5222 SHELLFISH/Commercially Important Molluscs
Commercially Important the mantle for shell secretion, gills for feeding and
Molluscs
P F Duncan, The University of the Sunshine Coast,
Maroochydore DC, Queensland, Australia
Copyright 2003, Elsevier Science Ltd. All Rights Reserved.
Background
0001 The commercially important molluscs belong to the
classes Bivalvia, Gastropoda, and Cephalopoda, with
this article covering the first two groups (for cephalo-
pods, see Marine Foods: Edible Animals Found in the
Sea). Bivalves and gastropods constitute a huge fish-
ery resource, amounting to over 12 million tonnes
from marine harvest fisheries and aquaculture in
1999, and over 550 000 tonnes from freshwater. The
production methods are as diverse as the species, and
products range from high value, such as abalone, to
low value, such as periwinkles. This article provides
details on the size and diversity of the commercially
important species and includes information on major
production methods and countries, product types,
and current issues.
0002 Vernacular terms to describe edible molluscs vary
around the world and may cause confusion, e.g.,
‘clam’ may refer to at least three distinct taxonomic
groupings. Vernacular names will be used only for
important species, with scientific taxonomy used
generally.
Bivalves
0003 There are approximately 20 000 bivalve species
worldwide, with the principal commercial species
belonging to three taxonomic orders: Mytiloida
(including mussels, scallops, and oysters), Veneroida
(true clams and cockles), and Myoida (soft-shell
clams and geoducks).
0004 Bivalves are generally characterized by paired shell
valves joined at a dorsal hinge and a sessile, filter-
feeding way of life. Their internal body form consists
of adductor muscles (usually paired) for shell closure,
tbl0001 Table 1 Principal commercial mussels, including main production sources and methods
Species Commonname
Mytilus edulis Blue
Mytilus galloprovincialis Mediterranean
Mytilus chilensis Chilean
Aulacomya ater Cholga
Perna viridis Green
P. perna South American rock mussel
P. canaliculus New Zealand or green lipped
A, aquaculture; F, fisheries.
respiration, a digestive gland, and seasonally variable
gonad tissue.
These features influence their commercial import- 0005
ance and use as food products. Generally, bivalves are
consumed whole, e.g., oysters and mussels, or pro-
cessed to adductor muscles (meat) and/or gonad (roe),
e.g., scallops. Muscle tissue is rich in carbohydrate
(glycogen) and protein, and mature gonad tissue is
high in lipid.
Mussels
Production Commercially important mussels be- 0006
long to the family Mytilidae and are predominantly
marine (see Table 1). Freshwater mussels involved in
pearl production belong to the Order Unionida. Com-
mercial Mytilid species come from wild harvest fish-
eries (global production of 237 823 tonnes in 1999)
and a greater proportion from aquaculture (1 451 032
tonnes in 1999). Global mussel aquaculture at first
sale was worth US$0.52 billion in 1999.
Harvest fisheries Harvest fisheries target three main 0007
species, Mytilus edulis (Blue) (121 964 tonnes in
1999), M. galloprovincialis (Mediterranean) (55 819
tonnes) and Perna viridis (Green) (21 520 tonnes).
The main countries harvesting M. edulis were Den-
mark (96 215 tonnes), Canada (11 565 tonnes), UK
(7972 tonnes), USA (4086 tonnes), and France (2098
tonnes). Production of M. galloprovincialis comes
from Italy (37 876 tonnes), Greece (15 860 tonnes),
and Turkey (1800 tonnes). Thailand dominates pro-
duction of P. viridis (21 500 tonnes). Mytilid mussels
are principally caught by dredging flat seabed areas
where the animals occur in dense aggregations,
connected by byssus, which are proteinaceous threads
used for attachment in many bivalves.
Aquaculture Aquaculture production is also dom- 0008
inated by M. edulis at 498 461 tonnes, worth
US$0.27 billion, and ranked as the 15th species
by quantity under cultivation. The principal produ-
cing countries are Spain (261 969 tonnes), The
Main production sources andmethod
Europe/North America/China (F/A)
Europe, South Africa (F/A)
Chile (F/A)
Chile, Peru (F)
South-east Asia (F/A)
Brazil, Venezuela (A)
New Zealand (A)

Netherlands (100 800 tonnes), France (51 600
tonnes) and Germany (37 912 tonnes).
