Mediterranean Marine Science Review Article Indexed in WoS (Web of Science, ISI Thomson) The journal is available on line at http://www.medit-mar-sc.net
Alien species in the Mediterranean Sea by 2010. A contribution to the application of 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, Italy 10 Institute of Oceanography, Hellenic Centre for Marine Research, P.O. Box 712, 19013 Anavissos, Hellas 11 Dipartimento di Scienze della Terra, University of Turin, via Valperga Caluso 35, 10125 Torino, Italy 12 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, Spain 16 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 Muelle Can~onero Dato S/N 51001, Ceuta, Spain 19 Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy
Corresponding author: [email protected]
Medit. Mar. Sci., 11/2, 2010, 381-493 381 Abstract
The state-of-art on alien species in the Mediterranean Sea is presented, making distinctions among the four subregions defined in the EU Marine Strategy Framework Directive: (i) the Western Mediterranean Sea (WMED); (ii) the Central Mediterranean Sea (CMED); (iii) the Adriatic Sea (ADRIA); and (iv) the Eastern Mediterranean Sea (EMED). The updated checklist (December 2010) 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 them having being introduced in the EMED (718), less in the WMED (328) and CMED (267) and least in the 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 introduced species 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 figure should not be directly read as an indication of higher biodiversity, as spreading of so many aliens within 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 for 88.4% of the introduced species in the EMED, 72.8% in the CMED, 59.3% in the WMED and 56.1% in the Adriatic. Cold water species, i.e. circumboreal, N Atlantic, and N Pacific, make up a small percentage of the introduced species, ranging between 4.2% and 21.6% and being more numerous 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.
Introduction In 1978, there were 128 proven Lessepsian immigrants and 76 doubtful The Mediterranean Sea is one of the cases (POR, 1978). Since the first review seas of the world most affected by biolog- of alien species in the Mediterranean ical invasions in terms of how long the (ZIBROWIUS, 1992), there have been invaders have been present (OCCHI- many papers and reviews dealing with this PINTI-AMBROGI, 2000; STREFTARIS subject. More recently, CIESM has pub- et al., 2005), in number of alien species lished four Atlases of Exotic Species in detected (COSTELLO et al., 2010) and in the Mediterranean (GALIL et al., 2002; the unprecedented rate of introduction GOLANI et al., 2002; ZENETOS et al., (ZENETOS, 2009; 2010). Triggered by 2004; VERLAQUE et al., in press). The warming waters and a newly improved most recent lists show 116 species of fish- route through the Suez Canal, es, 70 species of decapod and stomatopod tropical/subtropical marine species, some crustaceans, 137 species of molluscs and very invasive, have progressively moved 110 species of macrophytes. This into the Mediterranean, disrupting increased number of records, particularly ecosystem stability of the basin. over the past few years, has been partly
382 Medit. Mar. Sci., 11/2, 2010, 381-493 attributed to intensification of research emergence of the new generation of EU effort in this topic. ZENETOS & political actions covering major maritime POLYCHRONIDIS (2010), based on strategic objectives, such as the Marine data gathered, analysed for and adopted Strategy Framework Directive (MSFD) in the European Environment Agency (2008/56/EC), the European Strategy for (EEA) and UNEP MAP RAC/SPA Marine and Maritime Research (COM reports, estimated that nearly 1000 (2008) 534) encompassing the Marine species entered the Mediterranean dur- Spatial Planning and the Ecosystem ing the past century. Approach (ECAP) within the Barcelona A recent synthesis on Mediterranean Convention by UNEP/MAP1, as well as marine biodiversity (COLL et al., 2010) many initiatives of the UNEP RAC/SPA. described the Mediterranean Sea as a Alien species regulations are of major biodiversity hot spot hosting approxi- importance in those policies. In the cur- mately 17,000 marine species, of which rent MSFD the descriptor: "Non-indige- more than 600 (3.3%) are alien. These nous species introduced by human activi- most recent detailed biodiversity esti- ties are at levels that do not adversely mates for alien species show a discrepan- alter the ecosystems" is actually one of the cy from the figures provided by eleven qualitative descriptors for deter- ZENETOS et al. (2005; 2008). However, mining Good Environmental Status as COLL et al. (2010) have pointed out, (GES). It is also closely related to six the true numbers of alien species are cer- other GES descriptors because of the tainly biased downwards. The data are great variety of impacts that Invasive presumably accurate for large and con- Alien Species (IAS) may exert on native spicuous species that are easily distin- biodiversity, ecosystem functioning, and guished from the native biota and for seabed habitats as well as commercial species that occur along a frequently sam- marine resources (OLENIN et al., 2010). pled (or fished) coast and for which taxo- This territorial and institutional sce- nomic expertise is readily available, but nario raises the need for a multi-level are entirely absent for many of the small governance system with a quite complex members of invertebrates. This under- coordination system in which non-EU representation is common in many works Member States can also participate. This and efforts have been made to include is particularly important in the Mediter- less studied groups in recent reviews (see ranean, where the southern and eastern ZENETOS et al., 2008). coast belong to non-EU Member States An up-to-date inventory of the alien or countries with accession status (e.g. species in the Mediterranean, apart from Turkey and Israel). Furthermore, the its scientific merits, can fulfil the needs of Barcelona Convention of the UNEP/MAP the regulatory requirements and environ- supports the introduction of MSFD mental management options. This is of (2009) concepts that can be useful particular importance since the current throughout the basin despite the handi-
1 The Barcelona Convention of the UNEP/MAP supports the introduction of MSFD in the Mediterranean. To that end, they are following the ecosystem approach (including the concept of policy steps), which is the priority of the recently 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.
