DISEASES OF PEARL OYSTERS AND OTHER MOLLUSCS: A WESTERN

AUSTRALIAN PERSPECTIVE
Author(s): J. B. JONES and J. CREEPER
Source: Journal of Shellfish Research, 25(1):233-238. 2006.
Published By: National Shellfisheries Association
DOI: http://dx.doi.org/10.2983/0730-8000(2006)25[233:DOPOAO]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.2983/0730-8000%282006%2925%5B233%3ADOPOAO
%5D2.0.CO%3B2

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Journal of Shellfish Research, Vol. 25, No. 1, 233–238, 2006.

DISEASES OF PEARL OYSTERS AND OTHER MOLLUSCS: A WESTERN

AUSTRALIAN PERSPECTIVE

J. B. JONES* AND J. CREEPER

Department of Fisheries, Government of Western Australia, P.O. Box 20,
North Beach, WA 6920, Australia

ABSTRACT Mollusc culture, particularly the cultivation of pearl oysters, is an important component of the aquaculture industry in
Western Australia. As a result, there has been a long-term investment in surveys of commercial mollusc species for potential diseases
of concern. A number of pathogens, particularly haplosporidans, identified within wild-stock shellfish have the potential to adversely
affect mollusc populations. Others pose risks for translocations associated with aquaculture. The microsporidan Steinhausia mytilovum
(Field), found in ova of the blue mussel Mytilus galloprovincialis (Lamarck), poses intriguing questions about the origin and dispersal
of its host.

