Aquaculture International 8: 123–138, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

Review

Scallop introductions and transfers, from an animal

health point of view

S. MORTENSEN
Institute of Marine Research, Department of Aquaculture, PO. Box 1870, Nordnes, 5817
Bergen, Norway (E-mail: stein.mortensen@imr.no)

(Received 9 March 2000; accepted 14 March 2000)

Abstract. The effects of transfers and introductions of bivalue molluscs are always more
or less unpredictable. Moving scallops, there is a risk of introducing pathogenic agents or of
disturbing the balance between potentially pathogenic agents and host species in the recipient
ecosystem. Risk is not eliminated by merely following official regulations. Some factors which
may be important in order to assess risk, and to choose the level of risk we are willing to accept,
are:
The existence of “stowaways”, and the action of mechanical vectors. One organism may
carry another, and it is seems impossible to obtain a “clean” organism, in spite of long quar-
antines.
The biological vector function. There may be unknown reservoirs, intermediate or alternative
hosts of pathogenic agents in the “new” environment. An introduced species may also carry
potentially pathogenic agents into a new ecosystem.
The often lacking knowledge concerning epizootiology. Life cycles of many pathogenic
agents are poorly known, as are the infectious dose of agent, influence of environmental factors
on disease, etc.
The diagnostic tools, and the “human factor” in diagnostics. Methods are often not sens-
itive enough to detect a pathogenic agent in a carrier state, or immediately after the agent has
entered the host, or at a low prevalence/how intensity level. It is also important that personnel
are sufficiently trained to do the diagnostic work.
The official priority of disease control. There are no notifiable diseases of scallops.
EC regulations, and the “one area/one agent/one bivalve species” management.
National and international trade, including illegal transport of live bivalves.
Considering these factors enables a better focus on the risk factors and the assessment of risks
associated with introductions and transfers of scallops. Consideration may also help to clarify
the scientific level of knowledge and experience required to handle scallop diseases, and focus
on practical problems in scallop health control.

Key words: bivalves, disease, introductions and transfers, risk assessment, scallops
124

Introduction

A number of bivalue molluscs have been moved around the world, both
intentionally and unintentionally. Oysters may be accidentally transferred,
attached to ships hulls or hidden in consignments of other material, but
this is less likely to happen with scallops. Scallops are rather loosely or
not at all attached to their substrate, they usually live in deeper waters than
oysters, mussels and clams, and more separate from other bivalve species in
their natural habitants. Scallop transfers and introductions have so far only
involved species destined for aquaculture.
Reasons for moving may be that the selected scallop is a large species,
a fast growing species, a species which has high status in export markets,
or a species where there is knowledge on how to cultivate it. Examples of
scallops which have been moved during the last decades are the Bay scallop,
Argopecten irradians (moved from USA to China), the Japanese scallop,
Patinopecten yessoensis (moved from Japan to China, Ireland and France),
and the Great scallop, Pecten maximus (moved from Norway to Chile).
To minimise the risk of unwanted effects, considerations are normally
done prior to introductions and transfers. Both the ICES code of practice
(ICES, 1995), EC regulations (Anon, 1991), The Animal Health Code from
Office International des Epizooties (OIE, 1997), and of course common veter-
inary practice are designed in order to assess risk, and avoid introductions
of pathogenic agents and exotic species with the consignments. Even if all
guidelines and recommendations are followed, it is impossible to predict all
possible effects of transfers and introductions, and to predict which disease
problems may follow.
The spread of pathogens frequently occurs ahead of the diagnostics. Fortu-
nately for the scallop industry, relatively few serious disease conditions have
been described in scallops (Table 1). May the reason be that scallops in
general have been less affected by serious diseases than e.g. oysters? Or
perhaps that some of the scallop diseases have not yet been observed and
described? We also know little about which of the diseases described in
other bivalve species, may affect scallops. Hence, we often do not have
adequate scientific background knowledge in order to assess the relevance
of an observed microbe or parasite in a batch of scallops. Learning from
introductions and transfers of other bivalve species is therefore essential, to
enable a proper risk assessment. Having reviewed defined risk factors, and the
history of bivalve introductions and transfers, some risk factors still remain.
The aim of the present work is to point these out, to clarify the scientific
level of knowledge and experience to handle scallop diseases, and to discuss
some practical problems in scallop health control. Some points of particular
 125

relevance are briefly discussed below, using examples mainly from oyster
pathology.

Discussion

What is “disease”?