Medit. Mar. Sci., 11/2, 2010, 381-493 383 cap that only seven of 21 Mediterranean studied along the four subregions as countries are Member States of adopted and described under the MSFD. UNEP/MAP. The coastal areas of the countries and the Considering a) the high introduction regions included in these subregions are rate, b) recent publications and reviews listed in Table 1. This division imposes on little studied taxa and c) the need to some restrictions/difficulties in the case of assess the extent of the phenomenon at countries whose waters lie within the subregional level and inform the EU, boundaries of these subregions as in the UNEP/MED and other stakeholders case of Albania, Tunisia and Italy. accordingly, this work aims to present the The borders of the CMED are hereby 2010 state-of-art on the presence and defined as the Kythira-Anti-Kythira acclimatization status of alien species at Straits (Greece) and Libya-Egypt borders each MSFD Mediterranean subregion, to the east, Otranto Strait (Italy, Albania) namely: (i) the Western Mediterranean to the north, Cap Bon (Tunisia) and Sea (WMED); (ii) the Central Mediter- south-west Sicily to the west. ranean Sea (CMED); (iii) the Adriatic The list is updated based on valid Sea (ADRIA); and (iv) the Eastern species records up to December 2010. Mediterranean Sea (EMED). The latter Alien species are investigated within all is also commonly referred to as the four regions in terms of their establish- Aegean-Levantine basin, but in this work ment success (casual, established, ques- the Marmara Sea, bearing more similari- tionable, cryptogenic), as well as invasive ties to the Aegean than to the neighbour- success, their native range and geograph- ing Black Sea, is also included. ical expansion. The species lists are pre- sented for seven units which are systemat- ic groups namely: 1) Protozoa (including Methodology Foraminifera), 2) Macrophytes, 3) Poly- chaeta, 4) Crustacea, 5) Mollusca, 6) Mis- Biological invasions have become a cellaneous Invertebrata (Arthropoda hot issue at a broad geographical scale Pycnogonida, Bryozoa, Chaetognatha, such as that of Large Marine Ecosystems Chordata Ascidiacea, Cnidaria, (LME). Among European LMEs, biolog- Ctenophora, Echinodermata, Porifera, ical invasions are most pronounced in the Sipuncula, Nematoda and Platy- Mediterranean (EEA, 2007a; ZENETOS helminthes), and 7) Fish, in each part. et al., 2009). The introduction of alien species in the On the European scale, over a dozen Mediterranean Sea is hard to demonstrate partitions can be found dividing up the for marine microalgae as for microrgan- waters of the NE Atlantic, the Mediter- isms in general (WYATT & CARLTON, ranean Sea and the Black Sea on oceano- 2002). Bearing in mind that it can rarely be graphic or purely geometric bases, or a excluded that a suspected microalgal invad- combination of both, as well as divisions er was already present as part of the rare, based on national jurisdictions2 (Fig. 1). hidden and unsampled phytoplankton, we In this work the Mediterranean is being refrain from citing a detailed list of plank-
2 EEZs, Marine Regions, and Subregions of the EU Directive on Marine Strategy (2008/56/CE).
384 Medit. Mar. Sci., 11/2, 2010, 381-493 Fig. 1: European marine and coastal regions(source: SUREZ DE VERO et al., 2009).
Table 1 Subregions, areas and coast of the Mediterranean studied in this work.
Western Central Mediterranean Adriatic Sea Eastern Mediterranean Mediterranean 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 Sea Sardinia Albanian Ionian Sea Croatia Sea of Marmara Corsica Apulian Sea Montenegro South Turkey France South East Sicily Albanian Adriatic Sea Cyprus Monaco Malta Syria Spain South Tunisia Lebanon Baleares Libya Palestine Authority Gibraltar Israel Morocco Egypt Algeria North Tunisia West Sicily
Medit. Mar. Sci., 11/2, 2010, 381-493 385 tonic and benthic microalgae as in made crafts are not considered. ZENETOS et al. (2005, 2008). The acclimatization status of each The revised checklist has been as wide species was assessed, and is given here as possible. Contrary to the CIESM Atlas, according to the following terminology. we have taken into account alien species Established: Introduced or feral popu- introduced from elsewhere within the lation of species settled in the wild with Mediterranean when the introduction free-living, self-maintaining and self-per- event was evident (e.g. the mollusc petuating populations unsupported by and Siphonaria pectinata introduced from the independent of humans. Species with at WMED to Greece, or the algae Cla- least two records in the area spread over dosiphon zosterae, Desmarestia viridis, time and space (at least three records for Ectocarpus siliculosus var. hiemalis and fishes) are also classified as established, in Pylaiella littoralis from the Adriatic to the the sense of the CIESM Atlas series. Thau Lagoon, WMED). Casual: Casual species are those hav- Freshwater species occurring in estuar- ing been recorded only once (no more ine waters such as Acipenser gueldenstaedtii, than twice for fish) in the scientific and Acipenser baeri and Huso huso in the North grey literature and are presumed to be Aegean estuaries and Micropterus salmoides non-established in the area. In this paper in Ionian Sea estuarine waters are not includ- ‘casual’ is used in the same sense as ‘alien’ ed in our list. Similarly, the freshwater cray- in the CIESM Atlas series. fish Procambarus clarki, though it can stand Questionable: Species with insuffi- slightly brackish waters and has been report- cient information–‘suspect’. This includes ed in the inner part of the Varano Lagoon, old casual records that have not been sub- Adriatic Sea (FLORIO et al., 2008) and in sequently found despite appropriate the Palude di Torre Flavia, Tyrrhenian Sea investigation, and also new entries not (SCALICI et al., 2010), is not