KEY WORDS: abalone, ciliates, parasites, Perkinsus, Thraustochytridea, scallops

INTRODUCTION have now been overcome by improved management practices. The

The most valuable of the marine mollusc culture industries in
Western Australia is pearling. The main species cultured is the
golden-lipped pearl oyster Pinctada maxima (Jameson), but other
pearl shells grown commercially include P. margaratifera (L), P.
albina (Lamarck) and Pteria penguin (Röding). The pearling in-
dustry is supported by four operational hatcheries capable of sup-
plying all of the spat requirements of industry. Western Australia
also has a small mussel industry based on culture of Mytilus gal-
loprovincialis; an abalone culture industry based on hatchery pro-
duced Haliotis spp. (greenlip abalone, H. laevigata (Donovan),
brownlip abalone, H. conicopora Peron and Roe’s abalone, H. roei
Gray); a rock oyster farm hatching and growing Saccostrea sp.; a
hatchery producing scallops (Amusium balloti Bernardi) for re-
seeding and there is also some experimental reseeding of tropical
reefs with hatchery produced trochus (Tectus niloticus [L]).
Because of this aquaculture activity, the disease status of these
Western Australian molluscs is relatively well known. To date
there have been few diseases of concern. The health status of the
aquaculture industry in a State such as Western Australia is based
on two factors. The first is the presence of a disease reporting
procedure and a diagnostic capacity to identify causative agents.
The second is the existence of passive and targeted surveillance
programs aiming to identify endemic diseases not associated with
morbidity or mortality. In Western Australia a dedicated fish dis-
ease diagnostic unit of the Department of Fisheries has worked
with industry since 1988 to identify and solve disease problems.
There have also been 3 major mollusc disease surveys in the State;
one involved a wide range of molluscs around the coast (Hine &
Thorne 2000), one on P. maxima (Humphrey et al. 1998) and an
ongoing national survey of abalone diseases. Some additional in-
formation is available in isolated publications on individual para-
sites and further unpublished information is available from labo-
ratory records. Parasites that are known from shellfish in Western
Australia are listed in Table 1.
Pearl Oysters
The disease that has caused most economic loss to the P.
maxima industry is vibriosis (Pass et al. 1987). In the 1980s Vibrio
spp. bacteria caused significant losses to the pearling industry that
*Corresponding Author. E-mail: bjones@agric.wa.gov.au
presence of a very rare haplosporidan in the digestive tubules is
cause for concern (Hine & Thorne 1998) because very little is
known about the biology of this parasite. The haplosporidan has
been identified only three times and each time the oysters on the
infected farm site have been destroyed. During the second occur-
rence, in late December 1995, 4.6% of a sample of 150 farmed
juvenile pearl oysters was found infected within 6 weeks of being
set at a remote site in Western Australia. By the time the oysters
were destroyed 15 days later the infection prevalence had in-
creased to 10% (n ⳱ 238). The notifiable disease Perkinsus olseni/
atlanticus has been reported from P. maxima in the Torres Strait
(Norton et al. 1993a) but not from Western Australia, though the
disease agent does occur here in bivalve molluscs other than P.
maxima (Table 1).
Another parasite of concern in pearl oysters has only been
detected in spat from the Exmouth Gulf and outlying islands since
2001, despite comprehensive monitoring of oysters in the State and
the industry practice of harvesting and translocating shell to all
other oyster growing areas for over 50 y. It is an intracellular
ciliate similar in appearance to ciliates reported as nonpathogenic
commensals in Spanish Mytilus galloprovincialis (Villalba et al.
1997) and in Mytilus edulis (L) on the east and west coasts of
North America (Figueras et al. 1991a). The teardrop shaped baso-
philic ciliate (10–15 ␮m × 5 ␮m) has a dense polymorphic mac-
ronucleus and normally occupies an intraluminal or intraepithelial
location within the digestive gland in P. maxima spat (Fig. 1). The
ciliate is often associated with an inflammatory response in oysters
smaller than 70 mm with 20–50 mm shell being most affected. A
feature in smaller (20–40 mm) spat is the ciliate’s capacity to
penetrate the mucosal basal lamina and reside within hemolymph
spaces or free within interstitial connective tissue. Translocation of
infected pearl oysters beyond the Exmouth zone is not permitted.
The origin of the ciliate is unknown, and its relationship to
similar organisms seen in northern hemisphere mussels is un-
known. Either the ciliate has become much more prevalent in
recent years or it has been introduced into the region. Its infectivity
to other bivalves is also unknown.
During surveys to determine the distribution of the ciliate, a
single infected oyster was found with an enigmatic unidentified
proctistan parasite (Fig. 2). The putative sporoblasts develop
within the epithelial cells of the digestive tubules and fill the lumen
of the tubules, with an associated basophilic hemocyte inflamma-

