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Training manual -1st International Training Workshop on Taxonomy of Bivalve


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TRAINING MANUAL
ST
1 INTERNATIONAL TRAINING WORKSHOP ON
TAXONOMY OF BIVALVE MOLLUSCS
TRAINING MANUAL
ST
1 INTERNATIONAL TRAINING WORKSHOP ON
TAXONOMY OF BIVALVE MOLLUSCS

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
st
Training manual -1 International Training Workshop on Taxonomy of Bivalve Molluscs.

Editors
Bijoy Nandan
P. Graham Oliver
Jayachandran P.R.
Asha C.V.

© Copy rights reserved by the authors,


First published in May 2016

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
st
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
st
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
st
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

There is an increasing lacuna on taxonomists, taxonomic collections, and institutional facilities


worldwide, realizing this there is an urgent need for concerted efforts to alleviate the situation, so
as to facilitate and assist countries in implementing the Convention on Biological Diversity. For
this, a regional to sub regional network is to be established and strengthened among various
institutions at the national and international level among the developing and developed countries.
Such actions will provide a platform for capacity building for developing the skills and expertise
in analyzing the taxonomic collections at the grass root level, from the field based data collector
to the scientific personal. Further capacity-building for taxonomy should be linked to the
effective implementation of the Convention on Biological Diversity, particularly the national
identification of areas of high diversity to threatened taxa that are or may be of value to
humanity, and those with potential use as biological indicators for conservation and sustainable
use of biological diversity. Capacity-building in taxonomy is also critical to address invasive alien
species and achieve Aichi Biodiversity Target 9, and it supports all other elements of the Strategic
Plan for Biodiversity 2011-2020. This must necessarily include training of the next generation of
taxonomists, appropriate leadership for natural history collections and the involvement of
practicing taxonomists to change the perception of taxonomy by decision-makers.

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.

Ernakulam, Kerala Editors


May, 2016
ACKNOWLEDGEMENTS

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

Our knowledge of the living species on the earth is still dramatically


incomplete. Taxonomists have so far described less than 2 million species,
whereas, using various methods, the total number of species was estimated to
at least 7–8 million, but perhaps 10, 50, 100 million or even more
(Groombridge, 1992; Heywood & Watson, 1995; Reaka-Kudla et al., 1997) little
known about nature and ecological roles of species identified. This huge
taxonomic gap (Dubois, 2010a) is both quantitative and qualitative, as very little
is known of most of the species that have been “described” and named so far
(Dubois, 1998). If the work of increment of our database on specific diversity
continued at the same pace as in the past, centuries would be necessary to
complete our inventory of the planet’s species (González-Oreja, 2008).

We are degrading and destroying biodiversity in many parts of the


world, and these threats are increasing. Species are becoming extinct 100 to
1,000 times faster than they were before modern humans arrived on the earth
(the background rate), and by the end of this century, the extinction rate is
expected to be 10,000 times the background rate. Conservative estimates of
extinction = 0.01-1.0%. The rates of extinction are higher where there are more
endangered species however, the estimating extinction rates is not an easy
subject.

There is a taxonomic urgency (Dubois, 2010a), i.e., the imperious need of


an acceleration of our taxonomic inventory of the planet (Wheeler et al., 2004),
is currently far from having been identified by our governments, by the
decision-makers in matters of scientific research and by most scientific
institutions. It is of utmost importance to consider the discovery, collection,
storing in collections, study and description of specimens and tissues of the
still unknown species of the biosphere, before they turn extinct, as an absolute
priority for biology in our century of extinctions (Dubois, 2003).

Although there is no agreement among scientists about the estimation


of the number of unknown species, it is estimated that about 90% of the world

1 Former Director, Zoological Survey of India


E-mail: [email protected]

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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.

Godfray (2002) indicates that taxonomy, the classification of living


things, has its origins in ancient Greece (with the first basic classification of
Aristotle) and in its modern form dates back nearly 250 years, to when
Linnaeus introduced the binomial classification still valid today. Specific rules
have been established for recognising, naming and classifying species to
avoid redundant descriptions or the use of the same name for more than one
species. These rules were introduced in the late 19th century and are
continuously monitored by international commission scientists (Tautz et al.,
2003). The discipline of taxonomy traditionally covers three stages: alpha
(analytically phase), beta (synthetic phase) and gamma (biological phase)
taxonomy (Kapoor, 1998; Disney, 2000). Alpha taxonomy is the level at which
the species are recognised and described; beta taxonomy refers to the
arrangements of the species into a natural system of lower and higher
categories, and gamma taxonomy is the analysis of intra-specific variations,
ecotypes, polymorphisms, etc.

Etymologically, the word taxonomy is derived from Greek taxis,


meaning arrangement or division’, and nomos, meaning ‘law’. Taxonomy can
thus be understood as meaning ‘laws of arrangement and division’. A host of
other definitions of the word can be found on the web, e.g.:
– The science of classification according to a pre-determined system

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– The practice and science of classification (Wikipedia)


– The science of categorisation, or classification, of things based on a
predetermined system (www.webopedia.com).

Kapoor (1998) pointed out, taxonomy is essential in theoretical and


applied biology (agriculture and forestry, biological control, public health,
wild life management, mineral prospecting through the dating of rocks by
their enclosed fauna and flora, national defense, environmental problems, soil
fertility, commerce, etc). Biological taxonomy (taxonomy) may perhaps be
visualised as ‘the language of biodiversity research’ or even as ‘the language
of biology’. However, taxonomy also consists of the interpretation of the
names, the so-called taxonomic concepts (Franz et al. 2008), and of the way we
believe that the taxa are phylogenetically related to one another. And like any
language, taxonomy evolves: new taxa are discovered and are given names;
taxonomic concepts change, as do phylogenetic hypotheses (Henrik Enghoff,
2009). Taxonomy involves not only collecting, identifying, naming new
species and developing sound classification but also analysis of biological
variations, biogeography, evolutionary biology and host parasitic
relationships, thus taxonomy and biodiversity are so intimately connected
(Narendran, 2006).

According to Darwin’s Initiative: Taxonomy is the science of


discovering, describing and naming the individual species of plants and
animals, including microscopic forms, that make up the biota, and of working
out their relationships to provide a classification. An even broader definition
of taxonomy was proposed by Enghoff & Seberg (2006). According to these
authors, taxonomy consists of seven types of activities:

1. Recognition, description and naming of taxa (species, genera,


families etc., also revision of older descriptions, synonymisations,
etc.) (» á-taxonomy).
2. Comparison of taxa, including studies of relationship (phylogeny)
(» part of â- taxonomy).
3. Classification of taxa (preferably based on phylogenetic analyses) (»
part of â- taxonomy).
4. Study of (genetic) variation within species (» ã-taxonomy)

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5. Construction of tools for identification (keys, DNA barcodes).


6. Identification of specimens (referring them to taxa, using the tools).
7. Inventories of taxa in specific areas or ecosystems , using the tools
for identification)

The description and identification of species are fundamental to


biology. Without taxonomy, biologists in various disciplines would be unable
to report their empirical findings or to access available information on their
target organisms because they would not be sure of their identities.
Taxonomy lays the foundations for the construction of the tree of life, makes
baseline data available for conservation and ecological studies, and affords
humans the possibility to take advantage of the underutilized resources
offered by the earths’ biodiversity (Wilson 2004).

The primary step in evaluating the consequences of the changes that


man is bringing about in his environment is to document the nature of such
changes which include changes in the diversity, distribution and composition
of animal and plant communities. Identifying and compiling as complete an
inventory as possible of the world’s animal and plant species is the basic work
involved in the process of such documentation. Thus, taxonomy is not only
man’s natural pursuit for knowledge but also an essential science to
understand many processes in evolutionary biology, more importantly; it is
also an applied science basic to the human welfare. Taxonomic data are
essential to conserve biodiversity and taxonomists are needed to provide
conservationists with the tools to identify and therefore to monitor the
prevalence of species indicating which species are nearer to extinction and by
indicting areas of the world with high density that could be conserved
(Anonymous, 2002). Taxonomy is the basis for all meaningful research in
biology and it is absolutely essential to know the name of an organism before
undertaking any kind of research on it. It has great relevance in various areas
of research including biodiversity, conservation, ecology, agriculture,
fisheries, medicine etc. In classification and naming, a taxonomist employs a
variety of methodologies. However, in all these methodologies, the basic
tools of the taxonomist’s job are attributes of the organism that he studies
such as its structural features, external and internal morphology etc.

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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.

The science of systematics evolved during the post Darwinain period,


systematics encompassed the vast array knowledge on phylogeny based on
evolutionary theories, population of species, variability of species,
differentiation of populations, reproductive isolations and origin of species
and hybridization. It also includes the tools and methodologies, like phenetic
classifications, cladistic classifications and molecular systematic. Though
taxonomy in the traditional sense remains as the processes of classifications,
identification of taxa, naming of species and hierarchical arrangement of taxa
based on International Code of Zoological Nomenclature, the integrative
taxonomy which is also called systematics has broadened the scientific base
which includes phylogenetic or cladistic concepts, molecular systematics
using DNA sequences and phenetic tools would further broaden the scientific
base.

Systematists have come under a barrage of criticism because of the


alleged inadequacy of the ‘traditional’ taxonomic paradigm to curb the
‘biodiversity crisis’ and expeditiously make available the products of
systematic research—usually species names—to the professional biological
‘user’ community (including ecologists, physiologists, population geneticists,
and conservationists). The accusations leveled on systematists range from
being ‘slow’ to ‘incapable’ of furnishing these products at a rate considered
(by users) appropriate, especially given that the professional systematic

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community is portrayed as being in stark decline while operating in a quickly


deteriorating natural world (Marcelo R. de Carvalho et.al. 2008)

Dubois (2005) explains that taxonomy and nomenclature are different


disciplines. Taxonomy recognizes classificatory units or taxa, whereas
nomenclature attaches a given scientific name to each of these units.
Taxonomy is a scientific discipline, whereas nomenclature is a technique.

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.

 It aids species identification making it an essential tool in the


control, conservation and sustainable use of biodiversity.
 It leads to the understanding of relationships of species and so
helps to find new opportunities for product development in
food or medicines.
 It shows which organisms can act as indicators of environmental
quality or that are essential for the maintenance of healthy
ecosystems, including crop-bearing soils and water resources.
 It is used in the fight against pests, parasites, vectors of disease,
invasive species and extinction.
 The agricultural, fish-farming, forestry and horticultural
industries.
 The tourist industry which increasingly focuses on eco-tourism.
 Water authorities.
 The oil and mining companies, including their work on
environmental impact.
 Conservation agencies and voluntary bodies concerned with the
protection of nature.
 National customs authorities who protect against pests and
invasive species as well as trade in endangered species.
 Governments, local and regional authorities who are
implementing the conservation and bioprospecting.

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The “taxonomic impediment” is a term that describes the gaps of


knowledge in our taxonomic system, the shortage of trained taxonomists and
curators, and the impact these deficiencies have on our ability to conserve, use
and share the benefits of our biological diversity. This crisis is mainly
characterized by a lack of specialists in several groups and geographic areas,
and by insufficient funding for taxonomic work (Godfray 2002, Mallet &
Willmott 2003).

There is a tendency among young and upwardly mobile ecologists to


view museums and herbaria as “dusty” places with old people old-fashioned
working on them (Brooke, 2000). Taxonomists are depicted as postage-stamp
collectors (Gewin, 2002). Species description is seen as an old-fashioned way
of doing research. The results is that taxonomical experts retire and are not
replaced, zoology and botany disappear from university curricula and new
researchers in biodiversity end up being either molecular biologists or
ecologists (Boero, 2001). Today traditional taxonomists are criticized for their
morpho- species concept. It is pertinent to record and acknowledge that due
to their efforts during the last 200 years at least the processes of identification
of two million species of organisms are scientifically recorded with voucher
specimens and types all over the world.

Taxonomic collections have been developed though hard field oriented


efforts by professionals and amateur taxonomists. These collections serve as a
read reference material for systematic research and identification of the
organisms. The collections in the museum are believed to provide
fundamental information for agriculture, health, pollution control and
conservation aspects. Zoological collections also help in determining
ecotypes, biotypes, sibling species, ecological races, etc. Restrictions imposed
to collect material for research will have far reaching implications on
taxonomic and biodiversity research, thus causing impediment to taxonomy
(Narendran, 2006). Besides, with the advent of agriculture sciences and
forestry sciences, the utilitarian aspect of taxonomy is side lined and
taxonomy became an academic activity not economically and socially
relevant. This is the main problem and burden of taxonomy.

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The decade of the 1990s was supposed to be a turning point when


taxonomy would finally receive the support it deserved to lead the effort to
preserve Life on Earth. This was when the Biodiversity crisis became the
cause celebre and taxonomy was universally proclaimed the key to
understanding and overcoming the crisis. Taxonomy benefited from some
impressive initiatives during that time. Initiatives, such as the National
Science Foundation's PEET (Partnerships for Enhancing Expertise in
Taxonomy), Tree of Life, and Biotic Surveys and Inventories programs, the
World Bank Global Environmental Fund and some of its spin-offs like the
GBIF (Global Biodiversity Information Facility) project, were all products of
this decade. Finally, we had the Convention on Biological Diversity (the Rio
Convention; www.biodiv.org), which is about as close as you can get to an
international mandate for a vastly expanded taxonomic initiative (Wills
Flower, 2007).

According to Bio NET International, Taxonomic impediment refers to

 95% of existing taxonomic expertise resides in “developed”


countries
 95% of taxonomic information and collections reside in
“developed” countries
 95% of remaining biodiversity resides in developing countries!
 Regional imbalances
 Expertise is rapidly ‘greying’ and not being replaced

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.,

 Define the end-user needs of taxonomy and the capacity required to


meet these needs;
 Define the role of technical cooperation partnerships in building
capacity,
 Define the role of new tools and new technologies for enhancing
capacity building;
 Define the roles of all stakeholders in the capacity building process;

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 Define roles and opportunities in resourcing the identified capacity


building; and, most importantly:
 To come away with an agreed Strategy for a global partnership to
move activities forward to overcome the Taxonomic Impediment.

In a recently published essay, Evenhuis (2007) discussed a facet of the


‘taxonomic impediment’ that he believes has been neglected by those
concerned with the current ‘crisis’ affecting this branch of science—that many
taxonomists are not delivering the goods because they are simply
unproductive, and therefore apathetic about advancing the noble causes of
taxonomy.

Quentin D.Wheeler, Peter H.Raven and Edward O.Wilson in Science


2004 said “Our generation is the first to fully comprehend the threat of the
biodiversity crisis and the last with the opportunity to explore and document the
species diversity of the planet. Time is running out. Society's investment for
centuries in great natural history collections can now be repaid through a powerful
taxonomic research platform connecting researchers, educators and decision makers."

As a means to revitalize traditional taxonomy and help it rise above the


taxonomic crisis, alternative and complementary approaches have been
promoted, for example; molecular taxonomy (Tautz et al. 2003, Hebert et al.
2003), information technology, the development of investment funds
(Wheeler 2007) and increased utilization of cyber tools (Pyle et al. 2008, La
Salle et al. 2009). Among those proposals, DNA barcoding (Hebert et al. 2003)
has been particularly successful in the identification and delimitation of new
species from various groups (Hebert et al. 2004, Ward et al. 2005, Cywinska et
al. 2006, Smith et al. 2007,Borisenko et al. 2008, Kerr et al. 2009).

Taxonomy as a worthy scientific endeavor is undeniably at risk when


universities do not hire taxonomists and university collections are threatened
with dissolution. Many non-molecular systematists in particular have
experienced difficulty in obtaining funding for their research. This paradox
has resulted in a palpable sense of isolation affecting many taxonomists
nested in university departments wherein the majority of faculty are
biochemists, physiologists, molecular biologists, and other ‘big lab’

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professionals who usually have greater amounts of funding and frequently


work (and publish) in teams. The taxonomists have been referred to as the
‘invisible poor of taxonomy’, who often only ‘exist outside the inner circle of
large institutions and grants’ (Flowers 2007b, p. 5), and who may even form a
substantial part of the working taxonomic force (Lo¨bl and Leschen 2005).

