Versuriga anadyomene, a newly recorded scyphozoan jellyfi sh(Scyphozoa: Rhizostomae) in Chinese waters
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摘要
Scyphozoans are currently causing great concern worldwide due to the ef fects of their blooms on the marine environment. During a ?eld survey in Beibu Gulf, South China Sea, on 27 May 2015, we found specimens of a rhizostome jelly?sh that had not previously been recorded in Chinese seas. The morphological characters and DNA sequences of several specimens were examined in the laboratory. These specimens had the following morphological characters: hemispherical umbrella with reticular grooves; eight large semicircular velar lappets alternating with narrow lappets in each of the 8 octants; laterally ?attened mouth-arms with numerous ?at, membranous branches bearing many clubs and intermediate ?laments;non-anastomosed broad perradial canals and anastomosed narrow interradial canals. The morphological characters of these rhizostome jelly?sh were consistent with descriptions of Versuriga anadyomene.Furthermore, phylogenetic analysis of mitochondrial 16 S rDNA and nuclear 18 S rDNA gene fragments con?rmed that these specimens were in the same cluster as V. anadyomene collected in Palau. Therefore,this rhizostome jelly?sh was identi?ed as V. anadyomene. This is the ?rst record of the genus Versuriga and the species V. anadyomene in Chinese seas. The arrival of this species may be attributed to physical transport from the coastal waters of South East Asia.
Scyphozoans are currently causing great concern worldwide due to the ef fects of their blooms on the marine environment. During a ?eld survey in Beibu Gulf, South China Sea, on 27 May 2015, we found specimens of a rhizostome jelly?sh that had not previously been recorded in Chinese seas. The morphological characters and DNA sequences of several specimens were examined in the laboratory. These specimens had the following morphological characters: hemispherical umbrella with reticular grooves; eight large semicircular velar lappets alternating with narrow lappets in each of the 8 octants; laterally flattened mouth-arms with numerous flat, membranous branches bearing many clubs and intermediate ?laments;non-anastomosed broad perradial canals and anastomosed narrow interradial canals. The morphological characters of these rhizostome jelly?sh were consistent with descriptions of Versuriga anadyomene.Furthermore, phylogenetic analysis of mitochondrial 16 S rDNA and nuclear 18 S rDNA gene fragments con?rmed that these specimens were in the same cluster as V. anadyomene collected in Palau. Therefore,this rhizostome jelly?sh was identi?ed as V. anadyomene. This is the ?rst record of the genus Versuriga and the species V. anadyomene in Chinese seas. The arrival of this species may be attributed to physical transport from the coastal waters of South East Asia.
Scyphozoans are currently causing great concern worldwide due to the ef fects of their blooms on the marine environment. During a ?eld survey in Beibu Gulf, South China Sea, on 27 May 2015, we found specimens of a rhizostome jelly?sh that had not previously been recorded in Chinese seas. The morphological characters and DNA sequences of several specimens were examined in the laboratory. These specimens had the following morphological characters: hemispherical umbrella with reticular grooves; eight large semicircular velar lappets alternating with narrow lappets in each of the 8 octants; laterally flattened mouth-arms with numerous flat, membranous branches bearing many clubs and intermediate ?laments;non-anastomosed broad perradial canals and anastomosed narrow interradial canals. The morphological characters of these rhizostome jelly?sh were consistent with descriptions of Versuriga anadyomene.Furthermore, phylogenetic analysis of mitochondrial 16 S rDNA and nuclear 18 S rDNA gene fragments con?rmed that these specimens were in the same cluster as V. anadyomene collected in Palau. Therefore,this rhizostome jelly?sh was identi?ed as V. anadyomene. This is the ?rst record of the genus Versuriga and the species V. anadyomene in Chinese seas. The arrival of this species may be attributed to physical transport from the coastal waters of South East Asia.
引文
Akaike H.1992.Information theory as an extension of the maximum likelihood principle.In:Proceedings of the 2nd International Symposium on Information Theory.Academiai,Budapest,Hungaty.p.267-281.
