Medit. Mar. Sci., 11/2, 2010, 381-493 381

Alien species in the Mediterranean Sea by 2010. A contribution to the applicationof European Union’s Marine Strategy Framework Directive (MSFD).

Part I. Spatial distribution

A. ZENETOS1, S. GOFAS2, M. VERLAQUE3, M.E. CINAR4, J.E. GARCI’A RASO5,C.N. BIANCHI6, C. MORRI6, E. AZZURRO7, M. BILECENOGLU8, C. FROGLIA9,

I. SIOKOU10, D. VIOLANTI11, A. SFRISO12, G. SAN MART N13, A. GIANGRANDE14,T. KATA AN4, E. BALLESTEROS15, A. RAMOS-ESPLA’16, F. MASTROTOTARO17,

O. OCA A18, A. ZINGONE19, M.C. GAMBI19 and N. STREFTARIS10

1 Institute of Marine Biological Resources, Hellenic Centre for Marine Research, P.O. Box 712,19013 Anavissos, Hellas

2 Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Ma’laga,E-29071 Ma’laga, Spain

3 UMR 6540, DIMAR, COM, CNRS, Université de la Méditerranée, France 4 Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100 Bornova, Izmir, Turkey

5 Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Ma’laga,E-29071 Ma’laga, Spain

6 DipTeRis (Dipartimento per lo studio del Territorio e della sue Risorse), University of Genoa,Corso Europa 26, 16132 Genova, Italy

7 Institut de Ciències del Mar (CSIC) Passeig Mar tim de la Barceloneta, 37-49, E-08003 Barcelona, Spain 8 Adnan Menderes University, Faculty of Arts & Sciences, Department of Biology, 09010 Aydin, Turkey

9 c\o CNR-ISMAR, Sede Ancona, Largo Fiera della Pesca, 60125 Ancona, Italy10 Institute of Oceanography, Hellenic Centre for Marine Research, P.O. Box 712, 19013 Anavissos, Hellas11 Dipartimento di Scienze della Terra, University of Turin, via Valperga Caluso 35, 10125 Torino, Italy12 Department of Environmental Sciences, University of Venice, Calle Larga 2137, 30123 Venice, Italy

13 Departamento de Biolog a (Zoolog a), Facultad de Ciencias, Universidad Auto’noma de Madrid,calle Darwin, 2, 28049 Madrid, Spain

14 Department of Biological and Environmental Sciences and Technologies, University of Lecce,Complesso Ecotekne, Via Prov. le Lecce-Monteroni, 73100 Lecce, Italy

15 Centre d’Estudis Avanç ats de Blanes (CSIC), Acc. Cala, Sant Francesc 14, 17300 Blanes, Girona, Spain16 Centro de Investigacio’n Marina de Santa Pola (CIMAR), Universidad de Alicante, 03130 Santa

Pola, Alicante, Spain 17 Department of Biology, University of Bari, Via Orabona 4, 70125 Bari, Italy

18 Departamento Oceanograf a Biolo’gica y Biodiversidad, Fundacio’n Museo del Mar MuelleCan~onero Dato S/N 51001, Ceuta, Spain

19 Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy

Corresponding author: zenetos@ath.hcmr.gr

Review ArticleMediterranean Marine ScienceIndexed in WoS (Web of Science, ISI Thomson)The journal is available on line at http://www.medit-mar-sc.net

Introduction

The Mediterranean Sea is one of theseas of the world most affected by biolog-ical invasions in terms of how long theinvaders have been present (OCCHI-PINTI-AMBROGI, 2000; STREFTARISet al., 2005), in number of alien speciesdetected (COSTELLO et al., 2010) and inthe unprecedented rate of introduction(ZENETOS, 2009; 2010). Triggered bywarming waters and a newly improvedroute through the Suez Canal,tropical/subtropical marine species, somevery invasive, have progressively movedinto the Mediterranean, disruptingecosystem stability of the basin.

In 1978, there were 128 provenLessepsian immigrants and 76 doubtfulcases (POR, 1978). Since the first reviewof alien species in the Mediterranean(ZIBROWIUS, 1992), there have beenmany papers and reviews dealing with thissubject. More recently, CIESM has pub-lished four Atlases of Exotic Species inthe Mediterranean (GALIL et al., 2002;GOLANI et al., 2002; ZENETOS et al.,2004; VERLAQUE et al., in press). Themost recent lists show 116 species of fish-es, 70 species of decapod and stomatopodcrustaceans, 137 species of molluscs and110 species of macrophytes. Thisincreased number of records, particularlyover the past few years, has been partly

Medit. Mar. Sci., 11/2, 2010, 381-493382

Abstract

The state-of-art on alien species in the Mediterranean Sea is presented, making distinctionsamong the four subregions defined in the EU Marine Strategy Framework Directive: (i) the WesternMediterranean Sea (WMED); (ii) the Central Mediterranean Sea (CMED); (iii) the Adriatic Sea(ADRIA); and (iv) the Eastern Mediterranean Sea (EMED). The updated checklist (December2010) of marine alien species within each subregion, along with their acclimatization status and ori-gin, is provided. A total of 955 alien species is known in the Mediterranean, the vast majority of themhaving being introduced in the EMED (718), less in the WMED (328) and CMED (267) and least inthe Adriatic (171). Of these, 535 species (56%) are established in at least one area.

Despite the collective effort of experts who attempted in this work, the number of introducedspecies remains probably underestimated. Excluding microalgae, for which knowledge is still insuffi-cient, aliens have increased the total species richness of the Mediterranean Sea by 5.9%. This figureshould not be directly read as an indication of higher biodiversity, as spreading of so many alienswithin the basin is possibly causing biotic homogenization. Thermophilic species, i.e. Indo-Pacific,Indian Ocean, Red Sea, Tropical Atlantic, Tropical Pacific, and circum(sub)tropical, account for88.4% of the introduced species in the EMED, 72.8% in the CMED, 59.3% in the WMED and56.1% in the Adriatic. Cold water species, i.e. circumboreal, N Atlantic, and N Pacific, make up asmall percentage of the introduced species, ranging between 4.2% and 21.6% and being morenumerous in the Adriatic and less so in the EMED.

Species that are classified as invasive or potentially invasive are 134 in the whole of the Mediter-ranean: 108 are present in the EMED, 75 in the CMED, 53 in the Adriatic and 64 in the WMED.The WMED hosts most invasive macrophytes, whereas the EMED has the lion’s share in poly-chaetes, crustaceans, molluscs and fish.

Keywords: Biological invasions; Marine aliens; Biogeography; Biodiversity; Mediterranean Sea.

attributed to intensification of researcheffort in this topic. ZENETOS &POLYCHRONIDIS (2010), based ondata gathered, analysed for and adoptedin the European Environment Agency(EEA) and UNEP MAP RAC/SPAreports, estimated that nearly 1000species entered the Mediterranean dur-ing the past century.

A recent synthesis on Mediterraneanmarine biodiversity (COLL et al., 2010)described the Mediterranean Sea as abiodiversity hot spot hosting approxi-mately 17,000 marine species, of whichmore than 600 (3.3%) are alien. Thesemost recent detailed biodiversity esti-mates for alien species show a discrepan-cy from the figures provided byZENETOS et al. (2005; 2008). However,as COLL et al. (2010) have pointed out,the true numbers of alien species are cer-tainly biased downwards. The data arepresumably accurate for large and con-spicuous species that are easily distin-guished from the native biota and forspecies that occur along a frequently sam-pled (or fished) coast and for which taxo-nomic expertise is readily available, butare entirely absent for many of the smallmembers of invertebrates. This under-representation is common in many worksand efforts have been made to includeless studied groups in recent reviews (seeZENETOS et al., 2008).

An up-to-date inventory of the alienspecies in the Mediterranean, apart fromits scientific merits, can fulfil the needs ofthe regulatory requirements and environ-mental management options. This is ofparticular importance since the current

emergence of the new generation of EUpolitical actions covering major maritimestrategic objectives, such as the MarineStrategy Framework Directive (MSFD)(2008/56/EC), the European Strategy forMarine and Maritime Research (COM(2008) 534) encompassing the MarineSpatial Planning and the EcosystemApproach (ECAP) within the BarcelonaConvention by UNEP/MAP1, as well asmany initiatives of the UNEP RAC/SPA.Alien species regulations are of majorimportance in those policies. In the cur-rent MSFD the descriptor: "Non-indige-nous species introduced by human activi-ties are at levels that do not adverselyalter the ecosystems" is actually one of theeleven qualitative descriptors for deter-mining Good Environmental Status(GES). It is also closely related to sixother GES descriptors because of thegreat variety of impacts that InvasiveAlien Species (IAS) may exert on nativebiodiversity, ecosystem functioning, andseabed habitats as well as commercialmarine resources (OLENIN et al., 2010).

This territorial and institutional sce-nario raises the need for a multi-levelgovernance system with a quite complexcoordination system in which non-EUMember States can also participate. Thisis particularly important in the Mediter-ranean, where the southern and easterncoast belong to non-EU Member Statesor countries with accession status (e.g.Turkey and Israel). Furthermore, theBarcelona Convention of the UNEP/MAPsupports the introduction of MSFD(2009) concepts that can be usefulthroughout the basin despite the handi-

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1 The Barcelona Convention of the UNEP/MAP supports the introduction of MSFD in the Mediterranean. To thatend, they are following the ecosystem approach (including the concept of policy steps), which is the priority of therecently adopted 5-year programme. Presently they are working on assessing the state of the environment, includ-ing economic and social aspects, in the four subregions.

cap that only seven of 21 Mediterraneancountries are Member States ofUNEP/MAP.

Considering a) the high introductionrate, b) recent publications and reviewson little studied taxa and c) the need toassess the extent of the phenomenon atsubregional level and inform the EU,UNEP/MED and other stakeholdersaccordingly, this work aims to present the2010 state-of-art on the presence andacclimatization status of alien species ateach MSFD Mediterranean subregion,namely: (i) the Western MediterraneanSea (WMED); (ii) the Central Mediter-ranean Sea (CMED); (iii) the AdriaticSea (ADRIA); and (iv) the EasternMediterranean Sea (EMED). The latteris also commonly referred to as theAegean-Levantine basin, but in this workthe Marmara Sea, bearing more similari-ties to the Aegean than to the neighbour-ing Black Sea, is also included.

Methodology

Biological invasions have become ahot issue at a broad geographical scalesuch as that of Large Marine Ecosystems(LME). Among European LMEs, biolog-ical invasions are most pronounced in theMediterranean (EEA, 2007a; ZENETOSet al., 2009).

On the European scale, over a dozenpartitions can be found dividing up thewaters of the NE Atlantic, the Mediter-ranean Sea and the Black Sea on oceano-graphic or purely geometric bases, or acombination of both, as well as divisionsbased on national jurisdictions2 (Fig. 1).In this work the Mediterranean is being

studied along the four subregions asadopted and described under the MSFD.The coastal areas of the countries and theregions included in these subregions arelisted in Table 1. This division imposessome restrictions/difficulties in the case ofcountries whose waters lie within theboundaries of these subregions as in thecase of Albania, Tunisia and Italy.

The borders of the CMED are herebydefined as the Kythira-Anti-KythiraStraits (Greece) and Libya-Egypt bordersto the east, Otranto Strait (Italy, Albania)to the north, Cap Bon (Tunisia) andsouth-west Sicily to the west.

The list is updated based on validspecies records up to December 2010.Alien species are investigated within allfour regions in terms of their establish-ment success (casual, established, ques-tionable, cryptogenic), as well as invasivesuccess, their native range and geograph-ical expansion. The species lists are pre-sented for seven units which are systemat-ic groups namely: 1) Protozoa (includingForaminifera), 2) Macrophytes, 3) Poly-chaeta, 4) Crustacea, 5) Mollusca, 6) Mis-cellaneous Invertebrata (ArthropodaPycnogonida, Bryozoa, Chaetognatha,Chordata Ascidiacea, Cnidaria,Ctenophora, Echinodermata, Porifera,Sipuncula, Nematoda and Platy-helminthes), and 7) Fish, in each part.

The introduction of alien species in theMediterranean Sea is hard to demonstratefor marine microalgae as for microrgan-isms in general (WYATT & CARLTON,2002). Bearing in mind that it can rarely beexcluded that a suspected microalgal invad-er was already present as part of the rare,hidden and unsampled phytoplankton, werefrain from citing a detailed list of plank-

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2 EEZs, Marine Regions, and Subregions of the EU Directive on Marine Strategy (2008/56/CE).

Medit. Mar. Sci., 11/2, 2010, 381-493 385

Table 1Subregions, areas and coast of the Mediterranean studied in this work.

Western Central Mediterranean Adriatic Sea Eastern MediterraneanMediterranean including (the Aegean-Levantine -

the Ionian Sea Marmara Seas)

Tyrrhenian Sea Greek Ionian Sea Italian Adriatic Sea Greek Aegean Sea Ligurian Sea Italian Ionian Sea Slovenia Turkish Aegean SeaSardinia Albanian Ionian Sea Croatia Sea of MarmaraCorsica Apulian Sea Montenegro South Turkey France South East Sicily Albanian Adriatic Sea CyprusMonaco Malta SyriaSpain South Tunisia LebanonBaleares Libya Palestine AuthorityGibraltar IsraelMorocco EgyptAlgeriaNorth TunisiaWest Sicily

Fig. 1: European marine and coastal regions(source: SUREZ DE VERO et al., 2009).

tonic and benthic microalgae as inZENETOS et al. (2005, 2008).

The revised checklist has been as wideas possible. Contrary to the CIESM Atlas,we have taken into account alien speciesintroduced from elsewhere within theMediterranean when the introductionevent was evident (e.g. the molluscSiphonaria pectinata introduced from theWMED to Greece, or the algae Cla-dosiphon zosterae, Desmarestia viridis,Ectocarpus siliculosus var. hiemalis andPylaiella littoralis from the Adriatic to theThau Lagoon, WMED).

Freshwater species occurring in estuar-ine waters such as Acipenser gueldenstaedtii,Acipenser baeri and Huso huso in the NorthAegean estuaries and Micropterus salmoidesin Ionian Sea estuarine waters are not includ-ed in our list. Similarly, the freshwater cray-fish Procambarus clarki, though it can standslightly brackish waters and has been report-ed in the inner part of the Varano Lagoon,Adriatic Sea (FLORIO et al., 2008) and inthe Palude di Torre Flavia, Tyrrhenian Sea(SCALICI et al., 2010), is not included.

Alien [synonyms: Non-indigenousspecies (NIS), exotic, non-native,allochthonous] are species, subspecies orlower taxa, present in the wild, introducedoutside of their natural range (past orpresent) and beyond their natural disper-sal potential. This includes any part,gamete or propagule of such species thatmight survive and subsequently repro-duce. Their presence in the given regionis due to intentional or unintentionalintroduction resulting from human activi-ties. Natural shifts in distribution ranges(e.g. due to climate change or dispersal byocean currents) do not qualify a species asa NIS (OLENIN et al., 2010). Specimenskept in captivity and specimens stillattached to a ship’s hull or other man-

made crafts are not considered.The acclimatization status of each

species was assessed, and is given hereaccording to the following terminology.

Established: Introduced or feral popu-lation of species settled in the wild withfree-living, self-maintaining and self-per-petuating populations unsupported by andindependent of humans. Species with atleast two records in the area spread overtime and space (at least three records forfishes) are also classified as established, inthe sense of the CIESM Atlas series.

Casual: Casual species are those hav-ing been recorded only once (no morethan twice for fish) in the scientific andgrey literature and are presumed to benon-established in the area. In this paper‘casual’ is used in the same sense as ‘alien’in the CIESM Atlas series.

Questionable: Species with insuffi-cient information–‘suspect’. This includesold casual records that have not been sub-sequently found despite appropriateinvestigation, and also new entries notverified by experts or species with taxo-nomic status unresolved.

Cryptogenic: Species with no definiteevidence of their native or introduced sta-tus according to CARLTON (1996) andspecies whose probable introductionoccurred in ‘early times’ and has not beenwitnessed (e.g. prior to 1800).

Invasive: Species defined as establishedaliens that have overcome biotic and abiot-ic barriers and are able to disseminate awayfrom their area of initial introductionthrough the production of fertile offspringwith noticeable impact, such as threat to thediversity or abundance of native species, theecological stability of infested ecosystems,economic activities dependent on theseecosystems, and human health.

True aliens need to be separated from

Medit. Mar. Sci., 11/2, 2010, 381-493386

species with seemingly isolated records,which have naturally spread to regionsbeyond their usual range. BOUCHET &TAVIANI (1992) illustrated this scenariowith the case of deep-sea molluscs in theMediterranean, brought into the basin aslarvae carried by the inflowing surfacewaters, but unable to establish fertile pop-ulations. This is also the case of someEastern Atlantic crustacean species sel-dom recorded in the West Mediterraneanbut occurring in the Alboran sea (Penaeop-sis serrata, Merhippolyte ancistrota, etc.),and of the big-eye thresher shark, Alopiassuperciliosus, which is distributed world-wide in tropical and temperate seas andwas formerly classified as alien in theMediterranean (ZENETOS et al., 2008).Occurrences in this kind of context havebeen filtered out of alien species lists.

Regarding the native range, caution hasbeen taken in establishing the origin of alienspecies. The true origin for many species ismuddled after becoming widely dispersedover a long time. Precise localities will beknown for some, but for others only a gen-eral region may be known. This is particu-larly important in the case of the Mediter-ranean where the true origin of populationsof a species widely distributed in the Indo-Pacific Ocean may be its population in theRed Sea, or much further afield. Theexpression "Lessepsian migration" wascoined by POR (1978) for those species thatinched their way through the Suez Canalinto the Mediterranean. However it is farfrom straightforward that all the Indo-Pacif-ic species documented in the Mediterraneanare Lessepsian immigrants, and at least twomore pools of species can be individualized.One such pool would include tropical Indo-Pacific species that did not first show up onthe Levantine or Egyptian coasts, but fur-ther away from the Suez Canal.

Finally, attention was paid to recentnomenclatural updates. These are theresult of the latest taxonomic and/ormolecular studies, such those of LAI et al.(2010) on the Portunus pelagicus speciescomplex; of HUBER (2010) on Anadarakagoshimensis, formerly known as Anadarainaequivalvis; and of MALAQUIAS &REID (2008) on Bulla arabica, formerlyknown as Bulla ampulla. A list of allspecies catalogued under their currentlyvalid names along with the old names isprovided in the ANNEX.

Nomenclature adopted in this paperfollows the World Register of MarineSpecies (WoRMS <www.marinespecies.org>) and contributing databases(AlgaeBase, CLEMAM, FishBase…).

The time boundary is generally set fromthe date of the opening of the Suez Canal(1869): species established earlier are treat-ed as cryptogenic (e.g. Teredo navalis).

Results and Discussion

The core of this work is Table 2, whichprovides a full list of alien species withineach Mediterranean sub-basin, along withtheir acclimatization status and origin. Fur-thermore, this part is built around eightsections. The first four deal in detail withthe distribution of the seven systematicgroups within each MSFD area. They arestructured in a way so as to be autonomousand therefore understood by the independ-ent taxonomist reader. The fifth sectiondescribes the state of art of phytoplanktonwithin the Mediterranean LME. The lastthree sections are more general andattempt to discuss some trends from theelaboration of Table 2. A list of invasive orpotentially invasive species for each basin isincluded. The possible role of warming inthe observed trends is also addressed.

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Table 2List of species with origin and establishment success in all Mediterranean MSFD areas

(WMED=Western Mediterranean, CMED=Central Mediterranean, ADRIA=Adriatic Sea,EMED=Eastern Mediterranean). Species in alphabetic order within each taxon.

Establishment success abbreviated as est=established, cas=casual, cry=cryptogenic, que=questionable.One asterisk denotes planktonic form. Two asterisks denote recent name changes. P= Parasite.

Species Author Origin WMED CMED ADRIA EMEDP R O T O Z O A

Miscellaneous ProtozoaBonamia ostreae (Pichot et al., 1979) Circumboreal cas casMarteilia refringens Cavalier-Smith, 2002 unknown cas casPerkinsus olseni** Lester & Davis, 1981 Cosmopolitan cry cryPhotobacterium damselae - NE Atlantic est ? ? ?ForaminiferaAgglutinella arenata (Said, 1949) Indo-Pacific casAgglutinella compressa El-Nakhal, 1983 Indo-Pacific casAgglutinella robusta El-Nakhal, 1983 Indo-Pacific casAgglutinella soriformis El-Nakhal, 1983 Indo-Pacific/Atlantic casAmphisorus hemprichii Ehrenberg, 1840 Circumtropical estAmphistegina lessonii d'Orbigny, 1826 Circumtropical est? estAmphistegina lobifera Larsen, 1976 Circumtropical est? estAmphistegina d’Orbigny, 1826 Indo-Pacific est estmadagascariensisArchais angulatus (Fichtel & Moll, 1798) W Atlantic cry/casArticulina alticostata Cushman, 1922 Indo-Pacific estAstacolus insolitus (Schwager, 1866) Indo-Pacific estAstacolus sublegumen (Parr, 1950) Indo-Pacific estBorelis sp. Circumtropical estBrizalina simpsoni (Heron-Allen & Earland, 1915) Indo-Pacific casClavulina angularis d' Orbigny, 1826 Circumtropical cry/cas cry/estClavulina cf. multicamerata Chapman, 1907 Indo-Pacific estCoscinospira hemprichii Ehrenberg, 1839 Indo-Pacific cry/cas cry/estCushmanina striatopunctata (Parker & Jones, 1865) Circumtropical casCycloforina sp. Indian/Red Sea estCyclorbiculina compressa (d'Orbigny, 1839 ) Circumtropical estCymbaloporetta plana (Cushman, 1924) Indo-Pacific cry/est cry/cas cry/estEdentostomina cultrata (Brady, 1881) Indo-Pacific estElphidium cf. charlottensis (Vella, 1957) Indo-Pacific estElphidium striatopunctatum (Fichtel & Moll, 1798) Indo-Pacific estEuthymonacha polita (Chapman, 1904) Indo-Pacific casHaddonia sp. Chapman, 1898 Indo-Pacific estHauerina diversa Cushman, 1946 Circumtropical estHeterocyclina tuberculata (Moebius, 1880) Indian est

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDHeterostegina depressa d’Orbigny, 1826 Circumtropical estMiliolinella cf. hybrida (Terquem, 1878) Indo-Pacific estNodophthalmidium (Cushman, 1922) Circumtropical estantillarumOperculina ammonoides (Gronovius, 1781) Red Sea casPegidia lacunata McCulloch, 1977 Indo-Pacific estPeneroplis antillarum d’Orbigny, 1839 Caribbean or SW cas

AtlanticPlanogypsina acervalis (Brady, 1884) Circumtropical cry cry cryPlanogypsina squamiformis (Chapman, 1901) Circumtropical estPlanorbulinella larvata (Parker & Jones, 1865) Indo-Pacific casPseudolachlanella slitella Langer, 1992 Indo-Pacific estPseudomassilina reticulata (Heron-Allen & Earland, 1915) Indo-Pacific estPulleniatina obliquiloculata* (Parker & Jones, 1865) Circumtropical casPyramidulina catesbyi (d'Orbigny, 1839) Indo-Pacific/Atlantic estPyrgo denticulata (Brady, 1844) Indo-Pacific estQuinqueloculina cf. Said, 1949 Indo-Pacific estmosharrafaiSchackoinella imperatoria d’Orbigny, 1846 Indo-Pacific cry/cas cry/cas cry/casSchlumbergerina (Brady, 1879) Circumtropical estalveoliniformisSorites orbiculus Forssk l, 1775 Circumtropical cry/est cry/cas cry/estSorites variabilis Lacroix, 1941 Indo-Pacific estSpiroloculina antillarum d'Orbigny, 1839 Circumtropical estSpiroloculina cf. angulata Cushman, 1917 Indo-Pacific casTriloculina fichteliana d'Orbigny, 1839 Circumtropical est

M A C R O P H Y T E SChlorophytaBatophora sp. Atlantic quesCaulerpa distichophylla Sonder Indo-Pacific est estCaulerpa mexicana Sonder ex Kützing Indo-Pacific estCaulerpa racemosa var. (Sonder) Verlaque, Huisman Indo-Pacific est est est estcylindracea & BoudouresqueCaulerpa racemosa var. (Montagne) Weber-van Indo-Pacific est estlamourouxii f. requienii BosseCaulerpa racemosa var. (J. Agardh) Eubank/ Indo-Pacific cry/ques cry/questurbinata /uvifera (C. Agardh) J. AgardhCaulerpa scalpelliformis (Brown ex Turner) C. Agardh Indo-Pacific estCaulerpa taxifolia (Vahl) C. Agardh Indo-Pacific est est estCladophora cf. patentiramea (Montagne) Kützing Indo-Pacific estCladophora herpestica (Montagne) Kützing Indo-Pacific est

a

Medit. Mar. Sci., 11/2, 2010, 381-493390

Table 2 (continued)

Cladophora hutchinsioides Hoek & Womersley Pacific casCodium fragile subsp. (Suringar) Hariot NW Pacific est est est estfragile**Codium parvulum (Bory ex Audouin) P.C. Silva Indo-Pacific estCodium taylorii P.C. Silva Indo-Pacific est estDerbesia boergesenii (Lyengar & Ramanathan) Indo West Pacific cas

MayhoubDerbesia rhizophora Yamada NW Pacific estNeomeris annulata Dickie Indo-Pacific estUlva fasciata Delile Cosmopolitan est cry/ques cry/ques cry/quesUlva pertusa Kjellman Indo-Pacific estUlvaria obscura (Kützing) Gayral NW Pacific est est estFucophycaeAcrothrix gracilis Kylin Circumboreal estBotryella cf. parva (Takamatsu) H.-S. Kim Pacific estChorda filum (Linnaeus) Stackhouse N Atlantic/N Pacific est estCladosiphon zosterae (J. Agardh) Kylin Atlantic est cry/ques est cry/quesColpomenia peregrina Sauvageau Indo-Pacific est est estDesmarestia viridis (O.F. Müller) J.V. Lamouroux Atlantic/Pacific est cry/quesEctocarpus siliculosus (P.L. Crouan & H.M. Atlantic est cry/quesvar. hiemalis Crouan) FoslieFucus spiralis Linnaeus Atlantic casHalothrix lumbricalis (Kützing) Reinke N Atlantic/N Pacific est cry/ques cry/ques cry/quesLeathesia marina (Lyngbye) Decaisne Cosmopolitan est est estMicrospongium tenuissimum (Hauck) A.F. Peters Atlantic estPadina antillarum (Kützing) Piccone Indo-Pacific casPadina boergesenii Allender & Kraft Indo-Pacific est est estPadina boryana Thivy in W.R. Taylor Indo-Pacific est estPunctaria tenuissima (C. Agardh) Greville NE Atlantic est estPylaiella littoralis (Linnaeus) Kjellman N Atlantic/N Pacific est cry/ques cry/ques cry/quesRugulopteryx okamurae (Dawson) I.K. Hwang, Pacific est

W.J. Lee & H.S. KimSaccharina japonica** (Areschoug) C.E. Lane, NW Pacific cas

C. Mayes, Druehl & G.W. Saunders

Sargassum muticum (Yendo) Fensholt NW Pacific est estScytosiphon dotyi Wynne NE Pacific est estSpatoglossum variabile Figari & De Notaris Indo-Pacific casSphaerotrichia firma (Gepp) A.D.Zinova NW Pacific est estStypopodium schimperi (Kützing) Verlaque Indo West Pacific est est

& Boudouresque

Species Author Origin WMED CMED ADRIA EMED

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Table 2 (continued)

Undaria pinnatifida (Harvey) Suringar Pacific est cas estMagnoliophytaHalophila stipulacea (Forssk l) Ascherson Red Sea est est est estPelagophyceaeChrysonephos lewisii (W.R.Taylor) W.R.Taylor W Atlantic estRhodophytaAcanthophora nayadiformis (Delile) Papenfuss Indo-Pacific cry/ques cry/ques cry/ques cry/quesAcrochaetium codicola B rgesen Atlantic/Pacific est est est estAcrochaetium robustum B rgesen Indo-Pacific casAcrochaetium spathoglossi B rgesen Indo-Pacific casAcrochaetium subseriatum B rgesen Indo-Pacific casAcrothamnion preissii (Sonder) E.M. Wollaston Indo-Pacific est est casAgardhiella subulata (C. Agardh) Kraft & Atlantic/Pacific est est est

M.J. WynneAglaothamnion feldmanniae Halos N Atlantic est estAhnfeltiopsis flabelliformis (Harvey) Masuda Pacific estAnotrichium okamurae Baldock NW Pacific cry/ques cry/ques cry/ques cry/quesAntithamnion amphigeneum A. Millar SW Pacific estAntithamnion hubbsii Dawson Indo-Pacific est estAntithamnionella boergesenii (Cormaci & G. Furnari) Indo-Pacific cry/ques cry/ques

AthanasiadisAntithamnionella elegans (Berthold) J.H. Price & unknown cry/ques cry/ques cry/ques cry/ques

D.M. JohnAntithamnionella (Schiffner) E.M. Wollaston Indo-Pacific est est est Requiresspirographidis confirmationAntithamnionella sublittoralis (Setchell & Gardner) Pacific est est

AthanasiadisAntithamnionella ternifolia (J.D. Hooker & Harvey) Lyle unknown estApoglossum gregarium (E.Y. Dawson) M.J. Wynne Pacific est estAsparagopsis armata Harvey SW Pacific est est est Requires

confirmationAsparagopsis taxiformis Delile Atlantic cry/ques cry/ques cry/quesAsparagopsis taxiformis invasive strain Indo-Pacific est est est estBonnemaisonia hamifera Hariot Indo-Pacific est est est estBotryocladia Feldmann-Mazoyer Indian est est est estmadagascariensisCaulacanthus okamurae** Yamada NW Pacific estCeramium bisporum D.L. Ballantine W Atlantic cry/ques cry/quesCeramium strobiliforme G.W. Lawson & D.M. John N Atlantic cry/ques cry/quesChondria coerulescens (J. Agardh) Falkenberg E Atlantic est cry/ques cry/ques cry/quesChondria curvilineata F.S. Collins & Hervey Atlantic est est

a

Species Author Origin WMED CMED ADRIA EMED

Medit. Mar. Sci., 11/2, 2010, 381-493392

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDChondria pygmaea Garbary & Vandermeulen Indo-Pacific est est estChondrus giganteus f. Mikami Pacific estflabellatusChrysymenia wrightii (Harvey) Yamada Pacific estDasya sessilis Yamada Pacific estDasysiphonia sp. Yendo NW Pacific est est(= Heterosiphonia japonica)Galaxaura rugosa (J. Ellis & Solander) Red Sea est