233
234
Molluscan disease-causing agents (other than bacteria) reported from Western Australia.
Agent
Virus
Virus-like inclusions
Papova-like virus
Rickettsia-like organisms
Haplosporidia
Bonamia sp.
Haplosporidium sp.
Haplosporidium sp.
Marteilia sydneyi
Mikrocytos roughleyi
Microspora
Steinhausia mytilovum
Marteilioides sp.
Apicomplexa
Heart apicomplexan
Perkinsus olseni
Ciliates
Ciliates on gills
Ciliates in gut
Ancistrocomids
Sphenophyra-like ciliates
Trematoda
Proctoeces sp.
Sporocysts
continued on next page
JONES AND CREEPER
TABLE 1.
Host Associated with Epizootic? Reference
Pinctada maxima No Pass et al. (1988); Humphrey et al. (1998)
Haliotis laevigata No Laboratory records
Isognomon isognomum No Hine & Thorne (2000)
Pinna bicolor No Hine & Thorne (2000)
Saccostrea cucullata No Hine & Thorne (2000)
Haliotis laevigata No Laboratory records
Pinctada maxima No Norton et al. (1993b); Humphrey et al. (1998)
Barbatia helblingii No Hine & Thorne (2000)
Haliotis laevigata No Laboratory records
Isognomon isognomum No Hine & Thorne (2000)
Pinctada maxima No Humphrey et al. (1998)
Pinna bicolor No Hine & Thorne (2000)
Pinna deltoides No Hine & Thorne (2000)
Pteria penguin No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
Saccostrea cucullata No Hine & Thorne (2000)
Stavilia horrida No Hine & Thorne (2000)
Ostrea sp. Yes Laboratory records
Saccostrea cucullata Yes Hine & Thorne (2000)
Pinctada maxima No Hine & Thorne (1998)
Saccostrea cucullata No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
Saccostrea sp. No Laboratory records
Mytilus galloprovincialis No Laboratory records
Saccostrea echinata No Hine & Thorne (2000)
Pinctada maxima No Humphrey et al. (1998)
Barbatia helblingii No Hine & Thorne (2000)
Isognomon isognomum No Hine & Thorne (2000)
Malleus meridianus No Hine & Thorne (2000)
Pinctada albina No Hine & Thorne (2000)
Pinna deltoides No Hine & Thorne (2000)
Saccostrea cucullata No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
Septifer bilocularis No Hine & Thorne (2000)
Spondylus sp. No Hine & Thorne (2000)
Haliotis laevigata No Laboratory records
Pinctada spp. No Humphrey et al. (1998)
Saccostrea sp. No Laboratory records
Pinctada maxima No Laboratory records
Haliotis laevigata No Laboratory records
Pinctada albina No Hine & Thorne (2000)
Pinna bicolor No Hine & Thorne (2000)
Saccostrea cucullata No Hine & Thorne (2000)
Saccostrea echinata No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
Haliotis laevigata No Laboratory records
Saccostrea glomerata No Hine & Thorne (2000)
Malleus meridianus No Hine & Thorne (2000)
Haliotis laevigata No Laboratory records
Isognomon isognomum No Hine & Thorne (2000)
Pinctada albina No Hine & Thorne (2000)
Pinctada maxima No Humphrey et al. (1998); Pass (1987); Hine & Thorne (2000)
Malleus malleus No Hine & Thorne (2000)
Saccostrea echinata No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
 MOLLUSC DISEASES IN WESTERN AUSTRALIA 235

TABLE 1.
continued

Agent Host Associated with Epizootic? Reference

Cestoda
Tylocephalum sp. Dendostrea folium No Hine & Thorne (2000)
Isognomon isognomum No Hine & Thorne (2000)
Malleus malleus No Hine & Thorne (2000)
Malleus meridianus No Hine & Thorne (2000)
Pinctada spp. No Hine & Thorne (2000); Humphrey et al. (1998)
Pinna bicolor No Hine & Thorne (2000)
Pinna deltoides No Hine & Thorne (2000)
Pteria penguin No Hine & Thorne (2000)
Saccostrea cucullata No Hine & Thorne (2000)
Saccostrea glomerata No Hine & Thorne (2000)
Saccostrea echinata No Hine & Thorne (2000)
Stavilia horrida No Hine & Thorne (2000)
Tetrabothriate cestodes Pteria penguin No Hine & Thorne (2000)
Nematoda
Nematode larvae Saccostrea cucullata No Hine & Thorne (2000)
Sulcascaris sulcata Amusium balloti No Lester et al. (1980)
Echinocephalus sp. Amusium balloti No Lester et al. (1980)
Gregarines
Nematopsis sp. Saccostrea cucullata No Hine & Thorne (2000)
Unidentified gregarine Pinctada maxima No Humphrey et al. (1998)
Copepoda
Anthessius pinctadae Pinctada maxima No Humphrey et al. (1998)

