Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Brachidontes pharaonis



Brachidontes pharaonis


  • Last modified
  • 14 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Brachidontes pharaonis
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Mollusca
  •       Class: Bivalvia
  •         Subclass: Pteriomorphia
  • Summary of Invasiveness
  • Well documented in the Mediterranean Sea, B. pharaonis entered via the Suez Canal, starting colonization in the eastern part and becoming abundant on the Israeli coast, and in Egypt, Syria, southern Turkey (nor...

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Brachidontes pharaonis;	voucher specimens. Scale: the width of the text on the species/QR-code label is 38 mm. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
TitleVoucher specimens
CaptionBrachidontes pharaonis; voucher specimens. Scale: the width of the text on the species/QR-code label is 38 mm. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
CopyrightPublic Domain - Naturalis Biodiversity Center/Wikimedia Commons - CC0 1.0 Universal (CC0 1.0)
Brachidontes pharaonis;	voucher specimens. Scale: the width of the text on the species/QR-code label is 38 mm. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
Voucher specimensBrachidontes pharaonis; voucher specimens. Scale: the width of the text on the species/QR-code label is 38 mm. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.Public Domain - Naturalis Biodiversity Center/Wikimedia Commons - CC0 1.0 Universal (CC0 1.0)
Brachidontes pharaonis;	closer view of voucher specimens. Note scale. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
TitleVoucher specimens
CaptionBrachidontes pharaonis; closer view of voucher specimens. Note scale. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
CopyrightPublic Domain - Naturalis Biodiversity Center/Wikimedia Commons - CC0 1.0 Universal (CC0 1.0)
Brachidontes pharaonis;	closer view of voucher specimens. Note scale. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.
Voucher specimensBrachidontes pharaonis; closer view of voucher specimens. Note scale. Source institute: Zoölogisch Museum Amsterdam, The Netherlands.Public Domain - Naturalis Biodiversity Center/Wikimedia Commons - CC0 1.0 Universal (CC0 1.0)


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Preferred Scientific Name

  • Brachidontes pharaonis (Fischer P., 1870)

Other Scientific Names

  • Brachidontes semistriatus Krauss, 1848
  • Brachidontes variabilis Krauss, 1848
  • Crenella ehrenbergi
  • Mytilus arabicus Jousseaume ms. In Lamy, 1919
  • Mytilus exustus Linnaeus, 1827
  • Mytilus pharaonis Fischer P., 1870
  • Mytilus senegalensis Lamarck, 1889
  • Mytilus variabilis Krauss, 1848

International Common Names

  • English: variable mussel

Summary of Invasiveness

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Well documented in the Mediterranean Sea, B. pharaonis entered via the Suez Canal, starting colonization in the eastern part and becoming abundant on the Israeli coast, and in Egypt, Syria, southern Turkey (northern Cyprus), Greece (Egeu Sea), Croatia (northern Adriatic) and the Sicily coast. The first record was in Port Said, Egypt in 1876 by Fuchs (1878). It is considered invasive because of its capacity of colonization, forming extensive mats in midlittoral sites, it also appears in patches in the infralittoral (subtidal), usually on vertical surfaces among barnacles, displacing native species. This mussel represents a potential resource and space competitor of its ecological equivalent, such as the Mediterranean species Mytilaster minimus. The mats can be seen occupying areas where previously there were benthic communities. Coral reefs near Suez were replaced by mats of B. pharaonis interspersed with coralline red algae and Enteromorpha sp. (Moshira, 2008).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Mollusca
  •             Class: Bivalvia
  •                 Subclass: Pteriomorphia
  •                     Order: Mytiloida
  •                         Unknown: Mytiloidea
  •                             Family: Mytilidae
  •                                 Genus: Brachidontes
  •                                     Species: Brachidontes pharaonis