0009 Also important is M. galloprovincialis at
162 179 tonnes, valued at US$100 million. Italy
(130 000 tonnes), Greece (16 912 tonnes), and France
(10 900 tonnes) dominate production. The green-
lipped mussel (Perna canaliculus) (not to be confused
with the green mussel, P. viridis) is also a significant
aquaculture species at 71 000 tonnes, worth US$21.3
million, with production dominated by New Zealand.
The other main species, P. viridis, provides 68 509
tonnes, is worth US$8.3 m, and is cultivated in Thai-
land (40 300 tonnes) and the Philippines (15 478
tonnes).
0010 It should be noted that China is the dominant
aquaculture producer of mussels at 608 115 tonnes,
worth US$90 million. International statistics do not
recognize individual species composition in Chinese
production, but other sources indicate that M. edu-
lis dominates, with additional supply from M. cor-
uscus, Musculus senhousei, and P. viridis. All are
local species. Mussel aquaculture is principally
based on suspended longline or raft culture,
although other methods exist. (See Shellfish: Aqua-
culture of Commercially Important Molluscs and
Crustaceans.)
0011 Product types Mussels are traditionally a chilled,
live product, particularly in Europe, where a post-
harvest shelf-life of around 5 days is expected. Re-
cently, new packaging and processing methods,
greater acceptance of convenience foods, and value
adding have led to new products. Mussels are now
frequently sold as individually quick frozen (IQF)
cooked in the whole or half shell, precooked ready-
to-eat meals and high value, luxury products such as
smoked mussels. Traditional pickled or brined pre-
served products are common.
tbl0002 Table 2 Principal commercial scallops, including main production sources and methods
Species Common name
Pecten maximus Great/king scallop
Pecten novaezelandiae New Zealand scallop
Pecten fumatus/meridionalis Commercial scallop
Placopecten magellanicus Sea scallop
Chlamys islandica Iceland scallop
Chlamys farreri Zhikong scallop
Patinopecten yessoensis Yesso scallop
Patinopecten caurinus Weathervane scallop
Argopecten irradians Bay scallop
Argopecten purpuratus Peruvian Calico scallop
Argopecten ventricosus Pacific Calico scallop
Aequipecten opercularis Queen scallop
Amusium balloti Saucer scallop
A, aquaculture; F, fisheries.
SHELLFISH/Commercially Important Molluscs 5223
Issues As coastal filter feeders, where the whole 0012
animal is consumed, food safety is important for
mussels and other bivalves. Accumulation of bacteria,
viruses, toxic chemicals, elements, and biotoxins is
associated with bivalves. Most edible bivalves, in-
cluding mussels, are generally subject to purification,
or depuration, which places the shellfish in a clean
water flow for 12 h or more. This enables expulsion
of many potentially harmful contaminants. Health
certification of live mussels is now common, as are
monitoring and certification of growout waters.
However, depuration does not remove biotoxins, 0013
produced by microalgae (more accurately phycotox-
ins), and their control and impact on mussel produc-
tion are generally managed by long-term monitoring
programs. These toxins may result in illnesses such as
paralytic, diarrhoetic and amnesic shellfish poison-
ing. In severe biotoxin outbreaks, area closures may
be necessary, and these have been well reported in
recent years. The protozoan parasites Marteilia spp.
affect Mytilus spp., although generally, mussels
appear to little affected by disease in culture systems.
Scallops
Production Commercial scallops belong to the 0014
family Pectinidae, and the principal species are listed
in Table 2. Global harvest fishery production in 1999
was 567 507 tonnes and from aquaculture 951 866
tonnes. Aquaculture production has increased from
340 807 tonnes in 1990, with an equivalent increase
in value from US$0.55 billion in 1990 to US$1.35
billion in 1999.