J.V. LamourouxGanonema farinosum (J.V. Lamouroux) K.C. Fan Indian cry/ques cry/ques cry/ques

& Yung C. WangGoniotrichiopsis sublittoralis G.M. Smith NW Pacific estGracilaria arcuata Zanardini Indo-Pacific/Red Sea Requires Requires

confirmation confirmationGracilaria vermiculophylla (Ohmi) Papenfuss NW Pacific estGrateloupia asiatica Kawaguchi & Wang NW Pacific estGrateloupia lanceolata (Okamura) Kawaguchi Pacific est casGrateloupia minima P.L. Crouan & H.M. Pacific est

CrouanGrateloupia patens (Okamura) S. Kawaguchi Pacific cas

& H.W. WangGrateloupia subpectinata Holmes Pacific estGrateloupia turuturu Yamada NW Pacific est est estGriffithsia corallinoides (Linnaeus) Batters Atlantic/Pacific est est estHerposiphonia parca Setchell Indo-Pacific estHypnea anastomosans** Papenfuss, Lipkin & Silva Red Sea cas estHypnea cornuta (Kützing) J.Agardh Cosmopolitan est estHypnea flagelliformis Greville ex J. Agardh Indo-Pacific Requires

confirmationHypnea flexicaulis Yamagishi & Masuda Indo-Pacific estHypnea spinella (C. Agardh) Kützing Pantropical est est est estHypnea valentiae (Turner) Montagne Red Sea est est cas estLaurencia caduciramulosa Masuda & Kawaguchi SW Pacific est estLaurencia okamurae Yamada Pacific estLithophyllum yessoense Foslie Pacific estLomentaria flaccida Tanaka Pacific casLomentaria hakodatensis Yendo Pacific est estLophocladia lallemandii (Montagne) F. Schmitz Indo-Pacific est est est estNemalion vermiculare Suringar NW Pacific estNeosiphonia harveyi (J. Bailey) M.-S. Kim, H.-G. NW Pacific est est est est

Choi, Guiry & G.W. Saunders

Medit. Mar. Sci., 11/2, 2010, 381-493 393

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDNitophyllum Okamura Pacific eststellato-corticatumOsmundea oederi (Gunnerus) G. Furnari Atlantic cry/ques cry/quesPlocamium secundatum (Kützing) Kützing tropical/subtropical est estPolysiphonia atlantica Kapraun & J.N. Norris N Atlantic/N Pacific cry/ques cry/ques cry/ques cry/quesPolysiphonia fucoides (Hudson) Greville N Atlantic est cry/ques cry/ques cry/quesPolysiphonia morrowii Harvey NW Pacific est est estPolysiphonia paniculata Montagne E Pacific est cas casPolysiphonia stricta (Dillwyn) Greville NE Atlantic estPorphyra yezoensis Ueda Pacific estPterosiphonia tanakae Uwai & Masuda Pacific estRhodophysema georgei Batters Atlantic/Pacific cas casRhodymenia erythraea Zanardini Indo West Pacific casSarconema filiforme (Sonder) Kylin Indo West Pacific cas estSarconema scinaioides B rgesen Indian estSolieria dura (Zanardini) F. Schmitz Indo-Pacific casSolieria filiformis (Kützing) Gabrielson N Atlantic est est estSpongoclonium caribaeum** (B rgesen) M.J. Wynne W Atlantic estSymphyocladia (Harvey) Falkenberg Indo-Pacific estmarchantioidesWomersleyella setacea (Hollenberg) R.E. Norris Indo-Pacific est est est est

P O L Y C H A E T AAmphicorina pectinata (Banse, 1957) Pacific cas casApoprionospio pygmaea** (Hartman, 1955) Pacific casBranchiomma bairdi (McIntosh, 1885) Atlantic/Pacific estBranchiomma boholensis (Grube, 1878) Indo-Pacific est est estBranchiomma luctuosum (Grube, 1869) Indo-Pacific est est est estCapitellethus dispar (Ehlers, 1907) Indo-Pacific/Red Sea quesCeratonereis mirabilis Kinberg, 1866 Indo-Pacific cas estChaetozone corona Berkeley & Berkeley, 1941 unknown cryCirriformia semicincta (Ehlers, 1905) Red Sea quesCossura coasta Litamori, 1960 unknown cry/quesDasybranchus carneus Grube, 1870 Red Sea quesDesdemona ornata Banse, 1957 Indo-Pacific est est estDispio magnus (Day, 1955) Indian quesDispio uncinata Hartman, 1951 W Atlantic cas casDodecaceria capensis Day, 1961 Indian ques quesDorvillea similis (Crossland, 1924) Indo-Pacific estEunice antennata (Savigny, 1820) Indo-Pacific est est estEunice cf. cariboea Grube, 1856 W Atlantic quesEunice floridana (Pourtalès, 1867) W Atlantic ques

Medit. Mar. Sci., 11/2, 2010, 381-493394

Table 2 (continued)

Epidiopatra hupferiana Augener, 1918 Tropical Atlantic cashupferinaEpidiopatra hupferiana Day, 1957 Indian casmonroiEunice indica Kinberg, 1865 Indo-Pacific quesErinaceusyllis serratosetosa (Hartmann-Schroder, 1982) Pacific estEuniphysa tubifex (Crossland, 1904) Indian quesEurythoe complanata (Pallas, 1766) Atlantic/Pacific ques quesEusyllis kupfferi Langerhans, 1879 Atlantic estExogone africana Hartmann-Schröder, 1974 Indo-Pacific casExogone breviantennata Hartmann-Schröder, 1959 Circumtropical estFabriciola qhardaqa Banse, 1959 Red Sea casFicopomatus enigmaticus (Fauvel, 1923) Subtropical est est est estGlycinde bonhourei Gravier, 1904 Indo-Pacific estHaploscoloplos kerguelensis Mc Intosh, 1885 Antarctic quesHesionura serrata (Hartmann-Schroder, 1960) Red Sea casHyboscolex longiseta Schmarda, 1861 Cosmopolitan ques ques quesHydroides albiceps Grube, 1870 Indo Pacific/Red Sea casHydroides branchyacanthus Rioja, 1941 Indo-Pacific estHydroides dianthus (Verrill, 1873) NW Atlantic est est est estHydroides diramphus Mörch, 1863 Circumtropical est est estHydroides elegans (Haswell, 1883) Circumtropical est est est estHydroides heterocerus (Grube, 1868) Indian/Red Sea estHydroides homocerus Pixell, 1913 Indian estHydroides minax (Grube, 1878) Indo-Pacific/Red Sea estHydroides operculatus (Treadwell, 1929) Indian estHydroides steinitzi Ben-Eliahu, 1972 Red Sea cas casJanua (Dexiospira) steueri (Sterzinger, 1909) Indo-Pacific casIsolda pulchella Muller, 1858 Circumtropical casLaonome elegans Gravier, 1906 Red Sea casLaonome triangularis Hutchings & Murray, 1984 SW Pacific estLeiochrides australis Augener, 1914 Pacific cas cas casLeocrates chinensis Kinberg, 1866 Pacific ques ques ques quesLeonnates decipiens Fauvel, 1929 Indo-Pacific estLeonnates indicus Kinberg, 1966 Indo-Pacific estLeonnates persicus Wesenberg-Lund, 1949 Indo-Pacific estLepidonotus carinulatus (Grube, 1870) Indo-Pacific/Red Sea quesLepidonotus tenuisetosus (Gravier, 1902) Indo-Pacific/Red Sea casLinopherus canariensis** Langerhans, 1881 Atlantic est estLongibranchium atlanticum (Day, 1973) Atlantic cas casLoimia medusa (Savigny, 1818) Cosmopolitan ques ques

Species Author Origin WMED CMED ADRIA EMED

Medit. Mar. Sci., 11/2, 2010, 381-493 395

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDLumbrineris acutifrons (Gallardo, 1967) Pacific ques casLumbrineris neogesae Miura, 1980 S Africa cas casLumbrineris perkinsi** Carrera-Parra, 2001 Indo-Pacific est est estLysidice collaris Grube, 1870 Pacific/Red Sea est est est estLysidice natalensis (Kinberg, 1865) Indo-Pacific quesMarphysa disjuncta Hartman, 1961 Pacific casMediomastus capensis Day, 1961 Indian ques ques quesMegalomma claparedei Gravier, 1908 Red Sea casMetasychis gotoi (Izuka, 1902) Indo-Pacific est est est estNaineris quadraticeps Day, 1965 Red Sea quesNeopseudocapitella Rullier & Amoureux, 1979 W Atlantic/ est est est estbrasiliensis Red SeaNeanthes agulhana (Day, 1963) S Africa est estNeanthes willeyi (Day, 1934) Indo-Pacific casNereis gilchristi Day, 1960 Red Sea casNereis jacksoni Kinberg, 1866 Indo-Pacific est estNereis persica Fauvel, 1911 Indo-Pacific ques estNotomastus aberans Day, 1957 Indian/Red Sea est est est estNotomastus mossambicus (Thomassin, 1970) Indian est estNotopygos crinita Grube, 1855 W Atlantic quesNovafabricia infratorquata Fitzhugh, 1983 W Atlantic est estOphryotrocha diadema kesson, 1976 Pacific estOphryotrocha japonica Claparède & Mecznikow, 1968 Pacific est est estOenone cf. fulgida (Savigny, 1818) Indo-Pacific/Red Sea ques quesOnuphis eremita oculata Hartman, 1951 W Atlantic estPalola valida (Gravier, 1900) Red Sea estParadyte cf. crinoidicola (Potts, 1910) Indo-Pacific quesParaehlersia weissmaniodes (Augener,1913) Indo-Pacific casParaprionospio coora Wilson, 1990 Pacific cry cryPerinereis nuntia (Savigny, 1818) Indian casPherusa parmata (Grube, 1878) Indo-Pacific casPherusa saldanha Day, 1961 Indian casPileolaria berkeleyana (Rioja, 1942) E Pacific estPisione guanche San Mart›n, Lo’pez & Tropical Atlantic cas

Nu’n~ez, 1999Pista unibranchia Day, 1963 Indo-Pacific est estPlatynereis australis (Schmarda, 1861) Pacific quesPodarkeopsis capensis (Day, 1963) Indo-Pacific cas quesPolycirrus twisti Potts, 1928 Suez Canal estPolydora colonia Moore 1907 W Atlantic cas casPolydora cornuta Bosc, 1802 Cosmopolitan est est

A

Medit. Mar. Sci., 11/2, 2010, 381-493396

Table 2 (continued)

Polydora spongicola Berkeley & Berkeley, 1950 Pacific quesPomatoleios kraussii (Baird, 1865) Indo-Pacific/Red Sea estPrionospio (Aquilaspio) Fauvel, 1929 Indo West Pacific estkrusadensisPrionospio (Aquilaspio) Augener, 1918 Atlantic/Pacific estsexoculataPrionospio (Prionospio) Imajima, 1990 NE Pacific estdepauperataPrionospio (Prionospio) Mackie & Hartley, 1990 Indo-Pacific estsacciferaPrionospio (Prionospio) Blake & Kudenov, 1978 Pacific estpaucipinnulataPrionospio (Minuspio) Imajima, 1990 Atlantic/Pacific caspulchraProtodorvillea egena (Ehlers, 1913) Indian/Red Sea quesProtodorvillea biarticulata Day, 1963 Indian ques quesProsphaerosyllis (Hartmann-Schröder, 1979) SW Pacific caslongipapillata**Pseudonereis anomala (Gravier, 1900) Indo-Pacific est estPseudopolydora Okuda, 1937 Pacific estpaucibranchiataScoletoma debilis (Grube, 1878) Indo-Pacific quesSigambra constricta (Southern, 1921) Indo-Pacific/Red Sea quesSigambra parva (Day, 1963) Indian casSpirobranchus tetraceros (Schmarda, 1861) Circumtropical estSpirorbis marioni Caullery & Mesnil, 1897 Atlantic/Pacific est estStreblosoma comatus** (Grube, 1856) Indo-Pacific est cas estStreblospio Rice & Levin, 1998 W Atlantic estgynobranchiataSyllis alosae San Mart›n, 1992 W Atlantic quesSyllis bella (Chamberlin, 1919) Pacific estSyllis hyllebergi (Licher, 1999) Red Sea estSyllis cf. mayeri Musco & Giangrande, 2005 W Atlantic quesSyllis pectinans Haswell, 1920 Pacific est estSynelmis rigida (Fauvel, 1919) Indo-Pacific quesSyllis schulzi (Hartmann-Schröder, 1962) Indian/Red Sea ques quesTerebella ehrenbergi Grube, 1870 Indo-Pacific/Red Sea ques quesTimarete anchylochaeta (Schmarda, 1861) Pacific quesTimarete caribous (Grube, 1859) W Atlantic casTimarete dasylophius (Marenzeller, 1879) Indo-Pacific quesTimarete punctata (Grube, 1859) Indo-Pacific est

Species Author Origin WMED CMED ADRIA EMED

Medit. Mar. Sci., 11/2, 2010, 381-493 397

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDCRUSTACEA

AmphipodaBemlos leptocheirus (Walker, 1909) Indian casCaprella scaura Templeton, 1836 Indian est est estCymadusa filosa (Savigny, 1816) Indo-Pacific cry cryElasmopus pectenicrus (Bate, 1862) Circumtropical cas? est est estGammaropsis togoensis (Schellenberg, 1925) Cosmopolitan cry/estLinguimaera caesaris** Krapp-Schickel, 2003 Indo-Pacific est estMonocorophium sextonae (Crawford, 1937) unknown cry cry cryParhyale explorator Arresti, 1989 NE Atlantic casPhotis lamellifera Schellenberg, 1928 Indian casRhabdosoma whitei Bate, 1862 Red Sea casStenothoe gallensis Walker, 1904 Circumtropical ques est estCirripediaAmphibalanus eburneus** (Gould, 1841) W Atlantic est est est estAustrominius modestus** Darwin, 1854 Tropical Pacific est estBalanus reticulatus Utinomi, 1967 Circumtropical ques estBalanus trigonus Darwin, 1854 Circumtropical est est est estHeterosaccus dollfusip Boschma, 1960 Red Sea estMegabalanus tintinnabulum (Linnaeus, 1758) Circumtropical ques ques estTetraclita squamosa Pilsbry, 1916 Indo-Pacific casrufotintaCopepodaAcartia centrura Giesbrecht, 1889 Indian casAcartia fossae Gurney, 1927 Indo-Pacific estAcartia tonsa Dana, 1849 W Atlantic / est est est est

Indo-PacificArietellus pavoninus (G. O. Sars, 1905) Tropical-subtropical cas estCalanopia biloba Bowman, 1957 W subtropical Atlantic casCalanopia elliptica (Dana, 1846) Indo-Pacific ques estCalanopia media Gurney, 1927 Indo-Pacific estCalanopia minor A. Scott, 1902 Indo-Pacific casCanuellina insignis Gurney, 1927 Indo-Pacific quesCentropages furcatus (Dana, 1852) Indo-Pacific/Atlantic estClavellisa ilishaep Pillai, 1962 Indian estEnhydrosoma vicinum Por, 1967 Indo-Pacific quesEuchaeta concinna Dana, 1849 Indo-Pacific cas casLabidocera agilis (Dana, 1849) Indo-Pacific casLabidocera detruncata (Dana, 1849) Indo-Pacific ques casLabidocera madurae (A. Scott, 1909) Indo-Pacific estLabidocera orsinii Giesbrecht, 1889 Red Sea cas

Medit. Mar. Sci., 11/2, 2010, 381-493398

Table 2 (continued)

Labidocera pavo Giesbrecht, 1889 Indo-Pacific estMetacalanus acutioperculum Ohtsuka, 1984 Pacific est estMitrapus oblongusp (Pillai, 1964) Indian estMyicola ostreaep Hoshina & Sigiura, 1953 Pacific estMytilicola orientalisp Mori, 1935 Pacific estNothobomolochus fradeip Marques, 1965 Atlantic/Indian estParacalanus indicus Wolfenden, 1905 Subtropical Atlantic, cry/ques

Red SeaParacartia grani G. O. Sars, 1904 Atlantic est est est estParvocalanus crassirostris** Dahl, 1894 Indo-Pacific/Atlantic cas estParvocalanus elegans Andronov, 1972 Indo-Pacific casParvocalanus latus Andronov, 1972 Indian casPseudocyclops xiphophorusp Wells, 1967 Indian cryPseudocalanus elongatus (Boeck, 1865) E Atlantic cry cry cryRobertsonia salsa Gurney, 1927 Indo-Pacific quesScaphocalanus amplius Park, 1970 Subtropical Atlantic casScaphocalanus brevirostris Park, 1970 Subtropical Atlantic/ cas

IndianScolecithrix valens Farran, 1926 Subtropical Atlantic casScottolana longipes (Thompson & Scott, 1903) Indo-Pacific quesSpinocalanus terranovae Damkaer, 1975 Antarctic ques quesStenhelia inopinata (A. Scott, 1902) Indo-Pacific quesStenhelia minuta (A. Scott, 1902) Indo-Pacific quesSubeucalanus subcrassus** Giesbrecht, 1888 Atlantic/Pacific ques ques quesTriconia hawii (Böttger-Schnack Red Sea est

& Boxshall, 1990)Triconia rufa (Boxshall & Böttger, 1987) Indian/Red Sea est quesTriconia umerus (Böttger-Schnack Red Sea est cas cas

& Boxshall, 1990)CumaceaEocuma rosae Corbera & Galil, 2007 Indo-Pacific/Red Sea casEocuma sarsii (Kossmann, 1880) Indo-Pacific est estScherocumella gurneyi (Calman, 1927) Red Sea casDecapodaActumnus globulus Heller, 1861 Red Sea casAlpheus audouini Coutiere, 1905 Indo West Pacific estAlpheus inopinatus Holthuis & Gottlieb, 1958 Indian/Red Sea cas estAlpheus migrans Lewinsohn & Holthuis, 1978 Red Sea estAlpheus rapacida de Man, 1908 Indo West Pacific estAshtoret lunaris (Forssk l, 1775) Indo West Pacific casAtergatis roseus (Rüppell, 1830) Indo-Pacific est

a

Species Author Origin WMED CMED ADRIA EMED

Medit. Mar. Sci., 11/2, 2010, 381-493 399

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDCalappa hepatica (Linnaeus, 1758) Indo-Pacific casCalappa pelii Herklots, 1851 E Atlantic cas cas casCallinectes danae Smith, 1869 W Atlantic casCallinectes sapidus Rathbun, 1896 W Atlantic cas cas est estCarupa tenuipes Dana, 1851 Indo-Pacific estCharybdis feriata (Linnaeus, 1758) Indo-Pacific casCharybdis japonica (A. Milne-Edwards, 1861) Indo-Pacific casCharybdis helleri (A. Milne-Edwards, 1867) Indo West Pacific estCharybdis longicollis Leene, 1938 Indian/Red Sea estCharybdis lucifera (Fabricius, 1798) Indo-Pacific casColeusia signata (Paulson, 1875) Indo West Pacific estCryptosoma cristatum Brulle, 1837 Tropical East Atlantic casDaira perlata (Herbst, 1790) Indo West Pacific casDorippe quadridens (Fabricius, 1793) Indo West Pacific estDyspanopeus sayi (Smith, 1869) N East Atlantic estEriocheir sinensis H. Milne Edwards, 1853 Pacific cas casEucrate crenata de Haan, 1835 Indo-Pacific est estEurycarcinus integrifrons de Man, 1879 Indian/Red Sea casFarfantepenaeus aztecus (Ives, 1891) W Atlantic estFenneropenaeus merguiensis (De Man, 1888) Indo West Pacific casGlabropilumnus laevis** (Dana, 1852) Indian cas casGonioinfradens paucidentata (A. Milne Edwards, 1861) Indo-Pacific est?Grapsus granulosus H. Milne-Edwards, 1853 Red Sea casHalimede tyche (Herbst, 1801) Indo West Pacific casHemigrapsus sanguineus (de Haan, 1835) Pacific cas casHerbstia nitida Manning & Holthuis, 1981 Tropical East Atlantic casHyastenus hilgendorfi de Man, 1887 Indo West Pacific estIxa monodi Holthuis & Gottlieb, 1956 Red Sea estLeptochela aculeocaudata Paulson, 1875 Indo West Pacific estLeptochela pugnax de Man, 1916 Indo West Pacific estLibinia dubia H. Milne Edwards, 1834 W Atlantic estLucifer hanseni Nobili, 1905 Indo West Pacific quesMacrophthalmus graeffei A. Milne Ewdards, 1873 Indo West Pacific estMarsupenaeus japonicus (Bate, 1888) Indo-Pacific cas cas cas estMelicertus hathor (Burkenroad, 1959) Indian estMenaethius monoceros (Latreille, 1825) Indo-Pacific casMetapenaeopsis aegyptia Galil & Golani, 1990 Indo-Pacific estMetapenaeopsis mogiensis (Nobili, 1904) Indo West Pacific estconsobrinaMetapenaeus affinis (H. Milne Edwards, 1837) Indo West Pacific estMetapenaeus monoceros (Fabricius, 1798) Indo West Pacific est est

Medit. Mar. Sci., 11/2, 2010, 381-493400

Table 2 (continued)

Metapenaeus stebbingi (Nobili, 1904) Indian est estMicippa thalia (Herbst, 1803) Indo West Pacific estMyra subgranulata Kossmann, 1877 Indian/Red Sea estNecora puber (Linnaeus, 1767) NW Atlantic est est casNotopus dorsipes (Linnaeus, 1758) Indo West Pacific casOgyrides mjoebergi (Balss, 1921) Indo West Pacific estPalaemonella rotumana (Borradaile, 1898) Indo West Pacific estPalinurus ornatus (Fabricius, 1798) Indo West Pacific casParalithodes camtschaticus (Tilesius, 1815) NE Pacific casPenaeus semisulcatus de Haan, 1844 Indo West Pacific estPercnon gibbesi (H. Milne Edwards, 1853) W Atlantic est est cas estPericlimenes calmani Tattersall, 1921 Indo-Pacific casPilumnopeus vauquelini (Audouin, 1826) Indian/Red Sea cas est estPilumnus minutus** de Haan, 1835 Indo-Pacific cas estPlagusia squamosa (Herbst, 1790) Indo West Pacific ques est cas casPortunus segnis** (Forssk l, 1775) Indian cas est estProcessa macrodactyla Holthuis, 1952 Tropical East Atlantic est? casRhithropanopeus harrisii (Gould, 1841) Noth Atlantic cas? est estRimapenaeus similis (Smith, 1885) W Atlantic estScyllarus caparti Holthuis, 1952 Tropical East Atlantic casScyllarus posteli Forest, 1963 Temperate-tropical cas

East AtlanticSirpus monodi Gordon, 1953 Tropical East Atlantic cas casSolenocera crassicornis (H.Milne Edwards, 1837) Indo West Pacific casSphaerozius nitidus Stimpson, 1858 Indo West Pacific casSternodromia spinirostris** (Miers, 1881) Tropical East Atlantic casSynalpheus tumidomanus Crosnier & Forest, 1966 Tropical East Atlantic cry cryafricanus**Thalamita gloriensis Crosnier, 1962 Indo West Pacific casThalamita indistincta Apel & Spiridonov, 1998 Indian casThalamita poissonii (Audouin, 1826) Indo West Pacific cry cryTrachysalambria (Steinitz, 1932) Red Sea est estpalaestinensisUrocaridella pulchella Yokes & Galil, 2006 Indo-Pacific estIsopodaAnilocra pilchardip Bariche & Trilles, 2006 Indo-Pacific casApanthura sandalensis Stebbing, 1900 S Africa est estCymothoa indicap Schioedte & Meinert, 1884 Indo-Pacific casMesanthura spp. Tropical/subtropical ques ques quesParacerceis sculpta (Holmes, 1904) Subtropical est est est estParadella dianae (Menzies, 1962) NE Pacific est est? est

a

Species Author Origin WMED CMED ADRIA EMED

Medit. Mar. Sci., 11/2, 2010, 381-493 401

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDSphaeroma venustissimum Monod, 1931 Tropical/subtropical est

E AtlanticSphaeroma walkeri Stebbing, 1905 Indian est est estStomatopodaClorida albolitura Ahyong & Naiyanetr, 2000 Indo West Pacific estErugosquilla massavensis (Kossmann, 1880) Indian/Red Sea est estErythrosquilla sp. (postlarvae) Indo-Pacific casTanaidaceaZeuxo (Parazeuxo) coralensis Sieg, 1980 Circumtropical est

M O L L U S C ABivalviaAcar plicata (Dillwyn, 1817) Indo-Pacific/Red Sea casAfrocardium richardi (Audouin, 1826) Red Sea estAlectryonella crenulifera (Sowerby, 1871) Pacific/Red Sea quesAnadara granosa (Linnaeus, 1758) Indo West Pacific casAnadara inflata (L. A. Reeve, 1844) NW Pacific casAnadara kagoshimensis** (Tokunaga, 1906) Temperate North est est est

PacificAnadara natalensis (Krauss, 1848) W Pacific/Red Sea estAnadara transversa** (Say, 1822) W Atlantic est est est estAngulus flacca (Roemer, 1871) Indo Pacific/Red Sea cas Antigona lamellaris Schumacher, 1817 Indo-Pacific/Red Sea casAtactodea glabrata (Gmelin, 1791) Indo Pacific/Red Sea cas casBrachidontes pharaonis (Fischer, 1870) Indian/Red Sea est est est estCallista florida (Lamarck, 1818) Indian/Red Sea cas casCardites akabana (Sturany, 1899) Red Sea casChama asperella Lamarck, 1819 Indo-Pacific/Red Sea casChama aspersa Reeve, 1846 Indo-Pacific estChama pacifica Broderip, 1834 Indo-Pacific estCirce scripta (Linnaeus, 1758) Indo West Pacific casCircenita callipyga (von Born, 1778) Red Sea casClementia papyracea (Gray, 1825) Indo-Pacific/Red Sea estCrassostrea gigas (Thunberg, 1793) NW Pacific est est est estDendrostrea frons (Linnaeus, 1758) Indo Pacific/Red Sea estDiplodonta bogii** Van Aartsen, 2004 Red Sea estDivalinga arabica Dekker & Gould, 1994 Persian Gulf/Red Sea casDosinia erythraea Römer, 1860 W Pacific/Red Sea estElectroma vexillum (Reeve, 1857) Indian/Red Sea estEnsiculus cultellus (Linnaeus, 1758) Indo West Pacific casFulvia australis (Sowerby G.B., 1834) Indo-Pacific/Red Sea casFulvia fragilis (Forssk l, 1775) Indian est est esta

Medit. Mar. Sci., 11/2, 2010, 381-493402

Table 2 (continued)

Gafrarium pectinatum (Linnaeus, 1758) Indo-Pacific/Red Sea estGlycymeris arabicus (Adams H., 1871) W Pacific/Red Sea casHiatula ruppelliana (Reeve, 1857) Red Sea casLaternula anatina (Linnaeus, 1758) Indo-Pacific/Red Sea estLimopsis multistriata (Forssk l, 1775) Indo-Pacific/Red Sea casMactra lilacea Lamarck, 1818 Indo-Pacific/Red Sea estMactra olorina Philippi, 1846 Red Sea estMalvufundus regula (Forssk l, 1775) Indo-Pacific/Red Sea est estMercenaria mercenaria (Linnaeus, 1758) W Atlantic est estModiolus auriculatus (Krauss, 1848) Indian/Red Sea estMusculista perfragilis (Dunker, 1857) Indo-Pacific/Red Sea estMusculista senhousia (Benson in Cantor, 1842) Temperate North est est est est

PacificMya arenaria Linnaeus, 1758 N Atlantic est est estMytilopsis sallei (Recluz, 1849) W Atlantic estNanostrea exigua Harry, 1985 Indo-Pacific estPaphia textile (Gmelin, 1791) Indo-Pacific/Red Sea estPedicirce sulcata (Gray, 1838) Red Sea quesPetricola hemprichi (Issel, 1869) Indo-Pacific estPetricola pholadiformis Lamarck, 1818 W Atlantic estPinctada margaritifera (Linnaeus, 1758) Indo-Pacific/Red Sea quesPinctada radiata (Leach, 1814) Indo-Pacific/Red Sea est est cas estPsammotreta praerupta (Salisbury, 1934) Indo-Pacific/Red Sea casPseudochama corbieri (Jonas, 1846) Red Sea estRuditapes philippinarum (Adams & Reeve, 1850) Temperate North est est est est

PacificSaccostrea commercialis (Iredale & Roughley, 1933) Australia cas Saccostrea cucullata (Born, 1778) Indo-Pacific ques estSeptifer bilocularis (Linnaeus, 1758) Indo-Pacific casSeptifer forskali Dunker, 1855 Red Sea estSphenia rueppelli A. Adams, 1850 Indian estSpondylus groschi Lamprell & Kilburn, 1995 Indian/Red Sea quesSpondylus cf. multisetosus Reeve, 1856 Indo-Pacific quesSpondylus nicobaricus Schreibers, 1793 Indo-Pacific casSpondylus spinosus Schreibers, 1793 Indo-Pacific/Red Sea estTellina valtonis Hanley, 1844 Indian/Red Sea estTeredo navalis Linnaeus, 1758 Circumtropical cry cryTheora lubrica Gould, 1861 Indo-Pacific est estTimoclea marica (Linnaeus, 1758) Red Sea casTrapezium oblongum (Linnaeus, 1758) Indo-Pacific/Red Sea quesXenostrobus securis (Lamarck, 1819) Tropical Atlantic est est

a

a

Species Author Origin WMED CMED ADRIA EMED

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Table 2 (continued)

