- 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
- Water Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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 InvasivenessTop of page
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 TreeTop of page
- 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 NomenclatureTop of page
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).
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).
DescriptionTop of page
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.
DistributionTop of page
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 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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Mediterranean and Black Sea||Widespread||Introduced||1876||Invasive||Fuchs Th, 1878||First record in Port Said, Egypt (absent in Black Sea)|
|Israel||Widespread||Introduced||Haas, 1937; Rilov et al., 2002; Rilov et al., 2004; Galil, 2006; Galil, 2007|
|Lebanon||Widespread||Introduced||Galil, 2006; Nakhlé et al., 2006|
|Syria||Widespread||2006||Introduced||Invasive||Galil, 2006||First collected in the Mediterranean in 1876. Successively it has been found in Syria. Constitutes large, stable populations in the Levant Sea|
|Turkey||Widespread||Introduced||Kinzelbach, 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|
|Egypt||Widespread||Introduced||Fuchs Th, 1878; Pallary, 1912; Oliver, 1992; Galil, 2006|
|Rodriguez Island||Widespread||Oliver et al., 2004||Coton 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 Africa||Widespread||Introduced||Invasive||Arcidiacono and Geronimo, 1976||Along the western African coasts|
|Croatia||Widespread||2006||Introduced||Invasive||Galil, 2006||Successively it has been found in Croatia. Constitutes large, stable populations in the Levant Sea|
|Cyprus||Widespread||2006||Introduced||Galil, 2006||Successively it has been found in Cyrus. Constitutes large, stable populations in the Levant Sea|
|Greece||Widespread||Introduced||Invasive||Pancucci-Papadopoulou et al., 2005; Zenetos et al., 2005; Galil, 2006|
|Italy||Widespread||Introduced||Galil, 2006; Sarà et al., 2006|
History of Introduction and SpreadTop of page
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 IntroductionTop of page
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 ListTop of page
|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 EcologyTop of page
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.
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 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).
ClimateTop of page
|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 TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|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 enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Stramonita haemastoma||Predator||Adult||not specific||N/A|
Notes on Natural EnemiesTop of page
Stramonita haemastoma, a whelk from the Muricidae, preferentially preys on B. pharaonis on the Israeli coast (Rilov et al., 2002).
Means of Movement and DispersalTop of page
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 CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Biodiversity (generally)||Positive and negative|
Economic ImpactTop of page
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 ImpactTop of page
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 SpeciesTop of page
Risk and Impact FactorsTop 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
- Altered trophic level
- Changed gene pool/ selective loss of genotypes
- 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
- Competition - monopolizing resources
- Competition - smothering
- Rapid growth
- Highly likely to be transported internationally accidentally
Similarities to Other Species/ConditionsTop of page
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 ControlTop of page
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 NeedsTop of page
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.
ReferencesTop of page
Çinar M; Cem A, 2007. A Preliminary Study on the Population Characteristics of the Lessepsian Species Pseudonereis anomala (Polychaeta: Nereididae) in Iskenderun Bay (Levantine Sea, Eastern Mediterranean). Turk. J. Zool, 31:403-410.
Dogan A; Önen M; Öztürk B, 2007. A new record of the invasive Red Sea mussel Brachidontes pharaonis (Fischer P., 1870) (Bivalvia: Mytilidae) from the Turkish coasts. Aquatic Invasions, 2(4):461-463. http://www.aquaticinvasions.ru/2007/AI_2007_2_4_Dogan_etal.pdf
Fishelson L; Bresler V; Abelsonb A; Stonea L; Gefena E; Rosenfelda M; Mokadyb O, 2002. The two sides of man-induced changes in littoral marine communities: Eastern Mediterranean and the Red Sea as an example. The Science of the Total Environment, 296:139-151.
Fuchs Th, 1878. [English title not available]. (Die geologische Beschaffenheit der Landenge von Suez.) Denkschriften der Kaiserkichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 38:25.
Galil BS; Zenetos A, 2002. A sea change - Exotics in the eastern Mediterranean Sea. In: Invasive species of Europe; distribution, impact and management [ed. by Leppäkoski E, Gollasch S, Olenin] Dordrecht, The Netherlands: Kluwer Academic Publishers, 325-326.
Gianguzza P; Chemello R; Riggio S, 1997. [English title not available]. (Segnalazione di Brachi- dontes pharaonis (P. Fischer, 1870) (Bivalvia, Mytilidae) nella salina di Marsala e considerazioni sulla distribuzione della specie in Mediterraneo.) Boll. Malac, 33:169-172.