Everyone knows that traditional taxonomy has serious problems that


hinder it progress (May 2004). Numerous species remain unknown because of
the lack of specialists in their groups. Others are only known from their
original descriptions, from just the holotype, or have had their type material
lost or destroyed. The amount of type material deposited in museums waiting
for a specialist to take interest in it is unaccounted for (Padial & De La Riva
2007). Why not, then, look for alternatives that will help develop the field of
taxonomy?. The discrepancy between the amount of unknown taxa and those
that need revision (the ‘staggering number of specifics’ of Vane-Wright 1996),
on one side, and the overall lack of professional taxonomists in universities
and even in museums, on the other, may contribute to the sense of isolation
felt by professional taxonomists (Wilson 2003a).

Tools for identification


The ‘classical’ identification tool is the dichotomous key, often forms part of â-
taxonomic papers, Anderson et al. (2005). Although identification keys are
very often illustrated, the latter is so to an extent that it borders on another
type of identification tool more typical of groups such as birds and but other
types of identification tools, such as DNA barcodes (see Meier 2008 for a
critical review), and automated identification based on image recognition
(e.g., McLeod 2007), Butterflies: the pictorial key.

Identification of a specimen may be regarded as the most basic


taxonomic activity. The results of identification are used for a wide scope of
purposes, in the purest form for lists of species from particular areas:
inventories (Henrik Enghoff, 2009). The description and identification of
species are fundamental to biology. Without taxonomy, biologists in various
disciplines would be unable to report their empirical findings or to access
available information on their target organisms because they would not be
sure of their identities. Taxonomy lays the foundations for the construction of

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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.

Conventional keys to animals usually have a strictly dichotomous,


indented format in which the user is confined to a sequence outlined by the
author. A dichotomous or branching key is one in which there are two
choices, or leads, at each branching point. The two choices constitute a
couplet, which is denoted usually by a number followed by a and b, such as
1a and 1b or 9a and 9b. There may be many couplets in a key, depending on
the number of species or taxa included. In order to construct a dichotomous
key, there are several guidelines to follow. Make sure there are two leads or
choices at each point. Keep each of the leads parallel in construction, that is,
start each lead with the same noun and describe the same feature in each (e.g.,
shorter or longer abdomen than thorax; with horms or without horns, spines
present or absent etc). Be sure to give alternate conditions or states of the
same character when constructing a couplet. Avoid the use of 'not as above' or
'animal otherwise' or the names of taxa in the leads. Give exact measurements

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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.

Taxonomists also prefer to use Indented key, however found confusing


and slightly difficult when a large number of species are considered (more
than 50 species). For smaller groups, taxonomists employ Branching key,
found useful for non specialists of for those use in the identification of
common ones. Besides above, taxonomists or specialists also employ Circular
key, Box key or Multi entry key while describing species.

Recently, there has been considerable research in developing computerized


keys that have multiple entries derived from a detailed data base of the taxa.
The interactive nature of computerized keys makes them highly desirable and
allows users to identify a animal when some characters are missing on a
specimen. As software programs improve, there likely will be increased use of
computer-generated keys.

Dichotomous Key for Identification of Unknown Animals


1. a. Bilaterally symmetrical body ……. 5
b. Body is asymmetrical or parts radiate from a central disc or cylinder…… 2
2. a. Body is asymmetrical with no true organs ……….. Phylum Porifera
b. Body has true organs and is not asymmetrical ……………. 3
3. a. Body has tube feet and is divided into five parts…. Phylum Echinodermata
b. Body does not have tube feet or is not divided into five parts…………

4. a. Body is spherical or ovoid with rows of ctenes……….. Phylum Ctenophora


b. Body is radially symmetrical with tentacles around the mouth… Phylum Cnidaria
5. a. Body is wormlike…………………………….. 6
b. Body is not wormlike ………………………………… 8

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6. a. Body is flattened and ribbon-like…………………… Phylum Platyhelminthes


b. Body is not flattened but cylindrical……………………….
7
7. a. Body is divided into numerous segments……………… Phylum Annelida
b. Body is not segmented, and is tapered at each end………. Phylum Nematoda
8. a. Body with paired appendages and an internal skeleton of cartilage or bone ………..
Phylum Chordata
b. Body without paired appendages or an internal skeleton of cartilage or bone….. 9
9. a. Body has a segmented, chitinous exoskeleton and has jointed appendages……..
Phylum Arthropoda
b. Body lacking a segmented exoskeleton and is without jointed appendages……….
Phylum Mollusca

Biological observations and experiments are only valid when the


organisms in question are accurately identified. In environmental and
ecosystem studies, taxonomic keys may be the most practical method of
classifying large numbers of organisms into taxonomic groups. The specimens
can then be optionally sent to an authority for species identification. More
specialized keys permit one to identify an organism to species by progressing
from general family or genus characters to species-specific traits. In other
words, the principal outcome of the taxonomic process is usually the
grouping of organisms in an embedded hierarchical structure. A hierarchy
reflects the relationships amongst different groups of elements. A group at
any level of the hierarchy is known as a taxon (plural: taxa). The grouping of
organisms into taxa is based on the similarities and differences observed
among organisms with respect to a series of zoological aspects or features,
known in the taxonomic literature as characters (Sokal, 1974).

It is also quite likely the keys prepared by the taxonomists found


difficult to identify. In such cases, specimens are compared with previously
identified/ authenticated specimens available in the museum, or help
obtained from specialists. Recently, web based identification is also of
prevalence. In such cases, the identified materials are sent to specialists for
confirmation.

Modern tools in Taxonomy


Tautz et al. [2003] propose to ‘solve’ the lack of adequate classifications and
effective identification tools (the so called ‘taxonomic impediment’) by

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replacing existing classifications with a system in which an infinitesimally


tiny fraction of an organism’s genome is sequenced and used both to classify
and identify the organism in question.

Some of the critics have proposed solutions to this ‘taxonomic


impediment’ in the form of a triumvirate adjoining a unitary taxonomic
cyberstructure + automated DNA barcoding + molecular phylogeny.
According to these critics, with the right equipment and software, and
making full use of one stop, web-based taxonomic shopping, ecologists or
other non-taxonomists can simply (and finally!) do away with the dusty,
sluggish, and idiosyncratic systematist and deal directly with species
identification and phylogeny—technology, after all, has rendered these
complex issues a simple matter of ‘cooking’ (Grant et al. 2003 for a critique of
‘point-and-click’ systematics). Ecologists, population geneticists, and other
biologists if they prefer to have quick species names and even quicker
phylogenetic trees at their disposal—and without a deeper appreciation for
their essential qualities as hypotheses— then, perhaps, they should simply not
rely on systematic information at all. It is believed that ‘DNA barcoding
generates information, not knowledge’—if not a taxonomic specialist, who,
then, will define the species in question? Therefore, does the implementation
of this method not require, beforehand, a robust taxonomy? (Ebach and
Holdrege 2005, p. 697)

Reducing taxonomy to identification alone makes it a technical task


rather than a hypothesis-driven science. There is no credible reason to give
DNA characters greater stature than any other character type. When species
descriptions are based on a broad range of data, they become interesting
scientific hypotheses making explicit predictions about the distribution of
attributes among organisms. We reconstruct phylogenies to explain patterns
of organismic diversity. Molecular data certainly contribute, but when
nothing is known about organisms except their DNA, there are no
evolutionarily interesting patterns to explain – just a tedious pattern of
sequence similarity. The advocates of Corresponding author: Diana Lipscomb
([email protected]). DNA taxonomy seem not to understand the peerless
intellectual content of taxonomy based on all available information, or the
hypothesis-driven basis of modern revisionary work.

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"Field work: the need to scale up and adjust to new constraints"


Philippe Bouchet (MNHN, France). Fieldwork conducted specifically for
taxonomical purposes essentially remains small-scale and low-tech, and not
infrequently squarely unlawful. Taxonomists must scale up the way they are
conducting field work and must learn how to address the Access and Benefit
Sharing (ABS) requirements of the CBD. Large expeditions require a
distinction between participation, right to study, and collection ownership.
The future of taxonomic fieldwork is threatened by over-regulation, and the
Buffon declaration called for a distinction between academic versus
commercial bioprospecting.

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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a part under pressure. The Convention on Biological Diversity


(http://www.cbd.int/) has launched a Global Taxonomy Initiative http://
www.cbd.int/gti/ to ameliorate the existing taxonomic impediment. Large
international/regional initiatives like Encyclopedia of Life (http://www.eol.org/),
Fauna Europaea (http://www. faunaeur.org/), Global Biodiversity Information
Facility (www.gbif.org), and Species2000 (http:// www.sp2000.org/) help to
make biodiversity information, including taxonomic information, more
readily available, using the internet. In the United States of America, large
projects like Assembling the Tree of Life (http://atol.sdsc.edu/) and Planetary
Biodiversity Inventories (http:// www.nsf.gov/news
/news_summ.jsp?cntn_id=103065) generate lots of new taxonomic
information. And worldwide, taxonomists continue their work, which has
been going on for centuries. Nevertheless, it is a long way till we can claim
sufficient knowledge of Earth’s species.

Faunistic Survey’s in India


The period of 18th – 19th century constitutes a part of the modern period of the
Indian history and is marked by the decline of the Moghul Empire, political
turmoil and establishment of British rule in India. This period is also
remarkable from the point of development and rise of the modern life
sciences in India brought about by the Europeans (Jain, 1980). The science of
Botany was first to appear on the Indian soil as elsewhere in Europe and rest
of the world, however, refuted by the scholars like Hora (1951), Srenivasa Rao
(1957), Maheswari1958) and Kapil (1970). According to these scholars,
‘Hindus were no less than Greeks in their contribution to biological science.
Prior to 18th century, life science flourished under Portuguese and Dutch
visitors. For eg., Garcia da Orta and Acosta published manuals on Indian
drug plants while the Governor of Malabar, Heinrich Von Rheede tot
Drakken Stein was the first to publish 23 volumes of Hortus Malabaricus. It is
believed that the Britisher’s, Danes and Dutch collected plants and animals
and sent to Europe for identification. In India in the early part of 18 th century
the botanical illustrations and monographs dominated the scene, dominated
by the Madras scientists in the name of Tranquebar mission and the Botanical
society headed by J G Koenig (1772) William Roxburgh (1778), Benjamin
Heyne (1792) and others. In reality, Botanical garden at Howrah was first to

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develop in India and Robert Kid (1746 – 1793) contribution is worth


remembering in the annals of history of Indian science.

The Royal Asiatic Society of Bengal was first to develop a synthetic


approach of science education in India incorporating Geography. Literature,
Botany, Medicine and Agriculture etc. The emphasis on zoology was
comparatively poor and it was Russel Patrick (1776 – 1805) a naturalist
ventured to collect, illustrate and describe the Indian snakes and fishes,
besides Bloch (1776) exclusively on fishes of India. It was Thomas Hardwicke
a major general in Bengal Artillery of East India Company who prepared
colour drawings of reptiles and amphibians in Gray’s Illustrations (Francis
Day. 1958). The modern systematics and taxonomy of the Indian fish fauna
had its beginning in the 19th Century when Hamilton Buchanan published his
work on the fishes of Ganges way back in 1822. Since then the pioneering
researchers like J. Mc Clelland, Col. W. Sykes, T. C. Jerdon, Blyth, etc.
contributed a great deal of information to our knowledge on the fish wealth of
the country, laying down a firm foundation for Indian systematic
ichthyology. Francis Day (1875-78) published an authoritative monograph
viz., “Fishes of India” wherein he provided a comprehensive, descriptive
account of 1418 species of fresh water and marine fishes, under 72 families
and 342 genera from the Indian region.

Faunistic and floristic surveys provide information to address both


biodiversity conservation and ecological function values of the country, based
on qualitative and quantitative species composition, threatened species and
the ecosystem that sustains, as well, identifying landscapes, biologically rich
and valuable areas. It also serves to understand the intrinsic value of the
species, not only based on the rarity and vulnerability in terms of geographic
distribution, but also on the basis of the key functions that a large group of
organisms occupy such as “Keystone species”, “Indicator species” and even
the “Flagship species”. The survey also provides a basis and benchmark for
ongoing species and habitat action plans to conserve and increase
biodiversity. This is especially true throughout the history the spread of
plants and animals and other organisms are governed by natural ecological
processes and have accompanied trade in goods and services and the
movement of humans, resulting in the introduction of species outside their

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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 voucher specimens collected during various faunistic field surveys


and explorations, preserved and deposited in natural history museums helps
in recognising multiple species in a complex of closely related species,
variation in traits of populations that affect morphology, ecology, behaviour
or physiology, errors or omissions in keys or guides used for identification.
Accessible voucher specimens are critical for accurate identification and
subsequent verification of species. Species are the raw material of biodiversity
research, whether the focus of that research is taxonomic, evolutionary,
ecological, genetic, behavioural or physiological. Research projects in
biodiversity require a significant investment of time, effort and money, but
without adequate documentation in the form of vouchers there is great
potential for that investment to be wasted. Indeed, a failure to protect the
currency of science inevitably leads, just as in business, to bankruptcy.
(Extract from Biological Survey of Canada)

Formative days (1700’s-1916)

Faunal diversity of India was seriously


explored during the last quarter of 18th
century. The Asiatic Society of Bengal,
founded by Sir William Jones in 1784 was
only organisation documenting culture and
biological wealth of the east during that time.
The society possessed a building in 1808 and
established a museum in 1814. Sir William
Jones was personally not in favour of
collecting zoological specimens; however
there was definitely a bias towards biological
specimens from the very beginning in the museum.

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.

Many distinguished naturalists such as John McClelland, Edward


Blyth, William Blanford, Henry Blanford, Theodore Cantor, Francis Day,
H.H.Godwin-Austen, Thomas Hardwicke, Brian Hodgson, Geoffrey Nevill,
Henry Nevill, Ferdinand Stoliczka, W.M.Sykes, William Theobald, S.R.Tickell,
John Anderson and James Wood-Mason significantly contributed in
documenting the fauna of Indian subcontinent in early years.

Andrewes, H. (1929) Carabidae 1. Carabinae [ 431 p - 62 figs - 9 pl (1 col) ]


Andrewes, H. (1935) Carabidae 2. Harpalinae [ 323 p - 51 figs - 5 pl ]
Arrow, G.J. (1910) Lamellicornia 1. Cetoniinae and Dynastinae [xiv + 322 p - 76 figs
Arrow, G.J. (1917) Lamellicornia 2. Rutelinae, Desmonycinae, Euchirinae [xiii + 387 p – 77]
Arrow, G.J. (1931) Lamellicornia 3. Coprinae [428 p - 61 figs - 13 pl (1 col.) - 1 map ]
Arrow, G.J. (1949) Lamellicornia 4. Lucanidae & Passalidae[vii + 274 pp., 23 pl]
Arrow, G.J. (1925) Clavicornia : Erotylidae, Languriidae & Endomychidae [xv + 416 p ]
Browne, J Balfour (never published) Dytiscidae, Gyrinidae and Haliplidae
Cameron, M. (1930) Staphylinidae 1. (Micropeplinae, Oxytelinae, Oxyporinae, Megalopinae,
Steninae, Enaesthetinae) [471 p - 134 figs - 1 map - 3 col. plates]
Cameron, M. (1934) Staphylinidae. 2. (Paederinae) 257 p - 2 col. pl.
Cameron, M. (1932) Staphylinidae 3. (Staphylininae, Trichophyinae, Termitodiscinae,
Pygosteninae, Tachyporinae) 443 p - 4 col. pl.
Cameron, M. (1939) Staphylinidae 4. Part 1. Subfam. Pseudopernthinae and Aleocharinae
Cameron, M. (1939) Staphylindae 4. Part 2. Aleocharinae.
Fowler, W. (1912) Coleoptera. General introduction and Cicindelidae to Paussidae xx + 529 p
Gahan, C. (1906) Coleoptera. Cerambycidae 329 p -107 figs
Jacoby, Martin & others (1908-1936) Chrysomelidae Volumes 1-4. (Jacoby died before the first
volume was published. See preface to the first volume by C T Bingham)
Jacoby, M. (1908) Chrysomelidae 1. Eupodes, Camptosomes, Cyclica 534 p - 172 figs - 2 pl
Maulik, S. (1919) Chrysomelidae Volume 2 Hispinae and Cassidinae xi + 439 p - 130 figs
Maulik, S. (1926) Chrysomelidae Volume 3. Chrysomelinae and Halticinae 442 p - 139 figs - 1
Maulik, S.(1936) Chrysomelidae Volume 4. Galerucinae 648 p - 144 fig - 1 map - 1 pl - hbk
Marshall, Guy A.K. (1916) Rhynchophora, Curculionidae 367 p - 108 figs
Brunetti, E. (1912) Diptera : Nematocera excluding Chironomidae & Culicidae 581
Brunetti, E. (1920) Diptera 2. Brachycera Volume 1 (Stratiomyiidae, Leptidae, Nemestinidae,
Cyrtidae, Bombyliidae, Therevidae, Scenopinidae, Mydaidae, Empidae, Lonchopteridae,
Platypezidae) 401 p.
Brunetti, E. (1923) Diptera 3. Pipunculidae, Syrphidae, Conopidae, Oestridae [424 p - 83 fig.