Appeltans W,Ahyong S T,Anderson G,Angel M V,Artois T,Bailly N,Bamber R,Barber A,Bartsch I,Berta A,B?a?ewicz-Paszkowycz M,Bock P,Boxshall G,Boyko CB,Brand?o S N,Bray R A,Bruce N L,Cairns S D,Chan T Y,Cheng L N,Collins A G,Cribb T,Curini-Galletti M,Dahdouh-Guebas F,Davie P J F,Dawson M N,De Clerck O,Decock W,De Grave S,De Voogd N J,Domning D P,Emig C C,Erséus C,Eschmeyer W,Fauchald K,Fautin D G,Feist S W,Fransen C H J M,Furuya H,GarciaAlvarez O,Gerken S,Gibson D,Gittenberger A,Gofas S,Gómez-Daglio L,Gordon D P,Guiry M D,Hernandez F,Hoeksema B W,Hopcroft R R,Jaume D,Kirk P,Koedam N,Koenemann S,Kolb J B,Kristensen R M,Kroh A,Lambert G,Lazarus D B,Lemaitre R,Longshaw M,Lowry J,Macpherson E,Madin L P,Mah C,Mapstone G,McLaughlin P A,Mees J,Meland K,Messing C G,Mills C E,Molodtsova T N,Mooi R,Neuhaus B,Ng P K L,Nielsen C,Norenburg J,Opresko D M,Osawa M,Paulay G,Perrin W,Pilger J F,Poore G C B,Pugh P,Read G B,Reimer J D,Rius M,Rocha R M,Saiz-Salinas J I,Scarabino V,Schierwater B,Schmidt-Rhaesa A,Schnabel K E,Schotte M,Schuchert P,Schwabe E,Segers H,SelfSullivan C,Shenkar N,Siegel V,Sterrer W,St?hr S,Swalla B,Tasker M L,Thuesen E V,Timm T,Todaro MA,Turon X,Tyler S,Uetz P,Van Der Land J,Vanhoorne B,Van Ofwegen L P,Van Soest R W M,Vanaverbeke J,Walker-Smith G,Walter T C,Warren A,Williams G C,Wilson S P,Costello M J.2012.The magnitude of global marine species diversity.Curr.Biol.,22(23):2 189-2 202.
Arai M N.2001.Pelagic coelenterates and eutrophication:a review.In:Purcell J E,Graham W M,Dumont H J eds.Jelly?sh Blooms:Ecological and Societal Importance.Springer,Dordrecht,Netherlands.p.69-87.
Bayha K M,Dawson M N,Collins A G,Barbeitos M S,Haddock S H D.2010.Evolutionary relationships among scyphozoan jelly?sh families based on complete taxon sampling and phylogenetic analyses of 18S and 28Sribosomal DNA.Integr.Comp.Biol.,50(3):436-455.
Bayha K M,Graham W M.2009.A new Taqman?PCR-based method for the detection and identi?cation of scyphozoan jelly?sh polyps.Hydrobiologia,616(1):217-228.
Bos A R,Cruz-Rivera E,Sanad A M.2017.Herbivorous?shes Siganus rivulatus(Siganidae)and Zebrasoma desjardinii(Acanthuridae)feed on ctenophora and scyphozoa in the Red Sea.Mar.Biodiver.,47(1):243-246.
Brotz L,Schiariti A,López-Martínez J,álvarez-Tello J,Hsieh Y H P,Jones R P,Qui?ones J,Dong Z J,Morandini A C,Preciado M,Laaz E,Mianzan H.2017.Jelly?sh?sheries in the Americas:origin,state of the art,and perspectives on new?shing grounds.Rev.Fish Bio l.Fish,27(1):1-29.
Cardona L,Martínez-I?igo L,Mateo R,González-Solís J.2015.The role of sardine as prey for pelagic predators in the western Mediterranean Sea assessed using stable isotopes and fatty acids.Mar.Ecol.Prog.Ser.,531:1-14.
?evik C,Erkol I L,Toklu B.2006.A new record of an alien jelly?sh from the levantine coast of turkey-Cassiopea andromeda(Forssk?l,1775)[Cnidaria:Scyphozoa:Rhizostomea].Aquat.Invasions,1(3):196-197.
Collins A G.2002.Phylogeny of Medusozoa and the evolution of cnidarian life cycles.J.Evol.Biol.,15(3):418-432.
Daglio L G,Dawson M N.2017.Species richness of jelly?shes(Scyphozoa:Discomedusae)in the Tropical Eastern Paci?c:missed taxa,molecules,and morphology match in a biodiversity hotspot.Invertebr.Syst.,31(5):635-663.