Harvest fisheries Fisheries are dominated by Pati- 0015
nopecten yessoensis (305 510 tonnes), Placopecten
magellanicus (131 962 tonnes), Pecten maximus
(35 411 tonnes), and Argopecten purpuratus (30 141
tonnes). Japan (299 628 tonnes, 98%) is the main
Main production sources andmethod
Europe (F/A)
New Zealand (F)
South-eastern Australia (F)
USA, Canada (F)
Iceland, Canada, USA (F)
China (A)
Japan, Russia, China (F/A)
Canada, USA (F)
China, USA (F/A)
Chile, Peru (F/A)
Mexico (F/A)
Europe (F/A)
Western and eastern Australia, Japan (F)
 5224 SHELLFISH/Commercially Important Molluscs
producer of P. yessoensis, with the Russian Feder-
ation (5764 tonnes, < 2%), providing the remainder.
P. magellanicus production is from the USA (77 179
tonnes, 59%) and Canada (54 756, 41%), and
P. maximus fishery production derives mainly from
the UK (19 108 tonnes, 54%) and France (12 745
tonnes, 36%). Peru provided all production of
A. purpuratus in 1999.
0016 Wild harvest of all the main commercial scallop
species usually involves a toothed (e.g., Pecten
spp.) or smooth-framed dredge (e.g., Placopecten
spp.), or occasionally conventional trawl gear (e.g.,
Amusium spp., Argopecten gibbus). There are several
large-scale diver fisheries for some species, e.g.,
Chile, China, and Mexico, and some small fisheries
that supply high-value, live product markets, e.g.,
Scotland.
0017 Scallop fisheries are noted for catch fluctuations, as
a result of exploitation and recruitment variability.
Several fisheries have consequently declined or
closed, although notably, the successful application
of stock enhancement and management programs
helped to revive Japanese and New Zealand produc-
tion during the 1970s and 1990s.
0018 Aquaculture Three countries and two species dom-
inate scallop aquaculture production. In 1999, China
was the largest producer of P. yessoensis at 712 330
tonnes, followed by Japan at 216 017 tonnes. Chile
produced 20 668 tonnes of A. purpuratus, with the
remainder from Peru (1585 tonnes). On a much
smaller scale, and using various species, other coun-
tries are developing pectinid aquaculture. For
example, in 1999, the UK produced 114 tonnes of
Aequipecten opercularis, whereas France, Spain,
Ireland and the UK produced over 700 tonnes of
P. maximus.
0019 Scallop production from aquaculture is based on
suspended longline culture or on some form of seabed
culture.
0020 Product types Scallops are sold in processed form;
fresh or frozen, meat (muscle) only or meat with
gonad (roe) attached, although there is a small, valu-
able market for live scallops, particularly in Western
Europe. Processing removes the meat (and gonad)
from the remaining viscera and shell, a procedure
known as shucking. This is usually a manual oper-
ation, although machines are available. Usable prod-
uct may be block frozen in multi-kilogram amounts
or IQF if the product is of a higher value and quality.
0021 In scallop marketing, the term Coquille St Jacques,
refers to the Genus Pecten and is specified under
French law. This category is important because of
France’s position as one of the principal scallop
markets. True Pecten species are sold as adductor
muscle and gonad together (roe-on) and command
higher prices. This premium relates partly to selling
gonad also, providing a greater return per scallop.
China is unique in predominantly producing a 0022
dried scallop meat product, although small quantities
are locally sold live.
Issues Because of the type of product (meat and/or 0023
gonad only), scallops are less affected by contamin-
ation and biotoxin problems, as the contaminated
portions (mainly mantle and digestive gland) are
discarded during processing. However, microalgae
biotoxins can concentrate in meat and gonad, and
have resulted in production closures in some coun-
tries (e.g., UK) and the widespread establishment
of precautionary monitoring programs (e.g., New
Zealand, UK, various states in the USA).
Diseases affecting scallops appear to be rare com- 0024
pared with other bivalves, with Rickettsia-like and
Chlamydia-like organisms occasionally associated
with mass mortalities of wild scallops and hatchery
mortalities. Outbreaks may be associated with scal-
lop density. The protozoan Marteilia spp. has been
implicated in mass mortalities of wild scallops in the
USA, although such events are rare.