Notes on Taxonomy and Nomenclature

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The systematic classification and phylogeny of the whole family, Mytilidae, are problematic and are not completely developed due to several reasons: many Mytilidae species and even genera inhabit deep-water habitats which makes it very difficult to collect them; there are many endemic species; shell shape may be extremely dependent on environmental conditions and on the age of the animal (Seed, 1968), it is not always evident that visible characters provide diagnostic for differentiation; in the Bivalvia, the homology of many morphological characters is difficult to determine due to convergent and/or parallel evolution (Steiner and Hammer, 2000). Previously, subfamilies and genera of Mytilidae have been assigned largely according to the shape and characteristics of the valves, sculpture, hinges and dentition; these assignments were rarely, ever guided by genetic data. Nowadays, genetic studies are identifying differences in populations previously classified as a single species. Further morphological studies should be initiated to identify diagnostic characters for cryptic species, DNA barcoding is now being used with traditional methods for identification of animal taxa (Hebert et al., 2003).

B. pharaonis, classified by Fischer P. (1870), was previously known as Brachidontes semistriatus (Krauss, 1848) and is known by many authors as Brachidontes variabilis. Different interpretations of the high variability of the shell characters led systematists to use different names, all of them considered as synonyms with B. variabilis (Krauss, 1848), adding to the confusion; Mytilus exustus Linnaeus, 1827, Mytilus (or Hormomya) variabilis Krauss, 1848, Brachidontes semistriatus Krauss, 1848, Mytilus (or B.pharaonis) (Fischer, 1870), Mytilus senegalensis Lamarck, 1889, Mytilus arabicus Jousseaume ms. in Lamy (1919) are the ones found in literature. The ambiguity is the greatest for the Mediterranean Sea and the Red Sea regions, where many authors (Arcidiacono and Di Geronimo, 1976; Chemello and Oliverio, 1995; Gianguzza et al., 1997; Rilov et al., 2002) have used B. pharaonis (Fischer, 1870) as a synonym of B. variabilis (Krauss, 1848), but never as a different species. All this synonymy for B. pharaonis, by different authors, can transform this species in an organism dispersed throughout the world, but new classification methods can differentiate them in different endemic or cryptic species with similar phenotypes.

Recently, genetic analyses of nuclear and mitochondrial DNA have identified many species, previously considered as synonymies, as separated species. Lee and Foighil (2004) revealed that B. exustus has species differentiation within populations and thus different from B. variabilis and B. semistriatus. In the Mediterranean Sea, a B. pharaonis population was described using the mtDNA variation to make inferences about its invasion. These studies did not reveal any geographical pattern or differentiation among local populations (Shefer et al., 2004; Terranova et al., 2007). The name B. pharaonis is most appropriate for the species in the Mediterranean Sea and the Red Sea, but it must be determined whether B. variabilis should be applied to the species in the Indian Ocean, or the species in the Pacific Ocean. The nucleotide distances and the phylogenetic analysis suggest that the most ancient divergence was between the clades currently in the Pacific and Indian Oceans. Subsequently, the clade currently in the Mediterranean Sea and the Red Sea diverged from the clade in the Indian Ocean (Terranova et al., 2007).


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B. pharaonis is a small bivalve with a 40 mm shell, externally dark brown-black and internally tinged violet-black. Shell is equivalve, inequilateral, attached to substrate by stout byssus. Sculpture of numerous fine radial bifurcating ribs, which become coarser posteriorly and margin crenulate. The hinge has dysodont teeth. Outline mussel-like with terminal umbones but variable in shape and in its height/length ratio; sometimes greatly expanded posteriorly, sometimes arcuate; occasionally subcylindrical with beaks not quite terminal.


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B. pharaonis originates from the Indian Ocean and is widely spread throughout the Red Sea (Oliver, 1992). It was among the first migrants noticed in the eastern Mediterranean.