included. verified by experts or species with taxo- Alien [synonyms: Non-indigenous nomic status unresolved. species (NIS), exotic, non-native, Cryptogenic: Species with no definite allochthonous] are species, subspecies or evidence of their native or introduced sta- lower taxa, present in the wild, introduced tus according to CARLTON (1996) and outside of their natural range (past or species whose probable introduction present) and beyond their natural disper- occurred in ‘early times’ and has not been sal potential. This includes any part, witnessed (e.g. prior to 1800). gamete or propagule of such species that Invasive: Species defined as established might survive and subsequently repro- aliens that have overcome biotic and abiot- duce. Their presence in the given region ic barriers and are able to disseminate away is due to intentional or unintentional from their area of initial introduction introduction resulting from human activi- through the production of fertile offspring ties. Natural shifts in distribution ranges with noticeable impact, such as threat to the (e.g. due to climate change or dispersal by diversity or abundance of native species, the ocean currents) do not qualify a species as ecological stability of infested ecosystems, a NIS (OLENIN et al., 2010). Specimens economic activities dependent on these kept in captivity and specimens still ecosystems, and human health. attached to a ship’s hull or other man- True aliens need to be separated from
386 Medit. Mar. Sci., 11/2, 2010, 381-493 species with seemingly isolated records, Finally, attention was paid to recent which have naturally spread to regions nomenclatural updates. These are the beyond their usual range. BOUCHET & result of the latest taxonomic and/or TAVIANI (1992) illustrated this scenario molecular studies, such those of LAI et al. with the case of deep-sea molluscs in the (2010) on the Portunus pelagicus species Mediterranean, brought into the basin as complex; of HUBER (2010) on Anadara larvae carried by the inflowing surface kagoshimensis, formerly known as Anadara waters, but unable to establish fertile pop- inaequivalvis; and of MALAQUIAS & ulations. This is also the case of some REID (2008) on Bulla arabica, formerly Eastern Atlantic crustacean species sel- known as Bulla ampulla. A list of all dom recorded in the West Mediterranean species catalogued under their currently but occurring in the Alboran sea (Penaeop- valid names along with the old names is sis serrata, Merhippolyte ancistrota, etc.), provided in the ANNEX. and of the big-eye thresher shark, Alopias Nomenclature adopted in this paper superciliosus, which is distributed world- follows the World Register of Marine wide in tropical and temperate seas and Species (WoRMS
Medit. Mar. Sci., 11/2, 2010, 381-493 387 Table 2 List 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 EMED PROTOZOA Miscellaneous Protozoa Bonamia ostreae (Pichot et al., 1979) Circumboreal cas cas Marteilia refringens Cavalier-Smith, 2002 unknown cas cas Perkinsus olseni** Lester & Davis, 1981 Cosmopolitan cry cry Photobacterium damselae - NE Atlantic est ? ? ? Foraminifera Agglutinella arenata (Said, 1949) Indo-Pacific cas Agglutinella compressa El-Nakhal, 1983 Indo-Pacific cas Agglutinella robusta El-Nakhal, 1983 Indo-Pacific cas Agglutinella soriformis El-Nakhal, 1983 Indo-Pacific/Atlantic cas Amphisorus hemprichii Ehrenberg, 1840 Circumtropical est Amphistegina lessonii d'Orbigny, 1826 Circumtropical est? est Amphistegina lobifera Larsen, 1976 Circumtropical est? est Amphistegina d’Orbigny, 1826 Indo-Pacific est est madagascariensis Archais angulatus (Fichtel & Moll, 1798) W Atlantic cry/cas Articulina alticostata Cushman, 1922 Indo-Pacific est Astacolus insolitus (Schwager, 1866) Indo-Pacific est Astacolus sublegumen (Parr, 1950) Indo-Pacific est Borelis sp. Circumtropical est Brizalina simpsoni (Heron-Allen & Earland, 1915) Indo-Pacific cas Clavulina angularis d' Orbigny, 1826 Circumtropical cry/cas cry/est Clavulina cf. multicamerata Chapman, 1907 Indo-Pacific est Coscinospira hemprichii Ehrenberg, 1839Indo-Pacific cry/cas cry/est Cushmanina striatopunctata (Parker & Jones, 1865) Circumtropical cas Cycloforina sp. Indian/Red Sea est Cyclorbiculina compressa (d'Orbigny, 1839 ) Circumtropical est Cymbaloporetta plana (Cushman, 1924) Indo-Pacific cry/est cry/cas cry/est Edentostomina cultrata (Brady, 1881) Indo-Pacific est Elphidium cf. charlottensis (Vella, 1957) Indo-Pacific est Elphidium striatopunctatum (Fichtel & Moll, 1798) Indo-Pacific est Euthymonacha polita (Chapman, 1904) Indo-Pacific cas Haddonia sp. Chapman, 1898 Indo-Pacific est Hauerina diversa Cushman, 1946 Circumtropical est Heterocyclina tuberculata (Moebius, 1880) Indian est
388 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Heterostegina depressa d’Orbigny, 1826 Circumtropical est Miliolinella cf. hybrida (Terquem, 1878) Indo-Pacific est Nodophthalmidium (Cushman, 1922) Circumtropical est antillarum Operculina ammonoides (Gronovius, 1781) Red Sea cas Pegidia lacunata McCulloch, 1977 Indo-Pacific est Peneroplis antillarum d’Orbigny, 1839Caribbean or SW cas Atlantic Planogypsina acervalis (Brady, 1884) Circumtropical cry cry cry Planogypsina squamiformis (Chapman, 1901) Circumtropical est Planorbulinella larvata (Parker & Jones, 1865) Indo-Pacific cas Pseudolachlanella slitella Langer, 1992 Indo-Pacific est Pseudomassilina reticulata (Heron-Allen & Earland, 1915) Indo-Pacific est Pulleniatina obliquiloculata* (Parker & Jones, 1865) Circumtropical cas Pyramidulina catesbyi (d'Orbigny, 1839) Indo-Pacific/Atlantic est Pyrgo denticulata (Brady, 1844) Indo-Pacific est Quinqueloculina cf. Said, 1949 Indo-Pacific est mosharrafai Schackoinella imperatoria d’Orbigny, 1846 Indo-Pacific cry/cas cry/cas cry/cas Schlumbergerina (Brady, 1879) Circumtropical est alveoliniformis Sorites orbiculus Forsska l, 1775 Circumtropical cry/est cry/cas