, and the management measures on
the short-neck clam fishery of Ashtamudi Lake, Kerala, there are no
regulations for effective utilization and conservation of these sedentary
marine resources. One of the major bivalve resources the short-neck clam (P.
malabarica) is well protected by the following regulations formulated by the
Government of Kerala based on recommendations made by Central Marine
Fisheries Research Institute (CMFRI): (a) ban on fishing activity during
breeding season (September to February), (b) use of gears with 30 mm mesh-
sized to avoid exploitation of smaller clam, (c) restrict the grade of export of
frozen clams meat to 1,400 nos/kg and above, and (d) initiate semi-culture or
relaying of small clams. This co-management model involving the policy
makers, fishers and researchers resulted in the short-neck clam (P. malabarica)
fishery in the Ashtamudi Lake in Kerala receiving India’s first Marine
Stewardship Council (MSC) certification.
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Summary of MSC Principle level scores for the Ashtamudi Estuary Short
Necked Clam Fishery:
MSC Fisheries Standards
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Ashtamudi Clam Fisheries: The First MSc Certified
Fishery from India
K.K. Appukuttan 1
Introduction
Ashtamudi lake (Lat. 8o 45’ – 9o 26’ N and Long 76o 28’ – 77o 17’E) is the
second largest and deepest wetland ecosystem in Kollam District of Kerala in
the South West Coast of India with a water spread of 61.4 km2 (6140 ha). The
estuary is palm shaped with eight prominent arms and the arms converge
into a single outlet at Neendakara to enter into Lakshadweep Sea. The Kallada
river which originate from the Western Ghats traverse through the forests
120km and empty the freshwater into the lake. The water spread area from
the Neendakara barmouth upto 6-7 km upstream is the estuarine part of the
lake with rich biodiversity (Fig.1).
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Fig.2. Landings and CPUE of P. malabarica from the Ashtamudi Estuary, 2002-2010.
[Source: Mohamed et al, 2012].
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Studies on age and growth show that it attain 30mm in 1st year, 38mm
in 2nd year and 41mm is 3rd year. (Appukuttan et al 1996). Individuals of 1st
year class followed by IInd year class dominate the catch and the maximum
size recorded is 52.3mm. The clam enter the fishery at 22 mm and sexual
maturity is attained at a mean size of 21mm and mature clams are found in
October-January period. Spawning commences in the post monsoon months
especially in the December-January period depending on the intensity and
duration of monsoon. The clams are profusely spawning in the first breeding
season and spawn twice in a year, the second being weak and low fecund.
The condition index of the clam (meat weight- shell weight index decline
sharply during the spawning season with peak condition index in June – July
period. Mohamed et al (2013) estimated the clam biomass as 24191.6 t of
which short neck clam, Paphia malabarica biomass is estimated as 21155.4 t
(87.4% of total biomas) followed by 3036.2 t (12.6%) by Meretrix casta. The
density of clam in different areas of the estuary varied considerably, the
maximum observed in areas nearer to barmouth. The length of clam in the
commercial catch varied from 25-37mm.
Fishing methods
Two major methods of clam harvesting is practiced by fishermen. They are
diving and gathering by hand or feet and second, hand dredging by one or
two persons using hand dredges. For both they use canoe to reach the clam
beds, except in shallow waters. In free diving and gathering the clam, they
use hand in collecting clams from shallow waters or use feet to dislodge clams
from the shell bed and gather it in the net bags with 30mm mesh size. When
they get appropriate quantity they climb back to the canoe, wash the net with
catch thoroughly and empty them to the canoe. This is repeated several times
and within 4-5 hours they collect approximately 200 kg a day Some of the
fishermen use googles to locate good shell beds (Fig 4&5). The hand dredges
is operated by two or three fishermen.
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Fig.6. Driving the hand dredge to the bottom Fig. 7. Fisherman dragging the bet bag
Fig.8. Washing the net bag Fig.9. A day’s catch.
The hand dredge has got an iron handle preferably of 5m long GI pipe
with a rectangular metal frame having a conical net bag with more than
30mm mesh size, attached to the stem. The frame has a toothless 60cm wide
blade in the lower side of the frame. The dredge is operated from the canoe by
one fisherman standing in the canoe and his collegue begin to haul the dredge
when the dredge strikes the bottom. The fisherman in the canoe begin to haul
the dredge by pulling the rope tied to the cod end of the net bag of the
dredge. This action drag the dredge through the bottom for a distance of 2-3
meters and when the dredge is full with the harvest it is thoroughly washed
repeatedly to discard the juvenile clams, mud and other organisms. The clam
which are retained in the bag is emptied to the canoe and the process is
repeated and a day’s catch can be 300-400 kg. (Fig.6,7,8 &9).
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World Wide Fund (WWF) since 1999 took special interest in applying Marine
Stewardship Council Certification as a conservation measure in small scale
fisheries. As an attempt towards certification, WWF-India took up few
community based MSC Certification programmes in India. Pre-analysis
model is one of the tool to assess the targeted fishery and select it for
Certification. Pre-assessment was carried out for shortneck clam Fishery
(Paphia malabarica) from Ashtamudi Estuary in the South West Coast of India
for certification.