tory response. With a prevalence of less than 0.005% it is not
feasible to attempt follow-up sampling, but it does indicate that
parasitic organisms can be present at extremely low prevalence in
a population.
Another proctistan was detected at a low prevalence in P.
maxima from the Exmouth Gulf during investigations into the
ciliate parasite. The parasite was elongated (30 ␮m × 20 ␮m) and
intimately associated with the digestive gland epithelial cells to
which it appeared to have a sessile attachment (Fig. 3). The tubule
epithelial cells beneath the site of attachment were multinucleated
and ultrastructural examination indicated the multiple nuclei were
of molluscan origin, suggesting the proctistan had induced this
change within the host.
A proctistan tentatively attributed to the Thraustochytridea was
identified in moribund, gaping P. maxima from a farm that had
experienced losses after a cyanobacterial (Trichodesmium sp.)
bloom. The affected oysters showed extensive necrosis of external
epithelial surfaces of the palps and mantle with invasion of the
underlying leydig tissues by brown pigmented and eosinophilic,
segmented unicellular organisms 10–15 ␮m in diameter, and
smaller dense basophilic 5-␮m diameter cells that appeared to be
embedded in a mucinous matrix (Fig. 4).
Rock Oysters
One of 411 Saccostrea glomerata (Gould) collected between
Carnarvon to the Dampier Archipelago during 1995 was infected