Disease may be widely defined as any state in which the organism in one
way or another is impaired. Diseases may be caused by numerous factors,
such as toxins, malnutrition, autoimmune disorders, tumours or pathogenic
agents. Those which receive most attention is aquaculture are associated with
the presence of pathogenic agents. Disease is then a result of a series of
intricately correlated factors, some concerning the host, some related to the
pathogen and some to the environment (e.g. discussions by Hedrick (1998)
and Reno, (1998)). Placing any species in a new environment will, to an
unknown degree, disturb the equilibrium between these factors. The change
may result either in less favourable conditions for the scallop, or in more
favourable conditions for potentially harmful organisms. The result may be
“disease” if it leads to impaired life functions or to the death of the scallop.

What is a “pathogenic agent”?

Any organism which causes disease of its host, either due to its own replica-
tion in the host (like countless microbes), or due to damage caused by contact
or mechanical action (like larger macroparasites such as worms and coped-
pods), may be defined as a pathogenic agent. The distinction between more
or less harmless commensals and pathogenic agents may however be unclear.
Marine life is a matter of coexistence. The presence of one organism will
always create space and opportunity for others. Agents which we know to
be potential pathogens from descriptive studies of disease outbreaks may be
present for a long time without causing pathology, but at any one moment,
triggered by a disturbed balanced by the above mentioned factors may cause
disease. This may be the case with several of the organisms listed in Table 1.
One good example might be the prokaryote Rickettsiales-like organisms
which have been found in a variety of bivalve species, but which are generally
not considered pathogenic (Getchell, 1991; Bower et al., 1994). Rickettsiales-
like organisms were however associated with mass mortalities of the scallop
Placopecten magellancius in the US in 1979–1980 (Gulka et al., 1983), and
of the great scallop, Pecten maximus in France in 1987 (LeGall et al., 1988).
126

The existence of “stowaways”, and the action of mechanical vectors

The presence of these “usually harmless – potentially harmful” organisms

lead us to another problem; the existence of “stowaways”, and the action
of mechanical vectors. One organism will always carry another, and it
seems impossible to obtain “clean” animals, in spite of long quarantines.
An example of stowaways is hidden organisms in a consignment of bivalve
spat. Frequently, batches contain more species than those they are supposed
to contain, even if the batches have been (roughly) inspected, cleaned and
graded. Mechanical vectors are passive carriers, which are not needed for the
propagation of the species being carried.
In many cases of introductions and transfers of bivalve molluscs for
cultivation, no serious attempt has been made to avoid unwanted organ-
isms. The export of half grown Pacific oysters, C. gigas, spat from France
to Ireland in 1993 is an outstanding example. Examination after arrival of
the oysters, which had been certified “free from other species”, revealed
numerous other species: Several fouling organisms, other bivalve species
(which may potentially carry pathogens or parasites) and 67 species of phyto-
plankton, including dinoflagellate cysts (O’Mahony, 1993; Minchin et al.,
1993). Most of these accompanying organisms will disappear or have no
or minor effects on cultivated species in their new environment. However,
sometimes we will get unwanted surprises, as the new species may cause
permanent or long-lasting fouling problems, competition for space or food,
or in extreme cases – disease.
While fouling macro-organisms may be relatively easy to find and identify
if in appropriate numbers, microbes will be more troublesome. Through
feeding, scallops will filter an unknown variety of protozoa, bacteria and
viruses. The protozoa and bacteria will in most cases be digested, but labor-
atory tank cohabitation experiments of Atlantic salmon, Salmo salar with
scallops, P. maximus challenged with the bacterium Aeromonas salmonicida
subspecies salmonicida, causing the disease furunculosis in salmonid fish
species, showed that furunculosis was transmitted to susceptible salmon via
the scallops (Bjørshol et al., pers. comm).
Viruses can be surprisingly inert. After the finding of the fish patho-
genic infectious pancreatic necrosis virus (IPNV) in scallops, P. maximus
(Mortensen et al., 1990), the subsequent study of the fate of IPNV in scal-
lops (Mortensen et al., 1992; Mortensen, 1993) showed that the virus was
taken up during filtration, persisted for long periods of time, and was shed
into the water by contaminated scallops. No viral propagation was found,
and in nature, the virus excreted from contaminated bivalves would rapidly
be diluted in seawater. Scallops and other bivalves should probably still be
considered potential vectors of fish pathogenic viruses. The results from the
Table 1. Reported specific and potentially disease-causing organisms of scallops, with some comments on their significance

Type of organism Host scallop Target tissue/ Range References

Significance

VIRUS Virus-like particles Pecten novazealandiae Lesions in digestive gland New Zealand Hine and Wesney, 1997
epithelium