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Step 2: Evaluate fishery against MSC’s principles and criteria for sustainable
fisheries and drafting the outcomes. Stake holders are given an opportunity to
input to the evaluation, client given the opportunity to respond to the draft
report, stakeholders given an opportunity to comment up on selection of peer
reviews. Peer review of draft report of the fishery and conditions,
stakeholders given an opportunity to give comments.
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V. Post Certification
Surveillance and enforcement of conditions of certification.
Surveillance audit required within 12 months after certification and annually
thereafter. The results of audit are to be made public via MSC website.
The present assessment used standard Assessment tree set out in MSC
certification requirement and for each performance indicator, the performance
of the fishery is assessed as a ‘score’. Some of the issues identified during
were stock status, harvest strategy, harvest control, and tools, information on
abundance of stock and monitoring (Principle-1) Information and
management strategy for retained non-targeted species and discarded
species, habitats and ecosystems (Principle 2) Long term fishery objectives,
decision making process, compliance and enforcement, research plan,
governance and management (Principle 3). These issues were addressed to
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References
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Taxonomy of Marine Molluscs of India: Status and
Challenges Ahead
Biju Kumar, A. 1 & Ravinesh, R.
Systematics
Mollusca represents the second largest animal phylum on our planet and
recent estimates show that the extant species diversity is around 45,000 to
50,000 marine 25,000 terrestrial and 5,000 freshwater (Appeltans et al., 2012;
Rosenberg, 2014; MolluscaBase, 2016). Originated in the early Cambrian
period almost 550 million years ago, molluscs entered almost every ecosystem
in the world, though their diversity is enormous in the marine realm,
occupying all habitats from the pelagic areas to the ocean trenches and
representing roughly one-quarter of the marine species described
(MolluscaBase, 2016). They are the morphologically megadiverse faunal
group in the marine ecosystem, exhibiting enormous diversifications in body
plan and habitat preferences and playing critical ecosystem roles, besides
forming a noticeable element in marine fisheries.
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other phyla (Halanych et al., 1995; Sigwart and Sutton, 2007). The existing
‘aculiferan’ model of molluscan phylogeny considers two subphyla,
Conchifera, the ‘shell-bearing’ molluscs, and Aculifera, ‘spiny’ molluscs
(Aplacophora and Polyplacophora), while ‘testarian’ model group
polyplacophorans together with conchiferans under Testaria (Sigwart and
Sutton, 2007). The consensus model considers the traditional concepts of
Aculifera and Conchifera, while Scaphopoda was considered in the clade
along with Gastropoda and Bivalvia and the details are given in Fig. 1 (Stöger
et al., 2013).
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During the latter half of the 19th century Abercrombie (1893a,b), Adam
(1939), Alder and Hancock (1864), Comber (1906), Crichton (1940, 1941), Eliot
(1903), Hornell (1921, 1949), Hoyle (1905), Melvill (1892, 1893, 1893-1894, 1898,
1909), Melvill and Ponsoney (1898), Melvill and Standen (1898), Prashad
(1930, 1932), Preston (1910, 1911), Robson (1926), Smith (1878, 1894, 1895,
1896,1899, 1903, 1904, 1906), Thurston (1890) and Winckworth (1927a,b, 1928,
1929, 1936, 1940) contributed knowledge to malacological studies. This was
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Phase 2: The beginning of the 20th century was the most productive and
significant period in the history of Indian malacology, with the lead role taken
up by Zoological Survey of India (ZSI), along with contributions from Central
Marine Fisheries Research Institute (CMFRI) and several maritime
universities of India. This helped consolidating information on marine
molluscs of India. Few general surveys on the taxonomy of the molluscan
fauna of Indian coasts are those of Appukuttan (1983), Appukuttan et al.
(1989), Apte (1993), Apte (1998), Bertsch and Attilio (1980), Hornell (1951),
Hornell and Tomlin (1951), Kohn (1978), Kundu (1965), Kurian (1948), Menon
et al. (1961 and 1967), Mookherjee (1985), Nagabhushanam and Rao (1972),
Narayanan (1968), Panicker (1977, 1978), Pinn (1990), Rajagopal and
Mookherjee (1978, 1982), Ramakrishna and Dey (2000), Rao (1970, 1977), Rao
(1980), Rao and Dey (1984, 1986), Rao and Rao (1981, 1991, 1993), Ray (1949 a,
b; 1951, 1956), Rockel et al. (1995), Santhakumaran (1973), Satyamurti (1952,
1956), Silas et al. (1985), Rao et al. (1991, 1992), Subrahmaniyan et al. (1952) and
Tikader et al. (1986). This phase witness good quality publications, especially
from Dr NV Rao and his team from ZSI based on exploratory studies on
various ecosystems of India.