Zygochlamys patagonica** (King & Broderip, 1832) W Atlantic cas casCephalopodaOctopus aegina Gray, 1849 Indo-Pacific/Red Sea estOctopus cyanea Gray, 1849 Indo-Pacific casSepia gibba Ehrenberg, 1831 Red Sea casSepia pharaonis Ehrenberg, 1831 Pacific/Red Sea quesSepioeuthis lessoniana Lesson, 1830 Indo-Pacific estTremoctopus gracilis (Eydoux & Souleyet, 1852) Indo-Pacific cas casGastropodaActeocina crithodes Melvill & Standen, 1907 Indo-Pacific/W Indian casActeocina mucronata (Philippi, 1849) Red Sea ques estAlvania dorbignyi (Audouin, 1826) Cosmopolitan cry cry cryAmathina tricarinata (Linnaeus, 1767) Red Sea estAngiola punctostriata (Smith E.A., 1872) Red Sea quesAnteaeolidiella foulisi** (Angas, 1864) Circumtropical cas casAplysia dactylomela Rang, 1828 Circumtropical est est estAplysia parvula Guilding in Mörch, 1863 Circumtropical cry cry cryAtys angustatus Smith, 1872 Red Sea cryAtys cylindricus (Helbling, 1779) Indo-Pacific/Red Sea casBostrycapulus odites** Collin, 2005 Tropical Atlantic estBulla arabica ** Malaquias & Reid, 2008 Indo-Pacific estBursatella leachii De Blainville, 1817 Circumtropical est est est estCaloria indica (Bergh, 1896) Indo-Pacific casCanarium mutabile** (Swainson, 1821) Indo-Pacific/Red Sea quesCantharus tranquebaricus (Gmelin, 1791) Indian casCellana rota (Gmelin, 1791) Indian/Red Sea cas cas estCerithidium diplax** (Watson, 1886) Persian Gulf estCerithidium perparvulum** (Watson, 1886) Pacific estCerithiopsis pulvis (Issel, 1869) Red Sea estCerithiopsis tenthrenois (Melvill, 1896) Indian estCerithium columna Sowerby 1834 Indo West Pacific quesCerithium egenum Gould, 1849 Indo-Pacific/Red Sea estCerithium litteratum (Born, 1778) W Atlantic casCerithium nesioticum Pilsbry & Vannata, 1906 Indian/Red Sea estCerithium nodulosum Bruguière, 1792 Indo West Pacific quesCerithium scabridum Philippi, 1848 Indian/Red Sea est est estChelidonura fulvipunctata Baba, 1938 Indo-Pacific est estChromodoris annulata (Eliot, 1904) Indian estChromodoris quadricolor (Rüppell & Leuckart, 1828) Red Sea cas cas estChrysallida fischeri Hornung & Mermod, 1925) Red Sea est estChrysallida maiae (Hornung & Mermod, 1924) Red Sea est

Species Author Origin WMED CMED ADRIA EMED

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Table 2 (continued)

Chrysallida micronana Öztürk & van Aartsen, 2006 Red Sea casChrysallida pirintella (Melvill, 1910) Red Sea estCingulina isseli (Tryon, 1886) Subtropical estClypeomorus bifasciatus (Sowerby G.B. II, 1855) Indo-Pacific/Red Sea cas estConomurex persicus** (Swainson, 1821) Persian Gulf est cas estConus arenatus Hwass, 1792 Indo-Pacific/Red Sea casConus fumigatus Hwass, 1792 Red Sea casConus inscriptus Reeve, 1843 Indian casConus rattus Hwass, 1792 Indo-Pacific casCoralliobia madreporarum (Sowerby, 1832) Pantropical cas casCrepidula fornicata (Linnaeus, 1758) NW Atlantic est cas estCuthona perca (Marcus, 1958) Tropical Atlantic casCycloscala hyalina (Sowerby, 1844) Indo-Pacific/Red Sea estCylichnina girardi (Audouin, 1826) Indo-Pacific estDendrodoris fumata (Rüppell & Leuckart, 1830) Indo Pacific/Red Sea casDiala semistriata (Philippi, 1849) Indo-Pacific/Red Sea estDiodora funiculata (Reeve, 1850) Indo-Pacific estDiodora ruppellii (Sowerby, 1834) Indo-Pacific/Red Sea cas estDiscodoris lilacina (Gould, 1852) Indo-Pacific/Red Sea cry cryDoxander vittatus** (Linnaeus, 1758) Indo-Pacific casEchinolittorina punctata (Gmelin, 1791) Tropical Atlantic Native est NativeElysia grandifolia Kelaart, 1858 Indian estElysia tomentosa Jensen, 1997 Indo West Pacific estEngina mendicaria (Linnaeus, 1758) Indo West Pacific casErgalatax contracta (Reeve, 1846) Red Sea casErgalatax junionae** Houart, 2008 Persian Gulf/Red Sea estErosaria turdus (Lamarck, 1810) Indian/Red Sea est estEthminolia hemprichi (Issel, 1869) Red Sea casFavorinus ghanensis Edmunds, 1968 Tropical Atlantic est Finella pupoides A. Adams, 1860 Indo-Pacific estFlabellina rubrolineata (O’Donoghue, 1929) Indo-Pacific/Red Sea estFusinus verrucosus (Gmelin, 1791) Indian estGastrochaena cymbium (Spengler, 1783) Indo-Pacific/Red Sea estGibborissoa virgata (Philippi, 1849) Indo-Pacific estGibbula albida (Gmelin, 1791) Adriatic Sea est NativeGodiva quadricolor (Barnard, 1927) Eastern Pacific casHalgerda willeyi Elliot 1904 Indo West Pacific casHaliotis pustulata cruenta Reeve, 1846 Indian/Red Sea cas casHaminoea cyanomarginata Heller & Thompson, 1983 Red Sea est estHaminoea japonica** (Pilsbry, 1895) Pacific estHinemoa cylindrica (de Folin, 1879) Indo-Pacific cas

Species Author Origin WMED CMED ADRIA EMED

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Table 2 (continued)

Hypselodoris infucata (Rüppell & Leuckart, 1828) Indo-Pacific/Red Sea estIolaea neofelixoides (Nomura, 1936) Pacific casLeucotina cfr eva Thiele, 1935 Indo-Pacific casLeucotina natalensis** Smith, 1910 Indo-Pacific/Red Sea estLienardia mighelsi Iredale & Tomlin, 1917 Pacific casLittorina saxatilis (Olivi, 1792) Atlantic cryMelibe viridis** (Kelaart, 1858) Indo-Pacific est est est estMetaxia bacillum (Issel, 1869) Red Sea estMitrella psilla (Duclos, 1846) Tropical Atlantic estMonetaria annulus (Linnaeus, 1758) Indo-Pacific quesMonotigma lauta** (A. Adams, 1853) Indo-Pacific/Red Sea estMurchisonella columna (Hedley, 1907) Indo-Pacific casMurex forskoehlii Roeding, 1798 Persian Gulf/Red Sea estNassa situla (Reeve, 1846) Indo-Pacific casNassarius arcularia plicatus (Roeding, 1798) Indian/Red Sea casNassarius concinnus (Powys, 1835) unknown casNassarius stolatus (Gmelin, 1791) Indo-Pacific casNerita sanguinolenta Menke, 1829 Red Sea cas estNotocochlis gualteriana** (Recluz, 1844) Indo-Pacific/Red Sea cas estOdostomia lorioli (Hornung & Mermod, 1924) Red Sea casOscilla jocosa Melvill, 1904 Persian Gulf casOxynoe viridis (Pease, 1861) Indo West Pacific estPalmadusta lentiginosa (Gray, 1825) Indian caslentiginosaParviturbo dibellai Buzzurro & Celalupo, 2006 unknown cryPatelloida saccharina (Linnaeus, 1758) Pacific casPhilinopsis cyanea (Martens, 1879) Indian/Red Sea casPlanaxis griseus (Brocchi, 1821) Red Sea casPleurobranchus forskalii Rüppell & Leuckart, 1828 Red Sea casPlocamopherus ocellatus Rüppell & Leuckart, 1830 Red Sea estPolycera hedgpethi Marcus Er., 1964 NE Pacific cas casPolycerella emertoni Verrill, 1881 Pantropical est estPseudominolia nedyma Melvill, 1897 Persian Gulf/Red Sea estPurpuradusta gracilis notata (Gill, 1858) Indian/Red Sea estPyrunculus fourierii (Audouin, 1826) Indo-Pacific/Red Sea estRapana rapiformis (Von Born, 1778) Indo-Pacific/Red Sea quesRapana venosa (Valenciennes, 1846) Temperate North est est est

PacificRetusa desgenettii (Audouin, 1826) Red Sea casRhinoclavis kochi (Philippi, 1848) Indo-Pacific/Red Sea estRhinoclavis sinensis (Gmelin, 1791) Indo West Pacific cas

Species Author Origin WMED CMED ADRIA EMED

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDRissoina ambigua (Gould, 1849) Indo-Pacific casRissoina bertholleti Issel, 1869 Indian/Red Sea estRissoina spirata Sowerby, 1825 Indo-Pacific cas casSabia conica (Schumacher, 1817) Indo-Pacific/Red Sea ques quesSiphonaria crenata de Blainville, 1827 Persian Gulf/ est

Arabian SeaSiphonaria pectinata (Linnaeus, 1758) Atlanto-Mediterranean Native estSmaragdia souverbiana (Montrouzier, 1863) Indo-Pacific/Red Sea estSticteulima cf. lentiginosa (A. Adams, 1861) Indo-Pacific casStomatella impertusa (Burrow, 1815) Indo-Pacific/Red Sea casSyphonota geographica (Adams & Reeve, 1850) Circumtropical est estSyrnola cinctella A. Adams, 1860 Indo-Pacific/Red Sea casSyrnola fasciata (Jickeli, 1882) Indo-Pacific estSyrnola lendix** (A. Adams, 1863) Red Sea estThais lacera (von Born, 1778) Indian cas estThais sacellum (Gmelin, 1791) Indian/Red Sea estTornus jullieni Adam & Knudsen 1969 W Africa quesTrivirostra triticum Schilder, 1932 Indo-Pacific casTrochus erithreus Brocchi, 1821 Persian Gulf estTurbonilla edgarii (Melvill, 1896) Indo-Pacific estVexillum depexum (Deshayes in Laborde, 1834) Indian/Red Sea ques Voorwindia tiberiana (Issel, 1869) Red Sea casZafra savignyi (Moazzo, 1939) Red Sea estZafra selasphora (Melvill & Standen, 1901) Indian/Red Sea estPolyplacophoraAcanthopleura gemmata (de Blainville, 1825) Indo-Pacific/Red Sea casChiton hululensis (Smith E.E. in Gardiner, 1903) Indo-Pacific/Red Sea ques

M I S C E L L A N E A I N V E R T E B R A T AArthropoda/PycnogonidaAmmothea hilgendorfi (Böhm, 1879) Circumboreal estAnoplodactylus californicus Hall, 1912 Circum(sub)tropical est estAnoplodactylus digitatus (Böhm, 1879) Circum(sub)tropical estPigrogromitus timsanus Calman, 1927 Circum(sub)tropical quesBryozoaAeverrillia setigera (Hincks, 1887) Circumtropical casArachnoidea protecta (Harmer, 1915) Indo-Pacific quesBowerbankia gracillima (Hincks, 1877) E Atlantic cas casBugula fulva Ryland, 1960 NW Atlantic est cas cas casBugula serrata (Lamarck, 1816) Indo-Pacific estCelleporaria aperta (Hincks, 1882) Circumtropical ques estCelleporaria brunnea (Hincks, 1884) NE Pacific est

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDCelleporaria pilaefera (Canu & Bassler, 1929) Indo-W Pacific quesCelleporella carolinensis Ryland, 1979 W Atlantic estCrepidacantha poissoni (Audouin, 1826) Circumtropical quesElectra tenella (Hincks, 1880) Circumtropical casHippaliosina acutirostris Canu & Bassler, 1929 Indo-Pacific quesHippopodina feejeensis (Busk, 1884) Indo-Pacific casParasmittina egyptiaca (Waters, 1909) Indian estParasmittina serruloides Harmelin, Bitar & Indo-Pacific est

Zibrowius, 2009Parasmittina spondylicola Harmelin, Bitar & Indo-Pacific cas

Zibrowius, 2009Pherusella brevituba Soule, 1951 NE Pacific cas casReteporella jermanensis (Waters, 1909) Red Sea casRhynchozoon lareyi (Audouin, 1826) Indo-Pacific estSchizoretepora hassi Harmelin, Bitar & Red Sea? ques

Zibrowius, 2007Scrupocellaria jolloisii (Audouin, 1826) Indo-Pacific estSmittina malleolus (Hincks, 1884) Circumtropical estTricellaria inopinata d'Hondt & Occhipinti, 1985 Indo-Pacific est estChaetognathaAidanosagitta neglecta* Aida, 1897 Indo-Pacific estFerosagitta galerita* (Dallot, 1971) Indian estChordata/AscidiaceaAscidia cannelata Oken, 1820 Indo-Pacific estBotrylloides violaceus Oka, 1927 NW Pacific estCystodytes philippinensis Herdman, 1886 Circumtropical cry cryDistaplia bermudensis Van Name, 1902 W Atlantic est estEcteinascidia styeloides (Traustedt, 1882) Circumtropical est estEcteinascidia thurstoni Herdman, 1890 Indo-Pacific estHerdmania momus (Savigny 1816) Indo-Pacific estMicrocosmus exasperatus Heller, 1878 Indo-Pacific est estMicrocosmus squamiger Hartmeyer & Michaelsen, 1928 Circumtropical est estPerophora multiclathtrata (Sluiter, 1904) Circumtropical estPhallusia nigra Savigny, 1816 Circumtropical estPolyandrocarpa zorritensis (Van Name, 1931) E Pacific est estRhodosoma turcicum (Savigny, 1816) Circumtropical cas cas estStyela clava Herdman, 1881 NW Pacific casSymplegma brakenhielmi (Michaelsen, 1904) Indo-Pacific estTrididemnum cf. savignii (Herdman, 1886) Circum(sub)tropical est casCnidaria/AnthozoaAcabaria erythraea (Ehrenberg, 1834) Indo-Pacific est

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDDiadumene cincta Stephenson, 1925 NE Atlantic est Haliplanella lineata (Verrill, 1870) Circumboreal estOculina patagonica De Angelis, 1908 ?SW Atlantic est estCnidaria/HydrozoaAequorea conica* Browne, 1905 Indo-Pacific casAmphogona pusilla* Hartlaub, 1909 Indo-Pacific casArctapodema australis* Antarctic/Indo-Pacific cas casBougainvillia niobe Mayer, 1894 (sub)tropical est

W AtlanticCampalecium medusiferum* Torrey, 1902 E Pacific/ Atlantic est casCirrholovenia tetranema* Kramp, 1959 Circumtropical est casClytia hummelincki* (Leloup, 1935) Circumtropical est est estClytia linearis** (Thornely, 1900) Circum(sub)tropical est est est estClytia mccradyi* (Brooks, 1888) Circumtropical est estCordylophora caspia (Pallas, 1771) Circumglobal cry cry cry cryCoryne eximia* Allman, 1859 Circum(sub)tropical ques quesDiphasia digitalis (Busk 1852) Circumtropical casDynamena quadridentata (Ellis & Solander, 1786) Circumtropical estEirene viridula* (Péron & Lesueur, 1810) Circumtropical est est est estEucheilota paradoxica* Mayer, 1900 Circumtropical cas cas cas estEudendrium carneum Clarke, 1882 Circumtropical est cas estEudendrium merulum Watson, 1985 Circumtropical est cas estEuphysora annulata* Kramp, 1928 Indo-Pacific cas casEuphysora bigelowi* Maas, 1905 Indo-Pacific casFilellum serratum* (Clarke, 1879) Circum(sub)tropical casGarveia franciscana (Torrey, 1902) Circum(sub)tropical cas estGonionemus vertens* A. Agassiz, 1862 Circumboreal est estKantiella enigmatica* Bouillon, 1978 Indian casLaodicea fijiana* Agassiz & Mayer, 1899 Indo-Pacific casMacrorhynchia philippina (Kirchenpauer, 1872) Circumtropical estMoerisia carina* Bouillon, 1978 Indo-Pacific estMoerisia inkermanica* Paltschikowa-Ostroumova, 1925 Circum(sub)tropical cas casNubiella mitra* Bouillon, 1980 SW Pacific casOctotiara russelli* Kramp, 1953 Indo-Pacific casOlindias singularis* Browne, 1905 Indian estParacytaeis octona* Bouillon, 1978 W Indian casRussellia mirabilis Kramp, 1957 Antarctic casScolionema suvaensis* (Agassiz & Mayer, 1899) Indo-Pacific estSertularia marginata (Kirchenpauer, 1864) Circum(sub)tropical est estSertularia thecocarpa (Jarvis, 1922) Indo-Pacific estTetrorchis erythrogaster* Bigelow, 1909 Circumtropical est

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDTrichydra pudica* Wright, 1858 Circumtropical cas casCnidaria/ScyphozoaCassiopea andromeda (Forssk l, 1775) Indo-Pacific est? estMarivagia stellata Galil & Gershwin, 2010 ?Indo-Pacific est

CircumtropicalPhyllorhiza punctata* von Lendenfeld, 1884 Circumtropical cas cas estRhopilema nomadica* Galil, Spannier & Red Sea cas est

Ferguson, 1990Stomolophus meleagris* (L. Agassiz, 1862) Atlantic/Pacific casCtenophoraBeroe ovata* Mayer, 1912 Circum(sub)tropical est estMnemiopsis leidyi* A. Agassiz, 1865 NW Atlantic est est est estSulculeolaria angusta* Totton, 1954 Indian estEchinodermataAcanthaster planci (Linnaeus, 1758) Indo-Pacific quesAmphiodia (Amphispina) Mortensen, 1940 Indo-Pacific estobtectaAmphioplus (Lymanella) (Lyman, 1874) Indo-Pacific caslaevisAquilonastra burtoni** (Gray, 1840) W Indian estAsterias rubens Linnaeus, 1755 NE Atlantic estDiadema setosum (Leske, 1778) Indo West Pacific casEucidaris tribuloides (Lamarck, 1816) (sub)tropical Atlantic: estOphiactis macrolepidota Marktanner-Turneretscher, Circumtropical est

1887Ophiactis savignyi (Müller & Troschel, 1842) Circumtropical cas est estProtoreaster nodosus (Linnaeus, 1758) Indo-Pacific casPrionocidaris baculosa (Lamarck, 1816) Indian casSynaptula reciprocans (Forssk l, 1775) Indo-Pacific estPoriferaCinachyrella australiensis Carter, 1886 Indo-Pacific quesGeodia micropunctata Row, 1911 Indo-Pacific quesHaliclona spinosella (Thiele, 1905) Indo-Pacific quesHaliclona viridis (Keller, 1881) Circum(sub)tropical quesHyrtios erecta (Keller, 1889) Indo-Pacific quesLissodendoryx (Ridley, 1844) Indo-Pacific quesschmidtiMycale erythraeana Burton, 1936 Indo-Pacific quesParaleucilla magna Klautau et al., 2004 SW Atlantic cry cry crySiphonophoraAbyla trigona* Quoy & Gaimard,1827 Indian cas

a

a

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDSipunculaApionsoma (Apionsoma) (Ikeda, 1904) Circumtropical ques quesmisakianumApionsoma (Apionsoma) (Sluiter, 1902) Circumtropical castricocephalusAspidosiphon (Akrikos) (Murina, 1967) Circumtropical ques cas casmexicanusAspidosiphon (Aspidosiphon) (Chamisso & Eysenhardt, Circumtropical estelegans 1821)Phascolion (Isomya) (Sluiter, 1902) Indo-Pacific est casconvestitumPhascolosoma scolops (Selenka & de Man, 1883) Indo-Pacific est estPlatyhelminthesAllolepidapedon fistulariaep Yamaguti, 1940 Indo-Pacific casGlyphidohaptor plectocirrap (Paperna, 1972) Red Sea estHirudinella ventricosap (Pallas, 1774) Atlantic casHysterolecitha siganip Manter, 1969 Indo-Pacific quesLecithochirium Fischthal & Kuntz, 1963 Red Sea casmagnicaudatumMonilicaecum ventricosump Yamaguti, 1942 Indo-Pacific casNeothoracocotyle (Meserve, 1938) Hargis, 1956 Atlantic/Pacific casacanthocybiip

Nosema ceratomyxap Diamant & Paperna, 1985 Red Sea estPolylabris cf. mamaevip Ogawa & Egusa, 1980 Indo West Pacific estTetrancistrum polymorphusp (Paperna, 1972) Red Sea cas casTetrancistrum Kritsky, Galli & Yang, 2007 Red Sea eststrophosolenump

Tetrancistrum suezicusp (Paperna, 1972) Red Sea cas estF I S H

ElasmobranchiiCarcharhinus altimus (Springer, 1950) Indo-Pacific est estCarcharhinus falciformis (Müller & Henle, 1839) Circumtropical estDasyatis marmorata (Smith, 1828) Tropical Atlantic ques quesGaleocerdo cuvier (Peron & Le Sueur, 1822) Tropical/subtropical cas casHimantura uarnak (Forssk l, 1775) Indo-Pacific estIsurus paucus Guitart Manday, 1966 Circumtropical casRhizoprionodon acutus (Rüppell, 1837) Circumtropical casSphyrna mokarran (Rüppell, 1837) Circumtropical casTorpedo sinuspersici Olfers, 1831 Indo-Pacific quesActinopterygiiAbudefduf vaigiensis (Quoy & Gaimard, 1825) Indo-Pacific cas cas

a

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Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDAcanthurus monroviae Steindachner, 1876 Tropical Atlantic est cas casAlepes djedaba (Forssk l, 1775) Indo-Pacific estAluterus monocerus (Linnaeus, 1758) Atlantic casAnarhichas lupus Linnaeus, 1758 N Atlantic casAnguilla japonica Temminck & Schlegel, 1847 Pacific quesApogon fasciatus (White, 1790) Indo-Pacific estApogon pharaonis Bellotti, 1874 Indo-Pacific estApogon queketti Gilchrist, 1903 Indian estApogon smithi (Kotthaus, 1970) Indian estArius parkii Gunther, 1864 Indo-Pacific casAtherinomorus forskalii** (Rüppell, 1838) Tropical Atlantic est estBeryx splendens Lowe, 1834 Circumtropical est estBregmaceros atlanticus Goode & Bean, 1886 Tropical Atlantic cas estCallionymus filamentosus Valenciennes, 1837 Circumtropical estCentrolabrus exoletus (Linnaeus, 1758) Tropical Atlantic casCephalopholis taeniops (Valeciennes, 1828) Subtropical Atlantic est casChampsodon nudivittis (Ogilby, 1895) Tropical Atlantic casChampsodon vorax Günther, 1867 Indo West Pacific casChaunax suttkusi Caruso, 1898 Indo-Pacific cas casCheilodipterus (Rüppell, 1838) W Indian casnovemstriatusCheilopogon furcatus (Mitchill, 1815) Circumtropical casChilomycterus reticulatus (Linnaeus, 1758) Circumtropical casChirocentrus dorab Forssk l, 1775) Indo-Pacific casCoryogalops ochetica (Norman, 1927) Red Sea estCrenidens crenidens (Forssk l, 1775) Indian est estCyclichthys spilostylus (Leis & Randall, 1982) Indo-Pacific estCyclopterus lumpus Linnaeus, 1758 Atlantic casCynoglossus sinusarabici (Chabanaud, 1913) Red Sea estDecapterus russelli (Rüppell, 1830) Indo-Pacific estDiodon hystrix (Linnaeus, 1758) Circumtropical cas quesDiplodus bellottii (Steindachner, 1882) Tropical Atlantic estDussumieria elopsoides Bleeker, 1849 Indo-Pacific estElates ransonetti (Steindachner, 1876) Tropical Pacific cas casEnchelycore anatina (Lowe, 1839) Tropical Atlantic estEpinephelus coioides (Hamilton, 1822) Indo-Pacific cas estEpinephelus malabaricus (Bloch & Schneider, 1804) Indo-Pacific estEquulites klunzingeri** (Steindachner, 1898) Indian cas cas estEtrumeus teres (DeKay, 1848) Subtropical est estFavonigobius (Fowler, 1934) Indo West Pacific casmelanobranchus**

a

a

a

Medit. Mar. Sci., 11/2, 2010, 381-493412

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDFistularia commersonii (Rüppell, 1835) Indo-Pacific est est est estFistularia petimba Lacepède, 1803 Tropical-Atlantic casGephyroberyx darwini (Johnson, 1866) Tropical Atlantic cas quesGlaucostegus halavi (Forssk l, 1775) Indo-Pacific cas quesGymnammodytes (Jourdain, 1879) N Atlantic estsemisquamatusHalosaurus ovenii Johnson, 1863 Tropical Atlantic estHemiramphus far (Forssk l, 1775) Indo-Pacific est cas estHeniochus intermedius Steindachner, 1893 Indian casHerklotsichthys punctatus (Rüppell, 1837) Red Sea estHippocampus fuscus Rueppell, 1838 Indian estHyporhamphus affinis (Gunther, 1866) Indo-Pacific casIniistius pavo Valenciennes, 1840 Indo-Pacific casKyphosus incisor (Cuvier, 1831) Tropical Atlantic casKyphosus sectator (Linnaeus, 1758) Tropical Atlantic estLagocephalus sceleratus (Gmelin, 1788) Indo-Pacific cas estLagocephalus spadiceus (Richardson, 1844) Indo-Pacific estLagocephalus suezensis Clark & Gohar, 1953 Red Sea estLiza carinata (Valenciennes, 1836) Indian estLiza haematocheila (Temminck & Schlegel, 1845) Indo-Pacific estLutjanus argentimaculatus (Forssk l, 1775) Indo-Pacific casLutjanus jocu (Bloch & Schneider, 1801) Atlantic casMakaira indica (Cuvier, 1832) Indo-Pacific cas casMicrochirus boscanion Chabanaud, 1926 Tropical Atlantic est casMicrochirus hexophthalmus (Bennet, 1831) Tropical Atlantic est estMonotaxis grandoculis (Forssk l, 1775) Indo-Pacific casMuraenesox cinereus (Forssk l, 1775) Indo-Pacific casMycteroperca fusca (Lowe, 1838) Tropical Atlantic casNemipterus randalli Russell, 1986 Indian estOmobranchus punctatus (Valenciennes, 1836) Indo-Pacific casOplegnathus fasciatus (Temminck & Schlegel, 1944) Pacific casOxyurichthys petersi (Klunzinger, 1871) Red Sea estPagellus bellottii Steindachner, 1882 Tropical Atlantic est casPagrus major (Temminck & Schlegel, 1843) NW Pacific casPampus argenteus (Euphrasen, 1788) Indo-Pacific casPapilloculiceps longiceps (Ehrenberg in Valenciennes, Indian cas

1829)Parexocoetus mento (Valenciennes, 1846) Indo-Pacific est est estParupeneus forsskali (Fourmanoir & Guézé, 1976) Indian ques casPelates quadrilineatus (Bloch, 1790) Indo-Pacific estPempheris vanicolensis Cuvier, 1831 Indo-Pacific est est

aa

a

a

a

Medit. Mar. Sci., 11/2, 2010, 381-493 413

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDPetroscirtes ancylodon Rüppell, 1838 Indian estPinguipes brasilianus Cuvier & Valenciennes, 1829 W Atlantic cas casPisodonophis semicinctus (Richardson, 1848) Tropical Atlantic est est casPlatax teira (Forssk l, 1775) Indo-Pacific casPlatycephalus indicus (Linnaeus, 1758) Indo-Pacific cas estPlotosus lineatus (Thunberg, 1787) Indo-Pacific estPomacanthus imperator (Bloch, 1787) Indo-Pacific casPomacanthus maculosus (Forssk l, 1775) Indo-Pacific casPomadasys stridens (Forssk l, 1775) Indian est estPriacanthus hamrur (Forssk l, 1775) Indo-Pacific ques quesPriacanthus sagittarius Starnes, 1988 Indo-Pacific casPsenes pellucidus Lutken, 1880 Tropical Atlantic est casPseudupeneus prayensis (Cuvier, 1829) Tropical Atlantic casPteragogus pelycus Randall, 1981 Indian estPterois miles (Bennet, 1803) Indian casRachycentron canadum (Linnaeus, 1766) Circumtropical cas casRastrelliger kanagurta (Cuvier, 1816) Indo-Pacific casRhabdosargus haffara (Forssk l, 1775) Indian estRhynchoconger trewavasae Ben-Tuvia 1993 Red Sea casSargocentron rubrum (Forssk l, 1775) Indo-Pacific cas estSaurida undosquamis (Richardson, 1848) Indo-Pacific est est estScarus ghobban Forssk l, 1775 Indo-Pacific estSciaenops ocellatus (Linnaeus, 1766) W Atlantic casScomberomorus commerson Lacepède, 1800 Indo-Pacific ? est estScorpaena stephanica Cadenat, 1943 Tropical Atlantic casSelene dorsalis Gill, 1862 W Atlantic casSeriola carpenteri Mather 1971 Tropical Atlantic estSeriola fasciata (Bloch, 1793) Tropical Atlantic est est casSeriola rivoliana Valenciennes, 1833 Tropical Atlantic casSiganus javus (Linnaeus, 1766) Indo-Pacific casSiganus luridus (Rüppell, 1829) Indian cas est estSiganus rivulatus Forssk l, 1775 Red Sea est cas estSilhouetta aegyptia (Chabanaud, 1933) Red Sea estSillago sihama (Forssk l, 1775) Indo-Pacific estSolea senegalensis Kaup, 1858 Tropical Atlantic est quesSorsogona prionota (Sauvage, 1873) Indian casSphoeroides marmoratus (Lowe, 1838) Atlantic cas casSphoeroides pachygaster (Müller & Troschel, 1848) Tropical Atlantic est est est estSphyraena chrysotaenia Klunzinger, 1884 Indo-Pacific est est estSphyraena flavicauda Rüppell, 1838 Indian est estSpratelloides delicatulus (Bennett, 1831) Indo-Pacific/Red Sea est

a

a

a

a

a

aaa

a

Medit. Mar. Sci., 11/2, 2010, 381-493414

Table 2 (continued)

Species Author Origin WMED CMED ADRIA EMEDStephanolepis diaspros Fraser-Brunner, 1940 Red Sea est cas estSynagrops japonicus (Doderlein, 1884) Indo-Pacific casSynaptura lusitanica Capello, 1868) Tropical Atlantic est estSyngnathus rostellatus Nilsson, 1855 N Atlantic cas quesTerapon jarbua (Forssk l, 1775) Indo-Pacific casTerapon puta (Cuvier, 1929) Indo-Pacific/Red Sea estTerapon theraps Cuvier, 1829 Indo-Pacific casTetrosomus gibbosus (Linnaeus, 1758) Indo-Pacific estTorquigener flavimaculosus Hardy & Randall, 1983 Indian estTrachurus indicus Necrasov, 1966 Indian casTrachyscorpia cristulata (Koehler, 1869) Tropical Atlantic est estechinataTridentiger trigonocephalus (Gill, 1859) Tropical Pacific casTrypauchen vagina (Bloch & Scheider, 1801) Indo-Pacific casTylerius spinosissimus (Regan, 1908) Indo-Pacific estTylosurus choram (Rüppell, 1837) Indo-Pacific casTylosurus crocodilus (Peron & Le Sueur, 1821) Indo West Pacific casUpeneus moluccensis (Bleeker, 1855) Indo-Pacific est estUpeneus pori Ben-Tuvia & Golani, 1989 Indian est estVanderhorstia mertensi Klausewitz, 1974 Indo-Pacific estZenopsis conchifera (Lowe, 1852) Atlantic/Pacific cas

a

1. THE WESTERN MEDITERRANEAN

The WMED occupies a key positionbecause it receives the influx of surfacewaters from the Atlantic, through the Straitof Gibraltar. It is further compartmental-ized into fairly isolated sub-basins with dif-ferent climatic and hydrologic conditions.These sub-basins have a different biogeo-graphic character, which may affect inva-sion and settlement of aliens. The AlboranSea, situated immediately east of Gibraltar,exhibits stronger Atlantic affinities, due tothe continued penetration of Atlantic floraand fauna with the incoming influx of water(HARMELIN & D'HONDT, 1993). Inreturn, most Mediterranean endemics arerare or missing. The incoming Atlantic

waters form a permanent clockwise gyre inthe Alboran Sea, which is separated fromthe remainder of the WMED by a well-marked hydrographic front between Oranand Almer a. As a consequence of theadmixture of fauna and flora of differentorigins, the westernmost part of the Albo-ran Sea, from the Ceuta region to Punta deCalaburras on the coast of Malaga, consti-tutes a hot spot of biodiversity. Its enrichedbenthic environment includes some specieslinked with the tropical periods of theMediterranean Sea (OCAN~A et al., 2009;GARC A RASO et al., 2010; URRA et al.,in press). The Tyrrhenian Sea is compara-tively isolated from the rest of the WMEDand is surrounded by mountains reducingthe impact of the meteorological events

that strongly influence the internal condi-tions of the other Western Mediterraneanbasins (ASTRALDI et al., 1995). The Gulfof Lions and the Ligurian Sea are the cold-est parts of the WMED. They are charac-terized by a severe reduction of ther-mophilic species, while some cold temper-ate species, not found to the south, arepresent (BIANCHI & MORRI, 1993,1994).