Oysters
Production Commercial oysters belong to the 0025
family Ostreidae, and principally to the genera
Ostrea and Crassostrea (see Table 3). The global
wild harvest in 1999 was 157 538 tonnes, and
aquaculture production 3 711 606 tonnes, valued at
US$3.4 billion. Aquaculture production has increased
from 1 251 660 tonnes since 1990 (value US$1.57
billion).
Harvest fisheries The dominant fishery species in 0026
1999 was the American oyster, Crassostrea virginica
(132 207 tonnes, 84% of total), followed by the Pa-
cific (cupped) oyster, C. gigas, at 12 271 tonnes (8%).
The genus Ostrea contributed an additional 5951
tonnes, the majority (2456 tonnes, 41%) from
O. edulis, the European flat oyster, the remainder
from O. lutaria and other minor species.
Fishery production of C. virginica in 1999 was from 0027
three countries: the USA (89 714 tonnes), Mexico
(39 268 tonnes), and Canada (3225 tonnes). Almost
all (11 609 tonnes) of C. gigas production comes from
China, Taiwan, and South Korea. Fishery production
of O. edulis derives from Turkey (840 tonnes), Ireland
(680 tonnes), Spain (443 tonnes), and the UK (407
tonnes). New Zealand produced 994 tonnes of the
dredge oyster, O. lutaria, and Mexico 2489 tonnes of
unnamed Ostrea spp. European Ostrea fisheries were

tbl0003 Table 3 Principal commercial oysters, including main production sources and methods
Species Commonname
Crassostrea gigas Pacific (cupped) oyster
Crassostrea virginica American oyster
Crassostrea iredalei Slipper oyster
Ostrea edulis European flat oyster
Ostrea lutaria Dredge oyster
Ostrea chilensis Chilean flat oyster
Saccostrea glomerata Sydney rock oyster
A, aquaculture; F, fisheries.
formerly more significant, but the protozoan, Bona-
mia ostreae, introduced from the USA, devastated
production in the 1980s. Commercial fishery
methods for oysters are predominantly dredge-
based, although hand picking occurs in some coun-
tries, e.g., USA, Australia.
0028 Aquaculture The dominant aquaculture oyster is
the Pacific (cupped) oyster, Crassostrea gigas, provid-
ing the greatest quantity of any aquaculture species in
1999 at 3 600 459 tonnes. This species accounted for
97% of all oyster aquaculture production with a
value of US$3.3 billion in 1999. China accounted
for 2 988 613 tonnes, with other major producers
including Japan 205 345 tonnes, Korea 177 259
tonnes, France 134 800 tonnes, and the USA 33 259
tonnes. Other aquaculture species are C. virginica
57 522 tonnes (54 037 tonnes, USA), C. iredalei
13 698 tonnes (Philippines), Ostrea edulis 6496
tonnes (Spain 3348 tonnes, France 2300 tonnes),
and Saccostrea glomerata (commercialis) 5104
tonnes (Australia). Production methods vary, al-
though most are based on the oyster’s natural attach-
ment to artificial structures, or seabed sowing.
0029 Product types Oysters are best known as live sea-
food, and most production is live, in-shell, or fresh
chilled in half-shell. Traditionally, oyster culture has
produced a bottled product, with the whole shucked
animal, washed and preserved in salt water. They
survive low-temperature storage for several weeks,
but half-shell and bottled oysters have a limited
shelf-life. Maintenance of quality and rapid transport
and sale are essential components of the industry.
Oysters are now sold as a frozen product, either in
whole or half shell. This maintains the traditional
appearance while limiting spoilage and health risk.
Other value-added products such as smoked oysters
and pre-cooked meals are becoming common.