Major publications during the phase include those of Apte (2004, 2009,
2012), Apte and Bhave (2014), Apte et al. (2010, 2012), Bhave and Apte (2011,
2013), Bijukumar et al. (2015), Carmona et al. (2014), Dey (2006), Dey and
Ramakrishna (2007), Franklin et al. (2009), Hylleberg and Kilburn (2002),
Lutaenko (2006), Modayil (2007), Mukhopadhyay et al. (2012), Pati and
Sharma (2012 a, b), Raghunathan et al. (2010), Ramakrishna and Dey (2000,
2003, 2010), Ramakrishna et al. (2007, 2010), Rao and Sastry (2005), Sreeja et al.
(2012, 2016), and Venkataraman et al. (2004, 2012).
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Gap Areas
Taxonomic impediment
The taxonomic impediment prevailing in other parts of the world is
effervescent in India as well and many institutions working on faunal surveys
and documentation lack the globally competent malacologists to carry out
extensive surveys and identification, not to speak of infrastructure facilities to
support such exploratory research. As revealed by the analysis of publications
on taxonomy from the country in the last two decades and analysing the
vision documents of marine research institutions, especially in the public
sector, taxonomy is not projected as a priority item. If at all proposals are
placed in paper, no strategies and action plans were suggested to overcome
the taxonomic impediment. Further, human resources in taxonomy for
satisfying the ever growing demands from various sectors, including marine
bioprospecting and biotechnology, is abysmally poor even in institutions
dedicated to biodiversity documentation.
Database
While analysing the recent works on molluscan taxonomy in India, the major
lacuna is the lack of a good quality updated data base on molluscs of India in
the public domain. For example, the available checklists including that of the
scheduled species in India document the synonyms and are not taxonomically
validated, further contributing to the chaos in taxonomy. As a foundation
element of biology, it is imperative that taxonomy is practised in a highly
professional manner, as dubious taxonomy destabilizes the foundation of
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science, with potentially serious setback in basic and applied research, and
therefore publications in predatory journals hamper development of
taxonomy in India (Rajeev et al., 2014). Therefore, publications that appear in
predatory journals, without even mentioning anything on voucher specimens
and accession numbers would not support taxonomic research. The existing
databases have to be strengthened by validating species identity of all the
collections by the research vessels of various organisations in India. Good
quality handbooks and field guides of various classes of Mollusca form
another requirement for strengthening taxonomic research in India.
Ecosystem/taxon based studies
In India majority of the molluscan studies were conducted in coral reef
ecosystems. Majority of the surveys were conducted as part of compilation of
data for general biodiversity data bases or all-phyla studies. Extensive
surveys are required along continental shelves, sea mounts and deep seas
along Indian coast. Ecosystem-based in-depth surveys are required to
document species diversity of coral reefs, lagoons, mud flats, sandy beaches,
estuaries and backwaters, intertidal and subtidal ecosystems.
biodiverse groups collected from Indian waters. Deep sea molluscs off Indian
coast is yet another priority area for consideration.
Integrative taxonomy
‘Integrative taxonomy’ is defined as the science that aims to delimit the units
of life's diversity from multiple and complementary perspectives
(phylogeography, comparative morphology, population genetics, ecology,
development, behaviour, etc.) (Dayrat, 2005). Molecular analyses play a very
important role in elucidating extent, origin and history of marine biodiversity,
and molecular techniques provide adequate information regarding the
phylogenetic relationships and divergence times of evolutionary lineages and
clades. Understanding the distribution and origin of diversity in the larger
marine, especially Indo-Pacific is a fundamental problem in biogeography.
Further, molecular studies would also facilitate identification of cryptic
species and speed up the process of biodiversity documentation. Integrative
taxonomic studies involving molluscan species should also be promoted to
fully realise the diversity of marine molluscs of India. There is a need to
develop specific course content focusing on ‘integrative taxonomy’ that needs
to be taught first before training in systematics (Pisupati, 2015).
‚Making taxonomy a combined study and science that brings on board non-
experts and non-biologists to support identification of species as a hobby,
passion and love for nature with support coming from trained scientists‛
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(Pisupati, 2015). In India the possibility of involving citizen scientists and civil
society in biodiversity documentation were not fully explored, though
opportunities for such an exercise are awesome. Long term biodiversity
monitoring studies and preparation of inventories can be tried by expanding
the network of local communities and civil societies.
Repositories
The depositions in the natural history museums and repositories reveal the
great natural history and biodiversity of the nation and a source material for
the taxonomists and biotechnologists to pursue their research. It also
provides identification services on natural objects and rich fauna, flora and
minerals resources to user groups. The priority therefore should be to prepare
a database of type materials available in each of the repository and to simplify
the procedure for sharing the data to practicing malacologists.
Conclusions
Despite their high diversity and importance for humankind marine molluscs
are not given due priority by researchers in India, as reflected in the lesser
number of practicing taxonomists involved in the process. This situation can
be improved only by taking concerted efforts in the following key areas: (i)
molluscs and their critical ecosystem roles should be brought to the attention
of general public in order to remove the public dilemma in this subject; (ii)
policymakers and stakeholders are mostly unaware of conservation problems
involving marine organisms and betterment of this political dilemma can be
done only by involving them in conservation thinking through practical
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Preston, H.B. (1910). Description of five new species shells from the Bay of
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Robin, A., 2008. Encyclopedia of Marine Gastropods. 480 pp.