cry/est Sorites variabilis Lacroix, 1941 Indo-Pacific est Spiroloculina antillarum d'Orbigny, 1839Circumtropical est Spiroloculina cf. angulata Cushman, 1917 Indo-Pacific cas Triloculina fichteliana d'Orbigny, 1839Circumtropical est MACROPHYTES Chlorophyta Batophora sp. Atlantic ques Caulerpa distichophylla Sonder Indo-Pacific est est Caulerpa mexicana Sonder ex Kützing Indo-Pacific est Caulerpa racemosa var. (Sonder) Verlaque, Huisman Indo-Pacific est est est est cylindracea & Boudouresque Caulerpa racemosa var. (Montagne) Weber-van Indo-Pacific est est lamourouxii f. requienii Bosse Caulerpa racemosa var. (J. Agardh) Eubank/ Indo-Pacific cry/ques cry/ques turbinata /uvifera (C. Agardh) J. Agardh Caulerpa scalpelliformis (Brown ex Turner) C. Agardh Indo-Pacific est Caulerpa taxifolia (Vahl) C. Agardh Indo-Pacific est est est Cladophora cf. patentiramea (Montagne) Kützing Indo-Pacific est Cladophora herpestica (Montagne) Kützing Indo-Pacific est
Medit. Mar. Sci., 11/2, 2010, 381-493 389 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Cladophora hutchinsioides Hoek & Womersley Pacific cas Codium fragile subsp. (Suringar) Hariot NW Pacific est est est est fragile** Codium parvulum (Bory ex Audouin) P.C. Silva Indo-Pacific est Codium taylorii P.C. Silva Indo-Pacific est est Derbesia boergesenii (Lyengar & Ramanathan) Indo West Pacific cas Mayhoub Derbesia rhizophora Yamada NW Pacific est Neomeris annulata Dickie Indo-Pacific est Ulva fasciata Delile Cosmopolitan est cry/ques cry/ques cry/ques Ulva pertusa Kjellman Indo-Pacific est Ulvaria obscura (Kützing) Gayral NW Pacific est est est Fucophycae Acrothrix gracilis Kylin Circumboreal est Botryella cf. parva (Takamatsu) H.-S. Kim Pacific est Chorda filum (Linnaeus) Stackhouse N Atlantic/N Pacific est est Cladosiphon zosterae (J. Agardh) Kylin Atlantic est cry/ques est cry/ques Colpomenia peregrina Sauvageau Indo-Pacific est est est Desmarestia viridis (O.F. Müller) J.V. Lamouroux Atlantic/Pacific est cry/ques Ectocarpus siliculosus (P.L. Crouan & H.M. Atlantic est cry/ques var. hiemalis Crouan) Foslie Fucus spiralis Linnaeus Atlantic cas Halothrix lumbricalis (Kützing) Reinke N Atlantic/N Pacific est cry/ques cry/ques cry/ques Leathesia marina (Lyngbye) Decaisne Cosmopolitan est est est Microspongium tenuissimum (Hauck) A.F. Peters Atlantic est Padina antillarum (Kützing) Piccone Indo-Pacific cas Padina boergesenii Allender & Kraft Indo-Pacific est est est Padina boryana Thivy in W.R. Taylor Indo-Pacific est est Punctaria tenuissima (C. Agardh) Greville NE Atlantic est est Pylaiella littoralis (Linnaeus) Kjellman N Atlantic/N Pacific est cry/ques cry/ques cry/ques Rugulopteryx okamurae (Dawson) I.K. Hwang, Pacific est W.J. Lee & H.S. Kim Saccharina japonica** (Areschoug) C.E. Lane, NW Pacific cas C. Mayes, Druehl & G.W. Saunders Sargassum muticum (Yendo) Fensholt NW Pacific est est Scytosiphon dotyi Wynne NE Pacific est est Spatoglossum variabile Figari & De Notaris Indo-Pacific cas Sphaerotrichia firma (Gepp) A.D.Zinova NW Pacific est est Stypopodium schimperi (Kützing) Verlaque Indo West Pacific est est & Boudouresque
390 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Undaria pinnatifida (Harvey) Suringar Pacific est cas est Magnoliophyta Halophila stipulacea (Forsska l) Ascherson Red Sea est est est est Pelagophyceae Chrysonephos lewisii (W.R.Taylor) W.R.Taylor W Atlantic est Rhodophyta Acanthophora nayadiformis (Delile) Papenfuss Indo-Pacific cry/ques cry/ques cry/ques cry/ques Acrochaetium codicola B rgesen Atlantic/Pacific est est est est Acrochaetium robustum B rgesen Indo-Pacific cas Acrochaetium spathoglossi B rgesen Indo-Pacific cas Acrochaetium subseriatum B rgesen Indo-Pacific cas Acrothamnion preissii (Sonder) E.M. Wollaston Indo-Pacific est est cas Agardhiella subulata (C. Agardh) Kraft & Atlantic/Pacific est est est M.J. Wynne Aglaothamnion feldmanniae Halos N Atlantic est est Ahnfeltiopsis flabelliformis (Harvey) Masuda Pacific est Anotrichium okamurae Baldock NW Pacific cry/ques cry/ques cry/ques cry/ques Antithamnion amphigeneum A. Millar SW Pacific est Antithamnion hubbsii Dawson Indo-Pacific est est Antithamnionella boergesenii (Cormaci & G. Furnari) Indo-Pacific cry/ques cry/ques Athanasiadis Antithamnionella elegans (Berthold) J.H. Price & unknown cry/ques cry/ques cry/ques cry/ques D.M. John Antithamnionella (Schiffner) E.M. Wollaston Indo-Pacific est est est Requires spirographidis confirmation Antithamnionella sublittoralis (Setchell & Gardner) Pacific est est Athanasiadis Antithamnionella ternifolia (J.D. Hooker & Harvey) Lyle unknown est Apoglossum gregarium (E.Y. Dawson) M.J. Wynne Pacific est est Asparagopsis armata Harvey SW Pacific est est est Requires confirmation Asparagopsis taxiformis Delile Atlantic cry/ques cry/ques cry/ques Asparagopsis taxiformis invasive strain Indo-Pacific est est est est Bonnemaisonia hamifera Hariot Indo-Pacific est est est est Botryocladia Feldmann-Mazoyer Indian est est est est madagascariensis Caulacanthus okamurae** Yamada NW Pacific est Ceramium bisporum D.L. Ballantine W Atlantic cry/ques cry/ques Ceramium strobiliforme G.W. Lawson & D.M. John N Atlantic cry/ques cry/ques Chondria coerulescens (J. Agardh) Falkenberg E Atlantic est cry/ques cry/ques cry/ques Chondria curvilineata F.S. Collins & Hervey Atlantic est est