RICKETTSIAE Rickettsiae Pecten maximus Necrosis of gills of wild France Le Gall et al., 1988
& CHILAMYDIAE scallops, associated with Le Gall et al., 1991
mass mortalities
Placopecten magellanicus Gills Atlantic U.S.A. Gulka et al., 1983
Gulka & Chang, 1984
Patinopecten yessoensis Gills Japan Elston, 1986
Argopecten irradians Kidney Atlantic U.S.A. Morrison and Shum, 1983
Gills & Canada Leibovitz et al., 1984

Chlamydiae Argopecten irradians In digestive epithelia. Atlantic North Leibovitz, 1989

Mortalities of hatchery America Morrison and Shum, 1982
reared spat

BACTERIA Vibrio spp. Vibriosis is a common and widespread problem in scallop larvae rearing Tubiash et al., 1970
Brown, 1981
Riquelme et al. 1995
Nicolas et al., 1996

127
 128
Table 1. Continued

Type of organism Host scallop Target tissue/ Range References

Significance

ALGAE Placopecten magellanicus Peripheral mantle tissues Newfoundland Naidu, 1971

Canada

FUNGI Homodendrum sp. ? Placopecten magellanicus Adductor muscle Maine, U.S.A. See Getchell, 1991

Sirolipidium zoopthorum Argopecten irradians Mortalities of cultivated larvae Atlantic U.S.A. See Getchell, 1991
Davis et al., 1954

PROTOZOA Lechnophora auerbachi Aequipecten opercularis Damage to eye Ireland Harry, 1977, 1980

Pseudoklossia pectinis Pecten maximus Damage to kidney tissue France See Getchell, 1991

Unidentified coccidia Argopecten irradians Damage to kidney tissue U.S.A. Leibovitz et al., 1984

Marteilia sp. Argopecten gibbus Digestive tissues. Associated Florida, U.S.A. Moyer et al., 1993
with mortalities of wild
scallops

Unidentified Argopecten irradians Connective tissues. Associated China Chu et al., 1996
haplosporidian with mortalities of cultured
scallops
Table 1. Continued

Type of organism Host scallop Target tissue/ Range References

Significance

Perkinsus karlsoni Argopecten irradians Connective tissues. Associated Atlantic North McGladdery et al., 1993
with mortalities of cultured America Goggin et al., 1996
scallops

Perkinsus qugwadi Patinopecten yessoensis Connective tissues. Mortalities B.C., Canada Bower et al., 1998, 1999
of bottom cultured introduced Blackbourn et al., 1998
scallops

Nematopsis spp. Widespread, in intermediate life stage, in different species See Bower et al., 1994

Trichodina spp. Widespread, in different species, no or little importance See Bower et al., 1994

SPONGES Cliona spp. Widspread, in different spcies. Grows and forms holes and tunnels in the See Bower et al., 1994
shell. Usually little impact on host.

HYDROIDS Hydractinia echinata Placopecten magellanicus Shell deformities Atlantic U.S.A. Merrill, 1967

CESTODES Larval forms may be found in digestive tract of bivalves. Presumably limited impact on host See Getchell, 1991

TREMATODES Bucephalus spp. and Pecten alba May damage reproductive Australia Sanders and Lester, 1981
others Pecten fumatus tissues Heasman et al., 1996
Chlamys asperrima

129
Table 1. Continued

130
Type of organism Host scallop Target tissue/ Range References
Significance

NEMATODES Larval forms reported from bivalves, including the scallops Argopecten irradians, A. gibbus, Amusium See Getchell, 1991
balloti, Argopecten aequisulcatus, Anachlamys leopardus and Chlamys asperrismus. Normally
presumed to have limited impact on host

POLYCHAETES Polydora spp. Widespread, in different species. Bores tunnels in host shell. Usually little Bergman et al., 1982
impact on host, but shell may be weakened. Associated with mortalities of Blake and Evans, 1973
cultured P. maximus spat. Evans, 1969
Skeel, 1979
Mortensen et al.,
this issue

Ceratonereis tridentata Arogpecten gibbus Causing shell blisters Atlantic U.S.A. Wells and Wells, 1962

CRUSTACEA Pectenophilus ornatus Patinopecten yessoensis Copepod attached to, and Japan Elston et al., 1985
Chlamys farreri causing damage to gills. Nagasawa et al., 1988

Pinnotheres spp. Arogpecten irradians, Pea crabs which live as Atlantic U.S.A. Kruczynski, 1972
Argopecten gibbus commensals in mantle cavity Japan Lauckner, 1983
Placopecten magellanicus may affect host
Chlamys nipponensis

MOLLUSCA Odostomia sp. Argopecten irradians Affects mantle during feeding Atlantic U.S.A. Leibovitz et al., 1984
 131

referred work have been taken into consideration when the Norwegian veter-
inary and aquaculture authorities decide on the management of production
and movements of cultured bivalves and fish in Norwegian coastal areas.