Eichhorst T.E. (2016). Neritidae of the world. Volume 1. Harxheim:
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Yonow, N. (2008). Sea slugs of the Red Sea. Pensoft Publishers, Sofia-Moscow,
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descriptions of two new species and ten new records (Mollusca,
Gastropoda). ZooKeys, 197: 1 129.
Huber, M. 2015 Compendium of Bivalves 2. A Full-Color Guide to the
Remaining Seven Families. A Systematic Listing of 8'500 Bivalve
Species and 10'500 Synonyms 907 pp.
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Promurex, Haustellum, Bolinus, Vokesimurex and Siratus. Harxheim:
ConchBooks. 197 pp.
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Gosliner, T.M., Valdés, A. & Behrens, D.W. 2015. Nudibranch and Sea Slug
Identification - Indo-Pacific. New World Publications, Jacksonville,
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Gosliner, T. M., Behrens, D. W., & Valdés, A. 2008. Indo-Pacific Nudibranchs
and Sea Slugs: A Field Guide to the World’s Most Diverse Fauna. Sea
Challengers, Gig Harbor, WA & California Academy of Sciences, San
Francisco, CA, 426 pp.
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Molecular approaches in Taxonomy with special
reference to Bivalve Molluscs
Hari Krishnan K.1 & Arjun J. K.
Introduction
Taxonomy, the study of classification of organisms, is the basis for all
meaningful studies on biodiversity, pest management, medicine,
bioprospecting and fisheries. Traditional taxonomy relies upon the
phenotypic characteristics of organisms to differentiate them in to different
groups and species. More than 1.5 million species of animals, plants and
microorganisms have been reported so far and it is estimated that the number
of undescribed living species could be more than 3 million. Taxonomists
claim that at least 50 million species of different organisms have become
extinct in this long course of evolution. Organisms possess unique attributes
to enhance their survival in a particular environment. The genetic variation
inherent in an individual or in a population will act as the driving force
behind this survival strategy. Molecular taxonomy is a relatively recent
branch of science, which contributed tremendously to modern taxonomy and
has improved a lot over the last decade with the advancement in DNA
sequencing technology.
The class Bivalvia consists of more than 20,000 species with a wide
distribution both in freshwater and marine environment. The bivalves are
economically important since they are important source of sea food and
precious pearls. Estimates shows that a vast majority of bivalve species are yet
to be unexplored in nature, its discovery with new molecular techniques will
certainly expand the knowledge on their biodiversity. The application of
DNA markers has allowed rapid progress in Bivalve taxonomy, such as in
investigations on genetic variability and inbreeding, parentage assignments,
species and strain identification and the construction of high-resolution
genetic linkage maps for aquaculture species.
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Protein markers
Genetic variation at protein level that is encoded by DNA can be identified by
Allozyme electrophoresis (Goodman, 1982). Allozymes are variant forms of
an enzyme that are coded by different alleles at the same locus and they are
highly conserved among various groups of organisms. An Allozyme study is
simple, cost effective and does not require specialized equipments. Genetic
variants of Allozymes differs slightly in electric charge that can be detected
through electrophoresis in an acrylamide or cellulose acetate gel. Thus
Allozyme analysis helps in the study of molecular phylogeny based on a
single locus genetic variation. Any protein which is soluble in nature can be
used for Allozyme analysis. Allozymes from tissues can be extracted with
suitable protocols and protein isolation kits. Individuals that are homozygous
show a single band where as heterozygous show two bands. The limitations
of this technique include requirement of a large amount of tissue and
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consequently this method could not be applied when the organisms are
small and also in case of larval forms.
DNA markers
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over 1400 different species of eukaryotes. In most cases, EST projects are used
either to complement existing genome projects or serve as low-cost
alternatives for purposes of gene discovery. However, with improvements in
accuracy and coverage, they are beginning to find application in fields such as
phylogenetics, transcript profiling and proteomics (Parkinson and Blaxter
2009). Fast and reliable analysis can be made for the genes expressed in
particular tissue types under specific physiological conditions or
developmental stages. Differentially expressed genes could be identified
using cDNA microarrays in a systematic way (Sukumaran and
Gopalakrishnan 2015).
Next Generation Sequencing (NGS) is a powerful tool for elucidating the true
diversity and species richness. Mitochondrial genomes comprise a small but
critical component of the total DNA in eukaryotic organisms. They encode
several key proteins for the cell’s major energy producing apparatus, the
mitochondrial respiratory chain. NGS of mitochondrial genome of higher
vertebrates like fishes, frogs and plants are widely used for molecular
taxonomy and phylogeny studies (Lloyd et al, 2012). NGS of 18S amplicons
and microsatellites reveals a previously hidden taxonomic richness, especially
for Copepoda and hard-to-identify meroplankton such as Bivalvia,
Gastropoda and Polychaeta. It also reveals rare species and parasites. While
this approach allows for broad diversity assessments of plankton it may
become increasingly attractive in future if sequence reference libraries of
accurately identified individuals are better populated (Lindeque et al, 2013).