Medit. Mar. Sci., 11/2, 2010, 381-493 391 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Chondria pygmaea Garbary & Vandermeulen Indo-Pacific est est est Chondrus giganteus f. Mikami Pacific est flabellatus Chrysymenia wrightii (Harvey) Yamada Pacific est Dasya sessilis Yamada Pacific est Dasysiphonia sp. Yendo NW Pacific est est (= Heterosiphonia japonica) Galaxaura rugosa (J. Ellis & Solander) Red Sea est J.V. Lamouroux Ganonema farinosum (J.V. Lamouroux) K.C. Fan Indian cry/ques cry/ques cry/ques & Yung C. Wang Goniotrichiopsis sublittoralis G.M. Smith NW Pacific est Gracilaria arcuata Zanardini Indo-Pacific/Red Sea Requires Requires confirmation confirmation Gracilaria vermiculophylla (Ohmi) Papenfuss NW Pacific est Grateloupia asiatica Kawaguchi & Wang NW Pacific est Grateloupia lanceolata (Okamura) Kawaguchi Pacific est cas Grateloupia minima P.L. Crouan & H.M. Pacific est Crouan Grateloupia patens (Okamura) S. Kawaguchi Pacific cas & H.W. Wang Grateloupia subpectinata Holmes Pacific est Grateloupia turuturu Yamada NW Pacific est est est Griffithsia corallinoides (Linnaeus) Batters Atlantic/Pacific est est est Herposiphonia parca Setchell Indo-Pacific est Hypnea anastomosans** Papenfuss, Lipkin & Silva Red Sea cas est Hypnea cornuta (Kützing) J.Agardh Cosmopolitan est est Hypnea flagelliformis Greville ex J. Agardh Indo-Pacific Requires confirmation Hypnea flexicaulis Yamagishi & Masuda Indo-Pacific est Hypnea spinella (C. Agardh) Kützing Pantropical est est est est Hypnea valentiae (Turner) Montagne Red Sea est est cas est Laurencia caduciramulosa Masuda & Kawaguchi SW Pacific est est Laurencia okamurae Yamada Pacific est Lithophyllum yessoense Foslie Pacific est Lomentaria flaccida Tanaka Pacific cas Lomentaria hakodatensis Yendo Pacific est est Lophocladia lallemandii (Montagne) F. Schmitz Indo-Pacific est est est est Nemalion vermiculare Suringar NW Pacific est Neosiphonia harveyi (J. Bailey) M.-S. Kim, H.-G. NW Pacific est est est est Choi, Guiry & G.W. Saunders
392 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Nitophyllum Okamura Pacific est stellato-corticatum Osmundea oederi (Gunnerus) G. Furnari Atlantic cry/ques cry/ques Plocamium secundatum (Kützing) Kützing tropical/subtropical est est Polysiphonia atlantica Kapraun & J.N. Norris N Atlantic/N Pacific cry/ques cry/ques cry/ques cry/ques Polysiphonia fucoides (Hudson) Greville N Atlantic est cry/ques cry/ques cry/ques Polysiphonia morrowii Harvey NW Pacific est est est Polysiphonia paniculata Montagne E Pacific est cas cas Polysiphonia stricta (Dillwyn) Greville NE Atlantic est Porphyra yezoensis Ueda Pacific est Pterosiphonia tanakae Uwai & Masuda Pacific est Rhodophysema georgei Batters Atlantic/Pacific cas cas Rhodymenia erythraea Zanardini Indo West Pacific cas Sarconema filiforme (Sonder) Kylin Indo West Pacific cas est Sarconema scinaioides B rgesen Indian est Solieria dura (Zanardini) F. Schmitz Indo-Pacific cas Solieria filiformis (Kützing) Gabrielson N Atlantic est est est Spongoclonium caribaeum** (B rgesen) M.J. Wynne W Atlantic est Symphyocladia (Harvey) Falkenberg Indo-Pacific est marchantioides Womersleyella setacea (Hollenberg) R.E. Norris Indo-Pacific est est est est POLYCHAETA Amphicorina pectinata (Banse, 1957) Pacific cas cas Apoprionospio pygmaea** (Hartman, 1955) Pacific cas Branchiomma bairdi (McIntosh, 1885) Atlantic/Pacific est Branchiomma boholensis (Grube, 1878) Indo-Pacific est est est Branchiomma luctuosum (Grube, 1869) Indo-Pacific est est est est Capitellethus dispar (Ehlers, 1907) Indo-Pacific/Red Sea ques Ceratonereis mirabilis Kinberg, 1866 Indo-Pacific cas est Chaetozone corona Berkeley & Berkeley, 1941 unknown cry Cirriformia semicincta (Ehlers, 1905) Red Sea ques Cossura coasta Litamori, 1960 unknown cry/ques Dasybranchus carneus Grube, 1870 Red Sea ques Desdemona ornata Banse, 1957 Indo-Pacific est est est Dispio magnus (Day, 1955) Indian ques Dispio uncinata Hartman, 1951 W Atlantic cas cas Dodecaceria capensis Day, 1961 Indian ques ques Dorvillea similis (Crossland, 1924) Indo-Pacific est Eunice antennata (Savigny, 1820) Indo-Pacific est est est Eunice cf. cariboea Grube, 1856 W Atlantic ques Eunice floridana (Pourtalès, 1867) W Atlantic ques
Medit. Mar. Sci., 11/2, 2010, 381-493 393 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Epidiopatra hupferiana Augener, 1918 Tropical Atlantic cas hupferina Epidiopatra hupferiana Day, 1957 Indian cas monroi Eunice indica Kinberg, 1865 Indo-Pacific ques Erinaceusyllis serratosetosa (Hartmann-Schroder, 1982) Pacific est Euniphysa tubifex (Crossland, 1904) Indian ques Eurythoe complanata (Pallas, 1766) Atlantic/Pacific ques ques Eusyllis kupfferi Langerhans, 1879Atlantic est Exogone africana Hartmann-Schröder, 1974 Indo-Pacific cas Exogone breviantennata Hartmann-Schröder, 1959 Circumtropical est Fabriciola qhardaqa Banse, 1959 Red Sea cas Ficopomatus enigmaticus (Fauvel, 1923) Subtropical est est est est Glycinde bonhourei Gravier, 1904 Indo-Pacific est Haploscoloplos kerguelensis Mc Intosh, 1885 Antarctic ques Hesionura serrata (Hartmann-Schroder, 1960) Red Sea cas Hyboscolex longiseta Schmarda, 1861 Cosmopolitan ques ques ques Hydroides albiceps Grube, 1870 Indo Pacific/Red Sea cas Hydroides branchyacanthus Rioja, 1941 Indo-Pacific est Hydroides dianthus (Verrill, 1873) NW Atlantic est est est est Hydroides diramphus Mörch, 1863 Circumtropical est est est Hydroides elegans (Haswell, 1883) Circumtropical est est est est Hydroides heterocerus (Grube, 1868) Indian/Red Sea est Hydroides homocerus Pixell, 1913 Indian est Hydroides minax (Grube, 1878) Indo-Pacific/Red Sea est Hydroides operculatus (Treadwell, 1929) Indian est Hydroides steinitzi Ben-Eliahu, 1972 Red Sea cas cas Janua (Dexiospira) steueri (Sterzinger, 1909) Indo-Pacific cas Isolda pulchella Muller, 1858 Circumtropical cas Laonome elegans Gravier, 1906 Red Sea cas Laonome triangularis Hutchings & Murray, 1984 SW Pacific est Leiochrides australis Augener, 1914 Pacific cas cas cas Leocrates chinensis Kinberg, 1866 Pacific ques ques ques ques Leonnates decipiens Fauvel, 1929 Indo-Pacific est Leonnates indicus Kinberg, 1966 Indo-Pacific est Leonnates persicus Wesenberg-Lund, 1949 Indo-Pacific est Lepidonotus carinulatus (Grube, 1870) Indo-Pacific/Red Sea ques Lepidonotus tenuisetosus (Gravier, 1902) Indo-Pacific/Red Sea cas Linopherus canariensis** Langerhans, 1881 Atlantic est est Longibranchium atlanticum (Day, 1973) Atlantic cas cas Loimia medusa (Savigny, 1818) Cosmopolitan ques ques