The biological vector function

The risk of disease transmission becomes greater when there are true biolo-
gical vectors, where a pathogenic agent maintains its normal function and
even propagates. Considering the above-mentioned coexistence between any
animal and its microbes, the microecological balance may be disturbed
during an introduction or transfer. From the introduced scallop’s point of
view, there may be unknown reservoirs, intermediate or alternative hosts of
pathogenic agents in its “new” environment. From the point of view of the
inhabitant of the recipient environment, the “newcomer” may pose a threat,
bringing new microbes, which are potentially pathogenic agents for them.
There are numerous examples, although some have never been scientifically
proven, and none concern scallops. One example is the virus causing gill
disease, which eradicated the susceptible populations of Portuguese oyster,
Crassostrea angulata, from the French coast, while the resistant Pacific
oyster, C. gigas remained only slightly affected by the disease (Comps et al.,
1976; Comps, 1988). It has been hypothesised that the Pacific oyster, which
was actually introduced to France just before the first outbreaks, was actually
the vector, being adapted to the virus through generations of coexistence in
Japan. Another example is the oyster pathogen protozoan Marteilia refrin-
gens which must be present in some unknown intermediate host or stage in
the environment on the French oyster beds. While flat oysters, Ostrea edulis,
may be kept free from Marteilia in tanks using water from the oyster beds,
oysters once moved out on the beds become infected.

The often lacking knowledge concerning epizootiology

The above example of Marteilia in oysters illustrates that we often lack know-
ledge on the life cycles of even the best known bivalve pathogenic agents.
Marteilia refringens seem to go through several stages in a complex life cycle
(Grizel et al., 1974; Perkins, 1976). Concerning the Marteilia sp. documented
from the Calico scallop Argopecten gibbus from the coast of Florida (Moyer
et al., 1992), knowledge is more scarce. Thus, we do not know which scallop,
or other bivalve species, may be susceptible, which species might be vectors,
in which stage the parasite may be dispersed, or which species might be
intermediate hosts. The most serious oyster pest in Europe, the protozoan
Bonamia ostreae also illustrates the problem. At first sight it seems not to
have a complicated life cycle like Marteilia. Bonamia propagates by binary
132

fission until the host cell, the oyster haemocyte, bursts. But despite almost
20 y of studies, there are unanswered questions. We do not know why small
oysters are unaffected, but die due to the parasite when they approach sexual
maturity. A life cycle with a phase in the ovarian cycle has been suggested
(van Banning, 1990), but it is still not fully understood. Also, the host range
of many agents is largely unknown, and extensive studies are necessary in
order to identify possible host species. We tend to link the pathogenic agents
to the species in which they are first described, but may often be wrong.
When the protozoan Microcytos mackini was identified as the causative agent
of Denman Island Disease of Pacific oysters, C. gigas in British Columbia,
Canada, the agent was first linked to this oyster species, but then similar
organisms were observed also in flat oysters, O. edulis, and Olympia oysters,
O. lurida, in the US, and Sydney rock oysters, Saccostrea commercialis,
in Australia. The causative agents were identified as two different Micro-
cytos species (Farley et al., 1988). Later challenge experiments showed that
M. mackini was pathogenic also for the oysters Crassostrea virginica, Ostrea
edulis and Ostreola conchaphila (Ostrea lurida) (Bower et al., 1997). The
example illustrates that what may seem as one disease in one species may
appear in different areas, and be caused by different, but related parasites –
parasites which themselves may be pathogenic for different host species. This
complicates the one disease-one host-one area management approach, which
is commonly applied (see below).
Even when we have documented that a specific agent is actually patho-
genic, there are often great uncertainties concerning the infectious dose of
agents, influence of environmental factors on disease, etc.

Diagnostic tools, and the “human factor”

In diagnostic work, there are some obvious problems. The main problem is
that we often have to know the parasite involved in order to be sure to detect
it during routine diagnostics. This has both human and technical aspects.
Firstly, is whether the personnel are sufficiently skilled and trained to do
the diagnostic work. Bivalve tissues, unlike vertebrate ones, undergo normal,
but significant alterations, due to reproductive and digestive cycles, and
general physiological state. This means that the people doing the diagnostic
work have to be trained – not only to detect the parasite in question, but also
to distinguish abnormalities due to disease from normal variation in tissues
due to the physiological state of the animal. Few practitioners have enough
expertise.
Secondly, is related to diagnostic methods. Classical histopathology with
microscopy of sectioned, paraffin-embedded tissues is still the basic tech-
nique in bivalve pathology. In addition to the “human factor” comes the
 133