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Christophers, S.R. (1933) Diptera Volume 4. Culicidae tribe Anophelini


Barraud, P.J. (1934) Diptera Volume 5. (Culicidae) tribes Megarhinini & Culicini
R Senior-White, Daphne Aubertin & John Smart (1940) Diptera Volume 6. Calliphoridae
Bingham, C. T. (1905) Butterflies Vol. 1
Bingham, C. T. (1907) Butterflies Vol. 2
George Francis Hampson & others (1892-1937) Moths. 5 volumes
Hampson, G. (1892) Moths 1. Saturniidae to Hypsidae 527 p - 333 fig
Hampson, G. (1894) Moths 2. Arctiidae, Agrostidae, Noctuidae 609 p - 325 figs
Hampson, G. (1895) Moths 3. Noctuidae (cont.) to Geometridae 546 p - 226 figs
Hampson, G. (1896) Moths 4. Pyralidae 594 p - 287 figs
Bell TRD & F B Scott (1937) Moths. Vol. 5. Sphingidae [537 p - 1 map - 15 pl]
Talbot, G. (1939) Butterflies. Vol. 1 Papilionidae, Pieridae xxix + 600 p - 184 figs - 1 map - 3
Talbot, G. (1947) Butterflies. Vol. 2 Danaidae to Acraeidae xv + 506 p - 104 figs - 2 col. pl.
Bingham, C.T. (1897) Hymenoptera. Vol. 1. Wasps and bees xxix + 579 pp.
Bingham, C. T. (1903) Hymenoptera, Vol. 2. Ants and Cuckoo-wasps 506 pp.
Morley, C. (1913) Hymenoptera Vol. 3. Ichneumones Deltoidei 531 p - 152 fig - 1 pl
Burr, M. (1910) Dermaptera (Earwigs) [217 p - 10 pl]
Distant,W.L. (1902) Rhynchota 1. Heteroptera. Pentatomidae, Coreidae, Berytidae (See
also Index to Rhynchota)
Distant,W.L. (1904) Rhynchota 2. Heteroptera. Family 4 to 16. (Lygaeidae - Capsidae)
Distant,W.L. (1906) Rhynchota 3. Heteroptera - Homoptera Heteroptera-family 17 to 24.
(Anthoceridae - Coricidae) / Cicadidae, Fulgoridae.
Distant,W.L. (1907-8) Rhynchota 4. Homoptera: Membracidae, Cercopidae, Jassidae &
Heteroptera: Appendix (initially published in two parts)
Distant,W.L. (1911) Rhynchota 5. Heteroptera: Appendix
Distant,W.L. (1916) Rhynchota 6. Homoptera. Appendix
Distant,W.L. (1918) Rhynchota 7. Homoptera:Appendix to Jassidae, Heteroptera:Addenda
Fraser, F.C. (1933) Odonata. 1 Introduction, Coenagriidae 423 p
Fraser, F.C. (1934) Odonata. 2 Agriidae, Gomphidae 398 p - 120 figs - 4 col. pl.
Fraser, F.C. (1936) Odonata. 3 Cordulegasteridae, Aeshnidae, Libellulidae. 46 p
Kirby, WF (1914) Acridiidae Orthoptera Taylor & Francis, London
Chopard L. (1969) The Fauna of India and the Adjacent Countries. Orthoptera. Vol.2:
Grylloidea. Calcutta: Baptist Mission Press.
Fauna of Arachnida :
Pocock, R.I. (1900) Arachnida Fauna of British India, including Ceylon and Burma, Pocock, R.
I. (Reginald Innes), 1863-1947 Taylor and Francis; [etc., etc.],1900.
Fauna of Mollusca:
Blanford W. T. & Godwin-Austen H. H. 1908. Mollusca. Testacellidae and Zonitidae. Taylor &
Francis, London. 311 pp.
Gude G. K. 1914. Mollusca.−II. (Trochomorphidae--Janellidae)''. xii + 520 pp., 164 figs.
Gude G. K. 1921. Mollusca.−III. Land operculates (Cyclophoridae, Truncatellidae, Assimineidae,
Helicinidae). 386 pp.

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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.

Sir Alfred Alcock, was working as


superintendent of the museum. At the Indian
Museum, Alcock worked on improving the
public galleries of Reptiles, Fishes and
Invertebrates. Sir George King who was the
chairman of the Trustees supported him,

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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."

Alcock protested to the Trustees that "it would be disgraceful to dismantle


a gallery of Invertebrates which included an exhibit of the recent mosquito-malaria
discoveries, at a moment when those discoveries seemed at last to have driven into the
thickest British skull the great truth that the study of zoology was of some use to
mankind." The gallery was spared but the library was to be cleared. These
experiences caused Alcock to quit and he returned home in 1906 writing to
the Government "telling him what an impossible post the superintendentship of the
Museum was and begging him to get it improved for the sake of the Science of
Zoology and of my successors." In the letter Alcock wrote that Zoology was "a
branch of pure science pregnant with human interest", important to the state "in
matters of education, in matters agricultural and veterinary, and in the vital matter
of public health”. He suggested the establishment of an Indian Zoological Survey
with a museum and laboratory administered by zoologists along the lines of
the Geological and Botanical Surveys (Wikipedia). The main task of the
Asiatic Society (1814- 1875) and Indian Museum (1875 – 1916), was to identify
and exhibit the zoological collections deposited by different explorers and
naturalists of British India when organised field studies for fauna were
limited. It was felt, that the role of survey was not purely of museum
taxonomy and without having detailed field studies, any conclusion drawn
regarding the habit and habitat of any species would be meaningless.

A representation made to the Government of India by the trustees of


the Indian Museum accepted the advice and separated the Zoological and
Anthropological section from the Indian Museum on 1st July, 1916 into a
separate department, the Zoological Survey of India. Later in 1945
Anthropological section of Zoological Survey of India was separated into
Anthropological Survey of India. The exploratory field studies gained
considerable importance after the official creation of the Zoological Survey of
India, as an independent organisation. The post of Surgeon Naturalists,
Marine Survey of India was transferred to Zoological Survey of India in 1920
and Major R B Seymour Sewell became an additional member of the survey in
the rank of Superintendent.

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Pre independence (1916-1947)

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.

According to Deepak Kumar (1980) zoology ranked quite low in the


priority list of the Government, during the pre independence, may be because
of the possibility of almost nil material gain as return. William Jones went on
to explain that “he would not patronize such studies because it is involved
pain to the living objects”. Hodgson summarized that Indian Zoology was

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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

Anonymous, 1980. History of Zoological Survey of India. Ed. Director,


Zoological Survey of India.

Deepak Kumar, 1980. Patterns of Colonial Science in India Indian Journal of


History of Sciecne 15 (1): 105 – 113.

Jain B.L. 1982. Development of Science in India in eighteenth –nineteenth


century Indian Journal of History of Science 17(1): 114 – 131.

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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.

In the Bay of Bengal and Arabian Sea bivalves are important


constituents of benthic fauna, forming 16% of the benthos and second to
polychaetes in dominance. They prefer muddy sandy substrate or fine
sediments with a high percentage of silt and clay in shallow regions up to 100-
meter depth. A few specialist species occur at depths beyond 100 meters.

1
(Rtd:) Chief Scientist, Zoological Survey of India
E-mail: [email protected]

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Molluscs were commercially exploited even in prehistoric times. A


shell is a gift of nature and a ’thing of beauty’. Molluscan shell has an
aesthetic appeal. Shells are used as raw material for shell craft, as source of
calcium and lime and in pearl production. Many of the bivalves are of edible
value. They play a major role in marine biodeterioration and useful as
biomonitars in checking ecosystem health. Recent researches have shown that
extracts prepared from certain mussels, clams and oysters were found to
contain high antiviral activity. The basic data for any malacological research is
provided by taxonomy. The correct identification of a species is the primary
requisite.

Hitherto identification of bivalves was based largely on shell


morphology. Many new species were described on the basis of shell
morphology. Shell shape, sculpture on the external surface of shell, its colour
the placement of umbones, the presence or absence of heart-shaped lunule
and escutcheon on the anterior and posterior sides of the umbo respectively
on the exterior and adductor muscle impressions, pallial line and pallial sinus,
hinge teeth on the interior of the shell are some of the characters used in
identification. Although these characters still hold good in the identification
of a species, more characters are added in latest taxonomic studies. Bieler et al.
(2014) have used a total of 210 characters in their analysis of the ‘Bivalvia Tree
of Life’ and 48% of the characters are used for the first time. The following
brief account gives an idea of the characters used by them (numbers in the
bracket are number of characters):

 Shell characters (21)


 Hinge characters (7)
 Shell microstructure (21)
 Mantle and sense organs (pifaunal7)
 Muscle, foot and pedal gland characters (6)
 Alimentary System Characters (60)
 Gill and labial palp characters (45)
 Development and larval characters (17)
 Endosymbiont characters (Symbiosis): Sulphide oxidizing bacteria-
absent/present
 Symbiotic zooxanthellae- absent/present
 Cellulolytic nitrogen fixing bacteria in the gland of Deshayes (3)
 Oxygen transport characters- absent /present
 Sperm characters (19)
 Molecular sequence data

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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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Annotated Classification of Indian Bivalves

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.

Bieler et al., (2014) combined molecular and novel morphological


characters and have shown that sperm ultrastructure features are among the
best morphological characters to diagnose bivalve clades, followed by the
characters of the shell, including its microstructure.

Another recent classification by Carter et al. (2011) is a synergistical


product of several minds. Fifty-one malacologists have applied their collective
wisdom and outlined “A Synoptical Classification of the Bivalvia “as a run-up
for the revision of Bivalvia in Treatise on Invertebrate Paleontology. Carter et al.,
(2011) assigned all taxa to their respective positions in the Linnaean hierarchy.
A total of eighteen Linnean ranks and eighteen orders to accommodate 107
were recognized. The phylogenetic relationships of molluscs have become

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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:

Class Bivalvia Linnaeus, 1758


Subclass Protobranchia Pelseneer, 1889
Synonym: Palaeotaxodonta Newell, 1965
Proto-primitive, branchia-gills.
Most primitive. Shell equivalved. Hinge with taxodont teeth. Labial palps
most primitive and possess filamentary extensions called proboscides to help
in food collecting. Mantle opens ventrally. Siphons not well developed. Gills
comprising two divergent rows of flat, short filaments. Byssus absent in
adults. Infaunal and detritus feeders. Mostly deep sea forms. Globally 700
species known (Huber, 2010).
The subclass encompasses three orders.
Order Nuculida Dall, 1889.
Shell nacreous or crossed lamellar. Hinge with taxodont teeth. Ligament
externa.Adductor muscle well developed and isomyarian. Labial palps large
and possess proboscides. Gills situated posterior to the foot at the back of the
mantle cavity. Foot flat, grooved and without byssus in the adult.
Superfamily Nuculoidea
Family 1. Nuculidae
Order Nuculanida D.C.Campbell & M.R. Campbell, 2000
Shell externally smooth or sculptured with concentric ribs and covered by
varnish-like periostrcum. Adductor muscles anisomyarian; anterior adductor
muscle larger than the posterior adductor muscle.Labial palps large and with
proboscides. Ctenidia more specialized than in Nuculida. Pallial line with
pallial sinus; siphons united. Hinge line a little arched and with taxodont
teeth. Ligament partially internal.
Superfamily Nuculanoidea H.Adams & A.Adams, 1858
Family 2. Malletiidae
Family 3. Neilonellidae
Family 4. Nuculanidae

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Family 5. Yoldiidae. Carter et al., (2011) recognized the family


name Sareptidae and includes Yoldiellinae and Yoldiinae as
subfamilies.
Order Solemyida Dall, 1889
Shell with weakly taxodont or edentulous hinge plate. Ctenidia large and
posteriorly situated. Shell covered by thick priostracum, which extend
beyond the ventral margin as frill. The clams harbour endosymbiotic, sulfur-
oxidising bacteria. Infaunal and highly modified deposit feeders.
A single family Solemyidae. In India known by types only.
Subclass Pteriomorphia Beurlen, 1944
Shell highly variable, many inequivalved, usually dimyarian and isomyarian,
but often anterior adductor muscle reduced or absent, leading to
monomyarian condition. Hinge with variable dentition. Many have reduced
anterior end and with auricles or wings. Mantle margins not fused; siphons
absent. Filibranch ctenidia with paired demibranchs of weakly united
filaments. Foot not well developed or even absent. Mainly marine, a few
estuarine. Epifaunal or semi-infaunal sedentary forms, usually attached by
byssus or cemented by one valve to the hard substratum. A few shallow
borers, some can swim. Intertidal suspension feeders. Globally 2001 species
are known (Huber, 2010)
The subclass comprises five orders.

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.

Superfamily Arcoidea Lamarck, 1809


Family 7. Arcidae Lamarck, 1809
Family 8. Cucullaeidae Stewart, 1930
Family 9. Glycymerididae Dall, 1908

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Family 10. Noetiidae Stewart, 1930. Recognized as subfamily of


Arcidae (Carter et al., 2011).
Superfamily Limopsoidea Dall, 1895
Family 11. Limopsidae Dall, 1895

Order Limida Waller,1978. File shells.


Shell usually elongate ovate or subtrigonal, equivalve, rather compressed or
inflated.Auricles usually absent or when present unequal. Umbones widely
separated. Byssal notch absent. Posterior adductor muscle of variable size.
One or two pairs of pedal retractors. Hinge short and straight, with well-
developed cardinal area, pseudotaxodonta denticles or lateral teeth. Ctenidia
plicate and with interlamellar connections. The ends of demibranch do not
lead into distal oral groove.

Superfamily Limoidea d’Orbigny, 1846


Family 12. Limidae d’Orbigny, 1846

Order Mytilida Ferussac, 1822. True Mussels


Shell more or less elongate with narrow and pointed anterior end. Shell
covered by conspicuous and strong periostracum of different colours.
Internally nacreous. Heteromyarian with a larger posterior adductor than the
anterior one and the latter absent insome. Two byssal retractor and one pedal
retractor muscle present. Labial palps rather recurved and normally elongate.
Pallial line entire and simple; siphons weakly developed. Hinge edentulous or
with weak crenulatios (dysodont teeth). Ligament mostly external or deep set
alvincular or pavincular. Mantle margin free. Ctenidia filibranchiate with stiff
ciliated disks connecting the filaments. Foot small with a byssal groove.
Mainly epibenthic sedentary and attach by strong byssus, a few nestlers. A
few make nests with byssal threads. Some liead a commensal life in tunicate.
Some specialized to bore into rocks and corals. Marine or estuarine.

Superfamily Mytilodea Rafinesque,1815


Family 13. Mytiliidae Rafinesque, 1815
The family has seven subfamilies. Carter et al., (2011) elevated
two subfamilies Crenellinae and Septiferinae to family ranks.