The first step in any molecular taxonomy work is the collection of appropriate
tissue sample from the whole specimen. Fresh tissue samples are preferred
instead of any samples preserved in alcohol or salt which will help in the
isolation of DNA in good quantity and quality. It is very difficult to isolate
high quality pure DNA from bivalves since the mollusc tissue has high
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Mitochondrial DNA evolves faster than nuclear DNA and has a higher
nucleotide mutation rate, excelling the evolution rate of the nuclear
genome DNA. This can be realized by analyzing haplotypes which
present unique genetic marker combinations in a chromosome. Haplotypes
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are different from each other for one or more nucleotides due to substitution,
insertion or deletion phenomena. It has been proven that the presence of
nuclear pseudogenes of mitochondrial origin (also named numts) should
be taken into consideration in mtDNA study (Ballard and Whitlock, 2003).
Because they are present in the majority of eukaryotes, numts shouldn’t
be interpreted real mitochondrial genes. Another disadvantage of
mitochondrial markers is that they may behave as a single molecule with a
unique evolutionary history, leading to an overestimation of the evolution
parameters (Ballard and Whitlock, 2003). Mitochondrial markers are not
representative for the evolutionary history of a species. Some of the most
frequently used regions of the mtDNA for phylogeny studies are
cytochrome b (cytb) and subunit I of the cytochrome c oxidase (COI)
(Feral, 2002). Cytochrome c oxidase represents the most used marker in
molecular studies and barcoding (Hebert et al., 2003).
ITS2 is situated between the 5.8S and 28S nuclear ribosomal genes and
constitutes a DNA fragment which evolves rapidly and found to be very
useful in the analysis of phylogenetic relations between closely related species
of bivalve mollusks (Freire et al., 2010). ITS regions can be sequenced directly
or can be cloned and sequenced. Majority of the studies on ITS1 and ITS2 in
bivalves usually involves cloning of the ITS2 region in order to separate two
or more individual (Freire et al., 2010; Vierna et al., 2010), while others
favoured direct sequencing (Flot et al., 2006; Ladhar – Chaabouni et al.,
2010). Direct sequencing is more accurate and cost effective compared to
cloning and sequencing.
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online bioinformatic tools available in the public domain that can be used to
analyse the sequences: Barcode of Life Data Systems (BOLD)
http://www.barcodeoflife.org and National Center for Biotechnology
Information (NCBI) Basic Local Alignment Search Tool (BLAST). After
sequence homology search, the sequences can be aligned and edited using
tools like BioEdit, ClustalW etc. Phylogeny analysis can be done by creating
phylogeny trees using softwares like MEGA, PHYLIP. Expressed Sequence
Tags (ESTs) and gene expression micro array data for marine organisms
which provides a unique collection for marine specific EST and micro
array data and is currently available at http://www.marinegenomics.org
which can also be employed for the systemic study of molluscs. Some global
and regional biodiversity catalogues for molluscs are available- Molluscs,
OBIS (http://www.amonline.net.au/invertebrates/) European Molluscs
database (http://www.mnhn.fr/base/malaco.html) which can also be
employed in the diversity studies of molluscs.
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References
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Espineira, M., González – Lavín, N., Vieites M and Santaclara, F.J. 2009.
Development of a method for the genetic identification of commercial
bivalve species based on mitochondrial 18S rRNA sequences. Journal of
Agricultural and Food Chemistry, 57,495-502.
Etter, R. J., Boyle, E. E., Glazier, A., Jennings, R. M., Dutra, E and Chase, M. R.
2011. Phylogeography of a pan-Atlantic abyssal protobranch bivalve
Implications for evolution in the Deep Atlantic. Molecular Ecology,
20,829-843.
Feral, J.-P. 2002. How useful are the genetic markers in attempts tounderstand
and manage marine biodiversity? Journal of Experimental Marine Biology
and Ecology, 268,121-145.
Flot, J.-F., Tillier, A., Samadi, S and Tillier, S. 2006. Phase determination from
direct sequencing of length variable DNA regions. Molecular Ecology
Notes, 6,627 - 630.
Freeland, J.R. 2005. Molecular Ecology. John Wiley and Sons, Chichester,
England. 388 pp.
Freire, R., Arias, A., Mendez, J. and Insua, A. 2010. Identification of european
commercial cockles (Cerastoderma edule and C. glaucum) by species-
specific PCR amplification of the ribosomal DNA ITS region. European
Food Research and Technology, 232,83-86.
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Giribet, G and Distel, D. L. 2003. Bivalve phylogeny and molecular data. In,
Molecular Systematics and Phylogeography of Mollusks (Lydeard, C
and D. R. Lindberg eds) Washington, Smithsonian Books. pp. 45-90.
Huff, S. W., Campbell, D., Gustafson, D.L., Lydeard, C., Altaba, C. R. and
Giribet, G. 2004. Investigations into the phylogenetic relationships of
freshwater pearl mussels (Bivalvia, Margaritiferidae) based on
molecular data, Implications for their taxonomy and biogeography.