quality of the histological sections. If not perfectly dissected, the cross

sections may be devoid of delicate parts like gills or parts of the mantle.
With scallops, the cross sections do not include all organs. To screen all soft
parts, more than one section is therefore needed, and the screening becomes
even more laborious. Additionally, a few 3 micron thick sections represents
a very small part of a big scallop. How sure are we to pick up a pathogen
in our sample specimens? If we search for unknown agents, we do not even
know in which organs they may be found. Concerning the methods, altern-
ative diagnostic tools have not been developed for scallop parasites – with an
exception of a monoclonal antibody against the Rickettsiales-like organisms
from P. maximus in France (Le Gall et al., 1992). Today, this antibody is
however not used.
Thirdly, is how to be able to detect parasites if the host is in a carrier state.
If the parasite is present in low prevalence, with low intensity, or immediately
after the agent has entered the host. Guidelines recommend sampling at a
level where 2% carriers in a population are detected on a 95% confidence
level. This means investigation of at least 147 specimens twice a year, given
that the “human factor” does not spoil the statistics. Health surveys on this
level are laborious and expensive, and are normally only carried out when
there are severe disease problems, and with notifiable diseases.

The official priority of scallop disease control

There are no notifiable diseases of scallops. As a result, no continuous health

survey is likely, as the surveys are expensive, and there will be no external
pressure or expectations of a health survey of non-notifiable diseases.
There is often also a tendency to give priority to health control of cultured
species supporting a commercial activity, as money still seems to be more
important than ecology. Very few of the worlds scallop species support such
an activity, but disease problems in large-scale scallop culture in the Far
East could clearly induce the development of scallop disease control and
diagnostic methods and routines.
Most commercially interesting scallop species support fisheries. It is more
difficult to keep continuous surveys on bivalves on the bottom than on
bivalves in the tidal zone or suspended cultures. Once scallops are harvested,
they are processed, and no longer available for examination. Unfortunately,
there are rarely health inspections and samplings on vessels. As one of the
exceptions, samplings of dredged scallops are carried out by pathologists
at the British Ministry of Agriculture, Fisheries and Food (S. Feist, pers.
comm.). Their investigations give baseline data on the health and parasite
fauna in wild scallops, and should be developed further.
134

EC regulations

The EC system, of replacing European borders with epizootiological zones,

is based on sound principles of using national information on the distri-
bution of animal diseases and geographical barriers. Movement of bivalves
for relaying is permitted only between zones of equal zoosanitary status, or
from uninfected zones to infected ones. The management is however applied
only on species in which notifiable diseases have been detected, which does
not include scallops. Consequently, EC legislation permits movement of live
animals carrying organisms, which potentially might cause a problem, as long
as they at the time being do not cause problems which are severe enough to
put them on the international disease lists. In addition, the regulations easily
create a “one area/one agent/one bivalve species” management in EC coun-
tries, as well as in countries trading with the EC. Due to the points concerning
epizootiology and vector functions stated above, these regulations are far
from a “no-risk” objective. EC regulations favour trade, rather than either
nature conservation or animal welfare.

National and international trade

There is different health status in different zones, and may also be in different
areas within a country which has not been divided into zones. The regula-
tions regarding production and trade of bivalve molluscs laid down in the
EC directives distinguish between bivalves for aquaculture (directive 91/67)
and for human consumption (directive 91/492). Bivalves meant for immediate
human consumption may be re-laid in water, or kept alive in tank systems, in
other areas than their origin. Even though scientists within the EU are aware
of this, and are trying to influence the authorities to change the regulations,
trade goes on. Changes within a large administrative system take time. In
addition to this, illegal import of both aquaculture products and bivalves for
consumption sometimes occur. People involved in the trade are frequently not
familiar with the health and disease problems discussed here.

Conclusions

1. History has shown that the effects of transfers and introductions are
always more or less unpredictable. There is still a risk, even after the
guidelines and regulations have actually been followed. Knowledge,
control and monitoring of all species in aquaculture is thus important,
in order to actually choose the level of risk we are willing to accept.
Consideration both of official regulations and guidelines, as well as the
 135

“unofficial” factors discussed in the present work, will enable us to make

better judgements.
2. There have been few recorded serious disease outbreaks in scallops, and
the published papers are nearly all descriptive. Due to this scarcity of
information, the risk associated with introductions and transfers of scal-
lops will be difficult to estimate. Monitoring of hatchery operations,
gathering of information from scallop aquaculture, and surveys of wild
populations are necessary, even though it is not officially required at
present.

Acknowledgements

My sincere thanks to Dan Minchin for useful background data, and to Lisbeth
Harkestad for excellent assistance during collection and systematisation of
the literature cited.