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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.

Superfamily Ostreoidea Rafinesque, 1815.


Family 14. Gryphaeidae Vyalov, 1936
Family 15. Ostreidae Rafinesque, 1815. A separate family,
Flemigostreidae Stengel, 1971 has been proposed with
Crossostreinae as its subfamily (Carter et al., 2011).
Superfamily Pinnoidea Leach, 1819
Family 16. Pinnidae Leach, 1819
Superfamily Pterioidea J.E.Gray, 1847
Family 17. Malleidae Lamarck, 1818. Isognamoninae is treated
as a subfamily of Malleidae by Carter et al.(2011) but placed in
the family Pteriidae by Bieler et al.(2010)
Family 18. Pteriidae J.E.Gray, 1847

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.

Superfamily Anomioidea Rafinesque, 1815


Family 19. Anomiidae Rafinesque, 1815
Family 20. Placunidae Rafinesque, 1815
Superfamily Pectinoidea Rafinesque, 1815
Family 21. Pectinidae Rafinesque, 1815

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Family 22. Propeamussiidae Abbott, 1954. It is placed in a


separate suborder Entoliidina Hautmann, 2011, in Carter et al.,
2011.
Family 23. Spondylidae J.E.Gray, 1826
Superfamily Plicatuloidea J.E.Gray, 1857
Family 24. Plicatulidae J.E.Gray, 1857

Subclass Palaeoheterodonta Newell, 1965. Freshwater mussels.


Hinge dentition variable, but generally hinge with schizodont teeth. Ctenidia
eulamellibranchiate. Mantle margins not fused. Complicated life cycle with
fish as intermediate host. Nonbyssate and infaunal. A total of 908 species, all
freshwater, are known globally (Huber, 2010).
Order Unionida Stoliczka, 1871(freshwater)

Superfamily Unionoidea Rafinesque, 1820


Family Unionidae Rafinesque, 1820
Superfamily Etherioidea Deshayes, 1830
Family Etheriidae Deshayes, 1830
(Syn.Mulleriidae, Pseudomulleriidae)

Subclass Heterodonta Neumayr, 1884


Shell shape variable. Shell interior never nacreous. Lunule and escutcheon
distinct. Adductor muscles one, two or three. Pallial line entire or with a small
sinus. In most cases ligament external, rarely sunken between the hinge teeth.
Resilium present or absent. Separate lithodesma absent. Hinge plate with a
subumbonal cardinal tooth. Ctenidia with interlamellar septa connecting
inner and outer demibranchs. Foot without byssus.

Heterodonta composed of two main lineages, Archiheterodonta and


Euheterodonta (Giribet & Distel, 2003). For detailed information of
Heterodonta and relationships of different clade please refer to Taylor et al.,
(2007), Giribet (2008) and Bieler et al., (2014).

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Infraclass Archiheterodonta Giribet in Taylor, Williams, Glover & Dyak,


2007
The name Archiheterodonta Giribet (in Press) was used in Taylor, Williams &
Dyak but the actual description of the clade was given by Giribet in 2008.
Archiheterodonta as the name suggests primitive forms and some of the
oldest known bivalves. Members without siphons and hence a pallial sinus
absent on the pallial line. Some members reported to possess haemoglobin.
Marine infaunal and byssate. Globally about 420 species known (Huber,
2010).

Order Carditida Dall, 1889. False cockles or Little heart clams


Shell quadrangular or subquadrangular or rounded triangular, equivalve or
inequivalve. Pedal retactor muscles well developed. Anterior and posterior
adductors subequal or isomyarian. Pallial line entire with only an impression
of the excurrent opening. Labial palps small to medium and trigonal.
Ligament external and set on nymphs. Hinge bears two or three strong,
transversely striate cardinal teeth and anterior and posterior lateral teeth. Foot
with a byssal groove and byssus, which may be present or absent in the adult.
Members filter feeders and burrow into soft substrates, some live attached by
byssus.

Superfamily Carditoidea Fleming, 1820v


Family 25. Carditidae Fleming, 1820
Superfamily Crassatelloidea Ferussac, 1822
Family 26. Crassatellidae Ferussac, 1822

Infraclass Euheterodonta Giribet & Distel, 2003.


This is the most speciose and widely distributed group.” Euheterodonta is
difficult to define based on morphological characters alone” (Giribet, 2008). It
includes tradional veneroidean families not included in Archiheterodonta,
Myoida and Anomalodesmata. Shell ultrastructure variable; nacre present in
the members of Anomalodesmata. Euheterodonta includes marine infaunal
groups and exhibit striking adaptation, such as bacterial symbiosis
(Lucinidae, Thyasiridae, Teredinidae) and harbor symbiotic zooxanthellae
(members of Cardiidae). A few acquired a secondary predatory life style.

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Superorder Anomalodesmata Dall, 1889


For a long time Anomalodesmata was treated as subclass, later it was
relegated to order. Now the latest classification accords superorder rank. Shell
internally nacreous and externally sculptured with radial ribs or radial or
concentric pustules. Labial palps small. Pallial line with a distinct sinus.Hinge
margin is thickened and with one weak obsolete tooth or edentulous and
corresponding socket in one or each valve, laterals absent. Ligament may be
absent or when present associated in many with separate resilium and
lithodesma. Resilium rests in spoon-shaped fossa. Mantle fused ventrally.
Ctenidia eulamellibranchiate or septibranchiate.

Superfamily Clavagelloidea d’Orbigny, 1844


Family 27. Clavagellidae d’Orbigny, 1844
Family 28. Penicillidae Bruguiere, 1789
Superfamily Cuspidarioidea Dall, 1886
Family 29. Cuspidariidae Dall, 1886
Superfamily Myochamoidea Bronn, 1862
Family 30. Myochamidae Bronn, 1862
Superfamily Pandoroidea Rafinesque, 1815
Family 31. Lyonsiellidae Dall, 1885
Family 32. Pandoridae Rafinesque, 1815
Superfamily Thracioidea Stoliczka, 1871
Family 33. Laternulidae Hedley, 1918
Family 34. Periplomatidae Dall, 1885
Family 35. Thraciidae Stoliczka, 1871
Superfamily Poromyidea Dall, 1886
Family 36. Cetoconchidae Ridewood, 1903
Superfamily Verticordioidea Stoliczka, 1871
Family 37. Euciroidae Dall, 1895

Caarter et al. (2011) recognized six of the seven superfamilies mentioned


above. The family Myochamidae is accommodated in the superfamily
Thracioidea. They have proposed three orders: Order Poromyida Ridewood,
1903 to include superfamilies Poromyoidea, Cuspidarioidea and
Verticordioidea; Order Pandorida Stewart, 1930 to include superfamilies

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Pandoroidea and Clavagelloidea and order Thraciida Carter, 2011, in Carter et


al. (2011) to include Thracioidea.

Superorder Imparidentia Bieler, Mikkelsen & Giribet, 2014


Impar-unequal, dens-tooth.
Hinge with unequal teeth. It is the traditionally known Heterodonta
excluding Anomalodesmata.
Order ?
Superfamily Chamoidea
Family 38. Chamidae
Superfamily Galeommatoidea. Carter et al. (2011) proposed suborder
Leptonidina Dall, 1889 to accommodate this superfamily.
Family 39. Galeommatidae
Family 40. Kelliidae
Family 41. Lasaeidae
Family 42. Montacutidae
Superfamily Gasrochaenoidea
Family 43. Gastrochaenidae
Superfamily Mactroidea
Family 44. Anatinellidae
Family 45. Cardiliidae
Family 46. Mactridae
Family 47. Mesodesmatidae
Superfamily Sphaerioidea (freshwater)
Family Sphaeriidae
Superfamily Ungulinoidea
Family 48. Ungulinidae

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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Order Adapedonta Cossmann & Peyrot, 1909.


Synonym: Hiatellida Carter, 2011, in Carter et al., 2011.
Shell without distinct hinge plate, with or without cardinal teeth but
lateral teeth always present. Ligament variable.

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

Order Cardiida Ferussac, 1822


Shell of variable size, equivalve, generally with radial sculpture. Ligament
external. Cardinals cone shaped, absent in the right valve, lateral teeth away
from the cardinals, seldom hinge teeth reduced. Siphons short or long. Pallial
line with an enlarged sinus or no sinus. In some anterior adductor muscle is
reduced. Gill lamina folded. A conspicuous character of presence of cruciform
muscles. On the support of molecular characters Cardioidea and Tellinoidea
are grouped together in this order.

Superfamily Cardioidea
Family 52. Cardiidae
Superfamily Tellinoidea
Family 53. Donacidae
Family 54. Psammobiidae
Family 55. Semelidae
Family 56. Solecurtidae
Family 57. Tellinidae

Order Lucinida Gray 1854


Shell equivalve, without gape. Shell margins smooth or feebly denticulate.
Antrior adductor scar narrowly elongate and posterior adductor rounded.
Pallial line entire and often with punctate pallial area. Ligament partly
internal and with or without resilium.

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Superfamily Lucinoidea
Family 58. Lucinidae
Superfamily Thyasiroidea
Family 59. Thyasiridae

Order Myida Goldfuss, 1820


Shell variable, in some it does not cover the entire animal. Lunule and
escutcheon absent. Shell with obsolete hinge, either edentulous or with
cardinal teeth. Siphons well developed. Ctenidia with interlamellar septa.
Burrowing or boring into wood, filter feeders.

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

Order Venerida H.Adams & A.Adams, 1856


Shell small to large, ovate to subtrigonal, thick and heavy, polished externally
and porcelaneous internally. Valves without gape. Lunule and escutcheon
well developed with well demarcated borders. In many ventral margin
denticulate. Anterior and posterior adductor muscles subequal and
demarcated into striate and smooth portions. Pedal retractors confluent with
the respective adductors. Pallial sinus of variable shape and size. Hinge with
three thickened or bifid cardinal teeth; lateral teth weak or absent. Ligament
external and placed in a groove. Formerly the order Veneroida was very
large and comprised 23 families. As per the latest classification it is reduced to
five families.

Superfamily Arcticoidea
Family 66. Trapezidae
Superfamily Cyrenoidea (partly freshwater)

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Family 67. Cyrenidae


Family 68. Glauconomidae
Superfamily Glossoidea
Family 69. Glossidae
Superfamily Veneroidea
Family 70. Veneridae
Superfamily ?
Family 71. Vesicomyidae

How many species are there?


It is generally estimated that there are 15,000 species globally; its undescribed
species diversity is more. Globally marine bivalves and gastropod are
estimated to be in 1:4 ratios (Rosenberg, 2014). But Huber in his Compendium
of Bivalves (2010) gave a complete breakup of the species superfamily- wise.
According to Huber the 46 super families encompassing 106 families contain
9000 species globally. In India, there is no comprehensive data base for
marine molluscs. Recently marine bivalves are dealt in detail and data on 71
families occurring in Indian Seas is presented (Subba Rao, in press). In
connection with the preparation of Bay of Bengal Biota, a checklist of Molluscs
was prepared (Subba Rao, 2012). As per that list 1610 gastropods and 575
bivalves were recorded from the Bay of Bengal and adjoining Andaman Sea.
The ratio of marine gastropods to marine bivalves is 2.8:1, which is generally
lower than at global level. The species diversity of bivalves is 760, which is
lower than that from the coasts of China (1172) and Taiwan (672) (Bernard,
Cai & Morton, 1998).
The current analysis of diversity and status of marine bivalves is based
on the comprehensive work cited above. The following is the break-up
families based on species diversity:

Total No. of marine bivalve families recorded in Indian Seas: 70


Most speciose families (each containing 2 or more genera and 10 or more
species): 18
Families with two or more genera, each containing less than 10 species: 22
Families with one genus, each with 2-5 species: 13
Families with one genus, each containing 8 or 9 species: 2
Families with one genus, each containing one species: 16

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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).

Table 1. No. of Species occurring in four important coral reef ecosystems

Andaman & Gulf of Gulf of Lakshadweep


Subclass Nicobar Mannar Kachchh
Islands
Prosobranchia 8 2 Nil 3
6 families
Pteriomorphia 127 88 27 29
18 families
Heterodonta 4 4 1 Nil
Archiheterodonta
2 families
Euheterodonta 220 151 42 50
Anomalodesmata
10 families
Imparidentia
34 families

Total no. of species 359 245 70 82


Source: Subba Rao (in press)

The estuaries and mangroves are also important ecosystems providing


shelter to many bivalves. The number of species in important estuaries and
mangroves are given in Table 2.

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Table 2. Taxa occurring in the major estuarine and mangrove ecosystems

Taxon Andaman& Hugli- Mahanadi Gautami- Krishna Vellar-


category Nicobar Matla Godavary Coleron
Families 11 24 6 12 2 11
Genera 26 40 13 21 6 19
Species 32 49 16 31 6 24

Gastropoda 60 55 16 33 9 16
Species
Source: Das & Dev Roy, 1989; Subba Rao et al. (1995); Bhoominathan et al.
(2012)

Recently Bhoominathan et al. (2012) have given a checklist of mangrove


molluscs recorded from India. The estuarine and mangrove fauna are difficult
to differentiate and often there is overlap. The authors reported 72 species. In
fact, they mentioned 77 species, but the list contains duplicates and synonyms
and excluding those species it comes down to 72 species. Further the inclusion
of Donacids, Donax cuneatus, Donax incarnatus and Donax lubricus as
mangrove species is not justified since they are marine sand dwelling forms.
Bhoominathan et al. (2012) included two species Donax cuneatus and Donax
lubricus following Das & Dev Roy (1989). The latter reported citing Daniel &
Rajagopal (1974), who collected the above mentioned two species from the
sandy shore of Galathea River mouth in Great Nicobar. Meretrix attenuata,
Batissa inflata, Batissa similis, and Geloina galathea are restricted to Great
Nicobar Island. The number of bivalves reported from estuaries and
mangroves of India is about 69 species. In the Indo-Pacific estuaries the
families Mytilidae and Veneridae are the most dominant and represented by
13 species each. But in Indian estuaries woodborers belonging to the family
Teredinidae is with more species diversity (19 species), followed by Mytilidae
(7 species) and Tellinidae (6 species) (Subba Rao et al., 1995).

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.

Sea Grass Ecosystem support a rich variety of molluscs. Two recent


studies on the composition and diversity of molluscs in Gulf of Mannar
(Paramasivam et al., 2015) and Minicoy Island (Susan et al., 2012). In species
composition bivalves seem to be less in number than gastropods. The names
of species, with their latest nomenclature, reported from the two locations are
given in Table 3.