Daly M,Brugler M R,Cartwright P,Collins A G,Dawson MN,Fautin D G,France S C,McFadden C S,Opresko D M,Rodriguez E,Romano S L,Stake J L.2007.The phylum Cnidaria:a review of phylogenetic patterns and diversity300 years after Linnaeus.Zootaxa,1668:127-182.
Dawson M N,Gupta A S,England M H.2005.Coupled biophysical global ocean model and molecular genetic analyses identify multiple introductions of cryptogenic species.P roc.Natl.Acad.Sci.USA,102(34):11 968-11 973.
Deidun A,Sciberras J,Sciberras A,Gauci A,Balistreri P,Salvatore A,Piraino S.2017.The?rst record of the whitespotted Australian jelly?sh Phyllorhiza punctata von Lendenfeld,1884 from Maltese waters(western Mediterranean)and from the Ionian coast of Italy.BioInvasions Rec.,6(2):119-124.
Dong Z J,Liu D Y,Keesing J K.2010.Jelly?sh blooms in China:dominant species,causes and consequences.Mar.Pollu t.Bull.,60(7):954-963.
Dong Z J,Liu D Y,Keesing J K.2014.Contrasting trends in populations of Rhopilema esculentum and Aurelia aurita in Chinese waters.In:Pitt K A,Lucas C H eds.Jelly?sh Blooms.Springer,Dordrecht,Netherlands.p.207-218.
Ender A,Schierwater B.2003.Placozoa are not derived cnidarians:evidence from molecular morphology.Mol.Biol.Evol.,20(1):130-134.
Folmer O,Black M,Hoeh W,Lutz R,Vrijenhoek R.1994.DNA primers for ampli?cation of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates.Mol.M ar.B iol.Biotech nol.,3(5):294-299.
Galil B S,Gershwin L A,Douek J,Rinkevich B.2010.Marivagia stellata gen.et sp.nov.(Scyphozoa:Rhizostomeae:Cepheidae),another alien jelly?sh from the Mediterranean coast of Israel.Aquat.Invasions,5(4):331-340.
Galil B S,Shoval L,Goren M.2009.Phyllorhiza punctata von Lendenfeld,1884(Scyphozoa:Rhizostomeae:Mastigiidae)reappeared of f the Mediterranean coast of Israel.Aquat.Invasions,4(3):481-483.
Gao S W,Hong H X,Zhang S M.2002.Fauna Sinica,Invertebrata Vol.27.Phylum Cnidaria,Class Hydrozoa,Subclass Siphonophorae.Class Scyphomedusae.Science Press,Beijing,China.(in Chinese)
Gershwin L A,Zeidler W,Davie P J F.2010.Medusae(Cnidaria)of Moreton Bay,Queensland,Australia.Mem.Qld.Mus,54(3):47-108.
Gershwin L A,Zeidler W.2008.Some new and previously unrecorded Scyphomedusae(Cnidaria:Scyphozoa)from southern Australian coastal waters.Zootaxa,1744:1-18.
Hamilton G.2016.The secret lives of jelly?sh:long regarded as minor players in ocean ecology,jelly?sh are actually important parts of the marine food web.Nature,531(7595):432-434.
Hong H X,Lin L M.2010.Study on the fauna of the scyphomedusae in the China Sea.J.Jimei Univ.(Nat.Sci.),15(1):18-24.(in Chinese with English abstract)
Hsieh Y P H,Leong F M,Rudloe J.2001.Jelly?sh as food.Hydrobiologia,45 1(1-3):11-17.
Kitamura M,Omori M.2010.Synopsis of edible jelly?shes collected from Southeast Asia,with notes on jelly?sh?sheries.Plankton Benthos Res.,5(3):106-118.
Kramp P L.1961.Synopsis of the medusae of the world.J.Mar.Biol.Assoc.UK,40:7-382.
Marques R,Bouvier C,Darnaude A M,Molinero J C,Przybyla C,Soriano S,Tomasini J A,Bonnet D.2016.Jelly?sh as an alternative source of food for opportunistic?shes.J.Ex p.Mar.Biol.Ecol.,485:1-7.
Mass O.1903.Die Scyphomedusen der Siboga-expedition.Leiden,Buchhandlung und Druckerei Vormals E.J.Brill.Siboga Exped.,11(9):1-91.