1.1. Alien protozoans in the WesternMediterranean

Four pathogenic protozoans havebeen recorded in WMED. Three of themare pathogenic to shellfish: Marteilia refrin-gens (RIERA et al., 1993), Bonamia ostreae(MONTES & LAMA, 1993), and Perkin-sus olseni, formerly Perkinsus atlanticus(SAGRIST et al., 1996). The fourth,Photobacterium damselae, formerly Pas-teurella piscicida, causes pasteurello-sis/photobacteriosis in fish: it was first iso-lated in mortalities occurring in naturalpopulations of white perch and stripedbass in 1963 in Chesapeake Bay, USA.From 1990 it has caused economic lossesin different European countries. Culturedgilthead seabream (Sparus aurata) andseabass (Dicentrarchus labrax) are the mostaffected species in the WMED (TO-RANZO et al., 1991).

Only four alien foraminiferal speciesoccur in the WMED. Planogypsina acer-valis (BLANC-VERNET, 1969, as Planor-bulina acervalis) and Cymbaloporetta planahave been previously cited in the other sec-tors of the Mediterranean. Very rare spec-imens of Sorites orbiculus have beenreported for the Gulf of Naples(HOFKER, 1930) and from the LigurianSea (BANCHETTI et al., 2009, as S. orbic-ularis). Schackoinella imperatoria, probablycryptogenetic/casual, has been recorded

from the Gulf of Naples, South Tyrrhenian(SGARRELLA & MONCHARMONT,1993), from the Ligurian Sea and northTyrrhenian (BANCHETTI et al., 2009)and from the coast of Vulcano, AeolianIslands (CIMERMAN & LANGER, 1991,as Conorbella imperatoria).

1.2. Alien macrophytes in the WesternMediterranean

A total of 91 alien macrophytes havebeen hitherto reported in the WMED,which represent 72% of the alien macro-phytes reported from the MediterraneanSea. The major vectors of introduction arethe shellfish aquaculture industry and shiptraffic (fouling, ballast waters). As much as75 alien macrophyte species are well estab-lished, seven are casual, and nine are cryp-togenic/questionable. Casual taxa (i.e.Cladophora hutchinsioides, Fucus spiralis,Grateloupia patens, Lomentaria flaccida,Rhodophysema georgei, Saccharina japoni-ca and Sarconema filiforme) might attainestablished status in the future. The cryp-togenic/questionable category includesAcanthophora nayadiformis, Anotrichiumokamurae, Antithamnionella boergesenii, A.elegans, the Atlantic strain of Asparagopsistaxiformis, Ceramium bisporum, Ganone-ma farinosum, Osmundea oederi andPolysiphonia atlantica.

On the 21 invasive or potentially inva-sive macrophytes introduced in theMediterranean (Table 3), 16 are present inthe western basin mainly in coastal lagoons.Codium fragile, invasive in the last century,has become less abundant in the open sea.

The northern coasts are the mostimpacted region, with a maximum of alienmacrophytes in France (78 taxa out of 91).On the other hand, the lowest number (23taxa) was recorded along the northernAfrica coast (Morocco and Algeria). Such

A

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a difference is probably due to insufficientinvestigations in the south and the hugeflood of introductions along with shellfishtransfers in the French coastal lagoonsharbouring shellfish aquaculture.VERLAQUE et al. (2007) listed up to 58alien macrophytes in the Thau Lagoon(64% of alien macrophytes of the WMEDand 46% of the total of alien macrophytesof the Mediterranean), which is the lead-ing site of shellfish aquaculture in theMediterranean Sea.

The majority of alien macrophytes ofthe WMED comes from temperate andcold regions (NE Atlantic and NW Pacific)in relation with the major vector of intro-duction (shellfish transfer). Only 10 puta-tive Lessepsian immigrants (i.e. Acan-thophora nayadiformis, Chondria pygmaea,Ganonema farinosum, Halophila stipu-lacea, Hypnea spinella, H. valentiae, Lopho-cladia lallemandii, Padina boergesenii, Sar-conema filiforme and Ulva fasciata) havebeen hitherto recorded, mainly out of thecoldest zones (Gulf of Lions and Gulf ofGenoa). However, the only WMED reportof Sarconema filiforme (France) requiresconfirmation. Acanthophora nayadiformis,Ganonema farinosum, Hypnea spinella,Padina boergesenii and Ulva fasciata couldalternatively originate from the AtlanticOcean. The strains of Ulva fasciata andHypnea valentiae introduced in the ThauLagoon most probably originate from thenorth-western Pacific (Japan or Korea).The invasive behaviour of Lophocladialallemandii in the Balearic Islands con-trasts with a non-invasive behaviour every-where else in the Mediterranean, and adifferent origin for the western straindeserves to be considered. Thus, there arehitherto only two unquestionable Lessep-sian macrophytes in the WMED: Chondriapygmaea and Halophila stipulacea.

1.3. Alien polychaetes in the WesternMediterranean

In the WMED, a total of only 49 alienpolychaete species have been reported todate, which is about 39% of the alien poly-chaete species reported from the Mediter-ranean Sea. Among them, 26 species areestablished in WMED (see Table 2), 12species are casual, 1 species (Paraprionospiocoora) is cryptogenic and 10 species arequestionable. Some species seem to haveinvasive character, forming dense popula-tions in semi- or heavily polluted areas,including harbours (Branchiomma luctuo-sum, Hydroides dianthus, H. elegans, Polydo-ra cornuta, and others), brackish waters(Desdemona ornata, Ficopomatus enigmati-cus, Polydora cornuta, etc.) or shallow-waterbenthic habitats (Eunice antennata, Lum-brineris perkinsi, Notomastus mossambicus,Prionospio (Aquilaspio) krusadensis, etc.).The WMED has a relatively low number ofalien species, in comparison with theEMED, especially the Levantine Sea, whichis densely colonized by alien polychaetes.The majority of the established alienspecies in the WMED are of Pacific, Indo-Pacific, Indian and/or Red Sea origin,whereas less originate in the tropicalAtlantic (six species) or West Atlantic (sixspecies).

A relatively low number of newcomersoriginating in tropical seas and anthro-pogenically introduced via the GibraltarStrait are true alien species. Most of themappear to have arrived via shipping, espe-cially in harbours, as also happens in theEMED. Although the polychaetes of theWMED are relatively well known, newreports are occurring continuously, mak-ing sometimes difficult to elucidate if theyare true invaders, or unrecorded or over-looked species. This is especially evidentwith small, meiobenthic species; for exam-

Medit. Mar. Sci., 11/2, 2010, 381-493416

ple, in a recent work (DEL-PILAR-RUSOet al., in press) two small syllids (Syllis mau-retanica and Parapionosyllis macaronesien-sis; not included in Table) are reported forthe first time in the Mediterranean; bothspecies were described from NE Africaand Canary and Madeira islands, relative-ly close to the Strait of Gibraltar. There isno way to know if these species were acci-dentally introduced, were overlooked inprevious papers, or simply never found.

1.4. Alien crustaceans in the WesternMediterranean

In total, 52 alien species of crustaceanshave been found in the WMED, of which20 have self-maintaining populations, 20are casual records, eight are questionableand four are classified as cryptogenic.These species belong to amphipods (4),cirripedes (5), decapods (20), stomatopods(1), cumaceans (1), planktonic and para-sitic forms of copepods (16), and isopods(5) (Table 2).

In the decapods, we have not consid-ered alien the Atlantic species found in theAlboran Sea and whose historical presenceis known in Morocco. For example,Brachynotus atlanticus, a tropical Africanspecies known in Morocco (FOREST &GANT S, 1960), was found in Europe forthe first time in the littoral of Malaga(GARC A RASO, 1984a), and later inthe Bay of Cadiz, in the Atlantic, where itforms a stable population (GARC ARASO, 1985). The Atlantic shrimpsPenaeopsis serrata and Hymenopenaeusdebilis, known in Moroccan waters (ZA-RIQUIEY, 1968; PÉREZ FARFANTE,1979), were recently found in the westernMediterranean Sea (perhaps due to theirdeep-water habitat); apparently, they arebeing found in more areas of the Mediter-ranean; the former in the Alboran Sea

(ABELL & TORRES, 1998) and Sar-dinia Channel (MURA et al., 2003), thelatter in Alboran Sea and Balearic Island(CARTES et al., 2000). A similar consider-ation can be made for Merhippolyteancistrota, a species occurring in the Albo-ran Sea but also known in the Gulf ofCadiz and Morocco (GARC A RASO,1996; UDEKEM D'ACOZ & URI ,1996), and Plesionika ensis (A. MilneEdwards, 1881), cited in the Alboran Sea(GARC A RASO, 1981) and Gulf ofCadiz (GARC A RASO, 1996). However,new references from the island of Crete,Aegean Sea (LABROPOULOU &KOSTIKAS, 1999) and its absence inintermediate Mediterranean areas raisedoubts about the status of the latterspecies in the EMED. There is an old cita-tion of the species Plagusia depressa(Fabricius, 1775) and Plagusia chabrus(Linnaeus, 1758), now Guinusia chabrus(Linnaeus, 1758), brought to the port ofMarseilles by an iron vessel in 1873(STEBBING, 1893), but they have notbeen included in Table 2, because theyhave never again been captured (there areno later references).

Half the species originate in the Indo-Pacific, Pacific, Indian Ocean or Red Sea,while seven species are known to have acircumtropical distribution. The remain-ders have a tropical/subtropical Atlanticorigin, many of them being known in thewestern Atlantic.

Two species, Synalpheus tumidomanusafricanus and Necora puber, not includedin earlier Mediterranean alien lists, areadded in Table 2. Synalpheus tumidomanusafricanus (syn. S. hululensis africanus) wasfirst captured in Israel (LEWINSOHN &HOLTHUIS, 1964, as Synalpheus hululen-sis) and considered as alien; later it wasfound in the Alboran Sea (GARC A

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RASO, 1984b; L PEZ DE LA ROSA &GARC A RASO, 1992; GARC ARASO, 1996), a characterization that sub-sequently changed following reclassifica-tion of the Mediterranean specimens with-in the subspecies S. tumidomanus africanus(or S. africanus?), known from Casablancaand Rabat, Morocco (LAGARD RE,1971). The Alboran populations may rep-resent a natural expansion of the distribu-tion range of the species, which enteredthrough the Strait of Gibraltar and colo-nized the southern coast of Spain (perhapswith the help of the Atlantic surface waterinflow). The references from Greece andTurkey (KOUKOURAS & KATTOU-LAS 1974; KOCATA , 1981) could indi-cate a separate advance in the EMED lit-toral from the Israeli populations. Recent-ly it has been captured in Sicily (BACCI etal., 2010). The unresolved question iswhether the EMED populations are alienor not. At this time we have classified it ascryptogenic.

Necora puber is included as ‘alien’species because (in agreement withZIBROWIUS, 1992) we believe that itspresence in several areas of the Mediter-ranean is due to introductions, most likelyrelated to its widespread food use ratherthan an entry through ballast waters. Onlythe specimens collected west of Malaga(GARC A RASO et al., 2010) could haveentered the Mediterranean through theStrait of Gibraltar by their natural meansof dispersion.

With regard to other groups of crus-taceans we should note that the availabledata are scarce and often difficult to assess.This is partly due to the smaller number oftaxonomic specialists and studies devel-oped on these groups in the differentMediterranean sectors. These limitationslead to a) a downwards-biased estimation

of aliens in the groups of amphipods, cirri-pedes, cumaceans, isopods andtanaidaceans and b) a high number ofquestionable and cryptogenic species.

The amphipod Elasmopus pectenicrushas a circumtropical distribution: AtlanticOcean, Red Sea, Indian Ocean, PacificOcean (described from New Guinea); ithas gradually colonized the EMED, theAdriatic and the CMED (ZAKHAMA-SRAIEB & CHARFI-CHEIKHROUHA,2010). The latest finding could be clearlythe result of increased scientific effort inthe area.

In the WMED the only speciesmarkedly invasive is Percnon gibbesi, whichshows a rapid expansion of its geographi-cal distribution range, e.g. in the east Span-ish coast (DEUDERO et al., 2005): firstrecorded from the Balearic Archipelago in1999 (GARCIA & REVIRIEGO, 2000),it established populations since 2002 inBarcelona, 2003 in Alicante and Murcia,2006 in Almeria.

Among alien planktonic copepodsreported in the WMED Sea, three areclearly Lessepsian immigrants (Triconiahawii, T. rufa and T. umerus) and werefound in the Gulf of Naples. Two otherLessepsian immigrants, Labidocera detrun-cata and Calanopia elliptica, were reportedfrom the Gulf of Naples (VIVES &SHMELEVA, 2007), but were never foundin the more than 20 years time series ofsamples collected bi-weekly to weekly inthe area (Mazzocchi, pers. comm.): there-fore, their presence is considered as ques-tionable. Pseudocalanus elongatus andParacalanus indicus seem to be cryptogenicin the area; according to RAZOULS et al.(2005-2010) the latter species could be con-founded with Paracalanus parvus, a verycommon native Mediterranean species.

Two copepods parasitic on oyster beds

Medit. Mar. Sci., 11/2, 2010, 381-493418

(Myicola ostreae and Mytilicola orientalis)are well established in the WMED, aftertheir accidental introduction into theFrench coast with the Pacific oyster Cras-sostrea gigas (CLANZIG, 1989).

1.5. Alien molluscs in the Western Mediter-ranean

The WMED has a low incidence ofalien Mollusca with 34 recorded species (22established, nine casual, two cryptogenicand one questionable) in a total of around1500 native species. Two of these (Cerithi-um scabridum and Brachidontes pharaonis)are found only very close to the Strait ofMessina and are merely outliers from theconfirmed area where these species haveestablished in the Ionian Sea. Others, likeChromodoris quadricolor, Pinctada radiata,Rissoina spirata and Thais lacera, had casu-al occurrences that are unlikely to persist,although they may be firmly established inthe EMED. Leaving aside the outliers andthese casual species, the cryptogenicspecies, and the questionable report of Sac-costrea cucullata for Tunisia, we are leftwith not more than 25 alien molluscs defi-nitely relevant to the basin.

The north coast of Tunisia representsa major step towards the colonization ofthe WMED. Two of the hardiest Lessep-sian immigrants are present there, havingmade their way across the WMED to thecoast of Spain and are locally invasive.These are Bursatella leachii(WEITZMANN et al., 2009; RAMOS-ESPL et al., 2010) and Fulvia fragilis(TAMAYO-GOYA, 2008). B. leachii alsoexists in the Atlantic, but the hypothesisthat WMED populations entered throughGibraltar does not hold since the speciesdoes not exist in Morocco.

The largest pool of alien molluscs inthe WMED includes species that are

actively cultured (for example Crassostreagigas and Ruditapes philippinarum, to alesser extent Mercenaria mercenaria) andspecies that were probably introducedaccidentally along with aquaculture(Crepidula fornicata, Gibbula albida, Mus-culista senhousia, Xenostrobus securis),thereby comprising one-fourth of thereported aliens. M. senhousia and X.securis are locally invasive in the basin, butC. fornicata, which is one of the most inva-sive aliens in the NE Atlantic, has difficul-ty in surviving where it has been intro-duced in the Mediterranean. Gibbula albi-da was originally an Adriatic endemic, butis now introduced in the Ebro Delta(TRIGO, 1981) and the French Mediter-ranean lagoons (CLANZIG, 1989), as wellas on the Atlantic coast of France.

A unique and certainly recent featureis the occurrence in the vicinity of Tunis(Tunisia) of two alien species (Favorinusghanensis, Mitrella psilla) introducedthrough shipping from tropical WestAfrica (BEN SOUISSI et al., 2004; ANTITet al., 2010). More generally, harbours area focus for established populations ofaliens, with random origins. Other exam-ples are the bivalve Theora lubrica inLeghorn, Italy, and the gastropodBostrycapulus odites in Alicante, Spain,which did not spread out to the neighbour-ing open sea.

On the whole, the incidence of alienmollusc species in the WMED is heavilyconcentrated in marginal marine environ-ments such as the lagoons of the FrenchMediterranean coast, the Spanish MarMenor and Ebro Delta, and the Italianlagoons of Caprolace and Fusaro on theTyrrhenian coast. With the exception ofthe recent report of the nudibranch Godi-va quadricolor from the large harbour ofAlgeciras (CERVERA et al., 2010), the

Medit. Mar. Sci., 11/2, 2010, 381-493 419

Alboran Sea is free of alien Mollusca, a sit-uation which may be related to its excep-tionally high species richness estimated asabout 1200 species. More generally, theopen sea in the WMED is hardly, if at all,impacted by alien species of molluscs.

Although a couple of alien molluscs inthe Mediterranean are of West African ori-gin, there is no species yet that has gradu-ally entered through the Strait of Gibraltarand become newly established in theMediterranean. As already noted byGOFAS & ZENETOS (2003), all theprevalently Atlantic species found in thewesternmost Mediterranean have a histor-ical range since at least the 19th century andare, therefore, to be considered nativespecies. The WMED is nevertheless asource area for such species as Siphonariapectinata, native to Algeria and Spain butintroduced in Greece and recently expand-ing its range towards northern Tunisia.Although not qualifying as ‘aliens’ and notincluded in the species counts, one mustkeep in mind the fact that such species asEastonia rugosa (see ALBANO, 2006) areexpanding their range.

1.6. Miscellaneous invertebrates in theWestern Mediterranean

Bryozoa include five alien species inthe WMED, out of 23 known from thewhole basin. Arachnoidea protecta is anIndo-Pacific species first recorded in theWMED by CHIMENZ GUSSO et al.(1998), further records being provided byD'HONDT & CHIMENZ GUSSO(2006). The morphological divergenceobserved between the Indo-Pacific andMediterranean specimens led Harmelin(in ZENETOS et al., 2005) to believe thatthey represent two distinct species. How-ever, OCCHIPINTI-AMBROGI et al.(2010) maintain A. protecta as established.

Alien Ascidiacea are represented byeight species (out of 16), most of themestablished. Five species are circumtropi-cal. Trididemnum cf. savignyi was firstrecorded by LAFARGUE (1972) at Port-Cros, France. LAFARGUE (1974) revisedthe Trididemnum tenerum complex, assign-ing the record by PÉR S (1954) fromsouthern Tunisia to T. cf. savignyi. Subse-quent records of this species include Cat-alonia and Columbretes Islands, Spain(RAMOS-ESPL , 1988). Perophora mul-ticlathrata and Ecteinascia styeloides havebeen reported from Corsica (MONNIOT,1983) and the northern Tyrrhenian Sea(MASTROTOTARO & TURSI, 2010),respectively. Microcosmus squamiger, firstrecorded in Bizerte (as M. exasperatus) iswidespread in the WMED (synthesis inTUR N et al., 2007). Cystodytes philip-pinensis, previously observed in southernTunisia (MÉLIANE, 2002), has beenrecorded recently in the Balearic Islands(D AZ-VALDÉS & RAMOS-ESPL ,2010). This cryptogenic species had proba-bly been confused in the past with its con-gener C. dellechiajei, widely distributed inthe Mediterranean Sea (IZQUIERDO-MUN~OZ et al., 2009). The WesternAtlantic species Distaplia bermudensis andthe Eastern Pacific species Polyandrocarpazorritensis have been introduced by shell-fish culture (TUR N & PEREA, 1988;BRUNETTI & MASTROTOTARO,2004; MASTROTOTARO & BRU-NETTI, 2006). Styela clava, originatingfrom the north-western Pacific, has beenspreading along the European Atlanticcoast since 1954 and has only recently beenrecorded in the WMED; however, itsabsence from harbours and marinas closedto Gibraltar may suggest that this specieshas also been accidentally transported intothe Mediterranean Sea by shellfish trans-

Medit. Mar. Sci., 11/2, 2010, 381-493420

fer (DAVIS & DAVIS, 2008).Cnidaria are an exception to most taxa

in that the majority of aliens occur in theWMED rather than in the EMED: 26species (out of 46 listed for the wholebasin). Alien Anthozoa are represented bytwo species, out of four reported for thewhole Mediterranean. Although firstrecorded in the Ligurian Sea(ZIBROWIUS, 1974), Oculina patagonicais abundant in the southern part of theWMED, both in Spain (ZIBROWIUS &RAMOS, 1983; BALLESTEROS, 1998;IZQUIERDO et al., 2007) and along theNorth African coast (SARTORETTO etal., 2008). Eighteen alien species (out of37) belong to Hydrozoa, a group well rep-resented in the fouling assemblages(MORRI & BOERO, 1986) that colonizethe large ports of the WMED. Cordy-lophora caspia has been reported only forthe brackish lake of Fondi, Tyrrhenian Sea(MORRI, 1979). Two alien Scyphozoawere found in the WMED: a single speci-men of Phyllorhiza punctata was seen inOctober 2009 off NE Sardinia (BOERO etal., 2009), while Stomolophus meleagris isthe only one recorded exclusively in theWMED: this may suggest it entered fromthe Atlantic, although the species is alsoknown from the Pacific (MORAND &DALLOT, 1985).

Both species of alien Ctenophorareported for the Mediterranean occur inthe WMED (BOERO et al., 2009;FUENTES et al., 2009; MILLS, 2009),while Sipuncula and Pycnogonida haveone alien species each (PANCUCCI-PAPADOPOULOU et al., 1999; BARTO-LINO & CHIMENZ GUSSO, 2010).

The only alien echinoderms recordedin the WMED are two seastars and oneophiuroid of Indo-Pacific origin: Acan-thaster planci, Protoreaster nodosus, and

Ophiactis savignyi. The former has beenreported from Port Cros, France, in 2000(ICES, 2006) with no further details(hence it is classified as questionable),whereas two individuals of Protoreasternodosus, probably released accidentallyfrom a private aquarium, have been caughtby trawling off Majorca, Balearic Islands,Spain, in 1981 (ALVARADO et al., 1986).Casual is considered the presence of Ophi-actis savignyi in Banyuls (GUILLE, 1969).

The only alien Porifera in the WMEDis Paraleucilla magna, which is proliferat-ing across the Mediterranean(GUARDIOLA et al., 2010). Platy-helminthes are represented by Allolepi-dapedon fistulariae, an endoparasite of Fis-tularia commersonii reported from Sar-dinia (PAIS et al., 2007).

1.7. Alien fish in the Western Mediter-ranean

MASSUTI et al. (2010) reported 38new fish in the WMED since the middle20th century. Here we report 45 species (20established, 25 casual) inventoried in the20th century. Most of them are of tropical(subtropical) Atlantic origin.

The entrance of alien fish in theWMED has drawn the attention on the so-called effect of ‘tropicalization'. This phe-nomenon has expanded geographically,reaching the EMED. Such is the case forthe Atlantic origin species Carcharhinusaltimus, Acanthurus monroviae, Pisodo-nophis semicinctus, Sphoeroides pachy-gaster, Pagellus bellottii and Seriola fasciata,which have recently entered the Mediter-ranean through the Strait of Gibraltar, andhave already reached the EMED.

Documentation on some species islimited so that those here reported as casu-als could have already established popula-tions. This could be the case of a) benthic

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species with limited swimming capacitysuch as Microchirus boscanion, which hasbeen captured at different localities andtimes (MASSUTI et al., 2010); b) confu-sion with others, such as the soleidsMicrochirus hexophthalmus and Synapturalusitanica known for the Iberian Peninsulaand the Gulf of Lions (MATALLANAS,1984). Some species, such as Anarhichaslupus, Aluterus monocerus, Scorpaenastephanica, Fistularia petimba and Zenopsisconchifera, are known only from singleobservations and are still limited to theWMED. The latest findings include Lut-janus jocu (VACCHI et al., 2010) andKyphosus incisor (ORSI-RELINI et al.,2010), both in the Ligurian Sea.

Within the alien Indo-Pacific species,Pomadasys stridens was reported for thefirst time in the WMED and then it wasknown in the EMED, where it is common;Abudefduf vaigiensis, a casual species in theLevantine basin, has been reported threetimes in the WMED could be similar. Bycontrast, Siganus luridus, and Fistulariacommersonii, which were first reported inthe EMED, where they are now wellknown and even common, have expandedwestwards and are considered established.Fistularia commersonii has expanded as farnorth as the Ligurian Sea (OCCHIPINTI-AMBROGI & GALIL, 2009) and as farwest as Algeria (HEMIDA & CAPAPÉ,2009) and Spain (S NCHEZ-TOCINOet al., 2007).

In the WMED, only a few speciesappear regularly in the catches of somefisheries. That could be the case of Soleasenegalensis, whose presence in theWMED was recorded in 1920 off the Iber-ian coast and has presently extended toAlgeria, Tunisia and the Gulf of Lions, andGymnammodytes semisquamatus reportedonly in the north-eastern Iberian coast,

where it is exploited in commercial fishery.Diplodus bellottii and Pagellus bellottii,reported at the Maghrebine and Iberiancoasts of the Alboran Sea, are occasionallycaptured by commercial fisheries(MASSUTI et al., 2010).

2. CENTRAL MEDITERRANEAN

The bulk of the CMED is representedby the Ionian Sea, the least known of allthe Mediterranean sub-basins(ZENETOS et al., 1997). Local endemics,mostly within molluscs, have been report-ed for its southern, non-European shores(SABELLI & TAVIANI, 1980) but alsofor Malta (EVANS et al., 2010). The Ion-ian is connected to the WMED throughthe narrow Strait of Messina, a micro-sec-tor that harbours a wealth of biogeograph-ic peculiarities, including Pliocene Atlanticremnants and local endemisms (FREDJ &GIACCONE, 1995), and the larger Straitof Sicily, the meeting point of nativeWestern and Eastern Mediterraneanspecies (BIANCHI, 2007), as well as ofaliens of either Atlantic or Indo-Pacificorigin (COLL et al., 2010). Reflecting thissituation at the crossroads, the composi-tion of the alien faunas is much more bal-anced between different sources than inthe EMED. Its situation at the transitionbetween the eastern and western basinsmake it a particularly sensitive place formonitoring the progression of the muchmore numerous aliens already establishedin the EMED.

2.1. Alien protozoans in the CentralMediterranean

Fish pasteurellosis from the protistPhotobacterium damselae was reportedfrom cultured seabass and seabreaminstallations in the Greek Ionian Sea and

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from Malta (BAKOPOULOS et al., 1995,1997).

Only 4 alien foraminiferal species arereported for the CMED: Amphistegina lob-ifera, which probably has established apopulation around the island of Malta(YOKES et al., 2007), A. lessonii and A.madagascariensis documented along thecoast of southern Tunisia (BLANC-VERNET, 1969; HOLLAUS &HOTTINGER, 1997). Planogypsina acer-valis seems to have been recorded in thisarea only by BLANC-VERNET (1969).

2.2. Alien macrophytes in the CentralMediterranean

In the CMED, 57 alien macrophyteshave been hitherto reported, which repre-sent 46% of the alien macrophytes knownfor the Mediterranean Sea as a whole. Themajor vectors of introduction are the shiptraffic (fouling, ballast waters), the SuezCanal, and, in the south Italian coastallagoons, the shellfish aquaculture. Amongthem, 35 alien macrophytes are well estab-lished, four are casual, 16 are crypto-genic/questionable, and two require con-firmation. Casual taxa (i.e. Grateloupialanceolata, Hypnea anastomosans, Polysi-phonia paniculata and Undaria pinnatifida)might turn into the established status inthe future. The occurrence of Batophorasp. and Gracilaria arcuata in the CMEDrequires confirmation.