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Table 3. Bivalve molluscs recorded from two important sea grass ecosystems

Mi Minicoy Island Gulf of Mannar


Cardium asiaticum Anomia achaeus Gray, 1850
=Vasticardium asiaticum (Bruguiere, 1789) Barbatia fusca Born, 1789 not
?Corculum impressum (Lightfoot, 1786) Lightfoot,1786
?Ctena delicatula (Pilsbry, 1904) =Barbatia amygdalumtostum
?Lunulicardia auricula (Nebuhr, 1775) (Roeding,1798)
Pinna muricata Linnaeus, 1758 Cardium asiaticum
?Lithophaga nigra (d‘Orbigny, 1853) =Vasticardium asiaticum (Bruguiere,1789)
Modiolus metcalfei (Hanley, 1843) Cardium flavum= Vasticardium flavum
= Modiolus modulaides (Roeding, 1798) (Linnaeus, 1758)
Mactra cuneata Gmelin, 1791 ?Cerastoderma edule (Linnaeus, 1758)
Tellina palatam= Quidinipagus palatam Mactra maculata Gmelin, 1791
(Iredale, 1929) ?Mactrellona exoleta (Gray, 1833)
?Myadoropsis brevispinosa Habe, 1962 Modiolus philippinarum (Hanley, 1843)
(family Myochamidae) Pinna bicolor Gmelin, 1791
Gafrarium divaricatum (Gmelin, 1791) Tellina staurella=
Periglypta puerpera (Linnaeus, 1771) Tellinella staurella (Lamarck, 1818
Tellina virgata=Tellinella virgata (Linnaeus,
1758)
Circe scripta (Linnaeus, 1758)
Gafrarium tumidum Roeding, 1798
Paphia undulata (Born, 1778)
Donax cuneatus (Linnaeus, 1758)

Source: Susan et al., 2012 Source: Paramasivam et al., 2015

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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relevant to India. In the absence of any manual for identification of Indian


bivalves, confusion in identification and classification is understandable.
Revisionary studies of the families Arcidae, Mytilidae, Ostreidae and
Tellinidae are needed to fill the gaps in our knowledge. The first mentioned
two families include 12 and 21 species, which are to be authenticated and
illustrated. The latter two families are most confusing and contentious. About
21 species of Tellinidae are reported from India and their identity has to be
verified. From the World Register of Marine Species, it is seen that the genus
Tellina has 63 synonyms or alternate representations. The family Cyrenidae,
which includes 60 species globally (of which 50% freshwater), is represented
in India by 10 species. Seven of these species have their type localities within
Indian territory. Cochin figures as a type locality for at least six species and
fresh attempt should be made to focus studies on molluscs of Cochin
backwaters. Application of novel morphological and molecular characters
may reveal the status of different species. Studies should be made by making
fresh collections and identifying them by using more characters of shell
morphology and also anatomical characters. Finally, efforts should be made
to prepare a Register of Indian Marine Bivalve Species, checked and verified
by a taxonomic expert.

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

Bhoominathan,M., G.Ravikumar, M.D. Subhas Chandran & T. V.


Ramachndran 2012. Mangrove Associated Molluscs of India, pp.1-11.
Lake 2012: National Conference on Conservation and Management of
Wetland Ecosystem

Bieler, R., J.G.Carter & E.V.Coan 2010. Classification of Bivalve families.


Malacologia 52: 113-133.

Bieler, R., P. M. Mikkelsen, T.M.Collins, E.A. Glover, V.L. Gonzalez, D.L.Graf,


E.M.Harper, J. Healy, G.Y.Kawauchi, P.P.Sharma, S. Staubach, E.E.
Strong, J.D.Taylor, I. Temkin, J.D.Zardus, S. Clark, A.Guzman, E.
McIntyre, P.Sharp & G.Giribet 2014. Investigating the Bivalvia Tree of
Life- an exemplar-based approach combining molecular and novel
morphological characters. Invertebrate Systematics 28:32- 115.

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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.

Newell, N. 1965. Classification of the Bivalvia. American Museum Novitates No.


2206: 1-25.

Newell, N. 1969. Classfiication of Bivalvia, N205-N224. In: Treatise on


Invertebrate Paleontology (Ed. R.C. Moore), Part N.Mollusca 6, 1.Bivalvia.
Geological Society of America and the University of Kansas Press,
Lawrence, Kansas.

Paramasivam, K., K. Venkataraman, C. Venkataraman, R.Rajkumar &


S.Srinivaasu 2015. Diversity and Distribution of Sea Grass Associated

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Macrofauna in Gulf of Mannar Biosphere Reserve, Southern India, pp.


137-160. In: Marine Faunal Diversity in India, Taxonomy, Ecology and
Conservation (Eds. K. Venkataraman & C. Sivaperuman), Academic
Press.

Raman, A.V. personal communication

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.

Taylor, J.D., S.T. Williams, E.A.Glover & P. Dyak 2007. A Molecular


Phylogeny of Heterodont Bivalves (Mollusca: Bivalvia: Heterodonta):
new analysis of 18S and 28S Rrna genes. Zoologica Scripta 36(6): 587-606.

Thiele, J. 1998. Handbook of Systematic Malacology (Scientific Editors of


Translation: R.Bieler & P.M. Mikkelsen),Parts 3 and 4. Class Bivalvia
p.1201-1528, Smithsonian Institution Libraries, Washigton
D.C.(Original in German, in 2 parts, 1929-1935)

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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by the fishermen by removing the substratum by hand-held tools such as a


metallic or wooden pole or by using their feet. The exposed clams are then
hand-picked individually and collected in net bags/ canoes. The hand-dredges
on the other hand are operated from a canoe for the exploitation of clams
from deeper waters. Mussel and oysters are chiseled off hard-substratum by
wading or free-diving during low-tide. Mussels are then declumped
manually to separate dead shells and epibionts. Oysters are separated from
the substratum or from other oyster shells to which they adhere by using
chisel or knife.

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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in the Ashtamudi Lake in Kerala has received India’s first Marine


Stewardship Council (MSC) certification in 2014, to encourage sustainable
fisheries and also to protect the ecosystem. The Vembanad waters are
important for the commercial exploitation of black clam (V. cyprinoides). Major
green mussel (P. viridis) fishing areas are located along the Malabar Coast, of
Kerala which are also important for V. cyprinoides and C. madrasensis fishery.
Brown mussel fishery is observed only along the Vizhinjam coast and
southern parts of Tamil Nadu.

The share of Karnataka to the average bivalve production was only


6.92% during 2009-2015. Important clam and oyster fishing areas are the
Aghanashini, Mulki, Coondapur estuaries. Clams were transported in bulk
quantities by road and by rail from estuaries of Kerala and transplanted in
Coondapur estuary prior to retailing in local markets of Karnataka and Goa.
Green mussel, fishery is also observed along the intertidal and subtidal waters
in the coastal region of Karnataka, catering to the domestic markets of Goa
and Kerala. Malpe, Nagoor, Kirimanjeshwara and Gangoli are important
green mussel fishing areas, where seasonal fishery occurs from December to
May.

Ennore, Karapad Bay, Cuddalore and Pondicherry are important


bivalve fishing areas along Tamil Nadu. Mussel fishery in Tamil Nadu mainly
targets in the domestic markets of north Kerala. In Andhra Pradesh, Anadara
sp. constitutes the major fishery from Kakinada Bay. Bivalves are fished and
utilized from the creeks and estuaries of Sindhudurg and Ratnagiri Districts
of Maharashtra. Bivalves are exploited from three major creeks, Shirgoan,
Sakhartar and Bhatye creeks in Ratnagiri district. The Indian rock oyster,
Saccostrea cucullata locally known as ‘Kalva’ is the major landing.
Windowpane oyster, Placuna placuna locally known as ‘Kachga’ is exploited
from Kuda creek, a bay along Rajapuri. The local fishermen hand-pick
windowpane oyster from specially fabricated fiber rafts during low tide.

Meretrix meretrix, M. casta, A. granosa and M. opima are regularly


collected by the fishermen and transported to the different places from
Chaumukha (Balasore, Odisha). In Soula (Contai West Bengal) bivalve fishery

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for M. opima, M. meretrix, A. granosa and Donax sp. M. meretrix occurs for
industrial purposes.

The Andaman and Nicobar Islands have several bivalve resources


among which the black lip pearl oysters (Pinctada margaritifera), the Giant
clams (Tridacna maxima, T. squamosa, T. crocea, and Hippopus hippopus) and the
Mabe or winged oyster Pteria penguin are fished for the tourism based
ornamental shell industry. In the Lakshadweep Island, giant clams are fished
by the locals. However, there is no information on the quantities fished and
exploitation rate.

Overexploitation and destruction of seed resource


Bivalve fishery is supported by 0-year and I-year class. The fishing season is
usually during the post- and pre-monsoon. The bivalve fishery in Karnataka,
Goa and Maharashtra occur during the monsoon season, during the
mechanized ban period. Indiscriminate exploitation of seed clams is seen in
Kerala and Andhra Pradesh where the clams are utilized in the lime shell
industry. In Kakinada Bay, the intensity of blood clam fishery has increased
and small-sized clams formed a major part of the total landing. In the mussel
fishery of Kerala, destruction of seed mussel has been observed as the fishers
discard the seed mussel after they are fished from the natural bed.

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.

India’s first Marine Stewardship Council [MSC] Certified short-necked


clam fishery
ICAR-Central Marine Fisheries Research Institute research leads to India’s
first Marine Stewardship Council [MSC] certified fishery in 2014. Ashtamudi
Lake is the second largest estuarine system in Kerala. Clam fishery in the
estuary dates back to 1981, supporting the livelihoods of around 3,000 fishers
who are involved in collection, cleaning, processing and trading the clams.
The growth of Ashtamudi clam fishery was driven by overseas demand in
Vietnam, Thailand and Japan in the 1980s and 1990s. By 1991, the catch
peaked at 10,000 t, but declined by 50% in 1993 due to overfishing. Based on
the participatory research by ICAR-CMFRI, a closed season and mesh size
restrictions for nets were introduced, along with a minimum export size and a
prohibition on mechanical clam fishing. These measures showed immediate
effects, and the clam fishery has sustained landings of around 10,000t
annually for the past decade.

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The CMFRI’s initiatives in studying and managing the short-necked clam


(Paphia malabarica) fisheries of Ashtamudi Lake since the 1990s helped the
WWF, India in identifying this small-scale fishery as an ideal candidate for
MSC certification. After three years of relentless hard work, the WWF and
CMFRI have together achieved a landmark in Indian fisheries by obtaining
MSC certification for the short-necked clam fishery of Ashtamudi Lake,
Kerala on November, 2014.
The Marine Stewardship Council (MSC) is a global, non-profit and
independent organization working to reverse the decline in global fish stocks
through the use of market incentives. The MSC’s fishery eco-labelling and
certification programme allows consumers to identify and support
environmentally responsible fishing practices through purchasing decisions.
Consumer support for sustainable fishing in the market place leads to
economic benefits for well-managed fisheries and long-term sustainability of
fisheries, ensuring secure livelihoods and continuous availability of fish for
food.
With its seal of approval on more than 6% of all wild-caught fish, MSC
is the world’s biggest eco-label for sustainable seafood. If a fishery is
successfully certified against the MSC Standard, products from the fishery can
be sold with the MSC’s unique ecolabel, which identifies seafood as coming
from a well-managed fishery. The short-neck clam (P. malabarica) fishery in
the Ashtamudi Lake in Kerala has received India’s first Marine Stewardship
Council (MSC) certification which will help boost sustainable fisheries and
also protect the ecosystem. It is the first MSC certification in India; it is only
the third in Asia (after Vietnam and the Maldives). Up to 1000 fishers rely on
this resource. MSC’s scoring system puts the fishery in
the best practice category on 29 of the 31 indicators,
with scoring greater than 80 out of 100. The MSC
certification was a joint effort by CMFRI, WWF, Kerala State Fisheries
Department and the local fishing community. The certification demonstrates
the power of collaboration between partners and the importance of a new
council-based management system for clam fishery governance. Benefits of
certification include potential for premium prices, access to new markets,
preferred supplier status, potential to attract ethical investment in the fishery
or funding for local community social and economic infrastructure,

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improvements in management of fisheries and public recognition of fishery


conservation efforts.

Summary of MSC Principle level scores for the Ashtamudi Estuary Short
Necked Clam Fishery:
MSC Fisheries Standards

Principle 1: Sustainable Fish Stocks


The Fishing activity must be at a level which ensures it can Score - 85.4 out of 100
continue indefinitely

Principle 2: Minimising environmental impact


Fishing operations must be managed to maintain the structure, Score - 83.7 out of 100
productivity, function and diversity of the ecosystem

Principle 3: Effective management


The fishery must comply with relevant laws and have a
management system that is responsive to changing
Score - 82.3 out of 100
circumstances

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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).

Fig.1. Location of Ashtamudi estuary in the state of Kerala, India.

1 (Rtd:) Principal Scientist, CMFRI, Kochi


E-mail: [email protected]

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Considering the unique biodiversity and the socio-cultural


characteristics of this wetland, the International Wetland Commission has
declared Ashtamudi Lake as one of the Ramsor Sites in India with effect from
Dec. 2002. The major resources of the lake are fin fishes, crabs and bivalves.
Among bivalves the shortneck clam, Paphia malabarica locally known as
Kallikakka is dominant with a distribution in 170-180 ha area in the estuarine
part and are exploited for export of clam meat since 1981. Followed by this
species Mercia opima, Meretrix casta and Villorita cyprinoides are available in
lesser saline to freshwater areas in the lake.

Considering the unique nature of fishing methods and fishery


management for sustainable yield of shortneck clam practiced by the clam
pickers of Ashtamudi estuary, World Wide Fund- India (WWF-India) made a
preliminary survey of the fishery in 2010 and requested Marine Stewardship
Council (MSC) for Certification of this fishery. Fishery Certification is a
voluntary process of assessment to determine whether the fishery qualify the
given standards so that the products from the fishery are usually entitled to
use eco - label in the market. MSC is the a non profit organization established
to promote sustainable fisheries and responsible fishing practices worldwide.
The MSC certified fisheries indicate that this product was caught from a well
managed and sustainable fishery. Added to that the concept of fisheries
certification/eco-labelling is becoming more popular in fisheries trade
worldwide.

History of Ashtamudi clam Fishery


Clam picking in Ashtamudi was age old and the demand for clam meat was
confined to the local area where it was eaten for generations. Commercial
fishing for shortneck clam started with the export of clam meat to Japan from
1981. The demand increased further when export of meat to customers in
Vietnam, Thailand, Indonesia and Malayasia in 1980’s – 1990’s. Along with
the increased export demand, the local fish processors in cooking, freezing
and exporting also increased considerably during this period. 200 t of meat
exported in 1981 has increased to 400 t in 1993 with a total catch of 10,000 t of
shell on clams. At that time the fishery was unregulated due to indiscriminate
exploitation and harvest of huge quantities of juvenile clams to meet export
demand and demand for shell for industrial purpose. The total catch declined
to 5000 t in 1993. Recognizing the threat of depletion in stock, the local

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fishermen approached the District Authorities requesting for immediate


conservation measures. The District Administration in consultation with the
Central marine Fisheries Research Institute and with the full co-operation of
the clam pickers of the estuary opted for conservation measures viz; closed
fishery season from November - December to January - February a three
month duration every year (When young clams recruit to the shell fish beds),
mesh size restriction for nets used in fishing (>32 mm), clam meat export
count restricted to <1400 nos per kilogram meat. These self imposed
regulations were followed by the fishermen for the last 23 years and these
fishery management measures showed immediate effects, and they clam
fishery has sustained landings of around 10,000t/year with stable CPUE.
(Fig.2)

Fig.2. Landings and CPUE of P. malabarica from the Ashtamudi Estuary, 2002-2010.
[Source: Mohamed et al, 2012].

Biology of Paphia malabarica


This species comes under phylum mollusks, class. Bivalvia, order Veneroida,
family veneridae sub family Tapetinae Genus Paphia popularly known as
shortneck clam. This species is a benthic filter feeder found in the estuarine
habitats on the east and west coast of India (Fig.3).

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Fig. 3. Photograph of Paphia malabarica

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.

Ecology of the Lake


Ecology of Ashtamudi Estuary has been well studied and recent study
indicate that salinity varied from 20.7 to 33.2 ppt, surface water temperature
varied from 29 to 30.4o c, dissolved O2 4.4 – 6.7 ml/l chlorophyll 2.1 to 6.1 mg/l.
and pH varied from 7.9 to 8.22 in the shortneck clam bed area. The sediment
texture showed that sand formed major component followed by clay and silt.
Presence of organic carbon ranged from 0.03 to 0.08%. Among the heavy
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metals mercury content was found to be below detectable level (Mohamed et


al, 2013). Since the fishing method for clam is selective, retained species in the
catches are oyster Crassostrea madrasensis, green mussel, Perna viridis, yellow
clam, Meretrix casta. Sanguinolaria spp and Arca sp. Since the catch is riddled
through the net with 30mm mesh before it is taken to boats, small clams and
other organisms like polychaetes are not retained in the catch.

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.

Fig. 4. Fishing by diving, Fig. 5. Catch emptied to the canoe

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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).