0030 Issues Oysters are subject to many significant dis-
eases affecting fishery and culture operations. These
include the protozoans Bonamia spp. (particularly
SHELLFISH/Commercially Important Molluscs 5225
Main production sources andmethod
China, Japan, Korea, France, USA, New Zealand (F/A)
USA, Canada (F/A)
Philippines (F/A)
Europe (F/A)
New Zealand (F)
Chile (F/A)
Australia (A)
affecting Ostrea spp.), Marteilia spp. (affecting
Ostrea and Saccostrea spp. (causing QX (Queensland
unknown) disease in the latter), Mikrocytos spp.
(affecting several Crassostrea, Saccostrea, and Ostrea
spp.), Haplosporidium spp. (affecting Crassostrea
spp.), and Perkinsus spp. (affecting Crassostrea and
Haliotis spp. (abalone)). An iridovirus, causing gill
necrosis, is also associated with hatchery-reared
C. gigas.
From a human health perspective, oysters are often 0031
associated with food poisoning events, because of
their intake of bacteria and viruses and subsequent
consumption as a whole, raw product. Such events
focus attention on the production environment and
require management of local sources of pathogenic
effluent. Consequently, water-quality monitoring
and health certification of production areas are
common, along with a certification requirement
from importing countries.
Other Bivalves
Production International fisheries statistics com- 0032
bine true clams and cockles (order Veneroida) with
arks (order Arcoida, Anadara spp.) and indicate total
world production from fisheries at 812 501 tonnes,
with a further 2 744 846 tonnes from aquaculture,
alone worth US$3.15 billion. This constitutes a
commercially significant group, despite including
numerous species. They are treated under broad
groupings, with taxonomic relationships indicated
where appropriate.
Order Veneroida This includes, amongst many 0033
others, the families Veneridae (venus clams), Cardii-
dae, (cockles), and Tridacnidae (giant clams). Several
Ruditapes spp. (carpet shells), support a modest
harvest fishery, mainly Japanese, producing around
58 874 tonnes in 1999. The most important carpet
shell is Ruditapes philippinarum (Manila clam),
which also provided 1 820 413 tonnes (worth
US$2.19b) from aquaculture, mainly from China.
Other cultivated Ruditapes spp. added 54 421 tonnes
and US$0.13 billion. The fishery for Chamelea
 5226 SHELLFISH/Commercially Important Molluscs
gallina (striped venus) was 45 012 tonnes in 1999,
and mainly from Italy (36 462 tonnes).
0034 Meretrix spp., known as hard clams, are fished in
South-east Asia, and provided 22 100 tonnes in 1999.
Indonesia is the largest supplier with 17 990 tonnes.
This genus is also cultivated in South Korea (Meretrix
lusoria, 17 tonnes), but more widely in Vietnam,
where 80 000 tonnes of M. meretrix and M. lyrata
are produced per annum. This product is sold into
South-east Asia, Japan, and Europe, although EU
regulations regarding biotoxin monitoring have
limited the European market.
0035 Arctica islandica (Ocean or black Quahog) is an
important fishery species in North America, with the
USA producing 147 933 tonnes in 1999. Iceland also
produces a small quantity (3501 tonnes) through har-
vest fisheries, most of which is exported to the USA.
0036 Mercenaria mercenaria (also known as the north-
ern Quahog or hard-shell clam) is also principally a
North American species, with Canada and the USA
traditional producers. In recent years, only Canada
has provided a harvest fishery product of 2536 tonnes
(1999), with a further 53 050 tonnes, worth US$63.4
million, from aquaculture, equally from China and
the USA. As with most clam culture, spat are sown on
to preselected sand/mud bottoms at densities appro-
priate for good growth and survival and with well-
exchanged phytoplankton rich water. Harvesting is
by dredge. To reduce growout periods, some clams
are cultured on trays, either suspended or stacked off
the bottom. This improves growth through increased
exposure to nutrients and higher temperatures.
0037 Fishery production of Paphia spp. (short-necked
clams) was 42 282 tonnes in 1999, from Thailand
and the Philippines. The North Atlantic surf clams,
Spisula solidissima and S. polynyma, are produced
from harvest fisheries in the USA (142 370 tonnes in
1999) and Canada (26 722 tonnes), respectively.