References

Anonymous (1991a). Council Directive of 28 January 1991 (91/67/EEC), concerning the

animal health conditions governing the placing on the market of aquaculture animals and
products. Official Journal of the European Communities, No. L 46/1.
Anonymous (1991b) Council Directive of 15 July 1991 (91/492/EEC), laying down the health
conditions for the production and the placing on the market of live bivalve molluscs.
Official Journal of the European Communities, No. L 288/1.
Bergman, K.M. Elner, R.W. and Risk, M.I. (1982) The influence of Polydora websteri borings
on the strengths of the shell of the sea scallop, Placopecten magellianicus. Canadian
Journal of Zoology 60, 2551–2556.
Blackbourn, J., Bower, S.M. and Meyer, G.R. (1998) Perkinsus qugwadi sp.nov. (incertae
sedis), a pathogenic protozoan parasite of of Japanese scallops, Patinopecten yessoensis,
cultured in British Columbia, Canada. Canadian Journal of zoology 76, 942–953.
Blake, J.A. and Evans, J.W. (1973) Polydora and related genera as bores in mollusks shells
and other calcareous substrates (Polychaetea: Spionidae). Veliger 15, 235–249.
Bower, S.M., Blackbourn, J. and Meyer, G.R. (1998) Distribution, prevalence, and pathogen-
icity of the protozoan Perkinsus gugwadi in Japanese scallops, Patinopecten yessoensis,
cultured in British Columbia, Canada. Canadian Journal of Zoology 76, 954–959.
Bower, S.M., Blackbourn, J., Meyer, G.R. and Welch, D.W. (1999) Effect of Perkinsus
qugwadi on various species and strains of scallops. Disease of Aquatic organisms 36,
143–151.
Bower, S.M., Hervio, D. and Meyer, G.R. (1997) Infectivity of Microcytos mackini, the causa-
tive agent of Denman Disease in Pacific oysters Crassostrea gigas, to various species of
oysters. Diseases of Aquatic Organisms 29, 111–116.
Bower, S.M., McGladdery, S.E. and Price, I.M. (1994) Synopsis of inectious diseases and
parasites of commercially exploited shellfish. Annal Review of Fish Diseases 4, 1–199.
136

Brown, C. (1981) A study of two shellfish – pathogenic Vibrio stains isolated from a Long
Island hatchery during a recent outbreak of disease. Journal of Shellfish Research 1, 83–87.
Chu, Fu-Lin E., Burreson, E.M., Zhang, F. and Chew, K.K. (1996) An unidentified haplospor-
idian parasite of bay scallop Argopecten irradians cultured in the Shandong and Liaoning
provinces of China. Diseases of Aquatic Organisms 25, 155–158.
Comps, M. (1988) Epizootic diseases of oysters associated with viral infections. In: Disease
Processes in Bivalve Molluscs. American Fisheries Society Special Publication 18 (ed.
W.E. Fisher), Bethesda, USA, pp. 23–37.
Comps, M., Bonami, J.-R., Vago, C. and Campillo, A. (1976) Une virose de l’huitre Portugaise
(Crassostrea angulata). C.R. Acad. Science, Paris 282 série D 22, 1991–1993.
Davis, H.C., Loosanoff, V.L. and Weston, W.H. and Matin, C. (1954) A fungus disease in clam
and oyster larvae. Science 120, 36–38.
Elston, R. (1986) Occurrence of branchial rickettsiales-like infection in two bivalve molluscs,
Tapes japonica and Patinopecten yessoensis, with comments on their significance. Journal
of fish Diseases 9, 69–71.
Elston, R.A., Wilkinson, M.T. and Burge, R. (1985) A rhizocephelan-like parasite of a bivalve
mollusc, Patinopecten yessoensis. Aquaculture 49, 359–361.
Evan, J.W. (1969) Borers in the shell of the sea scallop, Placopecten magellanicus. American
Zoologist 9, 775–782.
Farley, C.A., Wolf, P.H. and Elston, R.A. (1988) A long-term study of “micorcell” disease
in oysters with a description of a new genus, Microcytos (g.n.), and two new species,
Microcytos mackini (sp.n.) and Microcytos roughleyi (sp.n.). Fishery Bulletin 86(3), 581–
593.
Getchell, R.G. (1991) Diseases and parasites of scallops. In: Scallops: biology ecology
and aquaculture. Developments in Aquaculture and Fisheries Science No. 21 (ed. S.E.
Shumway), Elsevier press. NY, pp. 471–494.
Goggin, C.L., McGladdery, S.E., Whyte, S.K., Cawthorne, R.J. (1996) An assessment of
lesions in bay scallops Argopecten irradians attributed to Perkinsus karlssoni (Protozoa,
Apicomplexa). Diseases of Aquatic Organisms 24, 77–80.
Grizel, H., Comps, M., Cousserans, F., Bonami, J.-R. and Vago, C. (1974) Pathologie des
invertébrés. Etude d’un parasite de la glande digestive observée au cours de l’épizootie
actuelle de l’huitre plate. Comptes Rendus Academie des Science Paris, Series D. 279,
783–784.
Gulka, G. and Chang, P.W. (1984) Pathogenicity and infectivity of a rickettsia-like organism
in the sea scallop, Placopecten magellianicus. Journal of Fish Diseases 8, 308–318.
Gulka, G., Chang, P.W. and Marti, K.A. (1983) Procaryotic infection associated with a mass
mortality of the sea scallop, Placopecten magellanicus. Journal of Fish Diseases 6, 355–
364.
Harry, O.G. (1977) Observations on Licnophora auerbachi, a ciliate from the eyes of the queen
scallop Chlamys opercularis (Mollusca: Bivalvia). Journal of Protozoology 24, 48A.
Harry, O.G. (1980) Damage to the eyes of the bivalve Chlamys opercularis caused by the
ciliate Licnophora auerbachi. Journal of Invertebrate Pathology 36, 283–291.
Heasman, M.P., O’Connor, W.A. and Frazer, A.W.J. (1996) Digenean (Bucephalidae) infec-
tions in commercial scallops Pecten fumantus Reeve and doughboy scallops Chlamys
(Mimachlamys) asperrima (Lamarck) in Jervis Bay, New South Wales. Journal of Fish
Diseases 19, 333–339.
Hedrick, R.P. (1998) Relationships of the host, pathogen, and environment: Implications for
diseases of cultured and wild fish populations. Journal of Aquatic Animal Health 10, 107–
111.
 137