Journal of Molluscan Studies, 70, 379-388
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Lindeque PK, Parry HE, Harmer RA, Somerfield PJ, Atkinson A (2013) Next
Generation Sequencing Reveals the Hidden Diversity of Zooplankton
Assemblages. PLoS ONE 8(11): e81327
Liu, Z. J., & Cordes, J. F. (2004). DNA marker technologies and their
applications in aquaculture genetics. Aquaculture, 238(1), 1-37.
Lloyd, R. E., Foster, P. G., Guille, M., & Littlewood, D. T. J. (2012). Next
generation sequencing and comparative analyses of Xenopus
mitogenomes. BMC genomics,13(1), 1.
Mock, K. E., Brim-Box, J. C., Miller, M. P., Downing, M. E and Hoeh, W.R.
2004. Genetic diversity and divergence among freshwater mussel
(Anodonta) populations in the Bonneville Basin of Utah. Molecular
Ecology, 13,1085-1098.
Ni, L., Li, Q and Kong, L. 2011. Microsatellites reveal fine-scale genetic
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Mactridae) in Northern China. Marine Ecology, 1-10.
Olu-Le Roy, K., von Cosel, R., Hourdez, S., Carney, S. L and Jollivet, D. 2007.
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Ponder, W. F and Lindberg, D.R. 2008. Phylogeny and evolution of the Mollusca.
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Popa, O. P., Iorgu, E. I., Krapal, A. M., Kelemen, B. S., Murariu, D and Popa,
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Bivalvia, Cardiidae). Internaional Journal of Molecular Sciences, 12,456-
461.
Qin, Y., Liu, X., Zhang, H., Zhang, G and Guo, X. 2007. Identification and
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1819). Marine Biotechnology, 9,66-73.
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Wang, Y., Wang, X., Wang, A and Guo, X. 2010. A 16-microsatellite multiplex
assay for parentage assignment in the eastern oyster (Crassostrea
virginica Gmelin). Aquaculture. S28-S33.
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(Crassostrea virginica Gmelin) using AFLP and microsatellite markers.
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Ecology, 15, 639-651.
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Sampling Techniques for molluscan fauna
S. Bijoy Nandan 1, Jayachandran P. R., Asha C.V.
Introduction
Sampling techniques must be designed to take into account the fact that
benthic organisams are extremely patchy in distribution. Patchiness is caused
by processes external to the assemblage, particularly disturbances and
recruitment, in addition to processes operating within the existing
assemblage. Although anthropogenic disturbances are often very severe,
natural disturbances are common and important contributors to spatial
variability of populations. Numerous techniques have been developed for
collecting, sorting and handling for the study of molluscs. Live and dead
samples can be collected for studies, even though most collectors prefer the
live specimens. Live samples have less physical damage than ones that are
dead and exposed to the action of the surf, however dead shells are preferred
by some collectors since the animal is not to be killed and the preparation of
the shell requires less work. In deeper water and pelagic species forms are
most commonly found dead and found on the shore after storms or in dredge
spoil. Some groups require special methods of collection from various
substrates. Methods of sampling the bivalve population vary with the types of
substratum. Organisms living within the sediment must be dug out using
grabs or corers. Hard substratum living forms can usually be sampled by
divers or photographed using remote cameras or submersibles. Quantitative
studies of benthic population requires specially designed samplers which take
a standard bite of standard depth and area.
Hand collecting methods: Shell-rich portions of the sand are referred as shell
grit or shell sand. They can simply be scooped up and processed like any
other sediment samples. They usually have mainly empty shells and
sometimes can be the only source of certain species. The fine sand can be
removed from these samples by washing it in a test sieve or fine mesh bag.
While the bag method works well for the larger micromolluscs, it will lose
many of the smaller species.
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Algae associated molluscs: Algae (seaweeds, seagrass, algal mates etc.) are a
habitat of many molluscs. Certain molluscs are found only on particular algal
species. Larger algal species harbour many micromolluscs, particularly in
holdfasts. Larger algal species can be placed in a bucket, smaller species may
fit into a ‘zip-lock’ bag. For bulk collection, the simplest technique is to
vigorously wash algae on the shore in a bucket or bowl filled with seawater.
The algal material is then removed and the sample allowed to settle briefly.
The water can then be gently decanted, being run through a sieve to catch any
floating molluscs. Such samples are ideal for collecting living specimens for
later examination. More thorough washing can be carried out by vigorously
shaking algae in a 0.5–1 litre jar half filled with water from the environment
and with a tightly fitting lid. Samples may also be pre-treated with an irritant
or narcotic to ensure detachment of specimens, to be released from the
substrate.
Leaves and other litter: Rich organic material provides molluscan habitats
particularly in mangrove and other upper littoral and supralittoral habitats as
well as terrestrial habitats. Mangrove litter can be sieved, washed and to
retain the samples.