Short neck clam Fishery Management


The Government of Kerala has overall responsibility for fisheries
management within the territorial waters. Powers to manage these fisheries is
granted under the Kerala Fisheries Regulation Act 1980. Being an annual
renewable resource, clams has to be well managed for sustainable production.
Strict regulations are to be imposed in juvenile exploitation and mesh size
regulation for achieving this goal. The responsibility for management of

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Ashtamudi Clam is delegated to District Fisheries Office. To manage the


Ashtamudi clam fisheries there is a Village Clam Fishery Council with 20
members now representing all Stakeholders. (representatives of Local Self
Governments, Fishermen, Scientists, Police, Mining and Geology Department,
Fisheries Officials, Exporters, Marine Products Export Development
Authority) The District Collector is the Chairman of the Council and Deputy
Director of Fisheries, Govt. of Kerala is the Convener of the council. The
Council started functioning in 2013 and each quarter they meet and discuss
management issues. This management system provide opportunities for local
fishermen engaged in clam fisheries, a mechanism for regulating the fishery
and enforcing the regulation. This participatory management / co-
management system is the first of its kind in the country to manage an aquatic
resource. Added to that, informations on stock of clams and biological details
are collected regularly by the Central Marine Fisheries Research Institute
(CMFRI), the Central Institute of Fisheries Technology (CIFT) studied the
fishery gear technology and opportunity for economic development are
identified by Marine Products Export Development Authority (MPED), all the
three institution located at Kochi. Fishermen are well aware of fishery
regulation in force and the local police and coastguard are empowered to
enforce fishery regulations in this area. Violation of regulation leads to
punishment.

Marine Stewardship Council (MSC) Fisheries Certification Principles


Marine Stewardship Council (MSC) is a non-profit organization established to
promote sustainable fisheries and responsible fisheries practices worldwide.
Only fisheries certified to be sustainable can use MSC logo. (Fig-10) which
indicate that the product was caught from a well managed sustainable
fishery. The principles and criteria for sustainable fishery developed by MSC is
used as a standard by an independent third party for voluntary certification.
Certification based on the following principles.

Fig. 10. Marine Stewardship Council (MSC) Logo

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 Maintenance and reestablishment of healthy population of targeted


species
 The maintenance of the integrity of ecosystem
 The Fishery is subject to an effective management system that respects
local, national and international laws and standards. (Fig. 11)

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.

Fig. 11. Flow chart showing the MSC certification principles

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The methodology and associated actions for MSC Certification

I. Initial contact with client and Certification Body - MSC


The client has to provide, who might be the client, how to choose a
certification body, the unit of certification and possible certification costs and
an assessment of the fishing against the MSC’s principle and criteria of
sustainable fisheries. This is a pre requisite for full assessment.

II. Pre-assessment evaluation


This is the first stage of assessment to provide a client with an
evaluation of the possibility of fishery passing a more detailed certification
assessment, if necessary for assisting the certification body for full assessment.
A qualified individual or a team to conduct the pre-assessment evaluation, a
report by the client that indicate the issues generated by analysis of fishery
against MSC principles and criteria and identification of actions by client
prior to any announcement regarding full assessment are the major actions for
pre- assessment.

III. Full Assessment


Step 1: Full assessment is the formal and public evaluation of fishery against
MSC’s principles and criteria. MSC will advise the public the intension of
undertaking the Certification if possible stake holder consultation on
assessment team members, appointment of a team of experts, determination
of Assessment tree including performance indicators and scoring guideposts
for fishery.

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.

Step 3: Process involves public involvement and suggestion on whether


fishing can be certified or not or any special condition for certification by
certification body. MSC shall place final report on website for 21 days.

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IV. Certificate issued and Public Report


The certificate issue marks the end of the fishery assessment process.
MSC will place the completed final report on website.

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.

Ashtamudi Shortneck clam (Paphia malabarica) Fishery Certification


The pre-assessment made by the WWF- India following the principle and
criteria for sustainable fishery developed by MSC showed that the fishery is
not subject to any controversial unilateral exemptions to an international
agreement, no destructive fishing practices are used, there are mechanisms in
place to resolve disputes between fishery and management system, fishery
has not been assessed earliest for certification against MSC standards and
there are no non-targeted IPI species in fishery. Not an enhanced fishery and
this is not an introduced species. For full assessment default assessment tree
was used and for performance indicators of various characters Risk Based
Frame Work (RBF) was used. M/s. Intertek Fisheries, Certification (IFC), UK
did the full assessment. Evaluation process was done by site visits by
assessment team in September, 2013. Meetings were carried out during site
visit with stake holder and all aspects of fishery and management were
discussed. The assessment was announced through e-mail sent to all
stakeholders. Risk Based Frame work (RBF) for this fishery was done and RBF
workshop was attended by fishermen from the Ashtamudi estuary. All the
stake holders were invited to attend the meeting.

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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improve the performance within a period stipulated by the certification body.


An Action Plan was developed for meeting all conditions. The fishery could
achieve certification only if the overall score is 80 for all the three principles.
The scores achieved for Paphia malabarica fishery is Principle-1. Target species
85.4 out of 100, Principle 2, Ecosystem 83.7 out of hundred, & Principle 3.
Management system 82.3 out of 100 is the evaluation process. The Fishery
attained a score of 80 or more against each of the MSC principles and the
assessment team recommended Ashtamudi shortneck clam for MSC
Certification. Based on the entire fishery certification process, the Ashtamudi
clam Fishery was certified in November 2014 by Marine Stewardship Council.
The Ashtamudi Clam fishery is the first fishery certified in India and in the
developing world. MSC Certification could be useful in opening up new
markets for the sustainable resource. Traceability and further chain of custody
are benefits of certification.

References

Appukuttan K.K; Aravindan C.M; Yohannan, T.M and Balasubrahmanian,


N.K, 1996 Population dynamics of the exploited stock of the clam
Paphia malabarica in Ashtamudi Estuary (South India). Fourth Indian
Fisheries forum proceedings, 24-28. November 1996, Kochi p.31-34.

Mohamed, K.S, V. Venkatesan, V. Kripa, D. Prema, Mathew Joseph, P.S


Alloysious, Jenny Sharma, K.K. Valsala, K.K. Sajikumar, N. Ragesh,
Jose Bose and Anjana Mohan, 2013, Fishery Management Plan for
Ashtamudi Lake Clam Resources, CMFRI special publication No. 114,
48 p.

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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.

The ten classes recognised under phylum Mollusca are Caudofoveata,


Solenogastres (worm-like organisms in sea bed, 200-3000 meters),
Polyplacophora (chitons, rocky tidal and sea bed), Monoplacophora (cap-like
single shell, living forms discovered in 1952, generally deep sea form 1,800-
7,000 meters), Gastropoda (snails and slugs, in sea, land and freshwater),
Cephalopoda (marine), Bivalvia (marine and freshwater), Scaphopoda (tusk
shells, marine), Rostroconchia (marine), and Helcionelloida (marine); the
latter two classes are extinct and known only from their fossil records
(Haszprunar, 2001). Caudofoveata and solenogasters were also put into one
class, the Aplacophora (vermiform molluscs) by earlier workers (Healy, 2001).
Aplacophorans and polyplacophorans may represent the early evolved living
groups of molluscs (Sigwart and Sutton, 2007).

Of late, good quality molecular phylogenetic analyses support the


polyphyletic origin of Mollusca and place the molluscs in the
Lophotrochozoa, along with annelids, brachiopods, bryozoans and several

Department of Aquatic Biology and Fisheries


1

University of Kerala, Thiruvananthapuram 695581, Kerala


E-mail: [email protected]

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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).

Fig.1. Schematic representation of phylogenetic relationship of various


classes of Mollusca (modified from Stöger et al., 2013)

The total diversity of molluscs recorded from India is 5,169 species


(MoEF, 2014), representing around seven percent of the total global molluscan
diversity. With an extensive coastline of 8,129 kilometres, 0.5 million square
kilometres of continental shelf and 2.02 million square kilometres of EEZ,
marine biodiversity of India is impressive. Marine mollusan diversity of India
include 3,400 species (Rao, 1991, 1998; Venkataraman and Raghunathan,
2015). According to Rao (1991, 1998) 5,100 species of molluscs have been
recorded from freshwater (22 families, 53 genera 183 species), land (26
families, 140 genera and 1,487 species) and marine habitats (242 families 591
genera, 3,400 species) of India. There is no consensus among various authors
on the total number of marine molluscs from India. In a report on coastal
marine biodiversity of India, Venkataraman and Wafar (2005) considers 3,370
marine molluscs in India, while Tripathy and Mukhopadhyay (2015) report
2,300 species. As such there is no well-defined and updated checklist on

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marine molluscs of India. The hotspots of marine molluscan biodiversity in


India are Andaman and Nicobar islands (more than 1,000 species), Gulf of
Mannar (around 428 species) and Lakshadweep (around 424 species)
(Venkataraman and Wafar, 2005), all representing important coral reef
ecosystems of India.

Molluscan taxonomy research in India


A comprehensive review of molluscan taxonomy research is not attempted in
this note and the important studies are grouped into three phases, with Phase
1 listing major studies conducted before independence of India, Phase 2
compiling studies done after independence and Phase 3 documenting studies
after the year 2000 (excluding all single species records and publications in
predatory journals).

Phase 1: Taxonomic studies on molluscs of India were initiated by the Asiatic


Society of Bengal (1784) and the Indian Museum, Kolkata (1814). The first
scientific paper on Indian marine Mollusca was perhaps that of Benson (1830).
HMS Challenger put into sea by England in 1872 for oceanographic research
surveyed Indian Ocean and collected information on various groups of
organisms including molluscs. Collection of faunal samples by a series of
cruises by the Royal Indian Marine Ships (RIMS) since 1881 collected and
described marine molluscs from seas around India. Gardiner and Sewell
(1899-1900) carried out studies along Lakshadweep (Minicoy). Sewell carried
out investigations in seas around Andaman and Nicobar islands and led the
John Murray expedition (1933-34); primary objective of this expedition was
the study of the biology of the Arabian Sea, especially the benthic organisms
such as molluscs. Smith (1894) published notes on Mollusca dredged from
Bay of Bengal and Arabian Sea during the expedition of the H.M Indian
Marine Survey Steamer ‘Investigator’.

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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the initial phase of explorations and expeditions to record marine biodiversity


of seas around India.

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.

Phase 3: Recent studies on marine molluscan studies remain centred


primarily around certain charismatic groups such as opisthobranchs (Apte,
2009; Apte and Bhave, 2014; Apte et al., 2010; Apte et al., 2012; Bhave and
Apte, 2011, 2013; Carmona et al., 2014; Raghunathan et al., 2010; Ramakrishna
et al., 2010; Sreeraj et al., 2012). There was no systematic all taxa studies in any
of the specific 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.

One of the ways to circumvent the taxonomic impediment is to


promote co-ordinated taxonomic research involving practicing taxonomists.
Further, international collaboration in taxonomy should be promoted to
document the diversity of all marine taxa in seas around India, as
comprehensive data bases provide platform for advanced research and policy
making towards conservation and sustainable utilisation of resources.
Developing trained manpower in taxonomy is yet another priority to promote
taxonomy, besides reserving positions for taxonomists in all the marine
research institutions and universities to develop globally competent
taxonomists from the country. Further, the curricula should be framed in
schools and colleges involving taxonomy as a ‘joyful’ activity rather than a
‘cumbersome’ task, with more field oriented activities.

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.

Specific taxon based studies are also required to prepare


comprehensive data bases on molluscs. In a biodiverse group such as
Mollusca, developing taxonomic expertise in each family is a difficult task to
attain and in such cases services of ‘specialists’ should be sought in
collaboration with leading international museums and malacologists. Studies
on molluscs involved in various kinds of associations, invasive species and
planktonic molluscs are other areas that demand attention of malacologists in
India. Micro molluscs from India also did not receive much attention by
taxonomists.

Though marine molluscs consisting 242 families were recorded from


India, only few families such as Conidae, Cypraeidae, Muricidae, Strombidae
and Mitridae (Gastropoda), Donacidae, Tellinidae and Arcidae (Bivalvia)
received special attention of taxonomists of India. Further studies are required
in lesser known molluscan families from India, including Chilodontidae,
Cerithiidae, Naticidae, Ranellidae, Columbellidae, Velutinidae, Trividae,
Triphoridae, Volvatellidae, Vermatidae, Acteonidae, Buccinidae,
Calyptraeidae, Cerithidae, Columbellidae, Haminoeidae, Philinidae, etc.
(Gastropoda) and Galeommatidae, Nuculanidae, Yoldiidae, Spondylidae,
Limidae (Bivalvia). Family-wise revisions are also required in other
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biodiverse groups collected from Indian waters. Deep sea molluscs off Indian
coast is yet another priority area for consideration.

Species categorised as ‚taxon inquirendum‛ (name which is listed


from a literature source, but has not been recently re-evaluated for taxonomic
validity and/or generic or familial placement) and ‚nomen dubium‛ (name
which resists revision because the description and other supporting data are
deficient) also require priority attention by malacologists. Further, stock
assessments and ecosystem based studies are required for species included in
Redlist of IUCN and in various schedules of Wildlife (Protection) Act of India.

In the era where consumptive and non-consumptive values of molluscs


are held with much esteems, the services of taxonomists are all the more
important not only to confirm identification of species involved in various
economic benefits but also for preparing policy documents for conservation
and sustainable use of molluscan resources.

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).

Involving Citizen Scientists and Civil Society

‚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.

All the repositories should go for rampant modernisation, with the


help of latest science and technology inputs. For examples, leading museums
all over the world are in the process of digitalisation of collections, which has
not been initiated by national repositories in India. The digitalisation include
taking photographs of the type specimens and preparing 3 D images of the
specimens using modern software, preparing DNA fingerprints of type
specimens (as technology is now available for preparing DNA barcodes from
formalin-preserved specimens) and preparing collections details and maps in
GIS platform. The preparation of DNA barcodes has implications for
‚upstream sample collection and preservation methods, as well as
downstream implications for highlighting biorepository specimens available
for genetic and genomic research‛ (Hanner and Gregory, 2007).

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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examples of social and economic benefits arising out of marine biodiversity;


(iii) biodiversity studies, especially those involving economically
‘insignificant’ invertebrate taxa such as Mollusca are underfunded and better
funding options should be provided to the taxonomists to complete the
inventory of each taxa in every coastal and marine ecosystem of the country.
As suggested by Cardoso et al. (2011) this is all the more important since most
species are undescribed (the Linnean shortfall), the distribution of described
species is mostly unknown (the Wallacean shortfall), the abundance of species
and their changes in space and time are unknown (the Prestonian shortfall)
and species ways of life and sensitivities to habitat change are largely
unknown (the Hutchinsonian shortfall); and (iv) thinking beyond achieving
biodiversity targets fixed by the United Nations Convention on Biological
Diversity through Aichi Targets 2020, government should plan urgent
strategies and action plans to prepare a marine biodiversity data portal in the
public domain, public high quality field guides and monographs on marine
taxa, train a set of internationally competent taxonomists to cater to the future
demands in biodiversity science, ensuring positions for taxonomists in each
research institution involved in marine biology studies, nurturing young
generation of taxonomists through appropriate revisions in curricula, and
involving citizen scientists and local communities in biodiversity
documentation process.

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new records 1. Genus Strombus. Marine Biological Association of India,


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Robson, G.C. (1926). Notes on the Cephalopoda. No.1. Descriptions of two


new species of Octopus from Southern Indian and Ceylon. Ann. Mag. Nat.
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Smith, E.A (1903). Marine Mollusca pp 589-630. In: Gardiner; S.J. (ed.).Fauna
and Geography of the Maldive and Laccadive Archipelagos, 2(2): 119-222.
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octopus, Cistopus Gray, 1849 (Cephalopoda: Octopodidae) from the
southwest coast of India. Zootaxa. 4058(2): 244-256.
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Subrahmaniyan, T.V., K.R .Karandikar and N.N.Murti (1952). Marine


gastropoda of Bombay II. General charecters, habits and habitat of
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gastropoda of Bombay, I. Ibid., (8) 20: 50-81.
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Andaman and Nicobar Islands. Zoological survey of India, Calcutta,
188pp.
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Venkataraman, K and , Sivaperuman, C., Ed.) Marine Faunal Diversity
in India, Taxonomy, Ecology and Conservation, Academic Press is an
imprint of Elsevier, 32 Jamestown Road, London NW1 7BY, UK pp 39-74.
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Biodiversity of India pp303-372 In: Venkataraman, K. and C.
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of India. Zoological Survey of India; Kolkata; 296pp.
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pp 63-79 In, Venkataraman, K., C. Sivaperuman & C. Raghunathan
(eds.).Ecology and Conservation of Tropical Marine Faunal Communities.
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Winckworth, R (1927a). New species of chitons from Aden and south India.
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Winckworth, R (1927b). Marine mollusca from India and Ceylon. I.
Dentalium. Ibid., 17: 167-169.