Of the 21 invasive or potentially inva-sive macrophytes introduced in theMediterranean (Table 3), 13 have beenreported in the CMED. Undaria pinnatifi-da that was only reported from the MarPiccolo of Taranto (south Italy), seems tohave disappeared from the region(GRAVILI et al., 2010).

The Ionian coasts and Sicily are themost impacted regions, with 35 and 36

alien macrophytes respectively, followedby Tunisia (25) and Libya (12). Such dif-ferences are probably due to a lower inves-tigation effort, particularly in Libya.

The alien macrophytes of the CMEDcome from temperate/cold regions (NEAtlantic and NW Pacific) and from tropi-cal regions in almost equal proportions (51and 49%, respectively). A total of 17 puta-tive Lessepsian immigrants have beenhitherto recorded in the region.

2.3. Alien polychaetes in the CentralMediterranean

The CMED includes 36 alien speciesof polychaetes (17 established, nine casual,and ten questionable). Aliens alreadyestablished in the EMED have colonizedthis area to subsequently spread to theWMED: examples are provided by thethree Indo-Pacific species, and likelyLessepsian immigrants, Eunice antennata,Pseudonereis anomala (both established)and Protodorvillea egena (questionable).By contrast, Linopherus canariensis andOphryotrocha japonica, both established,could be indicative of a passage from westto east. The former is very abundant in theFaro Lake, on the Messina Strait(COSENTINO et al., 2009), together withanother alien species: Syllis hyllebergi,which is exclusively found in this area(COSENTINO, in press). Other exclusivespecies of the CMED are Eunice floridi-ana, Epidiopatra hupferiana hupferiana,Epidiopatra hupferiana monroi and Isoldapulchella, mostly collected in the Gulf ofNoto (Sicily), all considered casual, nonestablished taxa (OCCHIPINTI-AMBROGI et al., 2010). On the contrary,Ophryotrocha japonica is very abundant inthe Mar Piccolo of Taranto, an area con-sidered a hot spot of alien species diversitywithin the Italian coasts (OCCHIPINTI-

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AMBROGI et al., 2010). In this localityone of the most conspicuous populationsof Branchiomma luctuosum is present(LICCIANO et al., 2002), and it seemshere to compete with the native speciesSabella spallanzanii (Giangrande, pers.observ.). Most of the other establishedspecies in this area are the same as thosethat are common in the whole basin suchas Ficopomatus enigmaticus, Hydroidesdianthus, H. diramphus, H. elegans,Lysidice collaris, Metasychis gotoi, Neo-pseudocapitella brasiliensis, and Notomas-tus aberans.

2.4. Alien crustaceans in the CentralMediterranean

A total of 46 alien crustacean arereported in the CMED (26 established, 13casual, four questionable, three crypto-genic).

Three West Atlantic species are wellestablished at least in some regions of theCMED: Percnon gibbesi, Libinia dubia,and Rimapenaeus similis, which was discov-ered on the trawling grounds of the Gulf ofGabès (BEN HADJ HAMIDA-BENABDALLAH et al., 2010). Percnon gibbesiwas first observed in 1999 at Linosa Island(RELINI et al., 2000). It rapidly spread inother localities of the CMED: Pantelleria(GALIL et al., 2002), Malta (BORG &ATTARD-MONTALTO, 2002), theshores of southern and north-western Sici-ly (MORI & VACCHI, 2002), the Strait ofMessina (BELLANTONI & CORAZZA,2003) and the Gulf of Taranto (FACCIA& BIANCHI, 2007).

Several Lessepsian immigrants, suchas Trachysalambria palaestinensis, Eucratecrenata, Metapenaeus monoceros andMetapenaeus stebbingi, established in theEMED have spread westward as far asTunisian waters. Their apparent absence

along the Libyan shores may well be con-sequence of a lower research effort, butthey are also absent from Malta and thesouthern Italian shores. Another Indo-Pacific species is Plagusia squamosa.

The following species were recordedonly once and with a single specimen:Alpheus inopinatus, Dromia spinirostris, Sir-pus monodi, Callinectes sapidus, Grapsusgranulosus. The presence of one adultspecimen of Paralithoides camtschaticus, astrictly boreal species, in this sector of theMediterranean (FACCIA et al., 2009) isreally puzzling.

The number of alien copepods is low(five) in the CMED and only two areLessepsian immigrants (Euchaeta concin-na, Triconia umerus). The presence of theAntarctic to Subantarctic speciesSpinocalanus terranovae in the Malta areais questionable (SCIBERRAS &SCHEMBRI, 2007).

The bentho-planktonic calanoid cope-pod Pseudocyclops xiphophorus was previ-ously recorded only in coastal waters ofMozambique. The Mediterranean P.xiphophorus specimens were collectedfrom fouling attached to submerged moor-ing posts and ropes in the brackish LakeFaro, eastern Sicily. It is classified as cryp-togenic because according to ZAGAMI etal. ( 2005) it could represent a relict popu-lation of Tethyan origin. The genus Pseu-docyclops has a worldwide distributionfrom temperate to tropical shallow coastaland brackish waters.

2.5. Alien molluscs in the Central Mediter-ranean

The CMED has reports for 38 alienspecies of molluscs (19 established, 13 casu-al, two cryptogenic and four questionable)in a total of 1000 to 1500 native species.Among the species of Indo-Pacific origin,

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there are 12 species that qualify as Lessep-sian (e.g. Pinctada radiata, the earliestreported Lessepsian mollusc) and six thatare definitely non-Lessepsian (e.g. Melibeviridis which was first detected on the Ioniancoast of Greece), keeping the same propor-tion as in SE Turkey but far from the 90%of Lessepsians along the Levantine coast.

There are scattered reports of speciesfrom the temperate Atlantic (Crepidulafornicata, Polycerella emertoni, Anadaratransversa, Zygochlamys patagonica) andNorth Pacific (Crassostrea gigas purposelyintroduced for farming, Musculista senhou-sia, Ruditapes philippinarum) but most arefar less successful in this particular context,than in the northern part of the Mediter-ranean (Ligurian Sea, Adriatic Sea) wheresome of them are invasive (see Table 3).Considering the proximity of the EMED,the total numbers are neverthelessremarkably low. Few species are invasive,among them could be considered thebivalves Pinctada radiata and Fulvia fragilisand the opisthobranch Bursatella leachii inthe Gulf of Gabès.

Tunisia holds altogether 24 alienspecies (13 established, three casual, threecryptogenic, five questionable) but a dis-tinction must be made between the easternand southern coasts bordering the Gulf ofGabès, which are a continuation of the vir-tually unknown Libyan coast, and thenorth coast west of Cap Bon, which is con-sidered as part of the WMED. A cowry,Erosaria turdus, had a spectacular onset inrecent years and is so invasive that it willprobably displace the native Zonariapyrum. The limpet Cellana rota may be thenext spectacular invader in this part of theMediterranean.

Malta has possibly one of the beststudied molluscan faunas in the southernpart of the EU, and has reports for 17

species. The Ionian coast of Greece hasonly nine species reported, of which two(Bursatella leachii and Pinctada radiata)are among the most widespread Lessep-sian immigrants and five are non-Lessep-sian species of tropical Indo-Pacific origin.The Ionian coast of Italy holds 14 species.These three areas share very much of theiraliens including the most widespreadLessepsian species (Bursatella leachii andPinctada radiata, Cerithium scabridum andFulvia fragilis in Italy and Malta) whichhere are secondary introductions from theEMED. Also shared are some of the trop-ical Indo-Pacific species which startedtheir spread in this area (Aplysia dacty-lomela, Haminoea cyanomarginata, Melibeviridis). Some species, like Anadara trans-versa or Musculista senhousia, which areinvasive in the Adriatic, have spreadtowards the Italian Ionian coast.

2.6. Miscellaneous invertebrates in theCentral Mediterranean

Bryozoa are comprised of eightspecies (out of 23 known for the wholeMediterranean), of various origins (threecircumtropical, three Pacific, twoAtlantic). The records of Celleporaria aper-ta and C. pilaefera at Malta have been con-sidered questionable by SCIBERRAS &SCHEMBRI (2007).

Nearly half of the alien species ofAscidiacea inventoried in the Mediter-ranean (seven out of 16) are known for theCMED. These aliens are of various origins(Western Atlantic, circumtropical, Indo-Pacific, Eastern Pacific) and both the alienspecies of Microcosmus, the Indo-PacificM. exasperatus and the circumtropical M.squamiger, occur in the CMED(IZQUIERDO-MUN~OZ et al., 2009).Most records of alien ascidians come fromTaranto harbour (BRUNETTI &

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MASTROTOTARO, 2004; MASTRO-TOTARO et al., 2004; MASTROTO-TARO & BRUNETTI, 2006).

The CMED hosts nine out of 46 alienspecies of Cnidaria known for the wholebasin. Hydrozoa include six species, all witha wide distribution in warm waters. Thecryptogenic species Cordylophora caspiawas found abundant in the low salinitywaters of the ‘Palude del Capitano’, Gulf ofTaranto (BIANCHI et al., 1994). Scypho-zoa are represented by the three speciesCassiopea andromeda (SCHEMBRI et al.,2010a), Phyllorhiza punctata (ABED-NAVANDI & KIKINGER, 2007) andRhopilema nomadica (SIOKOU-FRANGOU et al., 2006).

The only alien species of Ctenophorais Mnemiopsis leidyi, recorded from Isola diCapo Rizzuto, Gulf of Taranto (BOEROet al., 2009), whereas the only alien speciesof Porifera is Paraleucilla magna, originat-ing from the SW Atlantic (ZAMMIT et al.,2009). In the case of Sipuncula, there arequestionable records of the circumtropicalAspidoshiphon mexicanus from Malta andLampedusa (PANCUCCI-PAPADO-POULOU et al., 1999; SCIBERRAS &SCHEMBRI, 2007).

Alien Echinodermata are representedby three species (out of 12 for the wholeMediterranean). The record of Eucidaristribuloides from Malta was also the first ofthis species for the Mediterranean(TANTI & SCHEMBRI, 2006).

2.7. Alien fish in the Central Mediter-ranean

In the CMED, there are records of 50alien fish. Of them, 25 seem to be estab-lished, 19 are casual, whilst the presence ofsix species is questionable. The bulk of theserecords comes from the Sicily Strait (seeGUIDETTI et al., 2010 and references

therein), especially from Pelagie Islands andMalta but also from Tunisia (BENSOUISSI et al., 2006a,b) and Libya(SHAKMAN & KINZELBACH, 2006;BEN ABDALLAH et al., 2007). A fewrecords are also available from the northernIonian Sea (TORCHIO, 1963; MASTRO-TOTARO et al., 2007) and the MessinaStrait, as in the case of Galeocerdo cuvier(CELONA, 2000), Pinguipes brasilianus(ORSI-RELINI, 2002), Platycephalus indi-cus (CASTRIOTA et al., 2009) and Psenespellucidus (NAVARRA et al., 2008).

Evidence of established populations inthe CMED are mostly related to the occur-rence of multiple records of the samespecies, such as for Cephalopholis taeniops(GUIDETTI et al., 2010) and Pisodonophissemicinctus (RAGONESE & GIUSTO,2000) while a few species such as Siganusluridus, Fistularia commersonii, Seriola car-penteri, Stephanolepis diaspros, Sphoeroidespachygaster have been collected or observedin large numbers (RAGONESE et al., 1997;PIZZICORI et al., 2000; AZZURRO &ANDALORO, 2004; BRADAI et al., 2004;AZZURRO et al., 2007). Latest recordsinclude Selene dorsalis (VELLA &DEIDUN, 2009) and Opleognathus fascia-tus (SCHEMBRI et al., 2010b).

The geographical partitioning ofCMED alien species was: 19 of Atlanticorigin, 25 of Indo-Pacific or Pacific origin,and three circumtropical. If we look onlyat the established species, the number ofIndo-Pacific or Pacific fish (15) is almostdouble than the number of Atlantic new-comers (eight) indicating unbalanced ratiotowards fish coming from the east. Indeed,the progressive penetration westwards ofLessepsian immigrants is a continuousphenomenon, which has accelerated dur-ing the last decades, as exemplified byUpeneus pori, Pempheris vanicolensis,

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Sphyraena chrysotaenia and Siganus rivula-tus (BRADAI et al., 2004; BEN SOUISSIet al., 2006a,b).

3. THE ADRIATIC SEA

The Adriatic Sea is a rather uniqueand differentiated area within theMediterranean, with a strong contrastbetween the predominantly linear sandyshores along the western (Italian) side, andthe opposite complex coasts of the easternside (Slovenia, Croatia, Montenegro andAlbania) forming a maze of islands andinlets with rocky shores. The hydrographicconditions are also peculiar, with very lowwinter temperatures in the northern part,which is also quite shallow (40 m depth),and very hot summers in the southern part,which is much deeper. All these featureslead to differentiation between the north-ern and southern Adriatic areas.

From a biogeographic standpoint, theAdriatic Sea is divided in three sectors,arranged more or less latitudinally. TheNorthern Adriatic Sea is perhaps the mostpeculiar sector of the whole Mediter-ranean. Strong winter cooling, low salinitydue to significant river input, and compar-atively great tidal range make it more sim-ilar to the Northern Atlantic than to therest of the Mediterranean. This is reflectedin the biota, which includes disjunctAtlantic-Adriatic species whose occur-rence contributes to what has been calledthe Northern-Adriatic ‘sub-Atlanticism’(SACCHI et al., 1985). Mediterraneanendemics are scarce and the overall diver-sity is the lowest of the Mediterraneanbasin, so that the phrase ‘North-Adriaticgap’ has also been coined (SACCHI,1983). In addition, peculiar to this sectorare some taxa with Black Sea affinity(SACCHI et al., 1985; BIANCHI et al.,

2004). The Central Adriatic is character-ized by the lack of both Northern-Adriaticendemics and Atlantic-Adriatic disjunc-tions. Mediterranean endemics and ther-mophilic species are still scarce, so that thissector is less diverse than the Gulf of Lionsand Ligurian Sea, to which it resembles.However, it is significantly richer in speciesthan the Northern Adriatic. Finally, theSouthern Adriatic exhibits a transitionalcharacter between the Adriatic and theIonian Sea, with which it communicatesthrough the Otranto Strait. Affinities withthe WMED are strongly reduced.

3.1. Alien protozoans in the Adriatic SeaThree protozoans pathogenic to shell-

fish, namely Marteilia refringens, Bonamiaostrea and Perkinsus olseni (formerlyPerkinsus atlanticus) have been reportedfrom shellfish cultures in the Adriatic Sea.Serious outbreaks of pasteurellosis causedby Photobacterium damselae (formerlyPasteurella piscicida) in the Adriatic arereported by CESCHIA et al. (1991).

Foraminiferal aliens from the Adriaticare few (six species). Moreover, all of themhave been reported based on single speci-mens, in only one publication or site andcould be considered as cryptogenic/casual.This is the case for warm water alienspecies such as Archais angulatus(LANGER & HOTTINGER, 2000),Clavulina angularis (BANCHETTI et al.,2009), Coscinospira hemprichii, Cymbalo-poretta plana (as Trethomphalus bulloides)and Sorites orbiculus (CIMERMAN &LANGER, 1991). A. angulatus is onlyreported from the Adriatic (LANGER &HOTTINGER, 2000). Schackoinellaimperatoria, described from the coast ofcentral Adriatic (CIMERMAN &LANGER, 1991), could be cautiously con-sidered a cryptogenic/casual form.

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3.2. Alien macrophytes in the Adriatic SeaA total of 49 alien macrophytes has

been hitherto reported from the AdriaticSea, which represent 39% of the alienmacrophytes reported from the Mediter-ranean Sea. The major vectors of intro-duction are the shellfish aquacultureindustry in northern coastal lagoons andthe ship traffic (fouling, ballast waters).Among them, 35 alien macrophytes arewell established, three are casual, and 11are cryptogenic/questionable. Casual taxa(i.e. Acrothamnion preissii, Hypnea valenti-ae and Polysiphonia paniculata) might turninto the established status in the future.The cryptogenic/questionable categoryincludes 11 species (Table 2).

Of the 21 invasive or potentially inva-sive macrophytes introduced into theMediterranean (Table 3), 14 are present inthe Adriatic Sea. Among them, cold tem-perate taxa (i.e. Codium fragile, Gracilariavermiculophylla, Grateloupia turuturu,Dasysiphonia sp., Sargassum muticum andUndaria pinnatifida) grow in north-westerncoastal lagoons, especially in the Lagoonof Venice and the lagoons of the Po Delta,while the warm temperate and tropicaltaxa (i.e. Asparagopsis spp., Caulerpa race-mosa var. cylindracea, Caulerpa taxifolia,Halophila stipulacea, Lophocladia lalle-mandii, and Womersleyella setacea) invadethe marine habitats of southern Italy andthe east coasts of the Adriatic (Croatia,Albania), with the exception of Acrotham-nion preissii. The invasive Gracilaria ver-miculophylla is hitherto known only in thelagoons of the Po Delta.

Northern Adriatic coasts are the mostimpacted, with 34 aliens out of a total of 49(69% of alien macrophytes of the Adriat-ic), while only 15 aliens were reported fromthe Central and Southern Adriatic.SFRISO et al. (2009 and unpublished data)

listed up to 33 aliens (67% of alien macro-phytes of the Adriatic) in the Lagoon ofVenice. Such a difference is probably dueto a flood of introductions along with shell-fish transfers (oysters, mussels, Manilaclams) on the north-Italian coasts whereaquaculture is extensively developed.

The majority of alien macrophytes ofthe Adriatic basin (35 taxa) come fromtemperate and cold regions (NE Atlanticand NW Pacific) in relation with the majorvector of introduction (shellfish transfer).Only seven putative Lessepsian immi-grants (i.e. Acanthophora nayadiformis,Chondria pygmaea, Halophila stipulacea,Hypnea spinella, Hypnea valentiae, Lopho-cladia lallemandii and Ulva fasciata) havebeen hitherto registered, mainly from thenorthern coasts. However, Acanthophoranayadiformis and Hypnea spinella couldalternatively originate from the AtlanticOcean. The strains of Hypnea valentiae andUlva fasciata found in the Lagoon ofVenice may originate from the north-west-ern Pacific as in the Thau Lagoon.

3.3. Alien polychaetes in the Adriatic SeaThe Adriatic hosts only 22 alien poly-

chaete species (11 established, six casual andfive questionable), which represent 17% ofthe Mediterranean polychaete alien species.At least four species appeared exclusive ofthe Adriatic: Fabriciola ghardaqa, Megalom-ma claparedei, Platynereis australis and Syllisalose, the latter two being considered asquestionable taxa.

The established species are amongthose well known and settled in the wholeMediterranean area, and most of them arealso invasive, such as Branchiomma luctuo-sum, Ficopomatus enigmaticus, Hydroidesdianthus, H. elegans, but also Lysidice col-laris, Metasychis gotoi, Neopseudocapitellabrasiliensis, and Notomastus aberans. The

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most recently introduced species is B. luc-tuosum, present with very large populationsalong the Mediterranean coasts. At presentthis species seems to have reached also theBrazilian Coast of Sa~o Paulo, in the Bay ofSantos, in the vicinity of the largest seaportin Latin America (DE MATOSNOGUEIRA et al., 2006), underlining itsability to invade new areas, possibly trans-ported by ships. An introduction via theSuez Canal and possible transfer via ship-ping, can be postulated not only for B. luc-tuosum, but also for the other two Red Seasabellid species, exclusive of this area andcollected on the hard bottoms of the South-ern Adriatic coast: M. claparedei and F.ghardaqa that were, however, found onlyonce with very few individuals and consid-ered therefore casual (GIANGRANDE &MONTANARO, 1999; GIANGRANDE& LICCIANO, 2008). By contrast, theother sabellid Novafabricia infratorquata isa Caribbean species that must have beenintroduced from the Strait of Gibraltar,appearing established in the WMED andcasual in the Southern Adriatic along theApulian coast (LICCIANO &GIANGRANDE, 2006).

3.4. Alien crustaceans in the Adriatic SeaA total of 24 alien crustacean are

reported from the Adriatic (11 established,11 casual, two cryptogenic).

The first records of alien species intro-duced in the Northern Adriatic by mar-itime traffic date back to the years beforeWorld War I, when STIASNY (1908)recorded the capture of Plagusia squamosa(Herbst, 1790), Thenus orientalis (Lund,1793) and Portunus sanguinolentus (Herb-st, 1796), respectively in the harbours ofTrieste and Fiume (Rijeka), at that timethe main harbours of the Austro-Hungari-an Empire (BABIC, 1913). The latter two

species are excluded from our list follow-ing GALIL et al. (2002), whereas P.squamosa is kept as questionable becausethriving populations were recently report-ed from elsewhere in the Mediterranean:Tunisia and Libya (ZAOUALI et al.,2007).

Only three crabs (Callinectes sapidus,Dyspanopeus sayi and Rhithropanopeusharrisii) are definitely established in thearea and all three originate from theAtlantic coast of USA. The spreading ofthe two panopeid crabs in the Adriatic Seain recent years (FROGLIA & SPE-RANZA, 1993; MIZZAN & ZANELLA,1996; ONOFRI et al., 2008) has been facil-itated by the development of mussel aqua-culture both in lagoons and open sea, withtransfer of mussels seed and half grownmussels among aquaculture plants.

The latest record is that of the Indo-Pacific crab Charybdis lucifera, caught sixmiles off the Venetian coast (MIZZAN &VIANELLO, 2009).

The following species were recordedonly once and with single specimens: Mar-supenaeus japonicus, Scyllarus caparti,Callinectes danae, Charybdis lucifera, Eri-ocheir sinensis, Hemigrapsus sanguineus,and Charybdis japonica (FROGLIA, inpress).

Only three planktonic copepod specieswere recorded as aliens in the AdriaticSea, originating from the Atlantic andIndo-Pacific oceans.

3.5. Alien molluscs in the Adriatic SeaThe Adriatic Sea has possibly the old-

est alien (treated as cryptogenic) marinespecies in the Mediterranean Sea, viz. Lit-torina saxatilis, originally described fromVenice (OLIVI, 1792) before it could besuspected of being introduced from theNE Atlantic (JANSON, 1985). The Adri-

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atic holds a few endemic species, one ofwhich (Gibbula albida) has been intro-duced into the WMED and elsewhere.

The Adriatic holds only 27 alienspecies (15 established, nine casual andthree cryptogenic) but the striking charac-teristic is the high proportion of themwhich have become invasive. Togetherwith the Levantine basin, the Adriatic maybe the part of the Mediterranean whichhas been most transformed by the onset ofalien species. The most invasive speciesinclude Anadara kagoshimensis (formerlyknown as A. inaequivalvis), Musculista sen-housia, Rapana venosa and Ruditapesphilippinarum, all originating from thetemperate North Pacific and thereforecomfortable despite seasonal lows in seawater temperature. Anadara kagoshimen-sis, now coupled to the possibly AtlanticAnadara transversa, has formed spectacu-lar accumulations on the NE Italian coast(RINALDI, 1985) and has profoundlyimpacted the sandy infralittoral bottomswhich are a habitat for commercial speciessuch as the venerid Chamelea gallina.Farmed species include Crassostrea gigasand Ruditapes philippinarum, and the activ-ity related to farming may be responsiblefor an important part of the introductionsto and from the Adriatic; the latter speciesis invasive and likely to displace or to drivelocally extinct the native clam Ruditapesdecussatus. The Sidney rock oyster Sac-costrea commercialis was introduced to theVenice Lagoon in the 1980s but seems tohave dwindled and has not been reportedrecently (MIZZAN, 1999).

Twenty-one of the alien species knownfrom the Adriatic are recorded in Italy,and only the few tropical species Brachi-dontes pharaonis, Cellana rota, Melibeviridis and Halgerda wileyi have recordsexclusively along the eastern coast.

3.6. Miscellaneous invertebrates in theAdriatic Sea

Three alien species of Bryozoa (out of23 known for the whole Mediterranean)have been recorded in the Adriatic. TheIndo-Pacific species Tricellaria inopinatawas originally described on material com-ing from the Lagoon of Venice byD'HONDT & OCCHIPINTI-AMBROGI(1985). It has been subsequently found inmany European seas (DYRYNDA et al.,2000). The alien status of Bugula fulva issupposed based on its patchy occurrencenearly restricted to harbours(HAYWARD & MCKINNEY, 2002).

The only alien species of Ascidiacea isBotrylloides violaceus, introduced in theLagoon of Venice by the north-westernPacific by shellfish culture (ZANIOLO etal., 1998).

Alien Anthozoa are represented onlyby the north-eastern Atlantic species Diad-umene cincta (BIRKEMEYER, 1996),which in a sense reinforces the Northern-Adriatic ‘sub-Atlanticism’.

Porifera are represented by Paraleucil-la magna only (LONGO et al., 2007).

Fourteen alien species of Cnidaria(out of 46 for the whole Mediterranean)are known for the Adriatic. Alien Hydro-zoa are comprised of 13 species (out of37). Most of them have circum(sub)tropi-cal origin, but two estuarine species(Cordylophora caspia and Gonionemusvertens) prefer temperate waters. The firstMediterranean record of Garveia francis-cana comes from the Venice Lagoon(MORRI, 1982). It is debatable whetherCoryne eximia can be considered as alienspecies since the Mediterranean occur-rence of this species seems to need re-con-firmation (BOUILLON et al., 2004).

Two of the three alien species ofCtenophora known for the Mediterranean

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are established in the Adriatic: Beroe ovataand Mnemiopsis leidyi (SHIGANOVA &MALEJ, 2009). These authors refer earli-er citations of Beroe ovata in the Mediter-ranean to the native species B. cucumis,and report the real B. ovata as a WesternAtlantic species first introduced to theBlack Sea and possibly from there to theAdriatic.

Sipuncula number two alien species:both are circum(sub)tropical (PANCUCCI-PAPADOPOULOU et al., 1999). Theonly alien pycnogonid is Ammothea hilgen-dorfi, first reported for the Mediterraneanfrom the Lagoon of Venice (KRAPP &SCONFIETTI, 1983).

3.7. Alien fish in the Adriatic SeaAdding Fistularia commersonii

(DUL IC’ et al., 2008), 11 Lessepsian fishspecies had reached the Adriatic Sea by2009 (DRAGI EVIC’ & DUL IC’,2010). The occurrence of Terapon therapsin 2007 (LIPEJ et al., 2008) is of particularinterest since its record was the first for theMediterranean; furthermore, it was foundat great distance from its usual distributionarea. Similar are the cases of Pampusargenteus and Epinephelus coioides.

The catch of Elates ransonnetii inMarch 2010 in the eastern Adriatic(DUL IC’ et al., 2010) brought the num-ber of Lessepsian fish immigrants thatwere recorded in the Adriatic Sea to 12.Sphoeroides pachygaster, of tropicalAtlantic origin, has undoubtedly formed awell established population in the Adriat-ic (Albania, Slovenia, Montenegro, Croat-ia, Italy).

4. EASTERN MEDITERRANEAN

The EMED includes two major bodiesof water: the Levant Sea and the Aegean

Sea, together with the smaller Sea of Mar-mara, which connects it to the Black Sea.The Levant Sea is warmer than the rest ofthe Mediterranean and harbours a signifi-cant number of circumtropical species.Atlantic-Mediterranean elements andMediterranean endemics are comparative-ly scarce (MORRI et al., 2009).

Since the construction of the SuezCanal, the Levant Sea is experiencing animportant influx of Red Sea species. POR(1990) defined the geographical limits tothe expansion of Red Sea immigrants inthe Mediterranean as the ‘Anti-Psara line’to the north (Anti-Psara being an island inthe Aegean) and the Strait of Sicily to thewest: these boundaries match the 15 ÆCsurface isotherm for February (BIANCHI,2007). The Aegean Sea has localendemics, but the typical Mediterraneanbiota is impoverished with respect to theWMED (BIANCHI & MORRI, 1983);this, however, may be partly due to insuffi-cient inventory effort (MORRI et al., 1999;KOUKOURAS et al., 2001). The Sea ofMarmara exhibits peculiar hydrologicalconditions, with low salinity waters comingfrom the Black Sea stratifying over salinewaters of Mediterranean origin on the bot-tom (ÜNLÜATA et al., 1990). This hydro-logical regime should facilitate the diffu-sion of Black Sea species into the NorthernAegean rather than vice-versa, but ourknowledge on the exchanges between thetwo areas is limited and their biotic affini-ty is low (KOUKOURAS et al., 2001). Inrecent times, climatic change favoured anincrease of biotic penetration from the Seaof Marmara into the Black Sea, whichtherefore has been undergoing a processof ‘mediterranization’ (TOKAREV &SHULMAN, 2007).

Some scientists (e.g., TORTONESE,1973; OLIVERIO & TAVIANI, 2003;

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POR, 2009) have argued that this basin,particularly in its eastern stretches, suf-fered from an ‘ecological vacuum’ that isnow being filled by Red Sea immigrants.This idea has been criticized by GALIL(2007).

4.1. Alien protozoans in the EasternMediterranean

Fish pasteurellosis caused by Photo-bacterium damselae was reported from cul-tured seabass (Dicentrarchus labrax) inTurkey (CANDAN et al., 1996).

Among the total of 50 alienforaminiferal species presently known forthe Mediterranean, all except one arepresent in the EMED (LANGER &HOTTINGER, 2000). All of them havetropical affinity. Despite a significant num-ber of species with circumtropical distribu-tion (KOUKOUSIOURA et al., 2010b),most have Indo-Pacific origin and theirpenetration in the EMED is likely to havebeen favoured by the opening of the SuezCanal (MERI et al., 2007a; LANGER,2008). Off the west Turkish coast, alienforaminifers showed high abundance athot submarine springs that allowed higherbottom water temperatures in winter(MERI et al., 2010). Similar observa-tions on alien macrophytes in the Aegean(DE BIASI & ALIANI, 2003) and in theTyrrhenian Sea (GAMBI et al., 2008) sug-gest that shallow water hydrothermal ventsin the Mediterranean Sea may act as step-ping stones for the penetration of tropicalaliens (BIANCHI et al., in press).

