Pinctada imbricata radiata (rayed pearl oyster)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Water Tolerances
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Pinctada imbricata radiata (Leach, 1814)
Preferred Common Name
- rayed pearl oyster
Other Scientific Names
- Avicula albina var. vaillanti Vassel, 1897
- Avicula chemnitzii Philippi, 1849
- Avicula occa Reeve, 1857
- Avicula radiata Leach, 1814
- Avicula radiata var. canarina Philippi, 1849
- Margaritifera vulgaris Schumacher
- Meleagrina conemenosi Monterosato, 1884
- Meleagrina occa Reeve, 1857 [Pallary, 1912; Issel, 1869]
- Meleagrina radiata (Deshayes) [Tiller and Bavay, 1905]
- Meleagrina savignyi Monterosato, 1884
- Meleagrina vulgaris (Schumacher, 1817)
- Pinctada aerata (Reeve, 1857)
- Pinctada badia (Dunker, 1852)
- Pinctada fimbriata (Dunker, 1872)
- Pinctada fuctata (Gould, 1850)
- Pinctada imbricata (Röding, 1798; Reeve, 1857; Jameson, 1901)
- Pinctada lacunata (Reeve, 1857)
- Pinctada longisquamosa (Dunker, 1872)
- Pinctada martensii (Dunker, 1872)
- Pinctada nebulosa (Conrad, 1837)
- Pinctada pernoides (Reeve, 1857)
- Pinctada perviridis (Reeve, 1857)
- Pinctada radiata (Leach, 1814)
- Pinctada squamulosa (Lamarck, 1819)
- Pinctada varia (Dunker, 1872)
- Pinctada vulgaris (Schumacher) [Tomlin, 1927]
International Common Names
- English: Ceylon pearl oyster; Persian Gulf pearl oyster
- Spanish: pintadina radiada
- French: pintadine radiée
- Arabic: balbal; mohar
Local Common Names
- Central America: Atlantic pearl oyster; Venezuela lingah
- Australia: Australian lingah; bastard shell
- Indian Ocean, Western: bil-bil
- Japan: Akoya pearl oyster
- Myanmar: pate goung
- USA/Hawaii: pipi; unahi pipi
Summary of InvasivenessTop of page
According to Streftaris and Zenetos (2006), P. imbricata radiata is considered to be one of the worst invasive species in the Mediterranean Sea, in terms of spread and impact. It is a relatively hardy species, tolerant to emersion (O’Connor et al., 2003) and to a wide temperature range (13-30°C) (DAISIE, 2009). It also has the ability to adapt to a changed environment (Mohamed et al., 2006) and its tolerance to chemical contamination has enhanced its expansion in enclosed polluted ecosystems (Katsanevakis et al., 2008). It is considered to be a habitat-modifying and gregarious bivalve, capable of impacting native fauna by forming oyster banks (DAISIE, 2009).
It was first recorded as invasive in Tunisia (Vassel, 1896). It is now established in all places where it has been introduced and it is steadily expanding its range (DAISIE, 2009). P. imbricata radiata is currently not on any alert list, but it is listed among the worst alien species in the Streamlining European 2010 Biodiversity Indicators on invasive alien species (European Environment Agency, 2007).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Mollusca
- Class: Bivalvia
- Subclass: Pteriomorphia
- Order: Pterioida
- Unknown: Pterioidea
- Family: Pteriidae
- Genus: Pinctada
- Species: Pinctada imbricata radiata
Notes on Taxonomy and NomenclatureTop of page
There is much confusion about the taxonomy and little agreement among authors about the correct scientific name of this important economic species. Over the years, this species has been described with over 20 names by different authors (i.e. Roding, 1748; Schumacher, 1817; Lamarck, 1819; Conrad, 1837; Phillippi, 1849, Gould, 1850; Reeve, 1857; Dunker, 1872; Monterosanto, 1884; Vassel, 1897). This large number of synonyms is due to the wide variety in shape, strength of sculpture, and colour of P. imbricata radiata shells (Zenetos et al., 2004a).
DescriptionTop of page
Adult P. imbricata radiata have a fragile, rather thin and compressed, small to medium size shell. The shell is inequivalve with the left valve more inflated, and has an almost quadrate outline. The dorsal margin is relatively long, and definitely longer than the body of the shell. The posterior margin is slightly concave and protrudes only slightly (or not at all) beyond the tip of the anterior ear. The beaks point anteriorly, and the hinge line is straight with no teeth present. The ligament is set in a single triangular depression. The common size of this oyster is usually 50-65 mm.