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Winckworth, R (1928). Marine mollusca from India and Ceylon. II. Limpets.
Ibid., 18: 133-136.
Winckworth, R (1929). Marine mollusca from India and Ceylon. I. Pinna with
an index to recent species of pinna. Ibid., 18: 206-208.
Winckworth, R (1929). New species of shell from Madras Ibid., 24: 41-43.
Winckworth, R. (1936). Marine Mollusca from South India and Ceylon. 4. A
New Indian Sepia. Proc. Molac. Soc. London, 22: 16-23.
Winckworth, R. (1940). New species of shells from Madras. Proc. malac. Soc.
London, 24 : 19-29.

Recent books
Lamprell, K., 2006. Spiny Oysters. A revision of the living Spondylus species
of the world. 119 p
Cossignani, T., 2006. Marginellidae and Cystiscidae of the world. Atlas with
over 2.600 photos. 408 p.
Fraussen, K. & Y. Terryn, 2007. A Conchological iconography. The family
Buccinidae: Genus Neptunea. 166 pp.
Debelius, H. & R.H. Kuiter, 2007. Nudibranchs of the world. 361 pp.
Robin, A., 2008. Encyclopedia of Marine Gastropods. 480 pp.
Eichhorst T.E. (2016). Neritidae of the world. Volume 1. Harxheim:
Conchbooks. 695 pp.
Huber, M. (2010). Compendium of bivalves. Germany: Conch books.pp. 795.
Yonow, N. (2008). Sea slugs of the Red Sea. Pensoft Publishers, Sofia-Moscow,
304pp.
Yonow, N. (2012). Opisthobranchs from the western Indian Ocean, with
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.
Houart R. (2014). Living Muricidae of the world. Muricinae. Murex,
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,
Florida.
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.

1 Environmental Biology Lab, Rajiv Gandhi Centre for Biotechnology


Poojappura, Thycaud. P.O, Thiruvananthapuram, Kerala, India - 695 014
[email protected]

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Approaches in Molecular Taxonomy


Molecular methods provide taxonomy, the tools to describe species as it relies
on genetic variability and uniqueness in organisms and also they can be
applied to all organisms and offer universal, quantifiable characters. Several
molecular markers are widely used in biological systematics. The general
approaches used in the molecular taxonomy is presented in Fig.1.

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

Mitochondrial DNA markers


Mitochondrial DNA is non nuclear, maternally inherited and are haploid.
They are physically separate from the rest of the DNA present in the cell
which makes them easy to isolate from any tissue. Due to the maternal
inheritance of mitochondrial DNA, the effective population size is
smaller than nuclear DNA and so mitochondrial DNA variation is more
sensitive to population bottlenecks and hybridizations.

The differences in the nucleotide sequence of DNA molecule in the


mitochondria can be determined directly or indirectly by several methods.
Many population genetic studies have employed RFLP (Restriction Fragment
Length Polymorphism) analysis of mitochondrial DNA for understanding
genetic variation in the population either by digesting the whole purified
mtDNA with restriction endonucleases or by DNA sequencing of small
segments of mtDNA molecule obtained by PCR amplification (Freeland,
2005).

The newly emerged sequencing technologies have enabled direct


sequencing of mitochondrial genes and several sets of universal primers
have been developed from conserved sequence regions. For inter species
comparison slow evolving gene regions in the mtDNA can be used while fast
evolving gene regions can be used for population comparisons. The only
non coding region of mtDNA is D-loop region and this region is fast
evolving and mostly used for population comparisons. Apart from that,
the cytochrome b, the region of gene coding for NADH dehydrogenases
(ND1, ND4 and ND-5/6), mitochondrial 16S rDNA and 12S rDNA are also
being used widely.

The mitochondrial cytochrome C oxidase I gene (CO I) is used as a


universal marker in molecular systematics of eukaryotes. CO I gene has
been considered as the universal barcode for species level identification
due to its conserved nature across a wide range of taxa. DNA barcodes are

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segments of approximately 600 base pairs of the mitochondrial COI


gene. Sequencing and homology analysis of COI gene is a fast, efficient and
inexpensive technique helpful in cataloguing the biodiversity.

 Nuclear DNA markers

Arbitrary DNA markers: DNA fragments of unknown functions are used as


arbitrary DNA markers. RAPD (Random Amplified Polymorphic DNA) and
AFLP (Amplified Fragment Length Polymorphism) are the most widely used
arbitrary nuclear DNA method. RAPD uses an arbitrary primer which can
amplify anonymous loci and this method does not require the knowledge of
genetic makeup of the organism (Williams et al., 1990). It is relatively simple,
cheap and shows very high amount of polymorphism. The major drawback of
this method is the lack of reproducibility. RAPD is a dominant marker and so
homozygous and heterozygous states cannot be differentiated and these
patterns are sensitive to slight changes in amplification conditions. RFLP
(Restriction Fragment Length polymorphism) is a non PCR based method
used in the past for molecular phylogeny studies based on restriction
enzymes that fragment the DNA sample (Botstein et al., 1980).

AFLP markers combine the benefits of both RFLP and RAPD. It is a


very useful technique in case of insufficiently characterized genomes (Kumar
et al., 2009). The total genomic DNA is digested using two restriction
enzymes. Double–stranded nucleotide adapters are ligated to the ends of
DNA fragments to serve as primer binding sites for PCR amplification.
Primers complementary to the adapter and restriction site sequence, with
additional nucleotides at the 3’-end, are used as selective agents to
amplify a subset of ligated fragments. The DNA fragments are detected on
polyacrylamide gels and thus polymorphisms are studied. Field Inversion Gel
Electrophoresis (FIGE) and Pulsed field gel electrophoresis (PFGE) can be
used for better resolution of the DNA fragments generated by AFLP, RFLP
and RAPD.

Specific DNA markers: Variable Number of Tandem Repeat (VNTR) is a


segment of DNA that is repeated tens or even hundreds to thousands of
times in nuclear genome of eukaryotes. They repeat in tandem; vary in
number in different loci and differently in individuals. There are two main
classes of repetitive and highly polymorphic DNA; minisatellite DNA
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referring to genetic loci with repeats of length 9-65 bp and microsatellite


DNA with repeats of 2-8 bp (1-6 bp) long. Microsatellites are much more
numerous in the genome of vertebrates than minisatellites. They are widely
used in population genetics studies of fishes and aquatic invertebrates (Selkoe
and Toonen, 2006). The PCR amplification and sequencing of the internal
transcribed spacers (ITS1 and ITS2) and the 5.8S rRNA is accepted as a
powerful method for species identification. (Korabecna, 2007).

Single Nucleotide Polymorphisms

Single nucleotide polymorphisms arise due to single nucleotide substitutions


(transitions / transversions) or single nucleotide insertions/deletions (Liu and
Cordes, 2004). These point mutations give rise to different alleles with
alternative bases at a particular nucleotide position. SNP’s are the most
abundant polymorphisms in the genome, both coding and non-coding
regions of any organism. These single nucleotide variants can be detected
using PCR, microchip arrays or fluorescence technology. These are considered
as next generation markers in molecular phylogeny and can be employed for
population genetics studies, genomics studies and for detection of diseases
(Sukumaran and Gopalakrishnan 2015).

DNA microarray consists of small glass microscope slides, silicon


chip or nylon membranes with many immobilized DNA fragments
arranged in a standard pattern. A DNA microarray can be utilized as a
medium for matching a reporter probe of known sequence against the
unknown DNA sequence isolated from the target sample. Species-specific
DNA sequences could be incorporated to a DNA microarray and this
could be used for identification purposes. DNA extracted from a target
sample should be labeled with a specific fluorescent molecule and hybridized
to the microarray DNA. When the hybridization is positive a fluorescent
signal is detected with appropriate fluorescence scanning/imaging equipment
(Sukumaran and Gopalakrishnan 2015).

Expressed Sequence Tags (ESTs)

Expressed sequence tags (ESTs) are fragments of mRNA sequences derived


through single sequencing reactions performed on randomly selected clones
from cDNA libraries. To date, over 45 million ESTs have been generated from

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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).

Oligonucleotides for the amplification of eukaryotic mitochondrial and


nuclear molecular markers are available at
http://web.archive.org/web/20120910031550/http://crandalllab.byu.edu/

Next Generation Sequencing

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).

Application of Molecular methods in Bivalve taxonomy

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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concentration of mucopolysaccharides. These compounds are usually isolated


along with the DNA, inhibiting the activity of multiple enzymes such as
ligases, polymerases, restriction enzymes. Sokolov (2000) proposed an
inexpensive method of DNA extraction that yielded high-quality DNA
suitable for different molecular analysis. Alternatively commercially
available tissue DNA isolation kits can be used with suitable modifications to
yield pure high molecular weight DNA.

DNA isolation and PCR amplification

Take approximately 50 -70 mg of sliced tissue preferably from muscle in a 2


ml plastic tube along with 1 ml lysis buffer, which consists of 50 mM Tris-
HCl, pH 7.5, 100 mM NaCl, 10 mM EDTA, 1% SDS and 0.2-0.4 mg/ml of
proteinase K. Mix the tube by vortexing and incubate at 55°C until the
complete digestion of the tissue. Add 100 ml of saturated KCl for
precipitating mucopolysaccharides and protein along with the insoluble
potassium dodecyl sulphate. Incubate for 15 minutes, centrifuge the tube at
maximum speed and collect the supernatant in another clean tube. Extract
the supernatant twice with phenol/chloroform/isoamyl alcohol (25:24:1).
Transfer the supernatant to another tube containing isopropanol and incubate
for 10-15 min at room temperature for precipitating the DNA. After
incubation centrifuge the tube at maximum speed for 20 min and discard the
supernatant. Wash the DNA pellet in 70 % alcohol, air dry and dissolve in 100
ml of TE buffer containing RNAse A. Amplify the DNA sequences using
primers of different genetic molecular markers, which will yield potential
information that enable to evaluate the processes that are at the basis of
population and evolutionary studies (Mulvey et al., 1998). Generally two
major categories of molecular markers are used for studying bivalves.

a. Mitochondrial (16S rDNA, 12S rDNA, COI)


b. Nuclear (ITS1, ITS2, 18S rDNA).

 Mitochondrial DNA markers

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).

 Nuclear DNA marker

The Internal Transcribed Spacer 2 (ITS2) represents a region of the


nuclear ribosomal gene group and regarded as a molecular marker in
systematics. The ribosomal gene group consists of 3 genes, small nuclear 18S
rDNA, 5.8S rDNA and large nuclear 28S rDNA which are transcribed into
RNA and are separated by ITS1 and ITS2 regions. After transcription, these
regions are eliminated. Consequently, these regions can rapidly accumulate
substitutions due to weak selection pressure and is useful in discriminating
species that are closely related (Cruickshank, 2002).

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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With the advent of PCR techniques, numerous DNA markers allowing


genetic comparison of bivalves at population level have been developed and
the markers were categorized as RAPD, AFLP and RFLP. All methods use gel
electrophoresis for the resolution of bands generated and contributed
valuable information on bivalve biology, ecology and conservation.

RAPD technique amplifies DNA sequences using single primers


(Williams et al., 1990) and has been successfully applied for the differentiation
of the bivalve Cerastoderma edule and C. lamarcki larvae (Andre et al., 1999).
RFLP, a non PCR based method which depends on the restriction digestion of
DNA fragments is another approach used in the past for genetic
fingerprinting (Botstein et al., 1980). It has been succesfully employed for
detecting larvae of Xenostrobus securis and Mytillus galloprovincialis from
Galician coast (Santaclara et al., 2007) and for distinguishing the cryptic
species of Cerastoderma sp. (Freire et al., 2010) and Ensis sp. (Fernadez-Tajez
and Mendez, 2007). The differences between the species in the family
Dreissenidae belonging to the class bivalvia have been resolved by RFLP and
also identified the cryptic invasive species Mytilopsis leucophaeata from north
America in to Black sea basin in Europe (Therriault et al., 2004). RFLP is a
potent method yielding reproducible results revealing heterozygosity
depending on the length of restriction digested DNA fragments. Specific
enzymes cut the DNA molecule at certain restriction sites which are similar
in homologus sequences, providing obvious discrimination both at
population as well as individual level and hence recommended for
phylogenetic studies (Kumar et al., 2009).

AFLP is a potential technique to assess insufficiently characterized


genomes. Compared with RAPD, the technique is highly reproducible
(Mueller and Wolfenbarger, 1999). Moreover, from technical perspective,
prior DNA sequence information is not necessary, several markers can be
analysed in a short time, and minimal amount of DNA is required. In order to
demonstrate the lack of nucleotide diversity, due to prolonged isolation in
fresh water mussel populations belonging to the genus Anodonta, Mock et al.
(2004) applied AFLP. Studies conducted by Yu and Guo (2004) on the genetic
variation of aquaculture species of Crassostrea virginica using both
microsatellites and AFLP did not find any significant variations in their
heterozygosity and hence recommend the use of AFLP markers when

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microsatellites are not available. The genetic control on morphological


characters in bivalves, such as shell colour was demonstrated through AFLP
in marine scallop Argopecten iradians iradians (Qin et al., 2007) emphasizing the
robustness of this method in that the number of polymorphisms per reaction
is much higher compared to RFLP and RAPD.
Microsatellites are short repeated DNA sequences with a length
between 1-6 repeated base pairs, like dinucleotides (CA)n, (AT)n, (GA)n
and (GC)n or mono, tetra and trinucleotide repeats They are also known as
simple sequence repeats (SSR), variable number tandem repeats (VNTR)
and short tandem repeats (STR). They are the most variable type of DNA
sequences in the genome and their polymorphisms derived mainly from the
variability in length rather than in the primary sequence. Above all, genetic
variation at many microsatellite loci is characterized by high heterozygosity
and the presence of multiple alleles, which is in sharp contrast to unique DNA
(Ellegren, 2004). With the advent of PCR technology, microsatellites became
the marker of choice in genome mapping, population genetics studies and
related areas. Features such as hypervariability and ubiquitous occurrence
have made them the most sought after molecular markers in addressing
questions in ecological research (Selkoe and Toonen, 2006).
Microsatellites have been successfully employed in the population
genetics studies of bivalves. The genetic structure of a bivalve Cerastoderma
glaucum, was studied from nuclear microsatellite loci (Tarnowska et al., 2010)
and observed the marked differences among populations due to the highly
fragmented distribution of the species. Microsatellite makers have also been
used, to know the genetic structure of the species population of endangered
bivalve species Hypanis colorata for establishing conservation strategies (Popa
et al., 2011), to reveal population bottleneck effects among flat oyster Ostrea
edulis (Launey et al., 2001), to observe the levels and patterns of genetic
variation in Pacific oyster Crassostrea gigas (Appleyard and Ward, 2006), to
study the genetic drift in commercially important Chinese surf clam Mactra
chinensis and observed that genetic differentiation is in accordance with the
pattern of isolation-by-distance (Ni et al., 2011) and parentage analysis of
oyster Crassostrea virginica (Wang et al., 2010). The advantages of using
microsatellite markers are the abundance and random distribution in the
genome of eukaryotes; high reproducibility; need only minimal amount of
template DNA. The disadvantages are their specific nature, laborious

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characterization involving cloning, PCR and selection; need to be develop


oilgos if universal primers are unavailable and null alleles (unamplified
DNA) resulting from mutations occurring at sites where primer anneals
(Kumar, 2009).