Among the nine new taxa added tothose recorded by ZENETOS et al. (2008),eight occur in the EMED (LANGER,2008) and 4 of them have been reportedonly from single or few localities: Brizalinasimpsoni and Euthymonacha polita fromthe coasts of Turkey (MERI et al., 2010),

Pegidia lacunata and Pseudolachlanellaslitella (as Quinqueloculina eburnea) fromnorth-eastern Africa (BLANC-VERNETet al., 1979) and Cyprus (ABU TAIR &LANGER, 2010).

The new recovery of alien foraminifersmainly comes from recent researches car-ried out on living coastal assemblages ofthe EMED, which testify to the increasingscientific interest in alien and Lessepsianimmigrants. These studies confirm thewide distribution of many large endosym-biont-bearing taxa, such as Amphiste-ginids, along the coasts of the Levantinebasin (LANGER, 2008), Greece(TRIANTAPHYLLOU et al., 2009;KOUKOUSIOURA et al., 2010a), Turkey(MERI et al., 2010) and Cyprus (ABUTAIR & LANGER, 2010). Amphisteginalobifera appears to have established suc-cessful populations in Greek coastalecosystems (TRIANTAPHYLLOU et al.,2009; KOUKOUSIOURA et al., 2010a,b).Taxa of Indo-Pacific origin, probably trueLessepsian forms, are represented byBorelis sp., Coscinospira hemprichii, Cyclo-forina sp., Heterostegina depressa, Pegidialacunata, Pseudolachlanella slitella andSorites orbiculus.

Taxa undetermined at the specificlevel (Borelis sp., Cycloforina sp.) havebeen included as alien forms only whendescribed and figured in previous work onRed Sea or Indo-Pacific areas(HOTTINGER et al., 1993; LANGER &HOTTINGER, 2000; LANGER, 2008with references) or (Haddonia sp.) if per-tain to endemic Indo-Pacific genera(LOEBLICH & TAPPAN, 1988).

At the present state of studies, it seemsprudent to exclude some very seldomrecorded species cited by ZENETOS et al.(2008), yet very rare or poorly preserved,but also known at various latitudes or as

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fossils in the Mediterranean area. Thesespecies are here represented by Acervulinainhaerens, Iridia diaphana, Cymbaloporettasquammosa, Pyramidulina perversa andTriloculina affinis.

Schackoinella imperatoria, a smallspecies originally described from fossilmaterial of the Tertiary Basin of Vienna,pertains to a genus now known only in theIndo-Pacific Ocean (LOEBLICH &TAPPAN, 1988). Rare specimens havebeen recorded along the eastern coasts ofNorth Africa (BLANC-VERNET, 1969)as Glabratella imperatoria. More data areneeded to verify its introduction by ship-ping through the Suez Canal. Its rareoccurrence suggests a cryptogenic/casualacclimatization status.

In addition to all these benthic species,planktonic foraminifers are representedonly by Pulleniatina obliquiloculata, a ther-mocline dwelling species characteristic ofequatorial Atlantic and Indo-PacificOceans (BE & TOLDERLUND, 1971;KENNETT & SRINIVASAN, 1983). Itsoccurrence has been reported only byMERI et al. (2004) along the Turkishcoasts and its introduction was inferredfrom the Atlantic Ocean via Gibraltar(ZENETOS et al., 2008). Recently, P.obliquiloculata has been reported fromCanary Islands, Eastern Atlantic (WILKEet al., 2009). It is absent in the Red Sea, butit has been frequently reported from theArabian Sea and Bay of Bengal (BE &TOLDERLUND, 1971; CHEN &FARRELL, 1991, inter alii). More data arerequired to interpret its occurrence and toexclude its transport within ballast waters.

4.2. Alien macrophytes in the EasternMediterranean

A total of 63 alien macrophytes havebeen hitherto reported in the EMED

which represent 50% of the alien macro-phytes reported from the MediterraneanSea. This high percentage is mainly due tothe proximity to the Suez Canal and denseinternational ship traffic in the area.Among them, 35 alien macrophytes arewell established, nine are casual, 15 arecryptogenic/questionable, and four requireconfirmation. Casual taxa (i.e.Acrochaetium spp., Derbesia boergesenii,Padina antillarum, Rhodophysema georgii,Rhodymenia erythraea, Solieria dura andSpatoglossum variabile) might turn into theestablished status in the future. The cryp-togenic/questionable category includesboth temperate taxa mainly registered inthe north of the basin, i.e. the Sea of Mar-mara and the Northern Aegean Sea(Anotrichium okamurae, Chondriacoerulescens, Cladosiphon zosterae, Ecto-carpus siliculosus var. hiemalis, Halothrixlumbricalis, Polysiphonia atlantica, P.fucoides and Pylaiella littoralis), and tropi-cal taxa distributed in the Aegean Sea andthe Levantine Basin (Acanthophora nayad-iformis, Antithamnionella elegans, theAtlantic strain of Asparagopsis taxiformis,Caulerpa racemosa var. turbinata, Cerami-um bisporum, Ganonema farinosum andUlva fasciata). The presence of Antitham-nionella spirographidis, Asparagopsis arma-ta, Gracilaria arcuata and Hypnea flagelli-formis in the EMED requires confirma-tion. The latest record is that of the sea-weed Codium parvulum, recent blooms ofwhich were observed on the northernshores of Israel (ISRAEL et al., 2010).

On the 21 invasive or potentially inva-sive macrophytes introduced into theMediterranean (Table 3), 11 are reportedin marine benthic habitats of the EMED.No data are available for coastal lagoons.Cladophora cf. patentiramea and Codiumparvulum are hitherto known only for the

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EMED, for Cyprus and Israel, respectively.The majority of alien macrophytes of

the EMED (63%) come from tropicalregions, mainly the Red Sea and the Indo-Pacific Ocean. The major vector of intro-duction is the Suez Canal. Hitherto 35putative Lessepsian immigrants (56% ofthe total) have been registered, of whichCladophora cf. patentiramea, Codiumparvulum, Halophila stipulacea, Lophocla-dia lallemandii and Stypopodium schimperiare considered invasive. Acanthophoranayadiformis, Ganonema farinosum andUlva fasciata were found in the Mediter-ranean before the opening of the SuezCanal in 1869, but the occurrence, at leastin the EMED, of exotic strains comingfrom the Red Sea and the Indian Oceanvia the Canal is highly probable today. Theother alien macrophytes reported from theEMED come from cold temperate regions(NE Atlantic and NW Pacific) and wererecorded along the north coasts of Greeceand Turkey (Sea of Marmara and North-ern Aegean Sea). Some of these lastmacrophytes, i.e. Ectocarpus siliculosusvar. hiemalis, Polysiphonia fucoides andPylaiella littoralis, also occur in the BlackSea (TA KIN et al., 2008): so, a native sta-tus cannot be excluded in the northernpart of the EMED.

4.3. Alien polychaetes in the EasternMediterranean

A total of 98 alien polychaete specieshave been reported to date from theEMED, almost 75% of the alien poly-chaete species reported from the Mediter-ranean Sea, mainly due to the proximity tothe Suez Canal and dense internationalship traffics in the area. Among them, 52species have been well established in habi-tats of the EMED, 16 species are casual,two species (Chaetozone corona and Para-

prionospio coora) are cryptogenic and 28species are questionable. 22 species seemto have invasive character (see Table 3),forming dense populations in semi- orheavily polluted areas (including har-bours) or shallow water benthic habitats.There is a relatively high number of ques-tionable species in the area. They includethe species only listed in ecological papers(i.e. Cossura coasta, Loimia medusa), thoseidentified as ‘cf.’ in the area (i.e. Oenone cf.fulgida, Syllis cf. mayeri) or whose descrip-tions based on the Mediterranean speci-mens were insufficient (i.e. Lepidonotuscarinulatus), hindering us in deciding theirreal taxonomic positions. Future studies tobe performed in the region would enableus to understand if these species reallyexist in the Mediterranean. A total of tencasual species (i.e. Laonome elegans,Nereis gilchristi) might turn into the estab-lished status if data regarding them areaccumulated. The native species, Chaeto-zone setosa (in part) and Paraprionospiopinnata, previously widely reported fromthe Mediterranean Sea in polluted anddeep waters, have been recently re-identi-fied as C. corona and P. coora, respectively( INAR & ERGEN, 2007; YOKOYA-MA et al., 2010). These species wereregarded as cryptogenic because the dataon their distributions are very limited; theyare relatively newly described species andcould have been in existence in the area formany years.

The Levantine Sea is densely colo-nized by alien polychaete species. In theregion, a total of 77 species (79% of thealien polychaetes from the EMED) havebeen hitherto reported. A total of 11species are categorized as questionable.The highest number of alien species (58species) are known from the Levantinecoast of Turkey, followed by the Israeli (29

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species) and Egyptian (28 species) coasts.On the Syrian coast, only two alien species(Spirobranchus tetraceros and Spirorbismarioni) were encountered. The majorityof species are Lessepsian immigrants (47species, 61% of total species), the otherswere introduced to the area via ships,except for Chaetozone corona which is acryptogenic species. A total of 66 species(86%) could have been originated in theRed Sea and Indo-Pacific areas. The oth-ers (11 species) were introduced to thearea from the Atlantic Ocean (mainlyfrom the western part).

The Aegean Sea is on the route of theLessepsian species that have become wellacclimatized to the Mediterranean envi-ronment and have a large dispersal capac-ity. Up to date, 39 alien polychaetes werereported from the Aegean Sea, of which 16species are Lessepsian immigrants. In thearea, 20 species have been well estab-lished, one species, Prionospio (Minuspio)pulchra, is casual, 16 species are question-able and two species, Paraprionospio cooraand Chaetozone corona, are cryptogenic.Ship-transported species dominate semi-or highly polluted areas, especially nearthe large international harbours. Theopportunistic species Malacoceros fuligi-nosus and Capitella spp. seem to have beenreplaced by the North Atlantic speciesPolydora cornuta and Streblospio gyno-branchiata and the Pacific species Pseu-dopolydora paucibranchiata ( INAR etal., 2005, 2006a). In the eastern part of theAegean Sea, 19 alien polychaete specieswere found, of which two species are ques-tionable. In the western part, 31 specieswere reported, of which 14 species arequestionable.

Polychaetes from the Sea of Marmarahave been recently reviewed by INAR(2010). He postulated that a total of 17

species of alien status were reported fromthe region. However, three species wereexcluded from the list, eight species arequestionable and six species (Nereis persi-ca, Paraprionospio coora, Polydora cornuta,Streblospio gynobranchiata, Desdemonaornata and Ficopomatus enigmatus) havebeen established. The report of theLessepsian species N. persica byRULLIER (1963) needs to be confirmed.The other species could have been intro-duced to the area by ships.

4.4. Alien crustaceans in the EasternMediterranean

A total of 119 alien crustaceans havebeen reported in the EMED among whichsix are cryptogenic species (Cymadusafilosa, Gammaropsis togoensis, Mono-corophium sextonae, Pseudocalanus elonga-tus, Synalpheus tumidomanus africanus andThalamita poissonii). Introduced crus-taceans belong to Amphipoda (10), Cirri-pedia (5), Copepoda (33), Cumacea (3),Decapoda (58), Isopoda (7), Tanaidacea(1) and Stomatopoda (2). These numbersinclude both benthic and pelagic forms ofcopepods as well as parasites among cirri-pedes, copepods and isopods. The vastmajority of them occur in the LevantineSea (e.g. 68% of the benthic forms, allplanktonic taxa and all parasites). Twothirds of the species have colonized theLevantine and are spreading to theAegean, while almost one third is knownfrom single records. Very few species, suchas the decapods Calappa pelii, Metape-naeus affinis and Sirpus monodi, are limit-ed to the Aegean Sea. Reasoning for therelatively high number of questionablecopepod species (ca. 10%) is provided inZENETOS et al. (2005, 2008).

With the exception of a few speciesoriginating in the Atlantic (Calappa pelii,

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Callinectes sapidus, Farfantepenaeusaztecus, Necora puber, Percnon gibbesi, Pro-cessa macrodactyla, Sirpus monodi,Synalpheus tumidomanus africanus andThalamita indistincta), the native range offour-fifths of the alien crustacean in theMediterranean is in the Indo-PacificOcean, Indian Ocean and Red Sea. Thepathway of introduction of decapodspecies is the Suez Canal, but penetrationhas been either unintentional (Lessepsianmigration) or ship-mediated. Shippingappears to be also responsible for theimmigration of species of Atlantic origin.However, spreading of self maintainingpopulations from the CMED appears tobe the mode of introduction of the westAtlantic crab, Percnon gibbesi now presentin Greece, Turkey, Syria, Cyprus, Israel(KATSANEVAKIS et al., in press and ref-erences therein) and Egypt (AZZURROet al., in press). Thalamita poissonii couldbe a Lessepsian species, but there arerecords of it before the Suez Canal open-ing (UDEKEM D’ACOZ, 1999), whichcould support a pre-Messinian Mediter-ranean presence. If so, the alien statusshould be downgraded to ‘questionable’.Perhaps at this moment it is best to con-sider it as ‘crypogenic’.

The latest additions to the inventory ofthe marine decapod species in the EMEDare the pilumnid crab Eurycarcinus inte-grifrons (ÖZCAN et al., 2010), the redswimming crab Gonioinfradens pauciden-tata (CORSINI-FOKA et al., 2010), bothof Indo-Pacific origin, and the westernAtlantic shrimp Farfantepenaeus aztecus(CENZIG et al., 2010).

The Indo-Pacific stomatopod Eru-gosquilla massavensis thrives along theLevantine coast and is expanding in theAegean (ÖZCAN et al., 2009).

Among the planktonic crustaceans,

information is available almost exclusivelyfor copepods, since it is the most studiedgroup (as in the entire MediterraneanSea). Most of the newcomers are Lessep-sian immigrants, two originating in theAtlantic (Calanopia biloba, Paracartiagrani), while the origin of Acartia tonsa,Arietellus pavoninus, Centropages furcatusand Parvocalanus crassirostris is question-able, since they are distributed in tropicalto subtropical waters both of the Atlanticand Indian oceans and even of the Pacific.The species Pontellina plumata, Corycaeusspeciosus and Subeucalanus crassus wereexcluded from the list of alien speciesgiven by ZENETOS et al. (2005) afterreconsideration of the available informa-tion on their distribution (RAZOULS etal., 2005-2010). The consideration ofSubeucalanus subcrassus as alien is ques-tionable because it is present in most areasof the Mediterranean Sea as well as in thetemperate to tropical waters of theAtlantic and Pacific oceans. The presenceof Spinocalanus terranovae in the EMEDseems questionable, given its restricteddistribution in Antarctic and subantarcticareas. Regarding the records of Triconiarufa (originating from the Red Sea) in theEMED (Lebanon) by MALT et al. (1989),BÖTTGER-SCHNACK & SCHNACK(2009) assumed that the previous authorsmay have confounded the specimens withTriconia sp. 8. Finally, Pseudocalanus elon-gatus should be considered as cryptogenicin the Mediterranean Sea: it has beeninterpreted as a boreal relict species(FURNESTIN, 1979) or an immigrant,either from the Black Sea (SIOKOU-FRANGOU, 1985) or the north-easternAtlantic (VIVES et al., 1981).

Two introduced cymothoid species ofIndo-Pacific origin, Anilocra pilchardi andCymothoa indica, typically parasitic of

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teleost fish, have been reported fromLebanon (BARICHE & TRILLES, 2006;TRILLES & BARICHE, 2006) parasitiz-ing mainly barracudas (Sphyraenidae).

Three parasitic copepods new to theMediterranean fauna were reported inwaters off the Egyptian coast near Alexan-dria by EL-RASHIDY & BOXSHALL(2009). Two of them, Mitrapus oblongusand Clavellisa ilishae, are of Indo-Pacificorigin and are considered to have co-invaded the Mediterranean through theSuez Canal on Red Sea immigrant hosts.The third parasite, Nothobomolochusfradei, was previously known from the Gulfof Guinea and the Arabian Gulf.

A rhizocephalan barnacle, Heterosac-cus dollfusi, followed its portunid hostcrab, Charybdis longicollis, from the RedSea through the Suez Canal to theMediterranean Sea.

4.5. Alien molluscs in the Eastern Mediter-ranean

The EMED has the lion’s share ofalien molluscs, building up impressivenumbers and by far the highest proportionworldwide. To date there are 190 alienmolluscs reported (105 established, 64casual, 15 questionable and six crypto-genic), to be added to around one thou-sand native species. This represents morethan 90% of all alien mollusc speciesreported in the Mediterranean as a whole,a percentage that remains remarkably sta-ble if we consider established species only.Six species are classified as cryptogenic(Alvania dorbignyi, Aplysia parvula, Atysangustatus, Discodoris lilacina, Parviturbodibellai, Teredo navalis), as there aredoubts on whether they are native or intro-duced or perhaps represent very old intro-ductions.

Predictably, the vast majority of these

species are of Indo-Pacific origin and theirpresence is related to the proximity of theRed Sea and to the Suez Canal connec-tion. Only six species (Siphonaria pectinata,Mya arenaria, Cerithium litteratum,Anadara transversa, Mytilopsis sallei, andPetricola pholadiformis) are Atlantic orAtlanto-Mediterranean in origin. Exam-ples of Indo-Pacific species yet non-Lessepsians, among the now well-estab-lished species, are the strombid Cono-murex persicus, first recorded in Turkeyand only later arrived on the Levantinecoast, or the opisthobranch Melibe viridis,first recorded in Greek waters. Some ofthese species, like Conomurex persicus, arenot even known from the Red Sea andtheir introduction must therefore haveinvolved maritime traffic at some stage.

POR (2009, 2010) expressed the viewthat the thermophilic species of Indo-Pacific origin should not be viewed asaliens but rather as a restoration of theTethyan faunal realm as it existed previ-ously to the closure of the communicationwith the Indian Ocean in the late Miocene.

Another quite distinct pool is made upby species which originate from the tem-perate North Pacific, rather than from thetropical areas, and whose introduction islinked with transfers of living molluscs foraquaculture and/or maritime traffic.

The EMED has comparatively limitedaquaculture activity and this categorywould hold, apart from the culturedspecies proper Crassostrea gigas and Rudi-tapes philippinarum, very few species suchas Rapana venosa, Anadara kagoshimensisor Musculista senhousia.

A particular aspect regarding molluscsis the high incidence of unsupportedrecords, often based on single shells andcarried perpetuated in subsequent check-lists. The rationale for their exclusion has

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been detailed in GOFAS & ZENETOS(2003). We have not considered in thisreport the records for 32 additionalspecies, including those found on an oil rig(MIENIS, 2004), among them oysters andother bivalves firmly attached to their sub-strate, considering that these have not yetbeen found in the wild.

The Levantine Sea is the area whichholds the highest number of reports, with149 species (83 established, 45 casual, 5cryptogenic, 16 questionable); historicallythis is the area where pioneer reports weremade on alien species of Indo-Pacific ori-gin (e.g. GRUVEL & MOAZZO, 1931;HAAS, 1937).

There are 48 of the species recordedfrom Israel, Lebanon and Syria, whichhave a record in the Suez Canal and maybe safely qualified as Lessepsian, againstonly seven (Melibe viridis, Murchisonellacolumna, Sticteulima cf. lentiginosa,Smaragdia souverbiana, Conomurex persi-cus, Zafra savignyi, Zafra selasphora)species of tropical Indo-Pacific originwhich are proven non-Lessepsians (whoseprogression started elsewhere than on theLevantine coast and have no Suez Canalrecord). The remainder cannot be posi-tively assessed but the ratio of Lessepsianto non-Lessepsian among them shouldprobably remain high.

The coasts of southern Turkey andCyprus are the next largest centres wherealien species have been reported, but therethe exploration is much more recent andthe fauna had remained virtually unknownuntil the 1980s. Here the totality of thespecies is of tropical Indo-Pacific origin(save the cryptogenic species) but theshare is more balanced between Lessep-sian and non-Lessepsian. For the coast ofsouthern Turkey, we have 32 provenLessepsians previously recorded in the

Suez Canal and along the Levantine coast,all established except the casual Psam-motreta praerupta versus 20 species of thetropical Indo-Pacific which are definitelynot Lessepsian and started their Mediter-ranean settlement in the area.

The Aegean Sea is the natural route ofprogression for the most successful Indo-Pacific invaders, both Lessepsian andthose introduced first in the southernTurkey/Cyprus area. Nevertheless it hasalso its own pools of species introducedthrough other pathways. Contrary to thesouth coast of Turkey, there are severalspecies that originate from the Atlantic(Anadara transversa, Crepidula fornicata,Cerithium litteratum, Petricola pholadi-formis, Mya arenaria) or the temperateNorth Pacific (Ruditapes philippinarum,Rapana venosa) and are involved neitherin the Lessepsian migration nor in the SETurkey focus of introductions. Siphonariapectinata, introduced in the Gulf ofSaronikos, is a clear example of intra-Mediterranean introduction.

As may be derived from this, theimportance of Lessepsian species is mod-erate, and even the proven Lessepsianspecies found there may be secondaryintroductions from populations settledelsewhere in the EMED.

Among the 18 species which couldhold as Lessepsian because they wererecorded in the Suez Canal, species likeCellana rota, Fulvia fragilis, Murexforskoehlii, first showed up in theSaronikos Gulf heavily impacted by mar-itime traffic: Cerithium scabridumappeared in Greece (ZENETOS et al.,2009) much later than in Sicily, where itwas obviously a secondary introduction.Only a few Lessepsian species such asPinctada radiata or Trochus erithreus mayhave arrived in the Aegean as a result of

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regular progression inside the Mediter-ranean.

4.6. Miscellaneous invertebrates in theEastern Mediterranean

Fourteen alien species of Bryozoa, outof the 23 recorded in the whole Mediter-ranean, are known for the EMED. Most ofthem are of Indo-Pacific origin, and mostare established. Three recently describedspecies from Lebanon have presumablypenetrated from the Red Sea: this seemssufficiently documented for Parasmittinaserruloides and P. spondylicola(HARMELIN et al., 2009) but the statusas a Levantine endemic cannot be exclud-ed for Schizoretepora hassi (HARMELINet al., 2007).

Seven of the 16 alien species of Ascidi-acea known for the Mediterranean areestablished in the EMED. They are inmajority Indo-Pacific, or alternatively cir-cumtropical. Rhodosoma turcicum,Ecteinascidia thurstoni, Phallusia nigra,Ascidia cannellata, Symplegma brakenhiel-mi, Herdmania momus, and probablyMicrocosmus exasperatus are regarded asRed Sea immigrants ( INAR et al.,2006b; BITAR et al., 2007; IZQUIERDO-MUN~OZ et al., 2009; KATSANEVAKISet al., 2009; SHENKAR & LOYA, 2009;KONDILATOS et al., 2010).

Cnidaria are represented by 32 alienspecies out of 46 known for the wholeMediterranean. Anthozoans are repre-sented by two species: the Indo-Pacificgorgonian Acabaria erythraea has beenfound in the harbour of Hadera, Israel(FINE et al., 2004), whereas the scleractin-ian coral Oculina patagonica, possibly orig-inating from the SW Atlantic, has beenrecorded from many sites through thewhole basin (BITAR & ZIBROWIUS,1997; FINE et al., 2001; INAR et al.,

2006b; SALOMIDI et al., 2006). Paradoxi-cally no Red Sea coral species has yet pen-etrated in the EMED (BIANCHI et al.,2010). In the case of the Hydrozoa, theEMED harbours two thirds of the alienspecies hitherto known for the Mediter-ranean (26 out of 37) in accordance withwhat is known for virtually all the othergroups of organisms, which typically showa preponderance of aliens in that basin.Among the 26 alien planktonic hydrozoa,17 were recorded in the EMED. Clytia lin-earis and Eirene viridula are the only estab-lished alien species in the entire Mediter-ranean Sea. Most of the alien Hydrozoa inthe EMED have a circumtropical distribu-tion, the remainders have Indo-Pacific ori-gin. Macrorhynchia philippina is the mostwidespread and abundant (MORRI,2008). Four Scyphozoa aliens known forthe Mediterranean supposedly came inthrough the Suez Canal and are now estab-lished along the coast of Israel (GALIL etal., 1990, 2010). Cassiopea andromeda hasextended its penetration range to Turkey(ÖZGÜR & ÖZTÜRK, 2008) andGreece (ZENETOS et al., in press). Abylatrigona is the single alien siphonophore inthe Mediterranean Sea, found in Egypt(ZAKARIA, 2006).

After having successfully invaded theBlack Sea, with severe impact on both thenative ecosystem-functioning and the fish-ery yield, the Western Atlantic combjellyMnemiopsis leidyi started to spread in theMarmara and North Aegean Sea andreached Israel in 2009 (GALIL et al.,2009). Its predator Beroe ovata followedthe same path and arrived in the AegeanSea (SHIGANOVA et al., 2007). Amongthe three alien ctenophore species in theentire Mediterranean Sea, Sulculeolariaangusta was found only in the Egyptiancoasts (ZAKARIA, 2006).

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Seven species of alien Porifera withIndo-Pacific origin have been traditionallyconsidered as Red Sea immigrants(TSURNAMAL, 1969). More recently,this has been questioned by VACELET etal. (2007), who evidenced taxonomic prob-lems with all of them except perhaps Geo-dia micropunctata. According toVACELET et al. (2007), sponges withtropical affinities hitherto known only forthe EMED are more likely to be remnantsof the thermophilic biota that thrived inthe Mediterranean during warmer periodsof the late Pliocene or Pleistocene and thatdisappeared from the rest of the basin dur-ing the Würmian cooling (BIANCHI et al.,2010).

All six alien species of Sipunculaoccurring in the Mediterranean have beenrecorded for the EMED. Phascolion(Isomya) convestitum and Phascolosomascolop are of Indo-Pacific origin, theremaining four are circumtropical(PANCUCCI-PAPADOPOULOU et al.,1999). All are known from a single or fewlocalities, where they can however beabundant (A IK, 2007; 2008).

KRAPP et al. (2008) reviewed the Pyc-nogonida of the EMED: three species arealien, although the status of Pigrogromitustimsanus is questionable. Anoplodactyluscalifornicus and A. digitatus are estab-lished.

The echinoderm fauna of the EMEDincludes eight alien species (out of twelvefor the whole Mediterranean). Most ofthem are likely to have penetrated throughthe Suez Canal, but Asterias rubens origi-nated from the North Atlantic and wasfirst introduced in the Black Sea(KARHAN et al., 2008). Two conspicuousIndo-Pacific species, Diadema setosum(YOKES & GALIL, 2006) and Synaptulareciprocans (ANTONIADOU &

VAFIDIS, 2009) might exert a consider-able impact on Mediterranean ecosystems.

The EMED hosts the vast majority ofthe alien platyhelminthes, all reported asfish parasites, all with an Indo-Pacificand/or Red Sea origin. Of the nine mono-geneids introduced into this basin, five areconsidered established. Both host and par-asite are Lessepsian immigrants that haveco-invaded the Mediterranean Sea via theSuez Canal. The first documented case ofa monogeneid invading a new biogeo-graphical region by ‘natural’ extension ofits host range is that of the gill ectoparasitePolylabris cf. mamaevi infecting the rabbit-fish Siganus rivulatus. The greater abun-dance of P. cf. mamaevi in the invading(Mediterranean) populations is probablydue to the changed, new environment,possibly impacting on host resistance tothe parasite and encouraging heavierinfections (PASTERNAK et al., 2007). Apossible explanation for the unexpectedcolonization success of immigrant fish par-asite species belonging to the class Myx-osporea, despite their putative complexlife cycles, is discussed in DIAMANT(2010).

4.7. Alien fish in the Eastern Mediter-ranean

A total of 106 alien fish (nearly 70% ofthe alien fish known in the Mediterranean)has been reported from the EMED, themajority of which originate from the Indo-Pacific and Red Sea regions. Details ontheir distribution per country, origin andmode of introduction are provided inGOLANI & APPELBAUM-GOLANI(2010). New data extending the distribu-tion of already reported alien species inthe Levantine basin include: Pisodonophissemicinctus and Pomadasys stridens(BILECENOGLU et al., 2009); Apogon

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smithi and A. queketti (GOREN et al.,2008); Enchelycore anatina, Lagocephalusspadiceus and Lagocephalus suezensis(KATSANEVAKIS et al., 2009).

The complicated flood of confamilialspecies is prominent and deserves specialinterest, as it is unlikely to be coincidental.Alien cardinal fishes represent a goodexample of this, since only a single specieswas known for over 60 years and fourspecies have been added to the fauna with-in the last five years (GOREN et al., 2009a;GOREN et al., 2010; TURAN et al., 2010).Two species of the family Champsodonti-dae were observed from Turkey (Champ-sodon nudivittis) and Lebanon (Champ-sodon vorax), interestingly both withoutrecords from the Red Sea ( I EK &BILECENOGLU, 2009; BARICHE,2010a). Among recent alien gobies, Van-derhorstia mertensi was found off southernTurkish coasts and became quite commonshortly after its first observation(BILECENOGLU et al., 2008; YOKES etal., 2009), while the far-east chameleongoby Tridentiger trigonocephalus is report-ed from Israel (GOREN et al., 2009b).Sudden occurrence of two alien angelfish-es, Pomacanthus imperator and Pomacan-thus maculosus, from the Israeli andLebanese coasts, respectively, is also note-worthy (BARICHE, 2010b; GOLANI etal., 2010). Among the latest records is thatof the Indo-Pacific goby Trypauchen vaginain Israel (SALAMEH et al., 2010).

The appearance of Mycteroperca fusca,a large Atlantic fish, in Israel in 2010, andwithout having been spotted on the way,raises the question of the route of itsarrival. A reasonable possibility is that M.fusca entered the Mediterranean throughthe Strait of Gibraltar, as many Atlanticspecies do, then expanded its distributionalong the North African coast but was

overlooked or confused with Mycteropercarubra. Alternatively, it was introduced inthe ballast waters of a ship (HEEMSTRAet al., 2010).

5. PHYTOPLANKTON

The introduction of marine microal-gae in the Mediterranean Sea is hard todocument. Theoretically, these microscop-ic organisms can easily be transported bycurrents, but also by the feet of migratorybirds, or introduced through human com-merce of marine animals, ship’s ballastwaters or fouling organisms. The conse-quences of a microalgal invasion are onlyevident if there are conspicuous changes inan area, e.g. water discoloration ormucilage, or where the species is impliedin toxicity events or other harmful impacts(ZINGONE & WYATT, 2005), whileinvasions go totally unobserved in case ofrare species, which are a conspicuous partof the phytoplankton biodiversity in allseas. In addition, to prove that a species isan alien requires very sound backgroundknowledge of the species of a given area.