The external coloration of the shell is variable. It can be uniform or with darker markings on radial rays, and is generally brownish or reddish. Sometimes, green and bronze coloration has been observed. The outer surface has densely set, appressed and flattened imbricating concentric lamellae, and often moderately small radially projecting spines, mostly preserved towards the margins. The internal side has a highly iridescent nacreous area, whereas the non-nacreous margin is glossy and light brown in colour, usually with dark brown or reddish blotches corresponding to the main external rays (Zenetos et al., 2004a).
DistributionTop of page
P. imbricata radiata is a bivalve with a very wide distribution. It is found in both hemispheres and in most oceans and seas around the world. It is present in the Atlantic, Pacific and Indian oceans, the Persian Gulf, the Red Sea, and more recently in the Mediterranean Sea.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Egypt||Present||Introduced||1874||Invasive||Native on Red Sea coast, Introduction in Mediterranean coast|
|United Arab Emirates||Present||Native|
|Ireland||Present||Introduced||on macro-litter; one shell; First reported: 2013-2014|
|Portugal||Present||Present based on regional distribution.|
|United Kingdom||Present||Introduced||1986||shells only/on macro-litter|
|Antigua and Barbuda||Present||Native|
|Saint Kitts and Nevis||Present||Native|
|Saint Vincent and the Grenadines||Present||Native|
|Trinidad and Tobago||Present||Native|
|U.S. Virgin Islands||Present||Native|
|Papua New Guinea||Present||Native|
|Atlantic - Northeast||Present, Localized||Introduced||1998||Azores Islands; Original citation: Àvila et al. (1998)|
|Atlantic - Southwest||Present||Native|
|Atlantic - Western Central||Present||Native|
|Indian Ocean - Eastern||Present, Widespread||Native|
|Indian Ocean - Western||Present, Widespread||Native|
|Mediterranean and Black Sea||Present, Widespread||Introduced||1874||Invasive|
|Pacific - Eastern Central||Present||Native|
|Pacific - Northwest||Present, Widespread||Native|
|Pacific - Southwest||Present||Native|
|Pacific - Western Central||Present||Native|
History of Introduction and SpreadTop of page
P. imbricata radiata was first reported outside its natural biogeographical distribution in 1874 in the Mediterranean Sea. It was collected on the shores of Alexandria, Egypt and was one of the very first marine species of Indo-Pacific origin to cross the Suez Canal into the Mediterranean. It was subsequently reported from Tunisia (1890), Israel (1899), Cyprus (1899), and Malta (1912).
In 1963 it was imported for aquaculture purposes to Greece, but its cultivation did not prove prosperous and was thus abandoned. However, P. imbricata radiata was naturalised and expanded in the Hellenic Seas (Pancucci-Papadopulou et al., 2005). Subsequent findings of this species in areas where aquaculture activities (historical or recent) are absent support a Lessepsian mode of introduction (Zenetos et al., 2005). Furthermore, molecular studies of P. imbricata radiata populations in the Saronikos area (Greece) have rather excluded shipping as the mode of transportation (Zenetos et al., 2004b).
In 1965 it was reported in Lebanon and in the early 1970s it was also reported in Libya (1973) and Syria (1975). In 1979, individuals of this species were found attached on the hull of a naval vessel in the port of Toulon, France (Zibrowius, 1979). It was reported in 1982 from Italy (Di Natale, 1982) and Turkey. More recently it was reported in the northern Adriatic Sea, in Croatia (2006). It is suspected that the populations in France and the northern Adriatic Sea are due to shipping transport (DAISIE, 2009). It is now well established in the eastern Mediterranean Sea and seems to be expanding (DAISIE, 2009). It is also reported to be established and spreading in the Central Mediterranean (Italy: Lodola et al., 2013; Stasolla et al., 2014. Malta: Evans et al., 2015).
In the Azores, (Northwestern Atlantic Ocean) P. imbricata radiata individuals were first found in 1998 attached to a ball-float near Faial. Later, there was a second report of occurrence in Vila Franca do Campo (São Miguel) (Àvila et al., 2000). The vector is still unknown but it is presumed that the introduction was not deliberate (Cardigos et al., 2006).
Winston et al. (1997) recorded Pinctada attached to marine litter off the coast of Florida. The species was also included in trans-Atlantic rafting mollusca on macro-litter: American molluscs on British and Irish shores (Holmes, 2015).
IntroductionsTop of page
Risk of IntroductionTop of page
P. imbricata radiata is currently spreading towards the north and western Mediterranean Sea (DAISIE, 2009) and it is suspected that it will slowly expand throughout the Mediterranean coastline. The relatively recent record of its finding in the Azores (northeastern Atlantic Ocean) (Àvila et al., 1998), indicates that there is a strong possibility of this species spreading towards the European Atlantic coast as well.