Mayer A G.1910.Medusae of the World.Vol III:Scyphozoa.Carnegie Institute,Washington,USA.
Medlin L,Elwood H J,Stickel S,Sogin M L.1988.The characterization of enzymatically ampli?ed eukaryotic16S-like rRNA-coding regions.Gene,71(2):491-499.
Morandini A C,Stampar S N,Maronna M M,Da Silveira F L.2017.All non-indigenous species were introduced recently?The case study of Cassiopea(Cnidaria:Scyphozoa)in Brazilian waters.J.Mar.Biol.Assoc.UK,97(2):321-328.
Oca?a-Luna A,Sánchez-Ramírez M,Aguilar-Durán R.2010.First record of Phyllorhiza punctata von Lendenfeld,1884(Cnidaria:Scyphozoa,Mastigiidae)in Mexico.Aquat.Invasions,5(S1):S79-S84.
Ohtsuka S,Koike K,Lindsay D,Nishikawa J,Miyake H,Kawahara M,Mulyadi,Mujiono N,Hiromi J,Komatsu H.2009.Symbionts of marine medusae and ctenophores.Plankton Benthos Res.,4(1):1-13.
Purcell J E.2012.Jelly?sh and ctenophore blooms coincide with human proliferations and environmental perturbations.Ann.Rev.Mar.Sci.,4:209-235.
Tamura K,Peterson D,Peterson N,Stecher G,Nei M,Kumar S.2011.MEGA5:molecular evolutionary genetics analysis using maximum likelihood,evolutionary distance,and maximum parsimony methods.Mol.Biol.Evol.,28(10):2 731-2 739.
Yahia M N D,Yahia O K D,Gueroun S K M,Aissi M,Deidun A,Fuentes V,Piraino S.2013.The invasive tropical scyphozoan Rhopilema nomadica Galil,1990 reaches the Tunisian coast of the Mediterranean Sea.BioInvasions Rec.,2(4):319-323.
Appeltans W,Ahyong S T,Anderson G,Angel M V,Artois T,Bailly N,Bamber R,Barber A,Bartsch I,Berta A,B?a?ewicz-Paszkowycz M,Bock P,Boxshall G,Boyko CB,Brand?o S N,Bray R A,Bruce N L,Cairns S D,Chan T Y,Cheng L N,Collins A G,Cribb T,Curini-Galletti M,Dahdouh-Guebas F,Davie P J F,Dawson M N,De Clerck O,Decock W,De Grave S,De Voogd N J,Domning D P,Emig C C,Erséus C,Eschmeyer W,Fauchald K,Fautin D G,Feist S W,Fransen C H J M,Furuya H,GarciaAlvarez O,Gerken S,Gibson D,Gittenberger A,Gofas S,Gómez-Daglio L,Gordon D P,Guiry M D,Hernandez F,Hoeksema B W,Hopcroft R R,Jaume D,Kirk P,Koedam N,Koenemann S,Kolb J B,Kristensen R M,Kroh A,Lambert G,Lazarus D B,Lemaitre R,Longshaw M,Lowry J,Macpherson E,Madin L P,Mah C,Mapstone G,McLaughlin P A,Mees J,Meland K,Messing C G,Mills C E,Molodtsova T N,Mooi R,Neuhaus B,Ng P K L,Nielsen C,Norenburg J,Opresko D M,Osawa M,Paulay G,Perrin W,Pilger J F,Poore G C B,Pugh P,Read G B,Reimer J D,Rius M,Rocha R M,Saiz-Salinas J I,Scarabino V,Schierwater B,Schmidt-Rhaesa A,Schnabel K E,Schotte M,Schuchert P,Schwabe E,Segers H,SelfSullivan C,Shenkar N,Siegel V,Sterrer W,St?hr S,Swalla B,Tasker M L,Thuesen E V,Timm T,Todaro MA,Turon X,Tyler S,Uetz P,Van Der Land J,Vanhoorne B,Van Ofwegen L P,Van Soest R W M,Vanaverbeke J,Walker-Smith G,Walter T C,Warren A,Williams G C,Wilson S P,Costello M J.2012.The magnitude of global marine species diversity.Curr.Biol.,22(23):2 189-2 202.