Unfortunately, the diversity of marinemicroalgae is scarcely known in wide areasof the Mediterranean Sea, e.g. the south-ern shores, where only a few sites havebeen investigated, or the offshore waters,where studies are limited to occasionalsampling during cruises. Even in the north-ern Mediterranean waters the knowledgeof the distribution of these unicellularorganisms in a given area is far from beingexhaustive, for several reasons. First, phy-toplankton is difficult to sample properly,as most species have an ephemeral occur-rence and a discontinuous spatial distribu-tion. The resolution of sampling programsrarely matches the space and time scale ofoccurrence of microalgae, and new species

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show up when more detailed or integrativesampling approaches are applied. Forexample, many cysts of the toxic dinofla-gellate Alexandrium andersonii were foundin sediment traps deployed in the Gulf ofNaples (MONTRESOR et al., 1998). Thespecies was not known from the Mediter-ranean Sea before, and has never beenrecovered in any other plankton study inthe area since. Probably Alexandriumandersonii appears in the plankton for veryshort periods, or it is restricted to specificdepths of the water column that are notcovered during investigations. Anothermajor problem concerns all organisms thatare difficult to preserve and either breakor loose their peculiar characteristics withcommon fixatives such as formalin andlugol. Micromonas pusilla, a 2-3 Ìm nakedprasinophyte, is easily identifiable when itgrows in cultures due to a very typicalswimming pattern. It can reach concentra-tions of up to 106 cells l-3 in coastalMediterranean waters in winter(THRONDSEN & ZINGONE 1994;ZINGONE et al., 1999). However, its cellsgenerally burst upon preservation. This isalso the case for much larger flagellates,e.g. the raphidophyte Chattonella subsalsa(30-40 Ìm) or naked dinoflagellates, whichexplode or lose their shape when fixed.Finally, many species require particularskills or methods to be identified. This isthe case for cryptomonads, generally veryabundant in inshore waters, which requirecultivation, electron microscopy and attimes molecular techniques to be identi-fied at the species level. While their abun-dance as a group can be traced with pig-ment analyses (HPLC) that allow alloxan-thin to be detected, or by flow-citometry,individual species are only identified bycombining a number of different tech-niques (CERINO & ZINGONE, 2006).

For the reasons stated above, it canrarely be excluded that a suspectedmicroalgal invader was already present aspart of the rare, hidden and unsampledphytoplankton. These factors are probablyat the origin of the extremely low numberof proven invaders among phytoplanktonspecies (WYATT & CARLTON, 2002).

Nonetheless, a number of cases of pos-sibly introduced microalgal species in theMediterranean Sea have been brought tothe general attention over the last fewyears. One of the first cases was providedby the toxic planktonic dinoflagellateAlexandrium catenella, appearing in highconcentrations in a French coastal lagoon,the Thau Lagoon, in 1998 (LILLY et al.,2002). Based on the molecular similarity ofthe species with strains from Japan, it wasproposed that the species had been trans-ported recently to the Mediterranean Sea.The species had been seen before in theBalearic Sea in 1983 (MARGALEF &ESTRADA, 1987), along the Spanish coastsince 1996 (VILA et al., 2001) and alongthe Sardinian coast in 1999 (LUGLI etal., 2003). Apparently its range is expand-ing across the Mediterranean Sea, as it hasbeen found recently on the Tunisian coast(TURKI & BALTI, 2007). On the otherhand, more recent studies using alternativemolecular tools have questioned the closerelationships between Japanese andMediterranean populations, and hence theidentification of Japan as the source area ofA. catenella (MASSERET et al., 2009).

While the knowledge of planktonicspecies distribution poses many problemsin terms of sampling and detection, benth-ic and epiphytic microalgae should be easi-er to collect. Yet these organisms havebeen studied very rarely in the Mediter-ranean Sea until the end of the last centu-ry, when they became popular mainly due

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to health and environmental problemscaused by a toxic species Ostreopsis cf.ovata, which forms conspicuous blooms inmany coastal areas (e.g., TOTTI et al.,2010; MANGIALAJO et al., in press). Alsoin this case, the sudden emergence of aconspicuous problem was initially attrib-uted to a new invasion, probably in relationto climate change, as Ostreopsis specieswere previously known mainly from tropi-cal areas. However, an Ostreospis speciesdid form a conspicuous bloom in Ville-franche Bay in 1972, where macroalgaeappeared to be covered by a brown mucouslayer (Taylor, pers. comm.), but it was justrecorded due to the curiosity of an experttaxonomist swimming in the area. Thespecies was reported as O. siamensis, theonly species known at the time (TAYLOR,1979), but it was likely to be the same as theone reported in more recent years in thesame area, i.e. O. cf. ovata. From themolecular point of view, the Mediter-ranean populations of this species wereknown to be similar only to Brazilian pop-ulations (PENNA et al., 2010), but compa-rable ribotypes have very recently beenfound also in Japanese waters (SATO et al.,2010). Molecular analyses on other mark-ers are required to establish where, amongthe three sites, the molecular diversity ofOstreopsis cf. ovata is the highest, whichwould indicate the source site for these dif-ferent populations around the world.

Species of the benthic dinoflagellategenus Gambierdiscus, which produce tox-ins that are responsible for the neurologi-cal syndrome ciguatera, had never beenrecorded in the Mediterranean Sea until2003, when specimens were found inGreek waters off Crete (ALIGIZAKI &NIKOLAIDIS, 2008). No other record isconfirmed from anywhere else in theMediterranean Sea, despite the intensive

microphytobenthos sampling carried outto study O. ovata at several sites over thelast decade. In this case, an introductionfrom the Indian Ocean could be hypothe-sised, but the source population would bemissing for the time being, as the taxo-nomic identity of the species is still uncer-tain. On the other hand, data on benthicmicroalgae are now more complete than10 years ago, which would allow one to testa predictable range expansion westward.Indeed, marginal range expansion havebeen observed for other species, such asAlexandrium catenella (LUGLI et al.,2003), G. catenatum (G MEZ, 2003), anda number of diatom species (G MEZ,2008), which could be related to the cli-matic changes observed in the Mediter-ranean Sea or to other unexplained long-term fluctuations in species abundance.

A sounder background knowledge onphytoplankton distribution is available forplaces where time series exist and accurateidentification is performed over a long-term period. At the LTER-MC station inthe Gulf of Naples, two species neverrecorded despite their relatively easy iden-tification, have suddenly appeared inrecent years. These were Skeletonema trop-icum (ZINGONE et al., 2003; KOOISTRAet al., 2008), which is easily identifiable asit is the only species in the genus havingmany chloroplasts, and Pseudo-nitzschiamultistriata (ZINGONE et al., 2003),which has a typical sigmoid shape. The twospecies had never been recorded before1995 and 2002, respectively, and haveshown increasing concentrations sincetheir first finding. In both cases, an intro-duction could be hypothesised but it can-not be excluded that these are other exam-ples of marginal range expansion from thesouthern Mediterranean Sea shore north-ward, or that these species were extremely

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rare for many years, suddenly findingappropriate environmental conditions toincrease their abundance. The latterhypothesis takes into consideration long-term fluctuations in species abundanceand is partially supported by the absenceof discoveries since 1992 of anotherPseudonitzschia species, P. subpacifica.This species was relatively common in the1984-1987 period in the Gulf of Naples,then it became rarer and eventually disap-peared. Although P. subpacifica could stillbe in the Gulf of Naples, this case indicatesthat there is always the possibility of unex-plained local extinctions, and more gener-ally points to important changes in speciesabundance over the long term.

Another relevant case is that of speciesshowing discontinuous geographicalranges. The dinoflagellate Alexandriumbalechii is only found in the Gulf of Mexicoand in Tyrrhenian waters, suggesting thatthe species originated in one of the twoplaces and was transported to the other,although there are no data to establishwhere the species should be considered analien. Several other dinoflagellates are onlyfound in the Indo-Pacific area and in theMediterranean Sea so far (G MEZ,2006), but the suspicion that they wereintroduced through the Suez Canal shouldbe substantiated when their absence corre-sponds to their actual absence in theAtlantic Ocean and not to our limitedknowledge of protist biogeography. Forsimilar reasons, the high number of flagel-late species that have been described in theMediterranean Sea and never recordedelsewhere cannot be considered as proof oftheir endemism in the Mediterranean Sea(THRONDSEN & ZINGONE, 1994).

In synthesis, in the Mediterranean Seaa number of planktonic and benthicmicroalgae has appeared over the last

decades and continue to do so, formingconspicuous blooms in places where theywere not recorded at all before. However,whether these species are really alienshould be tested effectively, considering atleast two alternative hypotheses: 1) thatthey belong to the rare component of theplankton and that they were missed in pre-vious studies; 2) that their range expandedfrom unexplored southern or easternMediterranean areas.

6. INVASIVE SPECIES

The ongoing incoming of immigrantsand their increasing success in theMediterranean basin imply a profoundand continuous alteration in the speciesdistribution patterns that seems to haveaccelerated in the last decade. Up torecent times the influence of these immi-grants was geographically limited to theareas close to the source of introduction,but today the idea of biogeographicalboundaries to Lessepsian invasions (POR,1981; QUIGNARD & TOMASSINI,2000) seems to be overcome.

The most invasive species per MSFDare listed in Table 3; their share per taxo-nomic group is depicted in Figure 2.

A total of 134 species are classified asinvasive or potentially invasive in theMediterranean. The present list is based ondata on worst invasive marine species col-lated from various sources for the Mediter-ranean (STREFTARIS & ZENETOS,2006; GALIL, 2007; STREFTARIS et al.,2008), European Seas (EEA, 2007b;DAISIE: VIL et al., 2008), and globally(Global Invasive Species Database), man-aged by the Invasive Specialist SpeciesGroup (ISSG: www.issg.org). The list wasenriched with species such as the bigfin reef-squid Sepioteuthis lessoniana, an invader

A

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Table 3Distribution of invasive (++) or potentially invasive alien species (+) in the Mediterranean MSFD

areas. (-) Species absent from the area, *marketable species.

SPECIES WMED CMED ADRIA EMEDF o r a m i n i f e r a

Amphistegina lobifera + + + ++M a c r o p h y t a

Acrothamnion preissii ++ + + -Asparagopsis armata ++ + + +Asparagopsis taxiformis ++ ++ + ++Caulerpa distichophylla - ++ - ++Caulerpa racemosa var. cylindracea ++ ++ ++ ++Caulerpa taxifolia ++ ++ ++ -Chrysonephos lewisii ++ - - -Cladophora cf. patentiramea - - - ++Codium fragile subsp. fragile ++ + ++ ++Codium parvulum - - - ++Dasysiphonia sp. + - + -Gracilaria vermiculophylla - - ++ -Grateloupia turuturu + + + -Halophila stipulacea + ++ + ++Lithophyllum yessoense + - - -Lophocladia lallemandii ++ + + +Sargassum muticum ++ - ++ -Stypopodium schimperi - + - ++Ulva pertusa + - - -Undaria pinnatifida ++ + ++ -Womersleyella setacea ++ ++ + +

P o l y c h a e t aBranchiomma bairdi - - - ++Branchiomma luctuosum + ++ + ++Ceratonereis mirabilis - - - ++Desdemona ornata + - ++ ++Dorvillea similis - - - ++Eunice antennata + + - ++Ficopomatus enigmaticus ++ ++ ++ ++Hydroides dianthus ++ ++ ++ ++Hydroides elegans ++ ++ ++ ++Hydroides operculatus - - - ++Leonnates indicus - - - ++Leonnates persicus - - - ++Laonome triangularis - - - ++Notomastus mossambicus + - - ++

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Table 3 (continued)

SPECIES WMED CMED ADRIA EMEDPolydora cornuta + - - ++Pomatoleios kraussii - - - ++Prionospio (Aquilaspio) krusadensis - - - ++Prionospio (Prionospio) saccifera - - - ++Pseudonereis anomala - + - ++Pseudopolydora paucibranchiata - - - +Spirorbis (Dexiospira) marioni ++ - - ++Streblospio gynobranchiata - - - ++

C r u s t a c e aCallinectes sapidus* + + ++ ++Charybdis helleri - - - ++Charybdis longicollis - - - ++Dyspanopeus sayi - - ++ -Erugosquilla massavensis - - - ++Heterosaccus dollfusi - - - ++Marsupenaeus japonicus* + + + ++Melicertus hathor - - - ++Metapenaeus monoceros* - + - ++Metapenaeus stebbingi* - + - ++Penaeus semisulcatus* - - - ++Percnon gibbesi ++ ++ + ++Portunus segnis* + + - ++Rhithropanopeus harrisii + + ++ -

M o l l u s c a / B i v a l v i aAnadara kagoshimensis + - ++ +Anadara transversa + + ++ ++Brachidontes pharaonis + + + ++Chama pacifica - - - ++Dendrostrea frons - - - ++Fulvia fragilis ++ ++ ++Musculista senhousia ++ + ++ +Pinctada radiata* + ++ + ++Ruditapes philippinarum* + + ++ +Septifer forskali - - - ++Spondylus spinosus - - - ++Xenostrobus securis ++ - ++ -

M o l l u s c a / C e p h a l o p o d aSepioeuthis lessoniana* - - - ++

M o l l u s c a / G a s t r o p o d aAplysia dactylomela ++ + ++Bursatella leachii ++ ++ ++ ++

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Table 3 (continued)

SPECIES WMED CMED ADRIA EMEDCellana rota + + ++Cerithium scabridum + + - ++Conomurex persicus* + + ++Crepidula fornicata + + - +Ergalatax junionae - - - ++Erosaria turdus - ++ - +Melibe viridis + ++ ++ ++Pseudominolia nedyma - - - ++Rapana venosa +? - + +Rhinoclavis kochi - - - ++

P o r i f e r aParaleucilla magna ++ + + -

C n i d a r i a / A n t h o z o aOculina patagonica ++ +

C n i d a r i a / H y d r o z o aClytia hummelincki ++ ++ ++ -Clytia linearis ++ ++ ++ ++Garveia franciscana + - ++ -Macrorhynchia philippina - - - ++

C n i d a r i a / S c y p h o z o aCassiopea andromeda - + - +Phyllorhiza punctata - + - +Rhopilema nomadica - + - ++

C t e n o p h o r aMnemiopsis leidyi + + + ++

B r y o z o aBugula fulva + + + +Tricellaria inopinata - + ++ -

E c h i n o d e r m a t aAquilonastra burtoni - - - ++Asterias rubens - - - +Synaptula reciprocans - - - ++

A s c i d i a c e aBotrylloides violaceus - - + -Distaplia bermudensis + + - -Microcosmus exasperatus - + - +Microcosmus squamiger ++ + - -Phallusia nigra - - - ++Polyandrocarpa zorritensis + + - -Rhodosoma turcicum + + + Styela clava + - - -

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Table 3 (continued)

SPECIES WMED CMED ADRIA EMEDSymplegma brakenhielmi - - - +

F i s hApogon pharaonis - - - ++Atherinomorus forskalii* - ++ - ++Diplodus bellottii* ++ - - -Etrumeus teres* - ++ ++Fistularia commersonii + ++ + ++Gymnammodytes semisquamatus ++ - - -Lagocephalus sceleratus - + - ++Pagellus bellottii* ++ - - +Parexocoetus mento - + + ++Pempheris vanicolensis - + - ++Plotosus lineatus - - - ++Pteragogus pelycus - - - ++Sargocentron rubrum* - + - ++Saurida undosquamis* - + + ++Scomberomorus commerson* - ++ - ++Seriola carpenteri* - ++ - -Seriola fasciata + + - +Siganus luridus* + ++ - ++Siganus rivulatus* - ++ + ++Sillago sihama* - - - ++Solea senegalensis* ++ + - -Sphoeroides pachygaster ++ ++ + ++Sphyraena chrysotaenia* - ++ + ++Sphyraena flavicauda* - + - ++Stephanolepis diaspros - ++ + ++Upeneus moluccensis* - + - ++Upeneus pori* - + - ++

of the last decade that presented a popula-tion explosion within 2010 (ZENETOS etal., in press).

A total of 108 species are locally inva-sive or merely present in the EMED, 75 inthe CMED, 53 in the Adriatic and 64 inthe WMED (Fig. 2). The WMED hostsmost macrophytes, whereas the EMEDhas the lion’s share in polychaetes, crus-taceans, molluscs and fish.

The share of invasive species does notsupport POR’s (2009, 2010) statement thatspecies from the tropical Indo-Pacificshould behave more like one more elementof the native fauna and be less aggressive intheir onset than real aliens, e.g. those orig-inating from the temperate Atlantic orPacific. Rather, it would conform to therule of thumb that one out of every tenestablished alien species becomes invasive.

In addition to documented impacts onbiodiversity, some of the invasive species,among decapods, molluscs and fish, haveturned to be beneficial to man and be com-mercially exploited in several countries(Table 3). Though no extinction of a nativespecies is known, sudden decline in abun-dance, and even local extirpations, concur-rent with proliferation of aliens, has beenrecorded (GALIL, 2007).

Yet, there is no way to predict invasi-bility of ecosystems. However, community‘invasibility' appears to be the result of therelationship between native species rich-ness and alien species ability to colonizenew habitats (BULLERI et al., 2008). Thisconcept implies that habitats with high lev-els of diversity are difficult to invade. Incontrast, species-poor communities, orstressed ecosystems, are arguably moreprone to invasion, primarily due to lack ofbiotic resistance (OCCHIPINTI-AMBROGI & SAVINI, 2003). AtMediterranean basin scale, this appears tobe true (see chapter 7).

Little has been done to evaluate therisk posed by the introduction and spreadof invasive species, or to minimize the riskof introducing additional alien species.Effective legislation on alien species existsonly in few countries and for selective vec-tors e.g. in EU countries for aquacultureintroductions; Egypt, France, Spain andSyria for introductions in ballast waters(IMO/ BWC, Ballast Water Convention).

6.1. MacrophytesMost spectacular has been the arrival

of several invasive exotic varieties of greenand red algae. These algae have flourishedat the expense of benthic Mediterraneanorganisms. Their dense and creepingmeadows - which feature up to 27,000erected blades per square meter in Cauler-pa racemosa var. cylindracea - overwhelmthe benthic assemblages. Their success isusually explained by a fast growth, perma-nent meadows and synthesis of secondarymetabolites active against natural enemies(herbivorous, epiphytes, pathogens, etc.).

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Fig. 2: Breakdown of invasive or potentially invasive species per taxonomic group.

Once they have spread, species like thisare impossible to eliminate.

Among the 125 alien macrophytes, 21are invasive or potentially invasive (17% oftotal taxa) (Table 3). The total number ofinvasive or potentially invasive alienmacrophytes decreases eastwards: WMED(16 taxa), ADRIA (14 taxa), CMED (13taxa), and EMED (11 taxa). Only six inva-sive alien macrophytes have colonized thefour basins: the Indo-Pacific strain ofAsparagopsis taxiformis, Caulerpa racemosavar. cylindracea, Codium fragile, Halophilastipulacea, Lophocladia lallemandii andWomersleyella setacea.

6.2. PolychaetesA total of 22 species (17% of the total

alien polychaetes) seem to have invasivecharacter, dominating shallow-water ben-thic habitats, estuarine areas, pollutedareas and harbour environments. All ofthem are invasive in the EMED, five arepresent in the CMED, six in the Adriaticand nine in the WMED.

Polluted soft bottoms especially nearharbours greatly facilitate the establish-ment of ship-mediated species such asPolydora cornuta, Streblospio gynobranchi-ata and Pseudopolydora paucibranchiata( INAR et al., 2005, 2006a; DAGLI &

INAR, 2008). Polluted environments,where competition among species is lowand food is ample, provided these specieswith an excellent opportunity to build updense populations. The maximum densityof P. cornuta was calculated as 3170 ind.m-2,that of S. gynobranchiata as 60480 ind.m-2

and that of P. paucibranchiata as 6180ind.m-2 in Izmir Bay ( INAR et al., 2005;DAGLI & INAR, 2008).

In a study by INAR (2006), alienspecies comprised more than 95% of theserpulids found on hard substrata (rocks,

molluscs, dock’s pilings, ropes and tires)along the Levantine coast of Turkey.Pomatoleios kraussii formed a densely pop-ulated belt in shallow-water areas inMersin Bay ( INAR, 2006). The invasivenereid species Pseudonereis anomala is adominant component of the algal commu-nities in the Levant Sea, attaining a popu-lation density of 2475 ind.m-2 on the algaJania rubens in Iskenderun Bay ( INAR& ALTUN, 2007). BEN-ELIAHU (1989)reported that a native nereidid species ofthe Mediterranean, Perinereis cultrifera(Grube), was excluded from the habitats ofthe Levant Sea by P. anomala. This speciesexpanded its distributional range to IzmirBay, North Aegean Sea ( INAR &ERGEN, 2005) and the coasts of Greece(KAMBOUROGLOU & NICOLAI-DOU, 2006). The eunicid worm Euniceantennata was found to be a commonspecies on the southern coast of Turkey(KURT SAHIN & INAR, 2009) andwas also previously reported from thecoasts of Italy (CANTONE, 1993).

6.3. CrustaceaThe Indo-Pacific prawns, in particular

Marsupenaeus japonicus, Metapenaeusmonoceros, and Penaeus semisulcatus, arehighly prized and are considered a boon tothe Levantine fisheries (DURUER et al.,2008). Portunus segnis, formerly known asP. pelagicus, is commercially exploited inEgypt (ABDEL-RAZEK, 1987). It is alsocommercially important but caught insmall quantities in Turkey (DURUER etal., 2008). The latest record of P. segnis isfrom the northern Tyrrhenian Sea(CROCETTA, 2006).

The invasive crab Percnon gibbesi isone of the most recent and successfulinvaders in the Mediterranean Sea. Its dis-tribution has been recently updated by

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KATSANEVAKIS et al. (in press) whoreport that the relative abundance of thespecies in the Ka -Kekova Marine Pro-tected Area (SE Turkey) reached 112 indi-viduals in 2010, from two individuals in2006.

The blue crab Callinectes sapidus wastransported into the Mediterranean in bal-last tanks from the north-east coast of theUSA. After the first Mediterranean record(Venice, Italy, 1949), this species has beenwidely recorded in different Mediter-ranean regions, especially in the EMED(Turkey, Greece, Syria, Lebanon, Israeland Egypt) where it became invasive and iscommercially exploited (STREFTARIS &ZENETOS, 2006; DURUER et al., 2008).

A decade after its introduction,Heterosaccus dollfusi, a parasite on Charyb-dis longicollis, became invasive in the southLevantine coast. Despite the high preva-lence of the parasite and its injuriousimpact on the host reproduction, the inva-sive host-parasite pair reached a modusvivendi with no noticeable reduction in thehost population (INNOCENTI & GALIL,2007).

6.4. MolluscaThe gastropods Cerithium scabridum,

Rhinoclavis kochi, Conomurex persicus andBursatella leachi and the bivalves Pinctadaradiata, Brachidontes pharaonis, Fulviafragilis, Chama pacifica and Spondylusspinosus have been reported as locallyinvasive in the EMED by ZENETOS et al.(2005, 2008). Conomurex persicus has beenreported to achieve densities of tens ofspecimens per m2 at some sites(FISHELSON, 2000) on the Israeli coast.To these may be added the limpet Cellanarota, reported as thriving to the point ofdisplacing the native limpet Patellacaerulea (MIENIS, 2002, 2003).

Rapana venosa, one of the most inva-sive species worldwide, has been rapidlyincreasing in the Adriatic Sea, thusenhancing the risk of invasion and furtherspreading. Maximum estimated popula-tion densities of > 500 ind..km-2 werereached near the shores of Cesenatico(SAVINI et al., 2007).

The spread of the alien species acrossthe Mediterranean is not easy. Only six ofthe ca. 50 Lessepsian species have crossedthe Sicily Strait westwards and only two(Bursatella leachii and Fulvia fragilis) havereached the Spanish coast so far.

None of the other species of Indo-Pacific origin, so successful in the EMED,have a real invasion success in the westernbasin. Some of the tropical Indo-Pacificspecies which started in the CMED(Aplysia dactylomela, Haminoea cyanomar-ginata, Melibe viridis) are now spreading toMalta (BORG et al., 2009; SCHEMBRI,2009) or have even reached the westernMediterranean. Several individuals ofMelibe viridis were recently observed inGulf Aranci, NE Sardinia (DONEDDU &TRAINITO, 2008).

The case history of the forerunnersindicates that the spread across theMediterranean involves a complex historyof natural spread with larvae and transportvia shipping from the original point ofintroduction. In any case the distributionof these aliens in the western and centralparts of the Mediterranean is patchy anddoes not suggest a continuous progression.The limiting factors may be on the onehand, the natural species richness of thewestern basin; on the other, the severe dif-ference in sea surface temperatures (POR,2009, 2010) which may make it inhos-pitable to thermophilic species such asPinctada radiata.

The impact of invasive species is com-

s

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parable within these two main poles ofalien molluscs in the Mediterranean, butthe pool of invasive species is not the same.To the south, an aggressive onset is seenfor some tropical Indo-Pacific species suchas Conomurex persicus, Cellana rota or (atthe beginning of the invasion) Pinctadaradiata, which fail to become massivelyestablished in other parts of the basin. Oneof the latest introductions is Sepioteuthislessoniana, a commercially importantsquid species for inshore fisheries through-out its distributional range, which wasobserved in Iskenderun Bay (SALMAN,2002) and has rapidly spread in the Levantand Aegean seas (KATSANEVAKIS etal., 2009; LEFKADITOU et al., 2009;ZENETOS et al., in press). In the northernparts (Northern Aegean, Adriatic, andFrench Mediterranean lagoons) invasivespecies like Anadara kagoshimensis, Rudi-tapes philippinarum or Rapana venosa orig-inate from temperate areas, and none ofthese are reported as invasive in the south-ern parts of the Mediterranean. Thereforethe climatic gradient in the Mediterraneanis likely to promote natural boundaries forthe alien species, which should reach a sta-ble range at some time.

6.5. Miscellaneous invertebratesThe ascidian Microcosmus squamiger,

first recorded at 1963 in Tunisia, is widelyspread in the Western Mediterranean(France, Spain, Italy, North Africa: seesynthesis in TUR N et al., 2007), andlocally in the CMED (Malta:IZQUIERDO-MUN~OZ et al., 2009).This species forms dense populationsreaching up to 2300 ind..m-2 and its pres-ence has strong implications in the struc-ture and functioning of the native com-munities (RIUS et al., 2009). Microcos-mus squamiger is considered a global

marine invader with a high invasivepotential.

A recent immigrant, Styela clava,among the worst invasives in Europe, hasbeen accidentally transported into theWestern Mediterranean by shellfish trans-fer (DAVIS & DAVIS, 2008). Styela clava,a solitary, hermaphroditic ascidian, is con-sidered to be an aggressive invaderthroughout the world; this species is amajor pest to the mussel farming industryof Canada (ARSENAULT et al., 2009). Itsrecent discovery in the Mediterranean Seacaused a considerable alarm to musselfarms.

Phallusia nigra is an old invader in theMediterranean Sea. The quite suddenappearance of the species in the AegeanSea and the extended distribution that itdemonstrated within a short time alongthe coasts of Rodos and Kriti(KONDILATOS et al., 2010; ZENETOSet al., in press) support the invasive charac-ter of the species, as already pointed out byIZQUIERDO-MUN~OZ et al. (2009).

Other species such as Botrylloides vio-laceus, Symplegma brakenhemi and possi-bly Polyandrocarpa zorritensis and Distapliabermudensis are locally invasive in alteredhabitats (harbours, aquaculture rafts, Poand Ebro deltas).

The scleractinian coral Oculina patag-onica, a species of uncertain, but probablyAtlantic origin, has taken advantage of thewarming sea and has explosively expandedaround the southern Mediterranean dur-ing the last years, building coral pavements(see latest updates in SARTORETTO etal., 2008).

Of the 37 alien hydrozoan speciesrecorded for the Mediterranean, four canbe considered invasive. The occurrence ofClytia hummelincki in the MediterraneanSea was only discovered in 1996 in the

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northern Ionian (BOERO et al., 1997),and within the subsequent decade furtherrecords came from the Southern Adriaticand the Tyrrhenian seas (BOERO et al.,2005; GRAVILI et al., 2008). At present,C. hummelincki is extremely abundant inthe summer in shallow rocks and is rapid-ly expanding its range thanks to the dis-persal capacity with currents of themedusa stage. Clytia linearis was recordedin the Suez Canal as early as in the 1930s(BILLARD, 1938) and rapidly spreadthrough the whole Mediterranean to getits north-western reaches by the 1950s(PICARD, 1951): thus, it is probably oneof the first and most successful Lessepsianimmigrants. The medusae of the twospecies of Clytia are relatively large andare produced in great number: their vora-cious predatory behaviour on coastal zoo-plankton may imply a significant impacton native Mediterranean ecosystems bythese two invasive species (GRAVILI etal., 2008).

The first Mediterranean record ofGarveia franciscana came from the Lagoonof Venice in 1978, where it showed amongthe most abundant hydroids (MORRI,1981). In subsequent years it was alsofound in other estuarine habitats of theNorth Adriatic and Catalonia, NE Spain(MORRI, 1982; GILI, 1986). Its spread inmany brackish environments world-wideand its occurrence in ship-hull fouling(MORRI & BOERO, 1986), suggest highinvasive potential for this species: lack ofrecent studies on Mediterranean brackish-water hydroids, however, prevents anysolid evaluation of its present degree ofsuccess.

Macrorhynchia philippina hasoccurred in the EMED since at least theearly 1990s, when has been found in pol-luted waters near Beirut (BITAR &

BITAR-KOULI, 1995). It is now wide-spread and frequent in the whole LevantSea ( INAR et al., 2006b). Its diffusionon coastal rocks from just below the sur-face to more than 40 m and the frequentreproductive status (MORRI et al., 2009)suggest a high invasive potential. Itscolonies, up to 30 cm tall and stinging tothe touch, are easily recognizable, so thatits further expansion in the Mediter-ranean should be easily tracked (MORRI,2008).