Gruvel A; Moazzo G, 1931. [English title not available]. (Contribution à la faune malacologique marine des côtes Libano-Syriennes.) In: Les états de Syrie. Richesses marines et fluviales [ed. by Gruvel, A.]. Paris, France: Société des Editions Géographiques, Maritimes et Coloniales, 437-456.
Haas G, 1937. Mollusca marina. Prodromus faunae Palestinae, Essai sur les Eléménts zoogéographiques et historiques du sud-ouest du sous-règne Paléarctique. Mémoires presents à l'Institut d'Égypte, 33 [ed. by Bodenheimer, F.]. 275-280.
Lee T; Foighil D, 2004. Hidden Floridian biodiversity: mitochondrial and nuclear gene trees reveal four cryptic species within the scorched mussel, Brachidontes exustus, species complex. Mol. Ecol, 13:3527- 3542.
Lee T; O´Foighil D, 2005. Placing the floridian marine genetic disjunction into a regional evolutionary context using the scorched mussel, Brachidontes exustus, species complex. Evolution, 59(10):2139-2158.
Moshira H, 2008. Biodiversity of benthic communities at high latitudes of the Red Sea. In: 11th International Coral Reef Symposium. Forte Lauderdale, Florida, USA. Convention Center, July 7-11, 2008. Cairo, Egypt: The American University in Cairo. http://www.google.com/url?sa=t&source=web&ct=res&cd=1&ved=0CA4QFjAA&url=http%3A%2F%2Fwww.nova.edu%2Fncri%2F11icrs%2Fabstract_files%2Ficrs2008-002585.pdf&ei=29X6SsLhCMWLnQeMuPD7DA&usg=AFQjCNETRHN83VAN8h88k83_Em8i1asekA&sig2=rM9HMOpwnmR2O-aumXhyuA
Nakhlé KF; Cossa D; Ghaby Khalaf; Beliaeff B, 2006. Brachidontes variabilis and Patella sp. as quantitative biological indicators for cadmium, lead and mercury in the Lebanese coastal waters. Environmental Pollution, 142(1):73-82.
Niederhofer; Enzenross L; Enzenross R, 1991. [English title not available]. (Neue Erkenntnisse uber die Ausbreitung von Lesseps'schen Einwanderern (Mollusca) an der turkischen Mittelmeerkuste.) Club Conchylia Informationen, 33(3-4):94-108.
Sarà G; Romano C; Caruso M; Mazzola A, 2000. The new Lessepsian entry Brachidontes pharaonis (Fischer P., 1870) (Bivalvia, Mytilidae) in the western Mediterranean: a physiological analysis under varying natural conditions. Journal of Shellfish Research, 19:967-977.
Sarà G; Romano C; Mazzola A, 2006. A new Lessepsian species in the western Mediterranean (Brachidontes pharaonis Bivalvia: Mytilidae): density, resource allocation and biomass. JMBA2 Biodiversity Records:7 pp.
Steiner G; Hammer S, 2000. Molecular phylogeny of the Bivalvia inferred from 18S rDNA sequences with particular reference to the Pteriomorphia. In: The Evolutionary Biology of the Bivalvia [ed. by Harper, E. M. \Taylor, J. D. \Crame, J. A.]. 11-29. [Geological Society, London, Special Publications 177.]
Streftaris N; Zenetos A; Papathanassiou E, 2005. Globalisation in marine ecosystems: The story of non-indigenous marine species across European seas oceanography and marine biology: An annual review, 43. 419-453.
Terranova MS; Lo Brutto S; Arculeo M; Mitton JB, 2006. Population structure of Brachidontes pharaonis (P. Fischer, 1870) (Bivalvia, Mytilidae) in the Mediterranean Sea, and evolution of a novel mtDNA polymorphism. Mar. Biol, 159:89-101.
Terranova MS; Lo Brutto S; Arculeo M; Mitton JB, 2007. A mitochondrial phylogeography of Brachidontes variabilis ( Bivalvia : Mytilidae ) reveals three cryptic species. Journal of Zoological Systematics and Evolutionary Research, 45(January):289-298.
OrganizationsTop of page
Monaco: CIESM - Commission Internationale pour l'Exploration de la Mer Mediterranee, Villa Girasole, 16 bd de Suisse, http://www.ciesm.org/atlas/
USA: OBIS - Ocean Biogeographic Information System, 71 Dudley Road, New Brunswick, NJ 08901, http://www.iobis.org
ContributorsTop of page
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
Distribution MapsTop of page
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