Its geographical range includes the western Pacific Ocean, the Indian Ocean, the Red Sea and the Mediterranean Sea (Taylor, 1971; Sasekumar, 1974; Barash and Danin, 1986; Morton, 1988), but the occurrence of this species in some areas is controversial. According to Barash and Danin (1986), B. variabilis occurs along eastern African coasts, from the Red Sea to southern Africa, in the Indian Ocean except for the Persian subregion and Malaysia, and in the western Pacific Ocean. In contrast, Sasekumar (1974) reported B. variabilis in Malaysia, while Arcidiacono and Di Geronimo (1976) supported the occurrence of B. variabilis along the western African coasts.

According to recent genetic studies (Terranova et al., 2007), the distribution of B. pharaonis is restricted to the Red Sea and the Mediterranean Sea.

Distribution Table

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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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Mediterranean and Black SeaWidespreadIntroduced1876 Invasive Fuchs Th, 1878First record in Port Said, Egypt (absent in Black Sea)


IsraelWidespreadIntroducedHaas, 1937; Rilov et al., 2002; Rilov et al., 2004; Galil, 2006; Galil, 2007
LebanonWidespreadIntroducedGalil, 2006; Nakhlé et al., 2006
SyriaWidespread2006Introduced Invasive Galil, 2006First collected in the Mediterranean in 1876. Successively it has been found in Syria. Constitutes large, stable populations in the Levant Sea
TurkeyWidespreadIntroducedKinzelbach, 1985; Kinzelbach, 1985; Aartsen and Kinzelbach, 1990; Tringali and Villa, 1990; Niederhofer et al., 1991; Buzzurro and Greppi, 1996; Ergen and Çinar, 1997; Çevik and Sarihan, 2004; Çinar, 2006; Çinar, 2006; Galil, 2006; Dogan et al., 2007


EgyptWidespreadIntroducedFuchs Th, 1878; Pallary, 1912; Oliver, 1992; Galil, 2006
Rodriguez IslandWidespreadOliver et al., 2004Coton Baie, Petite Butte and Ile Gombrani, can be found around all island and lagoon islets. One population with distinct characteristics was encountered only in the harbour at Point Monier
South AfricaWidespreadIntroduced Invasive Arcidiacono and Geronimo, 1976Along the western African coasts


CroatiaWidespread2006Introduced Invasive Galil, 2006Successively it has been found in Croatia. Constitutes large, stable populations in the Levant Sea
CyprusWidespread2006IntroducedGalil, 2006Successively it has been found in Cyrus. Constitutes large, stable populations in the Levant Sea
GreeceWidespreadIntroduced Invasive Pancucci-Papadopoulou et al., 2005; Zenetos et al., 2005; Galil, 2006
ItalyWidespreadIntroducedGalil, 2006; Sarà et al., 2006

History of Introduction and Spread

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B. pharaonis, originally from the Indo-pacific area, has colonized hard substrata and spread throughout the Red Sea (Oliver, 1992). It entered into Mediterranean marine ecosystems from the Red Sea, after the opening of the Suez Canal, in 1869. It was first recorded in Port Said, Egypt, in 1876, as an exotic species. Its introduction and consequent dispersion, by water circulation and human activities, favoured the formation of abundant populations on midlittoral and infralittoral rocky habitats, especially along the rocky shore of the Eastern Mediterranean.

It has spread along the Levant coast from Egypt to Turkey (Pallary, 1912; Kinzelbach, 1985; Zenetos et al., 2003, Rilov et al., 2004; Dogan et al., 2007). It was first recorded in Turkey, as B. semistriatus, in Iskenderun Bay, in 1978. Dense populations were recorded in Turkey by Çinar (2006). It was also recorded in northern Cyprus and Greece (Egeu Sea (Rhodes), Saronikos Gulf, Evoikos Gulf (Zenetos et al., 2005), Croatia (Kocatas and Bilecik, 1992) and the Sicily coast (Sara et al., 2000).