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Trawls and dredges (qualitative): Trawl nets are designed to skim over the
bottom and as they cover a large area, they have a good chance of collecting
widely dispersed species. A net much used for biological work is the Agassiz
trawl, which has the advantage of very easy handling because it does not
matter which side up it reaches the bottom. The mouth of this net is held open
by a metal frame, and it can be fitted with fine-mesh net to retain small
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creatures. It is simply dragged along the bottom. To capture animals that live
beneath the surface, the sampling device must be capable of digging into the
deposit. The naturalist’s dredge is a simple device which can be operated
from a small boat. It consists of a bag of strong sacking or wire mesh held
open by a heavy, rectangular metal frame. This can bite a few inches into a
soft sediment as it is hauled along, but tends to fill mainly with material lying
on the bottom and does not catch the deeper-burrowing creatures. An
example of an instrument that takes a considerably deeper bite is the Forster
anchor dredge. The dredge is lowered to the bottom and remains stationary
as the ship moves slowly as tern paying out a long length of cable, three to
five times the depth of the water. The winch brakes are then applied to the
cable, and the strain exerted as the ship is stopped causes the dredge to tilt
and bite deeply into the substrate. So instead of sliding along the bottom the
dredge digs in like an anchor to a depth of about 25 cm. Finally, the cable is
winched in until the dredge eventually breaks out, the contents being retained
within the net. For ease of use in deep water this type of dredge can be made
with digging-plates on both sides of the arm, so that it will bite whichever
way up it lands on the bottom. Beam, Agassiz and trawls used for qualitative
sampling of the epifauna. These nets are designed to skim over the surface of
the bottom, and because of the large area covered, are useful for collecting
scarcer members of the epifauna, species of mollucans, associated with the sea
bottom.
Bottom sledges (Epibenthic sledges): Epibenthic sledges are widely used for
sampling especially in the deep sea. The simplest sledge consists of a mesh
bag or bags mounted in a metal frame on runners. The collecting bag is
protected inside a steel mesh cage. The angle and height of the cutting plates
on the mouth of the frame can be adjusted and the sledge can operate either
way up. More sophisticated sledges have various instruments mounted on
the top and so must operate the right way up. Cameras photograph the
bottom area ahead of the sledge. Acoustic devices indicate the position of the
sledge relative to the bottom, and an odometer wheel coupled to a
potentiometer measures the distance the sledge has travelled over the sea-
bed.
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Box samplers and corers: Because of their reliability, box corers and samplers
have been extensively used in the deep sea. However, their use in shallower
water studies is equally relevant. Box corers provide a means of obtaining
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members of the epifauna. After storms, burrowing animals are often washed
in from shallow water onto the beach, and this may be the best means of
obtaining some deep-burrowing species not readily taken by ship-borne
samplers. Empty mollusc shells and other recent remains cast up on the beach
are usually some guide to the nature of the shallow-water offshore fauna.
There is still a need for the development of methods for their assessment,
although photographic methods give good results for some species. Other
methods: Floating light traps are used for collecting samples of Scissurellidae.
The use of attractants (light, bait) may be worth exploring, particularly. Small
grab samplers have been used for the collection of micromolluscs. Air-lift
pumps can be used as a very effective way of sampling both hard surfaces
and substrate and are also a means of obtaining, with careful and targeted
use, large quantities of living specimens. Dredging and benthic sledge can
provide significant amounts of material and sample a larger area than either
grab or air-lift pump. The benthic sledge is advantageous as it only skims the
top surface where most micromolluscs are found, but infaunal taxa will
largely be missed.
Pins and needles: Needles are probably the most important piece of
equipment for radular and many other micromollusc applications.
Forceps: The forceps are excellent for handling specimens. It can used for
routine sorting, including handling fragile specimens.
Hairs: For cleaning dust particles from mounted radulae it is often better to
use hairs (rather than a pin) glued to a small handle of wood or inserted into a
holder. Eyebrow hairs (if straight) and eyelashes are commonly used but
some animal hairs such as the pointed and stiff whiskers of a cat make very
good tools. The stiffness of hairs decreases with increasing length of the hair.
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Pliers: The smallest sizes of regular tool pliers are useful for cracking and
opening small shells. Locking vice grip pliers will prevent the specimens
being crushed.
Drills: Some power tools resembling a dentist’s drill have a flex-shaft
attachment and a variety of rotary tool bits, engraving cutters and high-speed
cutters that can be used to open shells with minimal damage.
Cavity slides: Cavity slides are useful for working on microscopic animals.
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Storage: The most commonly used storage medium is 70–80% ethanol. Borax,
powdered aragonite, or powdered calcite/ shells may be added to ethanol
solutions to safeguard against shell damage. Long-term storage in formalin is
generally avoided since formalin is on the list of suspected carcinogens and is
a well-known allergene. However, it has been successfully achieved, where
5% seawater formalin buffered with NaHCO 3 is used. For any preservative,
the pH needs to be kept below pH 8.5 to avoid tissue dissolution and above
pH 7 to prevent shells and other exoskeleton parts from dissolving. For field
storage, unbreakable plastic containers, ideally with screw tops, should be
used. Eppendorf tubes (1.5 ml) and Falcon tubes (10 or 50 ml) seal fairly well,
as long as the seal is free of dirt. Heat-sealed bags can leak when filled with
wet sediment samples, but are useful as secondary containers. Although
cheap, scintillation vials should be avoided, because ethanol evaporates
quickly from them.