Arai M N.2001.Pelagic coelenterates and eutrophication:a review.In:Purcell J E,Graham W M,Dumont H J eds.Jelly?sh Blooms:Ecological and Societal Importance.Springer,Dordrecht,Netherlands.p.69-87.
Bayha K M,Dawson M N,Collins A G,Barbeitos M S,Haddock S H D.2010.Evolutionary relationships among scyphozoan jelly?sh families based on complete taxon sampling and phylogenetic analyses of 18S and 28Sribosomal DNA.Integr.Comp.Biol.,50(3):436-455.
Bayha K M,Graham W M.2009.A new Taqman?PCR-based method for the detection and identi?cation of scyphozoan jelly?sh polyps.Hydrobiologia,616(1):217-228.
Bos A R,Cruz-Rivera E,Sanad A M.2017.Herbivorous?shes Siganus rivulatus(Siganidae)and Zebrasoma desjardinii(Acanthuridae)feed on ctenophora and scyphozoa in the Red Sea.Mar.Biodiver.,47(1):243-246.
Brotz L,Schiariti A,López-Martínez J,álvarez-Tello J,Hsieh Y H P,Jones R P,Qui?ones J,Dong Z J,Morandini A C,Preciado M,Laaz E,Mianzan H.2017.Jelly?sh?sheries in the Americas:origin,state of the art,and perspectives on new?shing grounds.Rev.Fish Bio l.Fish,27(1):1-29.
Cardona L,Martínez-I?igo L,Mateo R,González-Solís J.2015.The role of sardine as prey for pelagic predators in the western Mediterranean Sea assessed using stable isotopes and fatty acids.Mar.Ecol.Prog.Ser.,531:1-14.
?evik C,Erkol I L,Toklu B.2006.A new record of an alien jelly?sh from the levantine coast of turkey-Cassiopea andromeda(Forssk?l,1775)[Cnidaria:Scyphozoa:Rhizostomea].Aquat.Invasions,1(3):196-197.
Collins A G.2002.Phylogeny of Medusozoa and the evolution of cnidarian life cycles.J.Evol.Biol.,15(3):418-432.
Daglio L G,Dawson M N.2017.Species richness of jelly?shes(Scyphozoa:Discomedusae)in the Tropical Eastern Paci?c:missed taxa,molecules,and morphology match in a biodiversity hotspot.Invertebr.Syst.,31(5):635-663.
Daly M,Brugler M R,Cartwright P,Collins A G,Dawson MN,Fautin D G,France S C,McFadden C S,Opresko D M,Rodriguez E,Romano S L,Stake J L.2007.The phylum Cnidaria:a review of phylogenetic patterns and diversity300 years after Linnaeus.Zootaxa,1668:127-182.
Dawson M N,Gupta A S,England M H.2005.Coupled biophysical global ocean model and molecular genetic analyses identify multiple introductions of cryptogenic species.P roc.Natl.Acad.Sci.USA,102(34):11 968-11 973.
Deidun A,Sciberras J,Sciberras A,Gauci A,Balistreri P,Salvatore A,Piraino S.2017.The?rst record of the whitespotted Australian jelly?sh Phyllorhiza punctata von Lendenfeld,1884 from Maltese waters(western Mediterranean)and from the Ionian coast of Italy.BioInvasions Rec.,6(2):119-124.
Dong Z J,Liu D Y,Keesing J K.2010.Jelly?sh blooms in China:dominant species,causes and consequences.Mar.Pollu t.Bull.,60(7):954-963.
Dong Z J,Liu D Y,Keesing J K.2014.Contrasting trends in populations of Rhopilema esculentum and Aurelia aurita in Chinese waters.In:Pitt K A,Lucas C H eds.Jelly?sh Blooms.Springer,Dordrecht,Netherlands.p.207-218.
Ender A,Schierwater B.2003.Placozoa are not derived cnidarians:evidence from molecular morphology.Mol.Biol.Evol.,20(1):130-134.
Folmer O,Black M,Hoeh W,Lutz R,Vrijenhoek R.1994.DNA primers for ampli?cation of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates.Mol.M ar.B iol.Biotech nol.,3(5):294-299.
Galil B S,Gershwin L A,Douek J,Rinkevich B.2010.Marivagia stellata gen.et sp.nov.(Scyphozoa:Rhizostomeae:Cepheidae),another alien jelly?sh from the Mediterranean coast of Israel.Aquat.Invasions,5(4):331-340.