HabitatTop of page
P. imbricata radiata individuals are byssally attached to rocks, dead corals and various submerged objects, often forming large natural banks. They are most common on sub-littoral bottoms from depths of 5 to 25 m, but they can also be found on the littoral and shelf zones from low tide levels to a depth of about 150 m. On soft bottoms, they aggregate to one another (Carpenter and Niem, 1998).
Habitat ListTop of page
|Brackish||Inland saline areas||Principal habitat||Natural|
|Marine||Inshore marine||Principal habitat||Natural|
|Marine||Benthic zone||Principal habitat||Natural|
Biology and EcologyTop of page
The biology of pearl oysters is poorly understood, considering the importance of both natural and cultured pearl, and shell fisheries (Gervis and Sims, 1992). P. imbricata radiata is an epifaunal suspension feeder of the subtidal zone and a fouling species, living attached by its byssus to hard substrata. Its maximum lifespan is 8 years, although older specimens have been found. Growth studies have been conducted in populations of this species in the Red Sea (Yassien, 1998), Qatar (Mohammed and Yassien, 2003), India (Narayan and Michael, 1968; Jeyabaskaran et al., 1983), Japan (Wada, 1991), Taiwan (Hwang et al., 2007) and in the Mediterranean Sea (Yassien et al., 2000). Usually it attains a length of 50-65 mm. Maximum observed shell length was 93.2 mm in the Red Sea (Yassien, 1998), 64.0 mm in the Mediterranean Sea (Yassien et al., 2000), and 100.0 mm in Japan, on an individual with a ten year lifespan (Wada, 1990). Mohamed et al. (2006) concludes that the biotope and P. imbricata radiata’s interaction with the environment are important determinants of growth and shell dimensions.
Temkin (2010) found that there were low levels of molecular divergence between these populations and chose to regard this group as a single species with three subspecies; P. imbricata imbricata, P. imbricata fucata and P. imbricata radiata. The shells of this Pinctada complex are variable; differentiation of the subspecies can be difficult and are recognised by their location rather than morphology.
In contrast, according to Cunha et al. (2011), molecular analyses question the taxonomic validity of the morphological characters used to discriminate P. fucata and P. martensii that exhibited the lowest genetic divergence and are most likely conspecific as they clustered together. P. radiata and P. imbricata were recovered as monophyletic.
P. imbricata radiata is a protandric hermaphrodite species with sex inversion occurring in shells of 32-57 mm (Zenetos et al., 2004a). Sexual dimorphism is absent, and in contrast to other species of the same genus, the colour of the gonad is an unreliable aid in sex determination (Khamdan, 1998). The morphology of the gonad, which is not a discrete organ, is similar to other species of the genus Pinctada (Khamdan, 1998). Gonad maturity is controlled by temperature (Zouari and Zaouali, 1994).
Both oogenesis and spermatogenesis occur in similar timing, i.e. commence in winter and continue through spring. Thus, the breeding cycle in this species is seasonal with two peak spawnings in the summer and autumn (Khamdan, 1998; O’Connor et al., 2003). In the Persian Gulf and the Red Sea, the spawning of adult P. imbricata radiata is reported to be essentially continuous since there are always at least a few spawning individuals present throughout the year (Khamdan, 1998; Yassien, 1998). In other areas however, this is not the case. Microscopic examination of the gonad in P. imbricata radiata individuals from southeast Australia (O’Connor et al., 2003) indicated differences between the two peaks: samples collected following the summer peak showed a high proportion of empty gonads, consistent with spawning, while those taken in autumn suggested that the oysters were resorbing the gonad rather than spawning. The same study (O’Connor et al., 2003) showed a significant variation in the numbers of spat settling, and that settlement was restricted to the summer months. This is consistent with summer spawning and further suggests that the second, autumnal peak in reproductive activity does not contribute to oyster settlement (O’Connor et al., 2003).
Although salinity levels have been suggested for controlling spawning (Malpas, 1933), it seems that temperature is more likely to be the controlling factor (Khamdan, 1998).
As with temperature, salinity tolerances reported for P. imbricata radiata appear to be a function of a number of factors including geographic location and ontogeny(O’Connor et al., 2003). In Japan, long-term studies reported the minimum salinity optimum for spat to be 22.7 g kg-1 (Numaguchi and Tanaka, 1986b), while in India adult P. imbricata radiata have been found to be tolerant of salinities within the range 24-50 g kg-1 for 2-3 days (Alagarswami and Victor, 1976; Dharmaraj et al., 1987a). In Australia, P. imbricata radiata embryos failed to develop to D-veliger stage at salinities of 26 g kg-1 or less, byssal attachment by juveniles did not occur at salinities of 17 ppt or less, and high mortality occurred at salinities of 23 ppt or less within 7 days (O’Connor et al., 2003).