Specimen bottles: The specimens, even those collected as empty shells, must
be properly cleaned and free of salt. This can be done by soaking them in
clean water for a few hours and then drying thoroughly. The dried specimens
are best kept in small containers: Ideally gelatine capsules within larger high
sodium glass vials, or cardboard mounts made from acid free, archival quality
board.
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References
Daniel l. Geiger, Bruce A. Marshall, Winston W. Ponder, Takenori Sasaki &
Anders Waren, 2007. Techniques for collecting, handling, preparing,
storing and examining small molluscan specimens, Molluscan Research
27(1): 1–50
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Glossary
Accessory plate - calcareous and periostracal structure covering the soft parts in the
Pholadidae, in addition to the shell valves.
Adductor muscle - muscle connecting the 2 valves of a shell, tending to draw them
together.
Apophysis - finger-like shelly structure to which the foot muscles are attached in the
Pholadidae and Teredinidae.
Branchial - pertaining to the gills.
Branchial lamella - (see gill).
Byssus - clump of horny threads spun by the foot, by which a bivalve can anchor to a
hard substrate.
Cardinal area - surface of the shell extending between umbo and hinge margin.
Cardinal tooth - (see tooth).
Chomata - marginal crenulations in Ostreidae and Gryphaeidae, occurring all around
the inner side of valves or only near the hinge, composed of small tubercles or
ridge lets on the right valve, and corresponding pits on the left valve.
Commissure - line of junction of the valves.
Concentric - parallel to lines of growth.
Cruciform muscles - crossed muscles connecting valves and serving to retract the
siphons, leaving 2small scars near the posteroventral end of pallial line in some
bivalves (e.g.Tellinidae).
Ctenidial axis - (see gill).
Ctenolium - a row of small teeth on lower side of byssal notch in some Pectinidae.
Demibranch - (see gill).
Denticle - a small tooth.
Ear - lateral expansion of the dorsal part of a shell.
Equilateral - the condition of a valve when growth on either side of umbo is
symmetrical.
Equivalve - the condition of a shell when valves are of the same shape and size.
Escutcheon - differentiated area extending along dorsal margin of valves, behind the
umbones.
Eulamellibranchiate type - gill demibranchs composed of 2 lamellae. Branchial
filaments and lamellae always connected by tissular junctions (e.g., Veneridae).
Filibranchiate type - gill demibranchs composed of 2 lamellae. In addition to the ciliary
junctions between branchial filaments, anastomosed tissular junctions may unite
lamellae of each demibranch (e.g., Mytilidae,Pectinidae).
Foot - mobile and extensible muscular organ, used for locomotion or for attachment to
substrate by means of byssal threads.
Gape - opening or gap remaining between margins of valves, when shell is closed.
Gill - respiratory organ generally composed of 2 thin leaf-like structures (demibranches)
suspended to a dorsal axis (ctenidial axis); each demibranch may be either simple
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or bent back upon itself and then formed of 2 sheets (branchial lamellae). A
lamella is constituted of many ciliated filaments parallel to each other and
interconnected by more or less complex junctions. Four main types of gill
structures are currently recognized among bivalves: the protobranchiate,
filibranchiate, eulamellibranchiate, and septibranchiate types (see these terms).
Growth marks - (see sculpture).
Hinge - structures in the dorsal region of the shell, along which the valves meet, and
that function in the opening and closing of the shell.
Hinge line - shell margin adjacent to the hinge.
Hinge plate - in folding of dorsal shell margin bearing hinge teeth and sockets, and
lying in each valve in a plane parallel to that of junction of valves.
Imbricate - overlapping like tiles or shingles on a roof.
Inequilateral - the condition of a valve when growth on either side of umbo is
assymmetrical.
Inequivalve - the condition of a shell when valves are not alike in shape or size.
Keel - a prominent, angular ridge.
Lamellate - with thin, flattened plates.
Lateral tooth - (see tooth).
Lenticular - shaped like a biconvex lens.
Ligament - horny, elastic structure joining the 2 valves dorsally.
Ligamental area - part of cardinal area occupied by the ligament.
Lunule - differentiated area extending along dorsal margin of valves, just in front of
umbones.
Mantle - fleshy sheet surrounding vital organs and composed of 2 lobes, one lining and
secreting each valve.
Muscle scar - impression marking the place of attachment of a muscle inside the shell.
Nacreous - pearly, often with multi-coloured hues, as in mother-of-pearl.
Nymph - narrow plateform extending behind umbo along dorsal margin, to which the
external ligament is attached.
Opisthogyrate - the condition of a shell when umbones are directed posteriorly.
Orbicular - disk-shaped, nearly circular.
Orthogyrate - the condition of a shell when umbones are perpendicular to the hinge line
(directed neither anteriorly nor posteriorly).
Pallet - small paddle-shaped or feather-like calcareous and periostracal structure, a pair
of which closes the burrow opening when siphons are retracted in the
Teredinidae.
Pallial - pertaining to the mantle.
Pallial line - a line near internal margin of valve, marking the site of attachment of the
mantle edge.
Pallial sinus - posterior indentation of pallial line, marking the site of attachment of
muscles allowingsiphons to retract within the shell.
Pedal - pertaining to the foot.
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