Razor clam (Solen spp.) aquaculture in China is sig-
nificant with 479 252 tonnes, worth US$383 million,
produced in 1999. Additional sources indicate that
this species may also be reported as Sinonovacula
constricta.
0038 The European edible cockle, Cerastoderma edule,
is dredge-fished in The Netherlands (50 888 tonnes)
and the UK (14 123 tonnes), for a total production of
70 401 tonnes in 1999. Recent aquaculture, based in
Spain (3713 tonnes) and France (1300 tonnes), also
contributed 5169 tonnes to production, valued at
US$8.9 million. Aquaculture produced very small
quantities (2 tonnes, in 1998) of cockles (Family
Cardiidae) in the USA.
0039 Order: Myoida The soft-shell clam, Mya arenaria,
and Geoducks, Panopea abrupta are produced and
consumed in North America. In 1999, the USA
and Canada produced 5113 and 2643 tonnes of
M. arenaria, and 2451 and 1800 tonnes of P. abrupta,
respectively.
Order: Arcoida There are three main genera of Ark 0040
‘clams’ of commercial importance, Arca spp., Sca-
pharca spp., and Anadara spp. Fishery production
of Arca spp. and Scapharca subcrenata totalled
52 972 tonnes in 1999. Aquaculture of S. broughtonii
in South Korea produced 8550 tonnes, worth
US$49.8 million.
Dredge fisheries also produced 42 627 tonnes of 0041
Anadara spp. in 1999, with A. granosa (misleadingly
called the blood cockle) the main species. Aquacul-
ture of A. granosa was 315 811 tonnes, worth
US$277.4 million, mainly from China (188 355
tonnes) and Malaysia (79 912 tonnes).
Gastropods
The class Gastropoda is commercially less important, 0042
although the marine abalone, periwinkles, conchs
and whelks, and terrestrial snails are significant as
food products. Other species of gastropod are valued
as shell products, such as mother of pearl and speci-
men shells. Gastropods are generally characterized by
a single spiral shell and large muscular foot, enabling
greater motility than bivalves and forming the princi-
pal edible portion. Fishery production of gastropods,
as abalone, winkles, and conchs was 105 047 tonnes
in 1999. Equivalent aquaculture production was
2694 tonnes, worth US$32.4 million.
Abalone 0043
Production Abalone (family Haliotidae) is mainly a
temperate marine group, although they extend into
tropical areas. Approximately 50 species are recog-
nized, associated with rocky habitats off Australia,
New Zealand, Japan, Pacific North America, and
South Africa. The main commercial species are
shown in Table 4. Abalone production is from
divercollected harvest fishery or aquaculture.
Harvest fisheries The total fishery production was 0044
10 453 tonnes in 1999, the majority from Australia
(5593 tonnes of Haliotis rubra), followed by Japan
(2109 tonnes of H. gigantea). The value of combined
fisheries production has not been estimated, but, as
an indicator, the Australian fisheries production in
1999–2000 was worth approximately US$118 mil-
lion (A$235.6 million) (Australian Bureau of Agricul-
tural Resource Economics).
Commercial harvest is by diver using hookah or 0045
scuba equipment. Abalone attach to rocky substrates
 SHELLFISH/Commercially Important Molluscs 5227

tbl0004 Table 4 Principal commercial abalone, including main Other Gastropod species The following gastropod 0049
production sources and methods
molluscs are of minor commercial importance in
Species Commonname Main production terms of volume, but may be regionally or tradition-
sources andmethod ally important.
Haliotis gigantea Giant Japan (F)
Haliotis tuberculata Tuberculate Channel Islands, Periwinkles Periwinkles of the family Littorinidae 0050

or Ormer France (F/A) are small, intertidal algal grazers, associated with
Haliotis midae Perlemoen South Africa (F/A) rocky shores. They are a traditional source of food
Haliotis rubra Black lip Australia,
throughout their almost-worldwide distribution, par-
Solomon Islands (F/A)
Haliotis laevigata Green lip Australia (F/A) ticularly in Europe, where commercial production is
Haliotis roei Roe’s Australia (F) greatest.