While investigating the ecology of bivalves, the primary thing that


have to be considered is the interactions between organisms and their
environment (Gosling, 2003). The synergy was studied in the past through
morphology, direct observations and statistical interpretation of data. Despite
the great quantity of information obtained by ecologists, there are still many
queries left unanswered. The advent of molecular markers has greatly
enhanced the opportunity to fill this lacunae on bivalve ecology. In
general, molecular methods succeeded in providing information on
relationship between species (Huff et al., 2004; Mahidol et al., 2007; Espineira
et al., 2009; Vierna et al., 2010), on their evolutionary history (Stepien et al.,
1999; Cunha et al., 2011; Etter et al., 2011), on genetic variation within species
populations (Luttikhuizen et al., 2003; Zardus et al., 2006), on population
size, migratory events, and biodiversity conservation issues such as
hybridization events (Westfall and Gardner, 2010).

With respect to phylogeny of Bivalves, available molecular studies


have been focused on the taxonomic position of several groups. From an
evolutionary angle, a few investigations have been conducted regarding their
colonisation in freshwater or deep-sea environments, habits of wood or rock–
boring and symbiosis with lower life forms (Ponder and Lindberg, 2008). In
spite of all these, there are still many aspects need to be investigated, and
from a taxonomical point of view there may be many taxa still need to be
properly characterized. The molecular approach in bivalve studies generally
involves sampling, storage, DNA isolation, amplification and sequencing. The
methodology varies according to the specific aim of the study which involves
the choice of the right molecular marker to attain the goal. Several studies are
available combining morphological and molecular approaches on taxonomy
and phylogeny of Bivalvia (Giribet and Wheeler, 2002; Giribet and Distel, 2003;
Harper et al. 2006; Mikkelsen et al; 2006; Olu-Le Roy et al, 2007; Plazzi and
Passamonti 2010).

Sequences obtained through the studies based on molecular markers


can be analysed and interpreted with the Bioinformatics tools. There are two
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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.

In order to represent a study on molecular systematics of bivalves,


three samples collected from marine, brackish water and fresh water were
used. The Cytochrome oxidase gene I (COI) was amplified using universal
primers form the genomic DNA isolated from the bivalves. The amplicons
were cloned in a TA cloning vector and sequenced using vector specific
primers in an automated DNA sequencer. The sequences were subjected
homology search in the public database NCBI and the selected sequences
were used for phylogeny. Fig 2 represents the phylogenetic tree generated
using MEGA 5 through maximum likelihood method with 1000 bootstrap
analysis. The isolated species were identified as Crassostrea madrasensis, Perna
indica and Parreysia corrugata.

Molecular genetic methods provide taxonomy a novel perspective to


describe species as they apply to all living beings. Molecular taxonomy is
particularly effective in combination with classical taxonomy, which depends
on morphology of both external and internal features of an organism and
together they will provide a relatively full proof system for the phylogenetic
studies of Eukaryotes. Molecular markers find wide range of application in
systematics and the selection of the marker should be judicious to optimise
the quality of the output. No single molecular marker is superior to any other
and a combination of markers is always suitable. Developing additional

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markers and screening using next generation sequencing technologies will


lead to description of more novel organisms, augmenting the available
database.

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Giribet, G and Distel, D. L. 2003. Bivalve phylogeny and molecular data. In,
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T. T. Nguyen. 2007. Phylogenetic relationships among nine scallop
species (Bivalvia, Pectinidae) inferred from nucleotide sequences of one
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Mikkelsen, P. M., Bieler, R., Kappner, I and Rawlings, T. A. 2006. Phylogeny of


Veneroidea (Mollusca, Bivalvia) based on morphology and molecules. In, R.
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Mock, K. E., Brim-Box, J. C., Miller, M. P., Downing, M. E and Hoeh, W.R.
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Olu-Le Roy, K., von Cosel, R., Hourdez, S., Carney, S. L and Jollivet, D. 2007.
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Plazzi, F and Passamonti, M. 2010. Towards a molecular phylogeny of


Mollusks, Bivalves’ early evolution as revealed by mitochondrial
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Popa, O. P., Iorgu, E. I., Krapal, A. M., Kelemen, B. S., Murariu, D and Popa,
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markers and evolutionary relationships among invasive Dreissenoid
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Cerastoderma glaucum in a highly fragmented lagoon habitat. Marine


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1990. DNA polymorphisms amplified by arbitrary primers are useful
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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.

Sample collection methods

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.

1 Department of Marine Biology, Microbiology and Biochemistry


School of Marine Sciences, Fine Arts Avenue, CUSAT, Kochi-16, Kerala, India
[email protected]

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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.

Rock washing: Smooth rocks can be hand-washed with bare hands in a


bucket. Byssally attached bivalves, limpets, chitons and opisthobranch may be
tenacious and require some assistance to collect. Sculptured rocks or large
pieces of dead coral can be scrubbed with a brush. Rocks buried in sediment
often have an anoxic, blackish or rusty underside; certain molluscs occur
almost exclusively just at the oxic - anoxic border

Collection of intertidal fauna: Methods of sampling the bivalve population


vary with the types of bottom substratum. Quantitative studies require
samplers which take a standard bite of known area and depth. Collection of
intertidal bivalve fauna is easier than that of subtidal forms, but intertidal
areas are subjected to various environmental disturbances. On sandy or
muddy shores estimates of the standing crop of macrofaunal bivalves is made
by driving a square sheet metal frame of appropriate area (e.g. 0.25 m2) into
the substratum, the sediment within the frame being excavated to the desired

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depth. Initial excavations should be made to find a suitable sampling depth.


On some shores the majority of species and individuals occur in the top 15 cm
of depth, in others it may be necessary to excavate to >30 cm, or even deeper
for certain bivalves. Subsequent sorting is made easier if all traces of fine
material can be washed out of the sample when sieving. On muddy or fine
sandy shores it may be possible to use an aperture as small as 500µm, but on
coarse sand or gravel shores a coarser mesh of 1000µm may be necessary. The
selected mesh size (500µm) is indicative for the majority of benthic organisms.
On rocky shores, sampling is carried out by means of a square frame of heavy
gauge wire laid on the substratum, the animals within the frame being
counted directly, or estimated in terms of percentage cover/abundance scale
of the surface. Sometimes estimates may be made in situ, otherwise growths
must be scraped off for subsequent examination in the laboratory. For larger
organisms, a frame of 0.25 m2 is suitable, but for smaller organisms or where
the rock surface is irregular frames of 0.1 m2 (316 × 316 mm) should be used.
For counting small organisms, squares of 0.01 m2 (100 ×100 mm) are suitable.
For such counts a piece of thick (6 mm) Perspex exactly100 mm square and
etched with a grid of 10 mm squares is convenient, allowing smaller areas to
be counted and also minimizing the possibility of missing individuals.
Besides organisms living on the rock surface, estimates should be made of the
crevice-living and boring species, and of those sheltering among weeds. The
importance of photography for non-destructive recording of seashore species
and habitats must be emphasized. On sediment shores, positions may be
marked. It is advisable to position reference marks on rocks or other
permanent structures, from which locations are determined by tape measure,
and/or transect lines. The Global Positioning System (GPS), and its more
accurate sibling, Differential GPS (DGPS) which uses a fixed terrestrial
reference position, can give accuracies up to less than 5 m over the surface of
the planet.

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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Grabs: Quantitative samples of animals inhabiting sediments are usually


taken by grab. The grab, which is lowered vertically from a stationary ship,
captures slow-moving and sedentary members of the epifauna and infauna to
the depth excavated. The majority inhabit the top 5–10 cm, but some burrow
more deeply. Thus, a grab may be an adequate sampling instrument for some
grounds but may leave significant elements of the fauna unsampled below
the depth of bite. For sampling the macrofauna, grabs covering a surface area
of 0.1 or 0.2 m2 are commonly employed, several samples being taken to
aggregate to 0.5 or 1.0 m2 per station. Samples of this total size are usually
considered adequate for quantitative determinations of the commoner
species, measurements of population abundance and biomass, but do not
adequately sample scarcer animals, which are often members of the epifauna.
Moreover, some fast-moving species escape the grab altogether. It is,
therefore, advisable to supplement grab estimates of the epifauna by hauls
with an epifaunal bottom sledge, an Agassiz or beam trawl, or by underwater
photography, television or diving. Petersen grab, Okean grab, Van Veen grab,
Ponar grab, Hunter grab are also used for sampling. The Smith–McIntyre grab
was designed for sampling under difficult conditions often countered when
working from a small boat in the open sea. This grab has hinged buckets
mounted within a stabilizing framework and powerful springs to assist
penetration of the sediment. The Smith–McIntyre grab covers an area of 0.1
m2. The Baird grab was designed for sampling the epifauna of oyster beds,
but has also been used in the open sea. The Hamon grab samples an area of
about 0.29 m2 by means of a rectangular scoop rotating through 90◦. It appears
to be particularly effective in coarse, loose sediments, but, because of its mode
of action, may not sample entirely quantitatively. It appears to be particularly
effective in coarse, loose sediments, but, because of its mode of action, may
not sample. The Holme grab samples by means of a single semi-circular scoop
rotating through 180◦. This design minimizes loss of material while hauling.
The Grizzle–Stegner grab is another quantitative grab for shallow water
sampling which combines features of several grabs such as the design of a
van Veen grab mounted on a Smith–McIntyre frame.

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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deep and relatively undisturbed samples suitable for evaluation of


macrofauna from a variety of sediments, and are also suitable for deep-sea
use. The surface of the bottom, and supernatant water, appear to be taken
without undue disturbance, as evidenced by the persistence of animal tracks
and burrows and sessile forms in their living positions in the samples. Box
corer, Haps corer, gravity, piston or vibro corers have become standard
oceanographic samplers widely used in many benthic surveys.

Suction samplers: A number of samplers employ suction, either to force a


coring tube down into the substratum or to draw the sediment and its fauna
up into a tube leading to some form of self-sieving collector. Remote sampling
by suction devices on difficult substrata has also been designed and
successfully applied. A large number of suction samplers are designed for
sampling in the shallow waters of inshore areas in estuaries, lagoons and in
freshwater habitats. Some suction samplers are towed slowly along the
bottom, sediment and fauna being sucked into a collecting bag.

SCUBA diving: The advent of SCUBA diving has opened up a completely


new world for collectors. With this type of collecting, safety is of paramount
importance. Do not dive alone or in places where currents or turbulence.
Divers were used as carrier platforms even before the introduction of SCUBA
equipment. Because of their relative freedom of movement underwater, small
self-contained units are the most popular, especially camcorders. With recent
advances in mixed gas technology and decompression issues, divers can now
routinely go deeper and penetrate harsher environments. The relevance to
imaging is that pressure housings must take this into consideration and whilst
diving camera systems were once proofed to 30 m depth they must now be
routinely proofed to 100 m depth.

Other methods of sampling: Some species are not readily taken by


conventional sampling gear such as dredges, trawls or grabs, either because
they occur too sparsely to be represented in samples covering a limited area
or because they live in habitats inadequately sampled by the instrument
employed. Alternative methods, which do not necessarily sample other
members of the fauna, are available for such species. Techniques for under
water photography and television, of special value in estimating scarce

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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.

Laboratory Requirements for anatomical work

Microscope: A stereomicroscope is preferable for viewing the molluscs. For


dissection a good-quality dissecting microscope is essential. A minimum
magnification of 50x is desirable. In general, those with stepped
magnifications have better optical quality than those with zooms.

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.

Documenting and labeling


 In field book, record site, date, coordinates, ecological information and
other pertinent data (like air temperature, water temperature, time of
day, etc) for possible future use
 Use a different collection bag at each new site; if specimens are to be
processed later, a numbered tag can be added to each bag with its
number noted in the field book
 For small collections create a label with all collecting data (site, date,
collector, coordinates, habitat) and attach to specimen with a short
length of fishing line.
 Assign field number including - collector’s initials, specimen number
and year collected.

Cleaning and processing aquatic shells:


 Use a stiff brush to clean exterior of shells; take care not to scratch
surfaces; take care not to break small and fragile shells
 Boil a few at a time - choose like sizes for similar cooking requirements
- when bivalves open, they are ready - remove body while hot.
 If time is of little concern allow blow-fly larvae and other carrion
insects to clean shells; after a few weeks, rinsing is all that is necessary
 When only a few specimens are collected, shells may be opened at the
site, cutting the adductor muscles to remove; keep shell halves together
with rubber bands or by tying
 Small shells may be fixed in alcohol; do NOT boil
 If specimen is an operculate snail - boil and remove body using crochet
hook or similar tool - save operculum - fill shell with cotton or
crumpled paper - glue operculum to filler.

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 If preserving for anatomical purposes - do not boil- store in 60% Ethyl


Alcohol
 Specimens collected for shells only - place in bag; immerse bag into
boiling water - boiling time - 30 seconds for small shells; one minute or
more for larger shells. Boiling land snails may cause colour to fade; if
enough specimens were collected, give trial to test for colour fastness.

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.

Curating a Museum Collection


Specimens preserved in Ethanol: Long-term stability is determined by
 Biggest threats: inadequate preservation, loss of fluid preservative,
heat, light, UV light
 Quality of fixation or initial preservation
 Post-preserving process
 Quality and stability of the storage environment

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 Optimum is 65°F and 50% relative humidity


 Fluorescent lights produce significant UV- keep in dark when not in
use
 Do not put preserved specimens in water for examination
 Denatured ethyl alcohol is undesirable

Transfer of specimens - from field fixative to museum storage:


 Osmotic pressure rises steadily with ethanol concentrations below 75%,
so it is suggested that approximately equal concentration increments
are the most appropriate for stepping specimens to higher ethanol
concentrations.... 10% formalin-H2O; 20% ethanol; 40% ethanol; 60%
ethanol; 70% ethanol.
 Fill containers to the top leaving no air space.

Appropriate documentation and provide accession number


Collection records: Maintain a handwritten catalog and electronic data
storage system

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

Holme N. A. and A. D. McIntyre, 1984. Methods for the Study of Marine


Benthos. Second Edition. Oxford-London-Boston: Blackwell Scientific
Publications. ISBN 0-632-00894-6. 16-99 pp

Bakus, Gerald J, 2007. Quantitative analysis of marine biological communities:


field biology and environment, ISBN-13: 978-0-470-04440-7 (cloth/cd),
ISBN-10: 0-470-04440-3 (cloth/cd).

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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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Periostracum - layer of horny material covering the shell.


Plicate - folded or ridged.
Porcelaneous - with translucent, porcelain-like appearance.
Prosogyrate - the condition of a shell when umbones are directed anteriorly.
Protobranchiate type - gill demibranchs simple, formed of leaf-like filaments loosely
connected by sparseciliary junctions.
Radial - diverging from umbo, like the spokes of a wheel.
Rostrate - with a beak-like projection (rostrum).
Sculpture - relief pattern developed on the outer surface of the shell; sculpture is
overlain by concentricgrowth marks corresponding to various positions of shell
margins during growth.
Scabrous - rough, file-like.
Scale - localized projection on the outer surface of shell, commonly situated on a rib.
Septibranchiate type - gills absent, replaced by a muscular horizontal partition (the
“septum”) pierced bysmall pores. This structure enables a carnivorous nutrition
and is encountered in a group of predominantly deep-sea bivalves (e.g.,
Cuspidariidae).
Siphons - extensible, tube-like projections of the posterior marginal region of mantle,
forming 2 openings for water inflow (inhalant siphon) and outflow (exhalant
siphon).
Socket - recess of the hinge plate, for reception of a tooth of opposite valve.
Tooth - shelly projection from the hinge, received in socket of opposite valve; cardinal
teeth are close toumbo, whereas lateral teeth are set apart from these, anteriorly
or posteriorly.
Umbo (pl. umbones) - the first formed part of a valve, usually above the hinge.
Umbonal reflection - expansion of the internal dorsal margin which is folded over the
umbones in Pholadidae and Teredinidae.
Valve - one of the main shelly halves of a bivalve.

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