One of the top 100 worldwide invasivespecies, the comb jelly, Mnemiopsis leidyi,was first seen in the Saronikos Gulf(EMED) in 1990 (SHIGANOVA et al.,2001). Sporadic sightings ever since fromthe Turkish and Greek coasts raised noconcern about its notorious impacts onthe ecosystem and the fisheries, until 2009when swarms if it spread across theMediterranean, from Israel to Spain(FUENTES et al., 2010). Each summersince the mid-1980s swarms of the invad-ing jellyfish, Rhopilema nomadica, haveappeared along the Levantine coast.These swarms of voracious planktotrophscan stretch up to 100 km long. As theydraw nearer to shore, they adversely affecttourism, fisheries, and coastal installa-tions.

6.6. Fish Classically, Lessepsian species were

expected to be limited to the EMED, dueto hydrological conditions, but the spreadto the western basin is now a reality. Sofar, only a few Lessepsian fish species havebeen observed in the Western Mediter-ranean: Abudefduf vaigiensis, Siganusluridus, and Fistularia commersonii. Thislatter is probably the fastest-spreadingspecies in the Mediterranean: since thefirst record in Mediterranean ten years

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ago, F. commersonii is now reported moreor less in the whole Mediterranean (Table2). The very recent records of F. commer-sonii and S. luridus in the north-westernMediterranean (S NCHEZ-TOCINO etal., 2007; DANIEL et al., 2009) could thenbe the hint of future extensive spreadingof Lessepsian species. The opposite direc-tion was followed by Atlantic invaderssuch as Seriola fasciata (SONIN et al.,2009), Pagellus bellottii (SAAD &SBAIHI, 1995) and Sphoeroides pachy-gaster (GOLANI, 1996) that have reachedthe Levantine basin.

Today, some of these species consti-tute an important part of local fisherycatches; 16 such species are listed (Table3) among the most invasive species in theMediterranean, Siganus luridus andSiganus rivulatus though marketable, theyare classified as venomous species inFishBase. Lessepsian immigrants consti-tuted 19% of the total abundance of fishcaptured at the Lebanese rocky coast. Indecreasing order of importance, wereAtherinomorus lacunosus (11.5%), Siganusluridus (2.3%), S. rivulatus (2.3%), Pem-pheris vanicolensis (1.6%), Sargocentronrubrum (1.0%), Stephanolepis diaspros(0.2%), and Apogon nigripinnis (presentlyApogon pharaonis) (0.02%) (HARME-LIN-VIVIEN et al., 2005). Siganus luridusand S. rivulatus contributed 23% to thespecies composition captured in purseseine operating during the daytime inAbu-Qir and El-Mex Bays, Alexandria(Egypt) (AKEL, 2005).

Invasive species such as: Apogonpharaonis, Fistularia commersonii, Gym-nammodytes semisquamatus, Parexocoetusmento, Pempheris vanicolensis, Pteragoguspelycus, Sphoeroides pachygaster, Stephano-lepis diaspros, are just a nuisance to fish-eries. Others as the highly venomous

striped catfish, Plotosus lineatus, and thepufferfish Lagocephalus sceleratus are athreat to human health.

7. SYNTHESIS: STATUS ANDGEOGRAPHICAL ORIGIN

A total of 955 alien species occur inthe Mediterranean, the vast majority ofthem having being introduced in theEMED (718), fewer in the WMED (328)and CMED (267), and least in the Adriat-ic (171). Of these, 535 species (56%) havebeen established in at least one area. Wecalculated that the established speciesaccounted for 57.6% of the total numberof alien species in the WMED, 51.5%, inthe CMED, 52.6% in the Adriatic Sea and56.3% in the EMED. The casual recordsconstitute 27-29% of the alien species ateach basin. On the contrary, the number ofquestionable species in the EMED is threetimes as many as those of the WMED.Macrophytes, polychaetes and crustaceanshave the highest number of questionablespecies. The questionable status formacrophytes is generally due to the uncer-tainty regarding their origin. For poly-chaetes and crustaceans, the reason for thehigh percentage is the lack of precise taxo-nomic works in the area and the corollarythat these species were largely reportedfrom ecological studies and seldom docu-mented by museum specimens or illustra-tions.

The number of species per establish-ment success category along with the con-tribution of alien taxa distributed at eachMSFD area is illustrated in Figure 3. Fur-ther details per taxonomic group are pro-vided below.

Figure 4 shows the origin of alienspecies for each sub-basin. Species origi-nally distributed either in the Indo-Pacific

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Fig. 3: a: Establishment success per basin (all taxa); b, c, d: Partitioning of alien taxa per establishmentsuccess.

or Indian or Red Sea are clustered togeth-er. Similarly, species originating in eitherthe east or west of the tropical Atlantic aregrouped together, and so are those origi-nating from the North Atlantic or theNorth Pacific.

It is estimated that almost 60% of thealien species in the Mediterranean Seacome from the Suez Canal. By addingthose species introduced via shippingand/or aquaculture and trade, it is quiteclear that most of the alien species in theMediterranean are warm water species,originating in (sub)tropical seas. In partic-ular, thermophilic species, (Indo-Pacific,Indian Ocean, Red Sea, tropical Atlantic,tropical Pacific, circum(sub)tropical),account for 88.4% of the introducedspecies in the EMED, 72.8% in theCMED, 59.3% in the WMED and 56.1%in the Adriatic. The respective figures forIndo-Pacific species only are 47.5% forEMED, 24.9% for CMED, 22.8% for theAdriatic and 20% for WMED, whereastropical Atlantic species show a sharpdecrease from WMED (10.1%) to EMED

(2.7%) (Fig. 4). Cold water species, cir-cumboreal, N Atlantic, N Pacific, make upa small percentage ranging between 4.2%and 21.6%, being more pronounced in theAdriatic and less so in the EMED.

While maritime traffic and otherhuman activities such as aquaculture areimportant vectors for the introduction ofalien species worldwide (RUIZ et al.,2000), in the Mediterranean they are notthe main responsibles for the large differ-ences observed among the four basins. Inthe EMED the human interventionresponsible for most of the aliens is the re-establishment of the connection with theIndo-Pacific through the Suez Canal(1869), rather than the actual transfer ofthe invaders. In addition, with the presentclimate change (BELKIN, 2009), the trop-ical features and temperature of the watersare increasing more quickly in the EMED,implying dramatic modifications of thebiota (POR 2009, 2010). As a consequence,Indo-Pacific species (regardless of themode introduction) have found optimalenvironment for settlement in the EMED.

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Fig. 4: Origin of species per basin. IP-IO-RS: Indo-Pacific, Indian Ocean, Red Sea.

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On the contrary, the Adriatic Sea,which is topographically a dead end, is thearea with the lowest number of aliens,receiving them among those alreadyestablished in the EMED and CMEDthat spread northwards, or among thoseintroduced via shipping or aquaculture inhot spot areas such as the Venice Lagoon(OCCHIPINTI-AMBROGI et al., 2010).

A significant number of Indo-Pacificspecies reaches the WMED, which isenriched by ship-transferred species ofPacific origin mostly among macrophytes.The Strait of Gibraltar is essentially dif-ferent from the Suez Canal as a potentialpathway for alien species. First of all, it isan ancient waterway, believed to haveoriginated 5.33 million years ago(HILGEN & LANGEREIS, 1993), com-pared to the 142 years of the Suez Canal.Therefore, the current distribution ofAtlantic species, tropical or not, with partof their range in the Mediterranean is theresult of a natural process over a longtime; these species do not in any casequalify as aliens, even if their discovery inthe Alboran Sea comes later than theirfirst description in the Atlantic. Secondly,the Atlantic coast of Morocco is swept bya prevalently southward oceanic circula-tion that prevents many potential new-comers to approach the Strait of Gibral-tar.

7.1. Alien foraminiferans in the Mediter-ranean

A total of 50 foraminiferan speciesare reported in this work, all of thembeing warm water species. The high diver-sity of Indo-Pacific species in the EMEDprobably implies that many of them areLessepsian immigrants. Data on shallowwater foraminifers are very few and scat-tered. Most works concern deeper assem-

blages, almost totally made up by taxa ofAtlantic affinity, more useful for the bios-tratigraphical and paleobathymetricalapplications that make foraminifers soimportant for micropaleontologists. As aconsequence, it is very difficult to recog-nize new Atlantic immigrants enteringthe Mediterranean. On the basis of the(scarce) geological data the assumptionthat some more tolerant large fora-minifers, such as amphisteginids, couldhave survived during the Pleistocene inthe southern Mediterranean cannot beruled out.

7.2. Alien macrophytes in the Mediter-ranean

Compared to previous lists of alienmacrophytes in the Mediterranean Sea(RIBERA & BOUDOURESQUE, 1995;BOUDOURESQUE & VERLAQUE,2002; RIBERA SIGU N, 2002;CORMACI et al., 2004; ZENETOS et al.,2008), the present revised checklist hastaken into account all the possible intro-ductions of exotic genotypes. Severalspecies already represented in theMediterranean Sea by putative nativepopulations, such as Cladosiphon zosterae,Desmarestia viridis, Ectocarpus siliculosusvar. hiemalis, Pylaiella littoralis, have beenonly considered here if introductionevents are locally evident (e.g. ThauLagoon, Lagoon of Venice). A total of125 taxa (79 Rhodophyta, 24 Fuco-phyceae, 20 Chlorophyta, 1 Pelago-phyceae and 1 Magnoliophyta) has beenlisted on the basis of recent works(FURNARI et al., 2010; SFRISO et al.,2010; TSIAMIS et al., 2010; CECERE etal., in press; VERLAQUE et al., in press)and on-going researches. Among them,97 taxa (78% of alien macrophytes) areestablished in at least one of the four

Mediterranean basins, the remainders(22%) belong to casual and crypto-genic/questionable categories.

As far as the major vectors of intro-duction are concerned, the transfer ofshellfish stands the first (46% of alienmacrophytes), followed by the shippingand the Suez Canal (29% and 27%,respectively each). Some species havebeen probably introduced several times bydifferent vectors. The predominantlydonor regions are temperate and coldregions (54% of alien macrophytes) vstropical regions (46%). Such a result maybe an artefact due to the unequal effort ofinvestigation between the WMED and theEMED. Introductions along with shellfishtransfers from north-eastern Atlantic andnorth-western Pacific have been thor-oughly studied in the north-western basinand the Adriatic, while in the EMEDsmallest alien macrophytes remain proba-bly insufficiently studied (most of report-ed Lessepsian migrants are large-sizedspecies).

7.3. Alien polychaetes in the MediterraneanCompiling data on alien polychaete

species reported from the Mediter-ranean Sea revealed a total of 129species belonging to 26 families, ofwhich 59 species have become estab-lished in the region, 31 species are casu-al, 37 species are questionable and 2species (Chaetozone corona and Parapri-onospio coora) are cryptogenic. Themajority of alien polychaete species havebeen found in the EMED (97 species),followed by the WMED (49 species), theCMED (35 species) and the Adriatic Sea(22 species). In the Sea of Marmara, 14alien polychaete species are known, ofwhich 6 species have become establishedin the area and 8 species are question-

able ( INAR, 2010).The Mediterranean Sea is largely

under the siege of polychaete species,mainly introduced from the RedSea/Indo-Pacific areas (99 species, 77% oftotal species) and Atlantic Ocean (24species). The Suez Canal is one of themain vectors for the introduction of thespecies. The EMED habitats are largelydominated by the Red Sea immigrants.Polychaete species that, most probably,entered the Mediterranean Sea via theSuez Canal and expanded their distribu-tion to the WMED are Nereis jacksoni,Lysidice collaris, Lumbrineris perkinsi,Notomastus aberans, N. mossambicus,Metasychis gotoi, Pista unibranchiata, Stre-blosoma comatus and Branchiomma luctu-osum.

The knowledge about alien poly-chaete species in the Mediterranean Seahas greatly increased recently. A total of59 species were reported after 2005. Thisnumber also includes some species (i.e.Sigambra constricta, S. parva, Dasy-branchus carneus) that were not classifiedas alien species by ZENETOS et al. (2005,2008). Due to the lack of precise taxo-nomic works on polychaetes, there is ahigh number of questionable species.These species were largely reported fromecological studies. The future taxonomicalstudies to be held in the whole basinwould enable us to realize the real numberof alien polychaete species in the Mediter-ranean Sea.

7.4. Alien crustaceans in the MediterraneanA total of 153 alien crustacean species

have entered the Mediterranean. Themajority are decapods (78 species), fol-lowed by copepods (42), amphipod (11),isopods (8), cirripedes (7), cumaceans (3),stomatopods (3) and tanaids (1).

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Approximately 77% of all alienMediterranean decapod species have anIndo-Pacific/Indian/Red Sea origin, whileonly 23% are from the Atlantic. The dom-inance of these ‘eastern alien species’over the total is obvious; it mainly reflectsthe situation (environment characteristicsmainly) in the EMED and CMED where82% of all Mediterranean alien speciescould be found (only the following alienspecies are not present: Actumnus globu-lus, Charybdis feriata, Charybdis japonica,Charybdis lucifera, Cryptosoma cristatum,Dyspanopeus sayi, Eriocheir sinensis, Hem-igrapsus sanguineus, Herbstia nitida,Menaethius monoceros, Scyllarus caparti,Scyllarus posteli and Thalamita gloriensis).In the WMED, the situation is more bal-anced, where only 52.6% of the aliendecapods are Indo-Pacific and theremainders are Atlantic. The Adriatic is apeculiar sector with 13 alien species, six ofwhich (Callinectes danae, Charybdisjaponica, Charybdis lucifera, Dyspanopeussayi, Herbstia nitida and Scyllarus caparti)are exclusive to this sector. The EMEDand WMED share only 10 species(12.8%), three of which inhabit estuarineenvironments or are related to aquacul-ture or food (Rhithropanopeus harrisii,Marsupenaeus japonicus and Necorapuber) and other two are invasive(Callinectes sapidus and especially Perc-non gibbesi).

The contribution of Indo-Pacificspecies reaches 86% in copepods vs 11.6%of Atlantic origin and one debatable case:the Antarctic visitor (Spinocalanus terra-novae). Recent taxonomic studies haveshed some light on alien microcopepodsin the Mediterranean (BÖTTGER-SCHNACK & SCHNACK, 2009). Thecontribution of warm water species is alsohigh among alien amphipods, excepting

Parhyale explorator, a NE Atlantic speciesfound in 2005 on the coast of IskenderunBay (BAKIR et al., 2008). Yet, the lack ofstudies to determine the actual originaldistribution of species such as Cymadusafilosa, Gammaropsis togoensis and Mono-corophium sextonae has led to the assign-ment of the cryptogenic status to almost30% of the species.

7.5. Alien molluscs in the MediterraneanMolluscs are one of the major groups

in the marine fauna worldwide and con-tinue as the first contributors to the alienfauna in the Mediterranean Sea. The totalnumber of reported alien species amountsto 212, a small but steady increase ofabout 8% since the recent review ofZENETOS et al. (2005). More than halfof the species (115 in total) are establishedin at least one of the four basins, 74 arecasual and the remaining 24 are question-able or cryptogenic.

There is a clear decrease in the impor-tance of aliens from the Levantine coastwhere extra-Mediterranean species mayrepresent 10 to 20% of the local fauna,towards the Alboran Sea where only onespecies (Godiva quadricolor) is reported,i.e. less than 1% of the total. Neverthelessthe numbers drop dramatically if we disre-gard as aliens the Lessepsian immigrants,following POR’s (2009, 2010) view thatthese species have made their way by theirown means following the restoration of anancient waterway, and therefore shouldnot be regarded as aliens. In this perspec-tive, there are hardly some thirty aliens inthe Mediterranean which originate fromdonor regions other than the tropicalIndo-Pacific. These have arrived mainly asa consequence of shipping and transfers ofliving spat for aquaculture, and the mostheavily impacted areas are not the Levan-

Medit. Mar. Sci., 11/2, 2010, 381-493 459

tine basin, but the marginal marine envi-ronments of the northern part of thebasins, including the Adriatic and thelagoons of the WMED.

It is noteworthy that the Adriatic Seashares more established species of mol-luscs (12 out of 16) with the WMED thanwith any of the other parts of the Mediter-ranean, even the neighbouring Ionian Sea(8 shared established species and 2 casu-al). This reflects not only the climatic sim-ilarity, with colder sea water temperaturein the northern parts, but also the fact thatmost of these aliens are estuarine specieshaving some connections with aquacul-ture, therefore likely to be transportedfrom the Venice area to the FrenchMediterranean lagoons or vice versa.

7.6. Alien miscellaneous invertebrates inthe Mediterranean

The prevailing taxa among alieninvertebrates other than polychaetes,crustaceans and molluscs, are thecnidarians (46 species: 37 hydrozoans, 4anthozoans, 5 scyphozoans) followed bybryozoans (23), ascidians (16), echino-derms (12), flatworms (12), sponges (8),sipunculans (6), pycnogonids (4),ctenophores (3), chaetognaths (2) andsiphonophores (1). With the exceptionof ascidians, the remaining taxa werepoorly treated by COLL et al. (2010), ifaddressed at all.

The current information is alwaysproportional to the degree of expertise inthe area. Along this line, cnidarians andbryozoans appear to be the best studiedtaxa represented by many alien species,while sponges, although well studied inthe region, have relatively few represen-tatives, and the reported ones are eitherquestionable or cryptogenic forms.

Alien Anthozoa are represented by

four species, the most widespread ofwhich is Oculina patagonica. It is a speciesof presumably temperate SW Atlanticorigin, probably introduced into theMediterranean by shipping. Invasive innatural and artificial habitats, especiallyalong the southern WMED and EMEDcoasts (SARTORETTO et al., 2008). Fivespecies coming from the subtropical east-ern Atlantic which have recently expand-ed their range into the Mediterraneanand are presently restricted to the Albo-ran Sea (OCAN~A et al., 2007, 2009) areexcluded from our list; yet we keep somereservations on their mode of introduc-tion. Dendrophyllia laboreli is a coral inexpansion at the Canaries and also thecontinental Atlantic coast of Africa. Para-muricea grayi, Eunicella gazella and Euni-cella labiata are mainly known from theAtlantic coast of Africa including a widetropical area (GRASSHOFF, 1992), buthave not been recognized as part of theMediterranean fauna by SHUCHERT etal. (2003). According to recent observa-tions, boats have borne larvae of Eunicel-la spp. from the Atlantic coast of Africa intheir ballast waters. Antipathella wollas-toni is a typical Macaronesian faunal ele-ment with clear tropical affinities(BRITO & OCAN~A, 2004). Its recentsettlement in the Alboran Sea may havebeen favoured by the present climatewarming.

The 16 alien ascidian species hithertorecorded in the Mediterranean basin rep-resent about 7% of the known species inthe whole basin. In many cases (13 out of16 species) the introduced ascidian speciesform established populations in antago-nism with the endemic species of theMediterranean benthos (ascidian or not).The species Phallusia nigra, Herdmaniamomus, Symplegma brakenhielmi, Rho-

Medit. Mar. Sci., 11/2, 2010, 381-493460

dosoma turcicum, Ascidia cannellata andprobably Microcosmus exasperatus are con-sidered immigrants via the Suez Canal(IZQUIERDO-MUN~OZ et al., 2009),while species such as Botrylloides violaceus,Polyandrocarpa zorritensis, Distapliabermudensis and Styela clava have beenrecently introduced by shellfish culture(BRUNETTI & MASTROTOTARO,2004; MASTROTOTARO & BRUNETTI,2006; DAVIS & DAVIS, 2008;IZQUIERDO-MUN~OZ et al., 2009). Inmany cases the first record has beenreported when the species was alreadypresent with dense populations often inantagonism with native species (e.g.BRUNETTI & MASTROTOTARO,2004; MASTROTOTARO & BRU-NETTI, 2006; DAVIS & DAVIS, 2008).

The majority of echinoderms origi-nate in the Indo-Pacific or are circum-tropical species and their distribution islimited to the EMED; only Ophiactis savi-gnyi reaches the WMED. Asterias rubens,originating in the N Atlantic, is limited tothe Marmara Sea where it was introducedfrom an established population in theBlack Sea (KARHAN et al., 2008).Chaetognaths are represented by twospecies, both of Indo-Pacific origin, limit-ed to the innermost part of the EMED.

The presence of the parasitic nema-tode Anisakis simplex s.str., in some indi-vidual hosts (Gaidropsarus granti) fromthe WMED, which seemed to be theresult of immigration of these fish fromthe Atlantic into the WMED(MATTIUCCI & NASCETTI, 2006), hasbeen excluded, because, according toMANFREDI et al. (2000), it is a commonparasite in many Mediterranean fishes.

7.7. Alien fish in the MediterraneanIn terms of number of species, fish

form a significant portion of the Mediter-ranean marine alien species list. The rateof introduction is significantly high, where25 species were added since April 2008(ZENETOS et al., 2008), bringing thetotal number of alien fish species to 149.

This figure reflects the relevance ofMediterranean invasion that has no equiv-alents on other seas. The richness ofMediterranean alien fish species isimpressive if compared with native ones.Even if there is no general consensus onthe total number of Mediterranean fish -for example FREDJ & MAURIN (1987)listed 612 species while QUIGNARD &TOMASINI (2000) numbered 664 species– this review can give a more precise ideaof how the original identity of Mediter-ranean fish fauna has changed followingthe arrival of these immigrants.

So far, no species extirpation has beendocumented. Therefore the ingression ofnew species generates an increase ofspecies richness, the main routes con-tributing to Mediterranean fish diversitybeing the Suez Canal and the Strait ofGibraltar. Almost half of the alien fishspecies that have been recorded in theMediterranean have given origin to per-manent populations. The magnitude ofthe phenomenon of fish invasion in theMediterranean Sea raises concern on theecological and economic consequencesand solicits the urgent need of ongoingmonitoring, especially over the southernrim of the basin were research effort is lessintense.

8. THE STATE OF THE ART INMEDITERRANEAN MARINE ALIENSPECIES

Species of tropical/subtropical affini-ty, favoured by climate warming, are intro-

Medit. Mar. Sci., 11/2, 2010, 381-493 461

duced and colonize the Mediterranean ata fast rate, while cold water species aresettling at a lower rate. The net outcome isa jump in species richness.

Compared to the latest lists of alienbiota (COLL et al., 2010; COSTELLO etal., 2010) the present work includes a larg-er number (954) of species (Table 4). Thedivergence in numbers is more obvious infish (149 this work vs. 116 in COLL et al.,2010), polychaetes (129 this work vs. 75 inCOLL et al., 2010), bryozoans (23 thiswork vs. one in COLL et al., 2010), andcnidarians (46 this work vs three in COLLet al., 2010) (Table 4). Here, we alsoinclude aliens among Porifera, Platy-helminthes and Foraminifera, taxa not

treated by COLL et al. (2010). In contrastto ZENETOS et al. (2008), we do notaddress microalgal species (either pelagicor benthic) for the reasons described inchapter 5. This implies that despite thecollective effort of experts attempted inthis work, the number of introducedspecies remains underestimated. It is clearthat the introduced species have increasedthe biodiversity of the Mediterranean Sea,as an entity, by 5.9% excluding phyto-plankton and microzooplankton (Table4). The figure is even higher reaching27.9% in fish and 9% on average for thestudied taxa. While species richness isincreasing at whole basin scale, leading toa higher Á diversity, cases of local replace-

Medit. Mar. Sci., 11/2, 2010, 381-493462

Table 4The number of alien species in the Mediterranean by taxonomic group.

COLL et al., 2010 This work

Taxon all aliens native aliens % aliensProtozoa (excluding Foraminifera) 0 0 4Foraminifera >600 0 600 50 8.3Rhodophyta 657 73 584 79 13.5Phaeophyta & Pelagophyceae 277 23 254 24+1 9.8Chlorophyta 190 17 173 20 11.6Magnoliophyta 7 1 6 1 16.7Polychaeta 1172 75 1097 129 11.8Crustacea 2239 106 2133 153 7.2Mollusca 2113 200 1913 212 11.1Cnidaria 757 3 754 46 6.1Bryozoa 388 1 387 23 5.9Ascidiacea 229 15 214 16 7.5Echinodermata 154 5 149 12 8.1Porifera 681 0 681 8 1.2Platyhelminthes 1000 0 1000 12 1.2Other Invertebrates 2168 2 2166 16 0.7Fish 650 116 534 149 27.9Total 637* 954*Average % 3.3% 5.9%

*excluding monocellular algae

ment have been reported (GALIL, 2007),which may imply an alteration of · diver-sity. In addition, the spatial overlapbetween alien and native species causesbiotic homogenization (BEN RAISLASRAM & MOUILLOT, 2009), andhence a depression in ‚ diversity. Thus,the relationship between the entering ofalien species and threats to biodiversity isnot straightforward (BIANCHI et al.,2010).

While biodiversity is currently a keyindicator of an ecosystem's health, statis-tics tell too simple a story in the Mediter-ranean Sea. In a recent study (2008)assessing fish assemblages associated withPosidonia oceanica meadows in theRodos Island (EMED), the silverver-stripe blaasop, Lagocephalus sceleratus,ranked among the 10 most abundantspecies, totalling 2% of total biomass,while bluespotted cornetfish, Fistulariacommersonii, and reticulated leatherjack-et, Stephanolepis diaspros contributed0.6% and 0.3%, respectively(KALOGIROU et al., 2010). Of the near-ly 30000 specimens collected by commer-cial benthic trawler off the central Israelicoast at depths between 15 and 30 m, inOctober 2008 (OCCHIPINTI-AMBROGI & GALIL, 2010), only 9%were native Mediterranean species, therest consisted of Erythrean aliens, such asthe highly venomous striped catfish, Plo-tosus lineatus (11437 specimens), theblotchfin dragonet, Callionymus filamen-tosus (5745), the silver sillago, Sillagosihama (1423), the kuruma prawn, Marsu-penaeus japonicus (1154) and the velvetshrimp Metapenaeopsis consobrina(1138).

The faunal survey of large harbourshas been largely neglected and is likely toyield many more localized, but estab-

lished, aliens if undertaken seriously. The application of molecular meth-

ods recently developed will be of greathelp in the definition of the relation-ships between populations from differ-ent areas, and in the assessment of pos-sible routes of introduction and immi-gration. In addition, metagenomic tech-niques will presumably allow a muchmore complete if not exhaustive knowl-edge of the microbial species in givenareas through their molecular signa-tures. In the meantime, classical taxo-nomic studies should be intensified toinvestigate the still wide proportion ofunreported alien species of small-sizedtaxa in the Mediterranean Sea includingmicroalgae.

Acknowledgements

The work was initiated under the aus-pices of the SEBI2010 - StreamliningEuropean 2010 Biodiversity Indicators -Expert Group 5: Numbers and costs ofinvasive alien species.

Further research on the distributionof alien species in the Mediterranean Seahas been carried out in the frame of theIntegrated Project ‘SESAME’ (SouthernEuropean Seas: Assessing and ModellingEcosystem changes: Project no: 036949)funded by the EU [A. Zenetos, I. Siokouand N. Streftaris] and in the frame of theresearch project ‘The impacts of biologi-cal invasions and climate change on thebiodiversity of the Mediterranean Sea’(C.N. Bianchi and C. Morri), an Italy-Israel co-operation funded by the ItalianMinistry for the Environment. Specialthanks are due to C. Salas Casanova andA. Logan whose constructive criticismand suggestions have improved the man-uscript.

Medit. Mar. Sci., 11/2, 2010, 381-493 463

Medit. Mar. Sci., 11/2, 2010, 381-493464

ANNEX: Name changes

Current name Old /others name(s) used

ProtozoaPerkinsus olseni Lester & Davis Perkinsus atlanticus MacrophytesCaulacanthus okamurae Yamada Feldmannophycus okamuraeCodium fragile subsp. fragile (Suringar) Hariot Codium fragile subsp. tomentosoidesHypnea anastomosans Papenfuss, Lipkin & Silva Hypnea esperiSaccharina japonica (Areschoug) C.E. Lane, C. Mayes, Laminaria japonicaDruehl & G.W. Saunders Spongoclonium caribaeum (B rgesen) M.J. Wynne Pleonosporium caribaeumPolychaetaApoprionospio pygmaea (Hartman, 1955) Prionospio pygmaeaLinopherus canariensis Langerhans, 1881 Pseudeurythoe acarunculataLumbrineris perkinsi Carrera-Parra, 2001 Lumbrineris inflataProsphaerosyllis longipapillata (Hartmann-Schroeder, 1979) Sphaerosyllis longipapillataStreblosoma comatus (Grube, 1856) Streblosoma hessleiCrustaceaAustrominius modestus Darwin, 1854 Elminius modestus Amphibalanus eburneus (Gould, 1841) Balanus eburneusGlabropilumnus laevis (Dana, 1852) Heteropanope laevisLinguimaera caesaris Krapp-Schickel, 2003 Maera hamigeraParvocalanus crassirostris Dahl, 1894 Paracalanus crassirostrisPilumnus minutus De Haan, 1835 Pilumnus hirsutus Portunus segnis (Forssk l, 1775) Portunus pelagicus Subeucalanus subcrassus Giesbrecht, 1888 Eucalanus subcrassusSternodromia spinirostris (Miers, 1881) Dromia spinirostrisSynalpheus tumidomanus africanus Crosnier & Forest, 1966 Synalpheus hululensisMolluscaAnadara kagoshimensis (Tokunaga, 1906) Anadara inaequivalvis / Arca/ScapharcaAnadara transversa (Say, 1822) Anadara demiriAnteaeolidiella foulisi (Angas, 1864) Aeolidiella indicaBostrycapulus odites Collin, 2005 Crepidula calyptraeiformis

Bostrycapulus aculeatusBulla arabica Malaquias & Reid, 2008 Bulla ampullaDiplodonta bogii Van Aartsen, 2004 Diplodonta cf. subrotundaCanarium mutabile (Swainson, 1821) Strombus mutabilisCerithidium diplax (Watson, 1886) Clathrofenella fusca

Clathrofenella ferrugineaCerithidium perparvulum (Watson, 1886) Cerithiopsis tenthrenoisConomurex persicus (Swainson, 1821) Strombus persicus

Strombus decorus raybaudii

a

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