Haliotis conicopora Brown lip Australia (F) Production of wild-harvested Littorinids was 4828 0051
Haliotis iris Paua New Zealand (F) tonnes in 1999. The principal species is Littorina
Haliotis rufescens Red Pacific North
littorea, with 3144 tonnes, mainly derived from Ire-
America (F/A)
land (3018 tonnes) and Spain (126 tonnes). Other
A, aquaculture; F, fisheries. Littorinids, without species designation, contributed
1714 tonnes, 78% from the UK, with the remainder
from the USA and Canada. European production
with a muscular foot, and they are manually re- is by hand harvesting from intertidal areas and
moved with a blunt knife or similar implement. represents a source of income for both local
Owing to the slow growth rates and high value, and transient workers. Periwinkles are bagged and
most countries regulate harvesting. Size and catch sold as a live product, or processed as brined or
limits and closed seasons apply in Australia, New pickled meat only. There is interest in cultivation,
Zealand, South Africa and the USA, although with principally in France and Scotland. French aquacul-
variable management success. ture production of Littorina spp. in 1999 was 800
tonnes, worth US$2.08 million.
0046 Aquaculture World aquaculture production in 1999
was estimated at 1885 tonnes, valued at US$30.6 Conchs The family Strombidae includes algal or 0052

million. China dominated with 1799 tonnes (H.
discus hannai and H. diversicolor) and Chile with
48 tonnes (H. discus hannai). Variable production
since 1990 has originated from South Africa, the
USA, Australia, and New Zealand. The principal
culture methods include suspended net culture, race-
way culture, or tidal pond culture. (See Shellfish:
Aquaculture of Commercially Important Molluscs
and Crustaceans.)
0047 Product types Most Australian abalone production
is canned, with 2340 tonnes (US$60 million)
exported to Hong Kong (40%), Japan (27%), and
Taiwan (20%) in 1999–2000. Other product types
detrital feeders, mainly associated with inshore
sandy or muddy areas throughout tropical and sub-
tropical seas. Conchs are an important marine
resource in the Caribbean, where the Queen conch
(Strombus gigas) is particularly well known. Conch
fishery production was 15 487 tonnes in 1999. The
largest producer was Mexico (8591 tonnes), followed
by Jamaica (1366 tonnes) and the Dominican Repub-
lic (1257 tonnes). Aquaculture of Strombus spp. is
developing with 9 tonnes, valued at US$61 000, pro-
duced by Netherlands Antilles and Turks and Caicos
Islands in 1999.
Whelks Whelks are predatory or scavenging mol- 0053

for fished and cultivated abalone include; parboiled
and frozen on shell, frozen on shell, chilled, frozen, or
salted meat, and vacuum packing. Live abalone are
common as a value-added market product or mail
order.
0048 Issues The long growout period of most abalone, up
to 5 years, may affect aquaculture viability, although
their final value is significant and may offset cash
flow issues. Abalone are affected by potentially sig-
nificant diseases, notably the protozoan, Perkinsus
olseni, Rickettsia-like infections, and the bacterial
luscs, of boreal, temperate, and tropical seas. The
term ‘whelk’ is associated with several families, with
Buccinidae and Melongenidae being commercially
important. The common whelk, Buccinum undatum,
and Busycon spp. are the principal species with
17 988 and 6181 tonnes, respectively, in 1999. The
B. undatum fishery is dominated by France (7885
tonnes), the UK (4925 tonnes), and Ireland (4561
tonnes). The USA and Canada produce all Busycon
spp., with 4624 and 1557 tonnes, respectively.