Hine, P.M. and Wesney, B. (1997) Virus-like particles associated with cytopathology in
the digestive gland epithelium of scallops Pecten novaezelandiae and tohera Paphies
ventricosum. Diseases of Aquatic Organisms 29, 197–204.
ICES (1995) ICES Code of Practice on the Introductions and Transfers of Marine Organisms
1994. International Council for the Exploration of the Sea, Copenhagen, Denmark. ICES
Co-operative Research Report No. 204. Report of the ICES Advisory Committee on the
Marine Environment, 1994, Annex 3.
Le Gall, G., Chagot, D., Mialhe, E. and Grizel, H. (1988) Branchial rickettsiales-like infection
associated with a mass mortality of sea scallop Pecten maximus. Diseases of Aquatic
Organisms 4, 229–232.
Kruczynski, W.L. (1972) The effect of the pea crab, Pinnotheres maculates Say, on the growth
of the bay scallop, Argopecten irradians concentricus Say. Chesapeake Science 13, 218–
220.
Lauckner, G. (1983) Diseases of mollusca: bivalvia. In: Diseases of Marine Animals.
Biologische Anstalt Helgoland, Hamburg 2 (ed. O. Kinne), 477–961.
Le Gall, G., Miahle, E., Chagot, D. and Grizel, H. (1991) Epizootiological study of rickett-
siosis of the Saint Jacques scallop Pecten maximus. Diseases of Aquatic organisms 10,
139–145.
Le Gall, G., Mourton, C., Boulo, V., Paolucci, F., Pau, B. and Mialhe, E. (1992) Mono-
clonal antibody against a gill Rickettsiales-like organisns of Pecten maximus (Bivalvia):
application to direct immunofluorescence diagnosis. Diseases of Aquatic Organisms 14,
213–217.
Leibovitz, L. (1989) Chlamydiosis: a newly reported serious disease to larval and postmeta-
morphic bay scallops, Argopec;ten irradians (Lamarck). Journal of Fish Diseases 12,
126–136.
Leibovitz, L., Schott, E.F. and Karney, R.C. (1984) Diseases of wild, captive and cultured
scallops. Journal of the World Mariculture Society 15, 269–283.
McGladdery, S.E., Bradford, B.C. and Scarrat, D.J. (1993). Perkinsus karlssoni n. sp.
(Apicomplexa) in bay scallops Argopecten irradians. Diseases of Aquatic organisms 10,
127–137.
Merrill, A.S. (1967) Shell deformity of mollusks attributable to the hydroid, Hydractinia
echinata. Fisheries Bulletin, Fish and Wildlife Service U.S. 66, 273–279.
Minchin, D., Duggan, C.B., Holmes, J.M.C. and Neiland, S. (1993) Introductions of exotic
species associated with Pacific oyster transfers from France to Ireland. International
Council for the Exploration of the Sea. C.M. 1993/F: 27, 11 pp.
Morrision, C. and Shum, G. (1982) Chlamydia – like organisms in the digestive diverticula of
the bay scallop, Argopecten irradians (Lmk). Journal of Fish Diseases 5, 173–184.
Morrison, C. and Shum, G. (1983) Rickettsias in the kidney of the bay scallop, Argopecten
irradians (Lamarc). Journal of Fish Diseases 6, 537–541.
Mortensen, S.H. (1993) Passage of infectious pancreatic necrosis virus (IPNV) through
invertebrates in an aquatic food chain. Diseases of Aquatic Organisms 16, 41–45.
Mortensen, S.H., Bachere, E., LeGall, G. and Mialhe, E. (1992) Persistence of infectious
pancreatic necrosis virus (IPNV) in scallops (Pecten maximus). Diseases of Aquatic
Organisms 12, 221–227.
Mortensen, S.H., Hjeltnes, B., Rødseth, O., Krogsrud, J. and Christie, K.E. (1990) Infec-
tious pancreatic necrosis virus, serotype N1 isolated from Norwegian turbot (Scopthalmus
maximus), halibut (Hippoglossus hippoglossus) and scallops (Pecten maximus). Bulletin
of the European Association of Fish Pathologists 10(2), 42–43.
138