394 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Lumbrineris acutifrons (Gallardo, 1967) Pacific ques cas Lumbrineris neogesae Miura, 1980 S Africa cas cas Lumbrineris perkinsi** Carrera-Parra, 2001 Indo-Pacific est est est Lysidice collaris Grube, 1870 Pacific/Red Sea est est est est Lysidice natalensis (Kinberg, 1865) Indo-Pacific ques Marphysa disjuncta Hartman, 1961 Pacific cas Mediomastus capensis Day, 1961 Indian ques ques ques Megalomma claparedei Gravier, 1908 Red Sea cas Metasychis gotoi (Izuka, 1902) Indo-Pacific est est est est Naineris quadraticeps Day, 1965 Red Sea ques Neopseudocapitella Rullier & Amoureux, 1979 W Atlantic/ est est est est brasiliensis Red Sea Neanthes agulhana (Day, 1963) S Africa est est Neanthes willeyi (Day, 1934) Indo-Pacific cas Nereis gilchristi Day, 1960 Red Sea cas Nereis jacksoni Kinberg, 1866 Indo-Pacific est est Nereis persica Fauvel, 1911 Indo-Pacific ques est Notomastus aberans Day, 1957 Indian/Red Sea est est est est Notomastus mossambicus (Thomassin, 1970) Indian est est Notopygos crinita Grube, 1855 W Atlantic ques Novafabricia infratorquata Fitzhugh, 1983 W Atlantic est est Ophryotrocha diadema Akesson, 1976 Pacific est Ophryotrocha japonica Claparède & Mecznikow, 1968 Pacific est est est Oenone cf. fulgida (Savigny, 1818) Indo-Pacific/Red Sea ques ques Onuphis eremita oculata Hartman, 1951 W Atlantic est Palola valida (Gravier, 1900) Red Sea est Paradyte cf. crinoidicola (Potts, 1910) Indo-Pacific ques Paraehlersia weissmaniodes (Augener,1913) Indo-Pacific cas Paraprionospio coora Wilson, 1990 Pacific cry cry Perinereis nuntia (Savigny, 1818) Indian cas Pherusa parmata (Grube, 1878) Indo-Pacific cas Pherusa saldanha Day, 1961 Indian cas Pileolaria berkeleyana (Rioja, 1942) E Pacific est Pisione guanche San Mart›n, Lo’pez & Tropical Atlantic cas Nu’ ~nez, 1999 Pista unibranchia Day, 1963 Indo-Pacific est est Platynereis australis (Schmarda, 1861) Pacific ques Podarkeopsis capensis (Day, 1963) Indo-Pacific cas ques Polycirrus twisti Potts, 1928 Suez Canal est Polydora colonia Moore 1907 W Atlantic cas cas Polydora cornuta Bosc, 1802 Cosmopolitan est est
Medit. Mar. Sci., 11/2, 2010, 381-493 395 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Polydora spongicola Berkeley & Berkeley, 1950 Pacific ques Pomatoleios kraussii (Baird, 1865) Indo-Pacific/Red Sea est Prionospio (Aquilaspio) Fauvel, 1929 Indo West Pacific est krusadensis Prionospio (Aquilaspio) Augener, 1918 Atlantic/Pacific est sexoculata Prionospio (Prionospio) Imajima, 1990 NE Pacific est depauperata Prionospio (Prionospio) Mackie & Hartley, 1990 Indo-Pacific est saccifera Prionospio (Prionospio) Blake & Kudenov, 1978 Pacific est paucipinnulata Prionospio (Minuspio) Imajima, 1990 Atlantic/Pacific cas pulchra Protodorvillea egena (Ehlers, 1913) Indian/Red Sea ques Protodorvillea biarticulata Day, 1963 Indian ques ques Prosphaerosyllis (Hartmann-Schröder, 1979) SW Pacific cas longipapillata** Pseudonereis anomala (Gravier, 1900) Indo-Pacific est est Pseudopolydora Okuda, 1937 Pacific est paucibranchiata Scoletoma debilis (Grube, 1878) Indo-Pacific ques Sigambra constricta (Southern, 1921) Indo-Pacific/Red Sea ques Sigambra parva (Day, 1963) Indian cas Spirobranchus tetraceros (Schmarda, 1861) Circumtropical est Spirorbis marioni Caullery & Mesnil, 1897 Atlantic/Pacific est est Streblosoma comatus** (Grube, 1856) Indo-Pacific est cas est Streblospio Rice & Levin, 1998 W Atlantic est gynobranchiata Syllis alosae San Mart›n, 1992 W Atlantic ques Syllis bella (Chamberlin, 1919) Pacific est Syllis hyllebergi (Licher, 1999) Red Sea est Syllis cf. mayeri Musco & Giangrande, 2005 W Atlantic ques Syllis pectinans Haswell, 1920 Pacific est est Synelmis rigida (Fauvel, 1919) Indo-Pacific ques Syllis schulzi (Hartmann-Schröder, 1962) Indian/Red Sea ques ques Terebella ehrenbergi Grube, 1870 Indo-Pacific/Red Sea ques ques Timarete anchylochaeta (Schmarda, 1861) Pacific ques Timarete caribous (Grube, 1859) W Atlantic cas Timarete dasylophius (Marenzeller, 1879) Indo-Pacific ques Timarete punctata (Grube, 1859) Indo-Pacific est
396 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED CRUSTACEA Amphipoda Bemlos leptocheirus (Walker, 1909) Indian cas Caprella scaura Templeton, 1836 Indian est est est Cymadusa filosa (Savigny, 1816) Indo-Pacific cry cry Elasmopus pectenicrus (Bate, 1862) Circumtropical cas? est est est Gammaropsis togoensis (Schellenberg, 1925) Cosmopolitan cry/est Linguimaera caesaris** Krapp-Schickel, 2003 Indo-Pacific est est Monocorophium sextonae (Crawford, 1937) unknown cry cry cry Parhyale explorator Arresti, 1989 NE Atlantic cas Photis lamellifera Schellenberg, 1928 Indian cas Rhabdosoma whitei Bate, 1862 Red Sea cas Stenothoe gallensis Walker, 1904 Circumtropical ques est est Cirripedia Amphibalanus eburneus** (Gould, 1841) W Atlantic est est est est Austrominius modestus** Darwin, 1854 Tropical Pacific est est Balanus reticulatus Utinomi, 1967 Circumtropical ques est Balanus trigonus Darwin, 1854 Circumtropical est est est est Heterosaccus dollfusip Boschma, 1960 Red Sea est Megabalanus tintinnabulum (Linnaeus, 1758) Circumtropical ques ques est Tetraclita squamosa Pilsbry, 1916 Indo-Pacific cas rufotinta Copepoda Acartia centrura Giesbrecht, 1889Indian cas Acartia fossae Gurney, 1927 Indo-Pacific est Acartia tonsa Dana, 1849W Atlantic / est est est est Indo-Pacific Arietellus pavoninus (G. O. Sars, 1905) Tropical-subtropical cas est Calanopia biloba Bowman, 1957 W subtropical Atlantic cas Calanopia elliptica (Dana, 1846) Indo-Pacific ques est Calanopia media Gurney, 1927 Indo-Pacific est Calanopia minor A. Scott, 1902 Indo-Pacific cas Canuellina insignis Gurney, 1927 Indo-Pacific ques Centropages furcatus (Dana, 1852) Indo-Pacific/Atlantic est Clavellisa ilishaep Pillai, 1962 Indian est Enhydrosoma vicinum Por, 1967 Indo-Pacific ques Euchaeta concinna Dana, 1849 Indo-Pacific cas cas Labidocera agilis (Dana, 1849) Indo-Pacific cas Labidocera detruncata (Dana, 1849) Indo-Pacific ques cas Labidocera madurae (A. Scott, 1909) Indo-Pacific est Labidocera orsinii Giesbrecht, 1889Red Sea cas