Nutritional features of commercially important 0054

pathogen Vibrio. molluscs Table 5 provides information on the

 5228 SHELLFISH/Contamination and Spoilage of Molluscs and Crustaceans

tbl0005 Table 5 Proximate nutritional composition (per 100 g fresh weight) of principal commercial molluscs (sources: Nettleton (1985),
Ulieg (1988))

Species Protein (g) Total fat (g) Carbohydrate (g) Energy (kJ) Cholesterol (mg) Fatty acids Moisture (%)
(saturated/
monounsaturated/
polyunsaturated/o-3)

Abalone 20.8 1.0 0.9 415 111a 0.1/0.09/0.12/ 0.04a 75.7

Haliotis iris
Scallop 15.4 1.3 2.7 349 102b 0.1/0.06/0.24/0.18c 78.7
Pecten novaezelandiae
Oyster 11–13 2.7 (0.8–3.2) 5.8 360 47 0.49/0.36/0.90/0.71 79.7
Crassostrea gigas
Mussel 12.0 2.2 (1.2–2.1) 4.5 372 63 0.41/0.50/0.75/0.43 80.9d
Mytilus edulis
Periwinkle 18.2 3.0 (1.2–4.5) 2.3 114 0.45/0.83/0.90/0.44
Littorina littorea
Hard-shell clam 9.2 1.0 (0.2–2.0) 60 11.7 40e 0.16/0.16/0.33/0.24
Mercenaria mercenaria
a
Data from Haliotis japonica.
b
Data from Pecten fumatus.
c
Data from Placopecten magellanicus.
d
Data from Perna canaliculus.
e
Estimated from cooked product assuming 95% retention.

nutritional composition of edible molluscs. It should
be noted that, particularly for molluscs consumed
whole, there is considerable variation in content, as
gonadal tissue is highly seasonal.
See also: Marine Foods: Edible Animals Found in the Sea;
Shellfish: Aquaculture of Commercially Important
Molluscs and Crustaceans
Sheperd SA, Tegner MJ and Guzman del Proo SA (1992)
Abalone of the World. Oxford: Blackwell.
Shumway SE (ed.) (1991) Scallops: Biology, Ecology and
Aquaculture. Developments in Aquaculture and Fisher-
ies Science, vol. 21. Amsterdam: Elsevier.
Vlieg P (1988) Proximate Composition of New Zealand
Marine Finfish and Shellfish. Palmerston North, New
Zealand: Biotechnology Division, Department of Scien-
tific and Industrial Research.

Further Reading
FAO (2001a) FAO Yearbook, Fishery Statistics: Aquacul-
ture Production 1999. Fisheries Series No. 58, vol. 88/2.
Rome: FAO.
FAO (2001b) FAO Yearbook, Fishery Statistics: Capture
Production 1999. Fisheries Series No. 57, vol. 88/1.
Rome: FAO.
Gosling E (ed.) (1992) The Mussel Mytilus. Developments
in Aquaculture and Fisheries, vol. 25. Amsterdam
Elsevier.
Guo X, Ford SE and Zhang F (1999) Molluscan aquaculture
in China. Journal of Shellfish Research 18(1): 19–31.
Hallegraeff GM (1993) A review of harmful algal blooms
and their apparent global increase. Phycologia 32(2):
79–99.
Kraeuter JN and Castagna M (eds) (2001) Biology of the
Hard Clam. Developments in Aquaculture and Fisher-
ies, vol. 31. Amsterdam: Elsevier.
Nettleton J (1985) Seafood Nutrition: Facts, Issues and
Marketing of Nutrition in Fish and Shellfish. New
York: Osprey Seafood Handbooks.
OIE (Office International des Epizooties) (2001) Inter-
national Aquatic Animal Health Code, 4th edn. Paris:
OIE.
Contamination and Spoilage of
Molluscs and Crustaceans
D R Livingstone, Plymouth Marine Laboratory,
Plymouth, Devon, UK
S Gallacher, Fisheries Research Services, Aberdeen,
UK
Copyright 2003, Elsevier Science Ltd. All Rights Reserved.
Introduction
Seafood is of worldwide economic and nutritional 0001
importance, providing about 16% of the total animal
protein consumed globally. Of this, approximately
25% is shellfish, corresponding to an annual 17 mil-
lion tons taken from sea, coastal, and estuarine
waters, with a total value on landing of about £10
billion. Included in the shellfish are crustaceans
(e.g., crab, lobster, shrimp), and bivalve (e.g., mussel,
oyster, clam, scallop), gastropod (e.g., whelk,