Figure 2. Unidentified proctistan parasite (arrow) in pearl oysters

Figure 1. Ciliate in digestive tubules of Pinctada maxima spat from Pinctada maxima from Exmouth Gulf. The parasite fills the digestive
Exmouth Gulf (arrow). H&E, scale bar = 10 µm. gland. H&E, scale bar = 30 µm.
236 JONES AND
Figure 3. A large proctistan intimately attached to digestive tubule
epithelial cells by apparent cytoplasmic attachments with hyperplasia
of tubule nuclei, in Pinctada maxima. H&E, scale bar = 10 µm.
with a parasite histologically identical to Marteilia sydneyi Perkins
and Wolf, the cause of QX disease in Saccastera commercialis
(Iredale and Roughley) in Queensland and New South Wales. The
same pathogen was also found in S. glomerata by Hine and Thorne
(2000). On the eastern seaboard where QX occurs, it has caused
losses of over 90% (Witney et al. 1988) and has resulted in a
decline in New South Wales rock oyster production since the
1970s.
S. cucullata (Born) from Airlie Island, Rosily Island, Varanus
Island, Hermite Island, King Bay and East Lewis Island in the
Figure 4. Large 15 µm dia. segmented refractile eosinophilic cells and
smaller 5-µm basophilic cells within the leydig tissues of Pinctada
maxima, tentatively attributed to a thraustochytrid. H&E, scale bar =
20 µm.
CREEPER
north of Western Australia are infected by a second species of
pathogenic haplosporidan protozoan, Haplosporidium sp. The
parasite was identified as the cause of a significant mortality (up to
80%) of rock oysters at Airlie Island (Hine & Thorne 2000). The
presence of this haplosporidan will threaten the viability of any
rock oyster farm sited north of Exmouth.
Mussels
The ova of mussels in Cockburn Sound, Western Australia are
infected with the microsporidan Steinhausia mytilovum but the
parasite has not been found in mussels from other growing areas.
Heavily infected mussels are readily identified by the gross ap-
pearance of the mantle of cooked females. Infected mantle tissue
has an uneven surface with depressed cream white patches and
swollen tubercles forming spots against the orange-pink back-
ground color of healthy female gonad. Mussel samples in August
and October 1995 from sites around the Sound showed a preva-
lence of Steinhausia sp. ranging from 22% to 57%, or 44.4%
overall. Histology revealed that the patches and tubercles were
associated with a marked infiltration of circulating granulocytes
and large basophilic hemocytes into affected follicles with resorp-
tion of the germinal epithelium and ova (Fig. 5). Small basophilic
hemocytes were not prominent in the response. Also present in 2%
to 4% of the ova were parasitophorous vacuoles closely associated
with the nucleus. These vacuoles measured 13.27 ± 2.37 ␮m (n ⳱
20) and contained over 30 small spores 2.5-␮m diameter. There
was usually only one vacuole in an ovum, but occasionally up to
three parasitophorous vacuoles could be seen. In histological sec-
tions the presence of resorbing follicles and focal accumulation of
granulocytes among the developed ova was evidence of infection,
but infected oocytes were also observed in apparently healthy fol-
licles. Ova (normal, infected and degenerating), granulocytes,
large basophilic granular hemocytes and cell debris occurred in the
ciliated gonad ducts. Parasitophorous cysts containing spores also
occurred free in the lumen of the ducts. The spore walls and
contents stained negative to Feulgen, Ziehl-Neelsen and Grocotts,
were Gram positive and Periodic-Acid-Schiffs negative.
Figure 5. Steinhausia mytilovum in Mytilus galloprovincialis from
Cockburn Sound, Western Australia (arrows), main picture and in-
sert. Note inflammatory response in main picture. H&E, scale bar =
10 µm.
 MOLLUSC DISEASES
The relationship between this Steinhausia sp. and the one
previously described by Anderson et al. (1995) from cysts with-
in ova of the rock oyster S. commercialis in Queensland, Australia
is unknown. The biology of Steinhausia sp. is not well under-
stood. Field (1923) reported that infected eggs were shed along
with normal eggs. Sparks (1985) suggested that infected eggs
do not seem to become necrotic or degenerate though Rybakov
and Kholodkovskaya (1987) noted that Steinhausia sp. clearly dis-
torts the nucleus of the ovum and can also cause the destruction
of the egg, as has been seen in Western Australian mussels. It is
likely that the loose spores are released along with intact
eggs or through phagocytosis and subsequent diapediasis. Figueras
(1991) reported that the presence of the parasite is always ac-
companied by a strong hemocyte response and the impact on
the host has been described as severe (Rybakov & Kholod-
kovskaya 1987) to negligible (Maurand & Loubès 1979). In the
case of the S. mytilovum infection seen here, there is an absence
of the typical bivalve inflammatory response, which involves
invasion of the site of trauma primarily by small agranular hya-
linocytes (90%), granular basophils (8%) and granular acido-
phils (2%) (Bayne et al. 1979, Brereton & Alderman 1979).
Instead the major components are the phagocytic hemocyte and
the granular acidophil and the process closely resembles gonad
resorption. Prevalence of Steinhausia mytilovum does not increase
with the size of the mussel, suggesting that infection is annual
(Table 2).
The proctistan observed in the ova of Western Australian blue
mussel has the same measurements and appearance under the light
microscope as S. mytilovum from both European and American M.
galloprovincialis. The taxonomy of the blue mussel in Western