Medit. Mar. Sci., 11/2, 2010, 381-493 397 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Labidocera pavo Giesbrecht, 1889Indo-Pacific est Metacalanus acutioperculum Ohtsuka, 1984 Pacific est est Mitrapus oblongusp (Pillai, 1964) Indian est Myicola ostreaep Hoshina & Sigiura, 1953 Pacific est Mytilicola orientalisp Mori, 1935 Pacific est Nothobomolochus fradeip Marques, 1965 Atlantic/Indian est Paracalanus indicus Wolfenden, 1905 Subtropical Atlantic, cry/ques Red Sea Paracartia grani G. O. Sars, 1904 Atlantic est est est est Parvocalanus crassirostris** Dahl, 1894 Indo-Pacific/Atlantic cas est Parvocalanus elegans Andronov, 1972 Indo-Pacific cas Parvocalanus latus Andronov, 1972 Indian cas Pseudocyclops xiphophorusp Wells, 1967 Indian cry Pseudocalanus elongatus (Boeck, 1865) E Atlantic cry cry cry Robertsonia salsa Gurney, 1927 Indo-Pacific ques Scaphocalanus amplius Park, 1970 Subtropical Atlantic cas Scaphocalanus brevirostris Park, 1970 Subtropical Atlantic/ cas Indian Scolecithrix valens Farran, 1926 Subtropical Atlantic cas Scottolana longipes (Thompson & Scott, 1903) Indo-Pacific ques Spinocalanus terranovae Damkaer, 1975 Antarctic ques ques Stenhelia inopinata (A. Scott, 1902) Indo-Pacific ques Stenhelia minuta (A. Scott, 1902) Indo-Pacific ques Subeucalanus subcrassus** Giesbrecht, 1888 Atlantic/Pacific ques ques ques Triconia hawii (Böttger-Schnack Red Sea est & Boxshall, 1990) Triconia rufa (Boxshall & Böttger, 1987) Indian/Red Sea est ques Triconia umerus (Böttger-Schnack Red Sea est cas cas & Boxshall, 1990) Cumacea Eocuma rosae Corbera & Galil, 2007 Indo-Pacific/Red Sea cas Eocuma sarsii (Kossmann, 1880) Indo-Pacific est est Scherocumella gurneyi (Calman, 1927) Red Sea cas Decapoda Actumnus globulus Heller, 1861 Red Sea cas Alpheus audouini Coutiere, 1905 Indo West Pacific est Alpheus inopinatus Holthuis & Gottlieb, 1958 Indian/Red Sea cas est Alpheus migrans Lewinsohn & Holthuis, 1978 Red Sea est Alpheus rapacida de Man, 1908 Indo West Pacific est Ashtoret lunaris (Forsska l, 1775) Indo West Pacific cas Atergatis roseus (Rüppell, 1830) Indo-Pacific est
398 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Calappa hepatica (Linnaeus, 1758) Indo-Pacific cas Calappa pelii Herklots, 1851 E Atlantic cas cas cas Callinectes danae Smith, 1869W Atlantic cas Callinectes sapidus Rathbun, 1896 W Atlantic cas cas est est Carupa tenuipes Dana, 1851 Indo-Pacific est Charybdis feriata (Linnaeus, 1758) Indo-Pacific cas Charybdis japonica (A. Milne-Edwards, 1861) Indo-Pacific cas Charybdis helleri (A. Milne-Edwards, 1867) Indo West Pacific est Charybdis longicollis Leene, 1938 Indian/Red Sea est Charybdis lucifera (Fabricius, 1798) Indo-Pacific cas Coleusia signata (Paulson, 1875) Indo West Pacific est Cryptosoma cristatum Brulle, 1837 Tropical East Atlantic cas Daira perlata (Herbst, 1790) Indo West Pacific cas Dorippe quadridens (Fabricius, 1793) Indo West Pacific est Dyspanopeus sayi (Smith, 1869) N East Atlantic est Eriocheir sinensis H. Milne Edwards, 1853 Pacific cas cas Eucrate crenata de Haan, 1835 Indo-Pacific est est Eurycarcinus integrifrons de Man, 1879Indian/Red Sea cas Farfantepenaeus aztecus (Ives, 1891) W Atlantic est Fenneropenaeus merguiensis (De Man, 1888) Indo West Pacific cas Glabropilumnus laevis** (Dana, 1852) Indian cas cas Gonioinfradens paucidentata (A. Milne Edwards, 1861) Indo-Pacific est? Grapsus granulosus H. Milne-Edwards, 1853 Red Sea cas Halimede tyche (Herbst, 1801) Indo West Pacific cas Hemigrapsus sanguineus (de Haan, 1835) Pacific cas cas Herbstia nitida Manning & Holthuis, 1981 Tropical East Atlantic cas Hyastenus hilgendorfi de Man, 1887 Indo West Pacific est Ixa monodi Holthuis & Gottlieb, 1956 Red Sea est Leptochela aculeocaudata Paulson, 1875 Indo West Pacific est Leptochela pugnax de Man, 1916 Indo West Pacific est Libinia dubia H. Milne Edwards, 1834 W Atlantic est Lucifer hanseni Nobili, 1905 Indo West Pacific ques Macrophthalmus graeffei A. Milne Ewdards, 1873 Indo West Pacific est Marsupenaeus japonicus (Bate, 1888) Indo-Pacific cas cas cas est Melicertus hathor (Burkenroad, 1959) Indian est Menaethius monoceros (Latreille, 1825) Indo-Pacific cas Metapenaeopsis aegyptia Galil & Golani, 1990 Indo-Pacific est Metapenaeopsis mogiensis (Nobili, 1904) Indo West Pacific est consobrina Metapenaeus affinis (H. Milne Edwards, 1837) Indo West Pacific est Metapenaeus monoceros (Fabricius, 1798) Indo West Pacific est est
Medit. Mar. Sci., 11/2, 2010, 381-493 399 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Metapenaeus stebbingi (Nobili, 1904) Indian est est Micippa thalia (Herbst, 1803) Indo West Pacific est Myra subgranulata Kossmann, 1877 Indian/Red Sea est Necora puber (Linnaeus, 1767) NW Atlantic est est cas Notopus dorsipes (Linnaeus, 1758) Indo West Pacific cas Ogyrides mjoebergi (Balss, 1921) Indo West Pacific est Palaemonella rotumana (Borradaile, 1898) Indo West Pacific est Palinurus ornatus (Fabricius, 1798) Indo West Pacific cas Paralithodes camtschaticus (Tilesius, 1815) NE Pacific cas Penaeus semisulcatus de Haan, 1844 Indo West Pacific est Percnon gibbesi (H. Milne Edwards, 1853) W Atlantic est est cas est Periclimenes calmani Tattersall, 1921 Indo-Pacific cas Pilumnopeus vauquelini (Audouin, 1826) Indian/Red Sea cas est est Pilumnus minutus** de Haan, 1835 Indo-Pacific cas est Plagusia squamosa (Herbst, 1790) Indo West Pacific ques est cas cas Portunus segnis** (Forsska l, 1775) Indian cas est est Processa macrodactyla Holthuis, 1952 Tropical East Atlantic est? cas Rhithropanopeus harrisii (Gould, 1841) Noth Atlantic cas? est est Rimapenaeus similis (Smith, 1885) W Atlantic est Scyllarus caparti Holthuis, 1952 Tropical East Atlantic cas