Moyer, M.A., Blake, N.J. and Arnold, W.S. (1993). An ascetosporan disease causing mass
mortality in the Atlantic calico scallop, Argopecten gibbus (Linnaeus, 1758). Journal of
Shellfish Research 2, 305–310.
Nagasawa, K., Bresciani, J. and Lutzen, J. (1988) Morphology of Pectenophilus ornatus, new
genus, new species, a copepod parasite of the japanese scallop Patinopecten yessoensis.
Journal of Crustacean Biology 8, 31–42.
Naidu, K.S. (1971) Infection of the giant scallop Placopecten magellanicus from Newfound-
land with an endozoic alga. Journal of Invertebrate Pathology 17, 145–157.
Nicolas, J.L., Corre, S., Gauthier, G., Robert, R. and Ansquer, D. (1996) Bacterial problems
associated with scallop Pecten maximus larval culture. Diseases of Aquatic Organisms 27,
67–76.
OIE (1997) International Aquatic Animal Health Code, Office International des Epizooties,
2nd edition. ISBN 92-9044-420.
O’Mahony, J.H.T. (1993) Phytoplankton species associated with imports of the Pacific oyster
Crassostrea gigas, from France to Ireland. International Council for the Exploration of
the Sea. C.M. 1993/F: 26, 8 pp.
Perkins, F.O. (1976) Ultrastructure of sporulation in the European flat oyster pathogen,
Marteilia refringes – taxonomic implications. Journal of Protozzology 23, 64–74.
Reno, P.W. (1998) Factors involved in the dissemination of disease in fish population. Journal
of Aquatic Animal Health 10, 160–171.
Riquelme, C., Hayashida, G., Vergara, N., Morales, Y. and Chavez, P. (1995) Bacteriology of
the scallop Argopecten purpuratus (Lamarck, 1819) cultured in Chile Aquaculture 138,
49–60.
Riquelme, C., Hayashida, G., Toranzo, A.E., Vilches, J. and Chavez, P. (1995) Pathogenicity
studies on a Vibrio anguillarum-related (VAR) strain causing an epizootic in Argopecten
purpuratus larvae cultured in Chile. Diseases of Aquatic organisms 2, 135–141.
Sanders, M.J. and Lester, R.J.L. (1981) Further observations on a bucephalid trematode infec-
tion in scallops (Pecten alba in Port Phillip Bay, Victoria. Australian Journal of Marine
and Freshwater Reserach 32, 475–478.
Skeel, M.E. (1979) Shell – boring worms (Spionidae: polychaeta) Infecting cultivated bivalve
molluscs in Australia. Proceedings of the World Mariculture Society 10, 529–533.
Tubiash, H.S., Cilwell, R.R. and Sakazaki, R. (1970) Marine vibriosis associated with bacil-
lary necrosis, a disease of larval and juvenile bivalve mollusks. Journal of Bacteriology
103, 272–272.
van Banning, P. (1993) The life cycle of the oyster pathogen Bonamia ostreae with a
presumptive phase in the ovarian tissue of the European flat oyster, Ostrea edulis.
Aquaculture 84, 189–192.
Wells, H.W. and Wells, M.J. (1962) The polychaete Veratonereis tridentata as a pest of the
scallop Aequipecten gibbus. Biological Bulletin of the Marine Biological Laboratory,
Woods Hole 122, 149–159.