Australia is disputed. Still referred to as M. edulis planulatus (L),
electrophoretic studies have shown that the species is M. gallo-
provincialis and that M. galloprovincialis from Australasia, east-
ern Asia, Western Europe, the Mediterranean and California, and
M. edulis from eastern North America and Western Europe are
electrophoretically distinct species with an overlapping distribu-
tion (Koehn 1991, McDonald et al. 1991, Geller et al. 1993). There
is a fossil record of mussels in Australian waters and Koehn (1991)
hypothesized that M. galloprovincialis may have been an early
introduction into the Northern Hemisphere as a hull-fouling or-
ganism. Distribution of parasites often reflects the distribution of
their primary hosts so S. mytilovum may also be an introduction
from the Southern Hemisphere. Steinhausia mytilovum is reported
to infect the ova of M. edulis along the Atlantic coast of the USA
(Field 1923, Figueras et al. 1991a). Sparks (1985) reported that,
TABLE 2.
Prevalence of Steinhausia sp. in female Mytilus galloprovincialis
from Cockburn Sound, Western Australia.
Number Total number
Size of host (mm) infected examined % infected
<50 16 34
50–59 33 65
60–69 23 80
70–79 93 189
80–89 51 114
>90 7 10
Overall 223 492
There is no increase in prevalence of infection with mussel size.
IN WESTERN AUSTRALIA 237
based on examination of thousands of mussels, the parasite was
absent from California, Oregon and Washington and it was unre-
ported from Europe. It now occurs in M. galloprovincialis from
Spain (Figueras et al. 1991b), Italy (De Vincentiis & Renzoni
1963), Greece (Rayyan et al. 2004), the Black Sea (Rybakov &
Kholodkovskaya 1987, Gayevskaya & Machkevskiy 1991), north-
ern France (Comtet et al. 2004) and the west coast of the USA
(Hillman 1990, 1991). Hillman (1990) noted that M. galloprovin-
cialis had been accidentally introduced into southern California
and suggested that S. mytilovum had been introduced with the
mussels.
Scallops
There are two nematode larvae found in scallops (Amusium
balloti) in the Shark Bay area of Western Australia. The “com-
mon” nematode in scallops is Sulcascaris sulcata. This was re-
ported by Lester et al. (1980) to infect up to 64% of the landed
catch in Shark Bay and occurs in a brownish capsule 3–7 mm dia.
The adult nematodes of S. sulcata live in the loggerhead turtle
Caretta caretta (L) and have a wide geographic distribution and
range of molluscan hosts. The second species, Echinocephalus sp.,
forms small yellow-brown cysts 2–3 mm dia. Lester et al. (1980)
reported that only 2 of 10 scallops he examined were infected, but
in recent years the nematode has been much more common and in
2001 was the dominant nematode in A. balloti. The genus Echi-
nocephalus occurs widely in molluscs in warm waters, and prob-
ably matures in marine skates or rays.
Abalone
The Western Australian component of a national health survey
of abalone has recently been completed. In this survey, up to 25%
of wild-caught green-lipped abalone (Haliotis laevigata) were in-
fected with trematode metacercariae. Low prevalences of proc-
tistans in the lumen of the stomach and digestive gland, apicom-
plexans and putative viral inclusions in the intestinal tract were
observed. Abalone in Western Australia are free of the disease
perkinsosis, found in South Australia and New South Wales, but
the organism does exist on the south coast of Western Australia.
Perkinsus is a primitive fungus-like organism of uncertain taxo-
nomic status, probably in the phylum Labrinthulomycota. A
worldwide effort to understand the taxonomy of this organism (or
group of organisms) is underway. In South Australia and New
South Wales Perkinsus olseni/atlanticus affects abalone with yel-
low-green pus filled blisters (0.5–8 mm dia.) containing a creamy
brown deposit. Once processed the lesions appear as pale brown
circles. Perkinsosis occurs in a variety of shellfish in the north of
the State (Hine & Thorne 2000), however, a survey of 300 abalone
from six sites along the south and west coasts of Western Australia
in 1995 were negative for Perkinsus sp. by the thioglycolate media
method. Subsequently, in 2003 Perkinsus sp. was cultured from
the gill tissue of one clinically normal abalone (H. laevigata) from
the south coast.
47.0
50.7 FUTURE TRENDS
28.7
49.2 Whereas, overall, disease has not been a problem for the mol-
44.7 lusc industry in Western Australia, it is certain that many more
70.0
pathogenic organisms remain to be discovered, particularly as mol-
45.3
luscs become subject to aquaculture or are subject to environmen-
tal stresses associated with economic activity. Because of the age
238 JONES AND
of the Australian continent and the relative isolation of the coastal
fauna it is likely that many of these will prove to be unique to
Western Australia. Strict controls are therefore imposed to limit
translocation of parasites by aquaculture. Whereas these pathogens
may represent an economic threat, it is probable that they will also
provide new insights on the zoogeography and derivation of the
Western Australian mollusc fauna.
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CREEPER
ACKNOWLEDGMENTS
The authors thank the shellfish industries in Western Australia
for their assistance in providing specimens and Paul Hillier
and Tina Thorne for help in collecting mussels. Melanie Crock-
ford, Greg Maguire and Fran Stephens provided editorial com-
ment.
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C. C. Shelley & J. H. Creeper. 1998. Pearl oyster (Pinctada maxima)
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