Risk of Introduction

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Initially was assumed that the introduction of B. pharaonis was a Lessepsian migration (natural migration through the Suez Canal) (Streftaris et al., 2005). However, new techniques, molecular studies, may reveal a different mode of introduction than initially assumed suggesting that the mode of transport in Italy was by shipping and that ships transported this species through the Suez Canal originating from distant locations (Galil and Zenetos, 2002). Its introduction and consequent diffusion certainly is related with human economical shipping activity e.g. ballast waters, ship hulls. After its dispersion, normally B. pharaonis starts to generate intensive populations growing in dense clusters on midlittoral and infralittoral rocks, piers and debris (Barash and Danin, 1992), such as what happened in the west of Sicily (Zenetos et al., 2005; Sará et al., 2006; Dogan et al., 2007). It is important to note also that this organism can be introduced by aquaculture, mainly associated with other cultures.

In the western Mediterranean, B. pharaonis is confined to habitats with high temperature and high salinity, where it has established dense beds on hard substrata. It has been predicted that from these area, B. pharaonis will continue its migration towards North Africa and Gibraltar (Sarà et al., 2006). In recent decades the rapid warming of European Mediterranean waters (the mean temperature of the Mediterranean has increased at least 3ºC from 20-21ºC in less than 10 years) is creating favourable conditions for invasion by some exotic species (Root et al., 2005), possibly enhancing the invasive capacity of B. pharaonis, a species well-adapted to tropical temperatures (Sarà et al., 2006). Considering its invasive potential and the recent warming trend of the Mediterranean, in the future B. pharaonis may actively invade more habitats, threatening indigenous bivalve species which may be unable to compete with B. pharaonis in terms of reproduction effort and density. Large extensions of shallow waters that have warm water, like the western Sicily saltpan, may function as a stepping-stone to invasion of the western basin providing an oasis of favourable warm waters.

Habitat List

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Intertidal zone Principal habitat Harmful (pest or invasive)
Benthic zone Principal habitat Harmful (pest or invasive)
Coral reefs Secondary/tolerated habitat Harmful (pest or invasive)
Inshore marine Secondary/tolerated habitat Harmful (pest or invasive)

Biology and Ecology

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Reproductive Biology

B. pharaonis
has separate sexes. The gametes are delivered in sea water where the fertilization occurs. Its planktonic larval period has two phases: trochophore, lasting 24 h, and veliger, lasting some weeks.

Physiology and Phenology
B. pharaonis can colonize many environments quite easily, reaching a similar degree of fitness in all colonized environments like other filter-feeder molluscs that are considered generalists.
It has a gregarious survival strategy that prevents other species carry out settlements. Its presence and absence seem to be correlated with the environmental features of habitats. In the Aegean Sea it does not form dense populations because the waters are less saline and colder (Kocatas and Bilecik, 1992).
B. pharaonis is a suspension feeder. It is able to exploit almost all the predominant sources available in its colonized environment both directly (phytoplankton) and indirectly (via particulate organic matter (detritus)).
Environmental Requirements
In the Mediterranean adult mussels show large temperature tolerances (9-31°C), and occurs at salinities from 35 to 53 PSU. Lower winter temperatures limit their physiological activity (Galil, 2006). In the western Mediterranean, B. pharaonis is confined to high temperature and high salinity habitats, where it has established dense beds on hard substrata. Its abundance seems to be negatively associated with wave exposure. The animals colonize debris, fixing by byssus threads to hard substrates and may reach very high densities and completely cover rock shore to form a “mytilid bed” (Safriel et al., 1980).

B. pharaonis appears to be able to better exploit intertidal sites exposed to air, producing high densities. Sarà et al. (2006) found that when saltpans are occasionally filled with seawater by salt workers, in spring and autumn, the surface of all hard substrata is completely covered. This suggests that even short periods of submersion are sufficient to guarantee recruitment in the surface population. Nevertheless, a slightly different response to air exposure reflected a slight difference in resource (Sarà et al., 2006).