ClimateTop of page
|Am - Tropical monsoon climate||Preferred||Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Depth (m b.s.l.)||Optimum||0–150 tolerated|
|Water temperature (ºC temperature)||13||25||Optimum||7–29 (Wada, 1991)|
Notes on Natural EnemiesTop of page
Natural enemies of P. imbricata radiata include fish, predatory flatworms, sponges (Cliona spp.), and shell boring spionid polychaetes (Polydora and Boccardia spp.) (O’Connor et al., 2003). Reports attributing damage and mortality in P. imbricata radiata to spionid polychaetes are common and have arisen in areas such as Sri Lanka (Herdman, 1903), Japan (Mizumoto, 1964), India (Dharmaraj et al., 1987b), the Persian Gulf (Doroudi, 1996) and China. Spionids are thought to “fatigue” the host pearl oyster (Wada, 1991) and weaken their shells, increasing their susceptibility to predators.
Means of Movement and DispersalTop of page
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
P. imbricata radiata has long been in use for the production of pearls and is also an edible mollusc species. There are generally no negative economic impacts caused by P. imbricata radiata. The only relevant report is that it fouls mussel lines and commercial shellfish collectors (DAISIE, 2009).
Environmental ImpactTop of page
No specific impact of P. imbricata radiata on either habitats or biodiversity has been cited in the literature (Streftaris and Zenetos, 2006). However, there is insufficient information concerning its role in affected ecosystems. This bivalve is considered to be a habitat-modifying, gregarious species capable of impacting native fauna by forming oyster banks (DAISIE, 2009).
Studies conducted in Tunisia to test for a possible community shift from a substrate without Pinctata and a substrate with initial low density Pinctada settlement, were not conclusive. Results may not confirm that the community structure variability is due to the impact of Pincata invasion because the potential and subtle community shift may be masked by the overwhelming influence of just the local environmental gradients. In spite of this, the introduced oyster may play the role of an engineer species at high densities, contributing to the complexity of the benthic habitat and influencing the trophic pattern of its fauna (Tlig-Zouari et al. 2011).
Social ImpactTop of page
No impact of P. imbricata radiata on human activities has been documented.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Pioneering in disturbed areas
- Long lived
- Fast growing
- Has high reproductive potential
- Modification of natural benthic communities
- Reduced native biodiversity
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
UsesTop of page
Uses ListTop of page
Human food and beverage
- Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
Similarities to Other Species/ConditionsTop of page
This species is hard to distinguish from shell appearance only. It looks very similar to Pinctada margaritifera, from which it differs in size, colour and shape of the adductor muscle scar (Zenetos et al., 2004a). The shell of P. imbricata radiata is smaller and thinner, and its outer shell coloration differs from P. margaritifera in that it is tan-coloured (in contrast to greyish green) and its markings are reddish to black (in contrast to white or yellowish) (Oliver, 1992; Zenetos et al., 2004a).
Gaps in Knowledge/Research NeedsTop of page
Being an important economic species, the biology and ecology of P. imbricata radiata have been extensively studied. Also, because of its long history as an intensively cultured marine species, most aspects of its reproduction and growth are well documented. However, there is little information in the literature concerning its status as an invasive species. Its effect on habitats and biodiversity is poorly studied, and not yet fully understood. More research on its rate of invasion, especially in the Mediterranean Sea is needed. Furthermore, information concerning its interaction with native species of invaded areas, and its role in trophic relations is definitely required.
ReferencesTop of page
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Poutiers J M, 1998. Bivalves, Acephala, Lamellibranchia, Pelecypoda. In: The living marine resources of the Western Central Pacific. Volume 1: Seaweeds, corals, bivalves and gastropods. [ed. by Carpenter K E, Niem V H]. Rome, Italy: FAO. 123-362.
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Turgeon DD, Quinn JF, Bogan AE, Coan EV, Hochberg FG, Lyons WG, Mikkelsen PM, Neves RJ, Roper CFE, Rosenberg G, Roth B, Scheltema A, Thompson FG, Vecchione M, Willams JD, 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. In: American Fisheries Society Special Publication, 26 526 pp.
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13/02/16 Updated by:
Argyro Zenetos, Institute Marine Biological Resources and Inland Waters, Hellenic Centre for Marine Research, P.O. BOX 712, Anavissos 19013, Greece
08/07/09 Original text by:
Argyro Zenetos, Institute of Oceanography, Hellenic Centre for Marine Research, P.O. BOX 712, Anavissos 19013, Greece
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