Galil B S,Shoval L,Goren M.2009.Phyllorhiza punctata von Lendenfeld,1884(Scyphozoa:Rhizostomeae:Mastigiidae)reappeared of f the Mediterranean coast of Israel.Aquat.Invasions,4(3):481-483.
Gao S W,Hong H X,Zhang S M.2002.Fauna Sinica,Invertebrata Vol.27.Phylum Cnidaria,Class Hydrozoa,Subclass Siphonophorae.Class Scyphomedusae.Science Press,Beijing,China.(in Chinese)
Gershwin L A,Zeidler W,Davie P J F.2010.Medusae(Cnidaria)of Moreton Bay,Queensland,Australia.Mem.Qld.Mus,54(3):47-108.
Gershwin L A,Zeidler W.2008.Some new and previously unrecorded Scyphomedusae(Cnidaria:Scyphozoa)from southern Australian coastal waters.Zootaxa,1744:1-18.
Hamilton G.2016.The secret lives of jelly?sh:long regarded as minor players in ocean ecology,jelly?sh are actually important parts of the marine food web.Nature,531(7595):432-434.
Hong H X,Lin L M.2010.Study on the fauna of the scyphomedusae in the China Sea.J.Jimei Univ.(Nat.Sci.),15(1):18-24.(in Chinese with English abstract)
Hsieh Y P H,Leong F M,Rudloe J.2001.Jelly?sh as food.Hydrobiologia,45 1(1-3):11-17.
Kitamura M,Omori M.2010.Synopsis of edible jelly?shes collected from Southeast Asia,with notes on jelly?sh?sheries.Plankton Benthos Res.,5(3):106-118.
Kramp P L.1961.Synopsis of the medusae of the world.J.Mar.Biol.Assoc.UK,40:7-382.
Marques R,Bouvier C,Darnaude A M,Molinero J C,Przybyla C,Soriano S,Tomasini J A,Bonnet D.2016.Jelly?sh as an alternative source of food for opportunistic?shes.J.Ex p.Mar.Biol.Ecol.,485:1-7.
Mass O.1903.Die Scyphomedusen der Siboga-expedition.Leiden,Buchhandlung und Druckerei Vormals E.J.Brill.Siboga Exped.,11(9):1-91.
Mayer A G.1910.Medusae of the World.Vol III:Scyphozoa.Carnegie Institute,Washington,USA.
Medlin L,Elwood H J,Stickel S,Sogin M L.1988.The characterization of enzymatically ampli?ed eukaryotic16S-like rRNA-coding regions.Gene,71(2):491-499.
Morandini A C,Stampar S N,Maronna M M,Da Silveira F L.2017.All non-indigenous species were introduced recently?The case study of Cassiopea(Cnidaria:Scyphozoa)in Brazilian waters.J.Mar.Biol.Assoc.UK,97(2):321-328.
Oca?a-Luna A,Sánchez-Ramírez M,Aguilar-Durán R.2010.First record of Phyllorhiza punctata von Lendenfeld,1884(Cnidaria:Scyphozoa,Mastigiidae)in Mexico.Aquat.Invasions,5(S1):S79-S84.
Ohtsuka S,Koike K,Lindsay D,Nishikawa J,Miyake H,Kawahara M,Mulyadi,Mujiono N,Hiromi J,Komatsu H.2009.Symbionts of marine medusae and ctenophores.Plankton Benthos Res.,4(1):1-13.
Purcell J E.2012.Jelly?sh and ctenophore blooms coincide with human proliferations and environmental perturbations.Ann.Rev.Mar.Sci.,4:209-235.
Tamura K,Peterson D,Peterson N,Stecher G,Nei M,Kumar S.2011.MEGA5:molecular evolutionary genetics analysis using maximum likelihood,evolutionary distance,and maximum parsimony methods.Mol.Biol.Evol.,28(10):2 731-2 739.
Yahia M N D,Yahia O K D,Gueroun S K M,Aissi M,Deidun A,Fuentes V,Piraino S.2013.The invasive tropical scyphozoan Rhopilema nomadica Galil,1990 reaches the Tunisian coast of the Mediterranean Sea.BioInvasions Rec.,2(4):319-323.