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A - Tropical/Megathermal climate Tolerated Average temp. of coolest month > 18°C, > 1500mm precipitation annually
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) Optimum Shallow water preferred; 0-3 m tolerated
Salinity (part per thousand) Optimum 35-53 tolerated, >45 (Sara et al., 2003)
Water temperature (ºC temperature) Optimum 9-31 tolerated

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Stramonita haemastoma Predator Adult not specific N/A

Notes on Natural Enemies

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Stramonita haemastoma, a whelk from the Muricidae, preferentially preys on B. pharaonis on the Israeli coast (Rilov et al., 2002).

Means of Movement and Dispersal

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B. pharaonis was among the first exotic migrant mollusc species noticed in the eastern Mediterranean Sea and its current distribution and progressive penetration through the Suez Canal was considered obvious and hence its characterization as a Lessepsian migrant was not questioned. It has been predicted that from these areas, B. pharaonis will continue its migration towards North Africa and Gibraltar (Sarà et al., 2000, 2006).

Natural Dispersal (Non-Biotic)

B. pharaonis appears to have all the characteristics needed to reach and colonize many coastal environments. Physiological plasticity permits the species to tolerate highly stressed conditions including resistance to air exposure and variable temperatures and food availability. B. pharaonis appears to increase its density and size moving towards the western basin of the Mediterranean. Thus, its actual populations represent an important larval reservoir for this invasive species in the new areas, because this organism has a planktonic larval stage and their dispersal occurs via local water bodies.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
AquacultureRecorded in the north Aegean Sea, in most cases connected with anthropogenic activites i.e. shipping Yes Yes Zenetos et al., 2005
Interconnected waterwaysOpening of the Suez Canal Yes Yes Sarà et al., 2000; Sarà et al., 2006; Zenetos et al., 2005

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Ship ballast water and sediment Yes Yes Zenetos et al., 2005
Ship hull fouling Yes Yes CIESM, 2009
WaterConstant immersion Yes Yes

Impact Summary

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Biodiversity (generally) Positive and negative
Economic/livelihood Negative
Environment (generally) Negative
Native fauna Negative

Economic Impact

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B. pharaonis is a fouling organism and was observed on hulls of fishing boats in harbours, they have the potential to cause problems such as the fouling of sea-water intake pipes (Çinar, 2006; Streftaris and Zenetos, 2006). Species such as B. pharaonis may slow down boats/ships when they foul the hulls which would most certainly have an effect on the amount of gasoline used (A Gittenberger, GiMaRIS, The Netherlands, personal commuication, 2011).

Environmental Impact

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Impact on Biodiversity
B. pharonis is displacing native species on the Israeli coast (first recorded in 1935) (Haas, 1937) and forms dense populations on rocky substrates. However, sediments trapped among shells and byssuses threads could provide more micro-habitats for many species. B. pharaonis was considered in the early 1970s circa 250 times rarer than the native mytilid Mytilaster minimus that formed dense beds on intertidal rock shores along the Israeli coastline, with up to 26 specimens per cm2 at Palmahim (Safriel et al., 1980; Galil, 2007). By the end of the 1980s, Safriel and Sasson-Frosting (1988) recorded that B. pharaonis interfered with the recruitment of Mytilaster and detrimentally affected its survival and growth. In the late 1990s a survey conducted in some of the same sites showed a rapid shift in dominance, with some dense populations of up to 300 specimens per 100cm2 on rocky shores, where mussel beds were absent in the past (Rilov et al., 2004). More recently, the same locations where completely covered with B. pharaonis, while M. minimus was only rarely encountered (Mienis, 2003; Galil, 2006).

Coral reefs near Suez have been replaced by mats of the B. pharaonis interspersed with coralline red algae and Enteromorpha sp. A layer of barnacles, found under Brachidontes, indicates a succession after the reefs died. Several genera of live coral, which were heavily infested by borers, were present on other patches, together with mats of Caulerpa and Brachidontes, whereas oysters were the dominant bivalves farther south (Moshira, 2008).

The establishment of massive beds of Brachidontes has significant effects on the biota of the intertidal rocky shore. Habitats are clearly impacted when the density of mussels is high, as recorded by Sarà et al. (2006): 375 individuals/400cm2 in saltpans and 10,000 individuals/m2 in western Sicily.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Mytilaster minimusNo details No detailsIsraelCompetition - monopolizing resourcesGalil, 2007

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Capable of securing and ingesting a wide range of food
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Altered trophic level
  • Changed gene pool/ selective loss of genotypes
  • Conflict
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Infrastructure damage
  • Modification of natural benthic communities
  • Modification of successional patterns
  • Negatively impacts aquaculture/fisheries
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Transportation disruption
  • Negatively impacts trade/international relations
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - smothering
  • Filtration
  • Fouling
  • Herbivory/grazing/browsing
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally

Similarities to Other Species/Conditions

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B. pharaonisresembles the Red Sea species Septifer binocularis, from which it is distinguished by the absence of a septum beneath the beaks and by the darker colour (whereas S. binocularis is bright green with reddish spots). Variability in a Brachidontes complex or even a case of mistaken identity is known from the harbour at Pointe Monier on Rodrigues’s Island. One bivalve has shells more inflated with a much stronger median angle. The sculpture is stronger and more granular and the dysodont teeth behind the ligament are very strong. There are multiple teeth below the beak rather than the single tooth seen in typical B. pharaonis. These differences can be regarded as inconclusive when considered within the great range of variation recognized for B. pharaonis. However, the anatomical character of the papillation of the posterior inhalant mantle edge appears diagnostic. In B. pharaonis from Rodrigues the papille are large and terminate in small finger-like processes, whereas in Brachidontes sp. the papillae are simple pimple-like processes. Further research on both morphological and molecular characters is required to resolve the problem (Oliver et al., 2004).

Prevention and Control

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Ballast water and ship hulls are considered the main vectors of introduction and dispersal of B. pharaonis. Ocean ballast water exchange and antifouling paints are possible ways to control the invasion and dispersal of this mytilid. Other possibilities, proposed by Galil (2006) include the erection of a saline barrier for preventing Lessepsian migration through the Suez Canal. This barrier could reduce the number of Red Sea species arriving in the Mediterranean.
Chemical control

All size groups of B. variabilis showed a progressive reduction in physiological activities such as oxygen consumption, filtration rate, foot activity and byssus thread production, when chlorine residuals were increased from 0 to 1mg/l. B. variabilis was able to sense the presence of chlorine at levels as low as 0.25mg/l and responded by reducing its physiological activities by 41–49%. If chlorination is carried out on a continuous basis, mussels do not get an opportunity to compensate the loss incurred due to reduced food intake and oxygen consumption. Under such circumstances, a significant decline of the growth rate could be expected. Continuous dosing at a residual level of at least 1mg/l is necessary to force B. variabilis to close its shells, without allowing a recovery phase. Therefore, such residual levels should be maintained during peak settlement periods of B. variabilis to prevent fresh colonization (Rajagopal et al., 2005).

Gaps in Knowledge/Research Needs

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Further genetic studies are required to provide a better understanding of variation encountered in the B. pharaonis group.

Studies on the control of B. pharaonis using both natural enemies and chemicals also warrant attention.


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Monaco: CIESM - Commission Internationale pour l'Exploration de la Mer Mediterranee, Villa Girasole, 16 bd de Suisse,

USA: OBIS - Ocean Biogeographic Information System, 71 Dudley Road, New Brunswick, NJ 08901,


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14/06/10 Original text by:

Flavio da Costa Fernandes, Instituto de Estudos do Mar, Pesquisador Titular, Chefe da Divisão de Biologia, Almirante Paulo Moreira, Rua Kioto, 253 - Arraial do Cabo - RJ, Brazil

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