Scyllarus posteli Forest, 1963 Temperate-tropical cas East Atlantic Sirpus monodi Gordon, 1953 Tropical East Atlantic cas cas Solenocera crassicornis (H.Milne Edwards, 1837) Indo West Pacific cas Sphaerozius nitidus Stimpson, 1858 Indo West Pacific cas Sternodromia spinirostris** (Miers, 1881) Tropical East Atlantic cas Synalpheus tumidomanus Crosnier & Forest, 1966 Tropical East Atlantic cry cry africanus** Thalamita gloriensis Crosnier, 1962 Indo West Pacific cas Thalamita indistincta Apel & Spiridonov, 1998 Indian cas Thalamita poissonii (Audouin, 1826) Indo West Pacific cry cry Trachysalambria (Steinitz, 1932) Red Sea est est palaestinensis Urocaridella pulchella Yokes & Galil, 2006 Indo-Pacific est Isopoda Anilocra pilchardip Bariche & Trilles, 2006 Indo-Pacific cas Apanthura sandalensis Stebbing, 1900 S Africa est est Cymothoa indicap Schioedte & Meinert, 1884 Indo-Pacific cas Mesanthura spp. Tropical/subtropical ques ques ques Paracerceis sculpta (Holmes, 1904) Subtropical est est est est Paradella dianae (Menzies, 1962) NE Pacific est est? est
400 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Sphaeroma venustissimum Monod, 1931 Tropical/subtropical est E Atlantic Sphaeroma walkeri Stebbing, 1905 Indian est est est Stomatopoda Clorida albolitura Ahyong & Naiyanetr, 2000 Indo West Pacific est Erugosquilla massavensis (Kossmann, 1880) Indian/Red Sea est est Erythrosquilla sp. (postlarvae) Indo-Pacific cas Tanaidacea Zeuxo (Parazeuxo) coralensis Sieg, 1980 Circumtropical est MOLLUSCA Bivalvia Acar plicata (Dillwyn, 1817) Indo-Pacific/Red Sea cas Afrocardium richardi (Audouin, 1826) Red Sea est Alectryonella crenulifera (Sowerby, 1871) Pacific/Red Sea ques Anadara granosa (Linnaeus, 1758) Indo West Pacific cas Anadara inflata (L. A. Reeve, 1844) NW Pacific cas Anadara kagoshimensis** (Tokunaga, 1906) Temperate North est est est Pacific Anadara natalensis (Krauss, 1848) W Pacific/Red Sea est Anadara transversa** (Say, 1822) W Atlantic est est est est Angulus flacca (Roemer, 1871) Indo Pacific/Red Sea cas Antigona lamellaris Schumacher, 1817 Indo-Pacific/Red Sea cas Atactodea glabrata (Gmelin, 1791) Indo Pacific/Red Sea cas cas Brachidontes pharaonis (Fischer, 1870) Indian/Red Sea est est est est Callista florida (Lamarck, 1818) Indian/Red Sea cas cas Cardites akabana (Sturany, 1899) Red Sea cas Chama asperella Lamarck, 1819Indo-Pacific/Red Sea cas Chama aspersa Reeve, 1846 Indo-Pacific est Chama pacifica Broderip, 1834 Indo-Pacific est Circe scripta (Linnaeus, 1758) Indo West Pacific cas Circenita callipyga (von Born, 1778) Red Sea cas Clementia papyracea (Gray, 1825) Indo-Pacific/Red Sea est Crassostrea gigas (Thunberg, 1793) NW Pacific est est est est Dendrostrea frons (Linnaeus, 1758) Indo Pacific/Red Sea est Diplodonta bogii** Van Aartsen, 2004 Red Sea est Divalinga arabica Dekker & Gould, 1994 Persian Gulf/Red Sea cas Dosinia erythraea Römer, 1860 W Pacific/Red Sea est Electroma vexillum (Reeve, 1857) Indian/Red Sea est Ensiculus cultellus (Linnaeus, 1758) Indo West Pacific cas Fulvia australis (Sowerby G.B., 1834) Indo-Pacific/Red Sea cas Fulvia fragilis (Forsska l, 1775) Indian est est est
Medit. Mar. Sci., 11/2, 2010, 381-493 401 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Gafrarium pectinatum (Linnaeus, 1758) Indo-Pacific/Red Sea est Glycymeris arabicus (Adams H., 1871) W Pacific/Red Sea cas Hiatula ruppelliana (Reeve, 1857) Red Sea cas Laternula anatina (Linnaeus, 1758) Indo-Pacific/Red Sea est Limopsis multistriata (Forsska l, 1775) Indo-Pacific/Red Sea cas Mactra lilacea Lamarck, 1818 Indo-Pacific/Red Sea est Mactra olorina Philippi, 1846 Red Sea est Malvufundus regula (Forsska l, 1775) Indo-Pacific/Red Sea est est Mercenaria mercenaria (Linnaeus, 1758) W Atlantic est est Modiolus auriculatus (Krauss, 1848) Indian/Red Sea est Musculista perfragilis (Dunker, 1857) Indo-Pacific/Red Sea est Musculista senhousia (Benson in Cantor, 1842) Temperate North est est est est Pacific Mya arenaria Linnaeus, 1758 N Atlantic est est est Mytilopsis sallei (Recluz, 1849) W Atlantic est Nanostrea exigua Harry, 1985 Indo-Pacific est Paphia textile (Gmelin, 1791) Indo-Pacific/Red Sea est Pedicirce sulcata (Gray, 1838) Red Sea ques Petricola hemprichi (Issel, 1869) Indo-Pacific est Petricola pholadiformis Lamarck, 1818 W Atlantic est Pinctada margaritifera (Linnaeus, 1758) Indo-Pacific/Red Sea ques Pinctada radiata (Leach, 1814) Indo-Pacific/Red Sea est est cas est Psammotreta praerupta (Salisbury, 1934) Indo-Pacific/Red Sea cas Pseudochama corbieri (Jonas, 1846) Red Sea est Ruditapes philippinarum (Adams & Reeve, 1850) Temperate North est est est est Pacific Saccostrea commercialis (Iredale & Roughley, 1933) Australia cas Saccostrea cucullata (Born, 1778) Indo-Pacific ques est Septifer bilocularis (Linnaeus, 1758) Indo-Pacific cas Septifer forskali Dunker, 1855 Red Sea est Sphenia rueppelli A. Adams, 1850 Indian est Spondylus groschi Lamprell & Kilburn, 1995 Indian/Red Sea ques Spondylus cf. multisetosus Reeve, 1856 Indo-Pacific ques Spondylus nicobaricus Schreibers, 1793 Indo-Pacific cas Spondylus spinosus Schreibers, 1793 Indo-Pacific/Red Sea est Tellina valtonis Hanley, 1844 Indian/Red Sea est Teredo navalis Linnaeus, 1758 Circumtropical cry cry Theora lubrica Gould, 1861 Indo-Pacific est est Timoclea marica (Linnaeus, 1758) Red Sea cas Trapezium oblongum (Linnaeus, 1758) Indo-Pacific/Red Sea ques Xenostrobus securis (Lamarck, 1819) Tropical Atlantic est est
402 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Zygochlamys patagonica** (King & Broderip, 1832) W Atlantic cas cas Cephalopoda Octopus aegina Gray, 1849Indo-Pacific/Red Sea est Octopus cyanea Gray, 1849Indo-Pacific cas Sepia gibba Ehrenberg, 1831 Red Sea cas Sepia pharaonis Ehrenberg, 1831 Pacific/Red Sea ques Sepioeuthis lessoniana Lesson, 1830 Indo-Pacific est Tremoctopus gracilis (Eydoux & Souleyet, 1852) Indo-Pacific cas cas Gastropoda Acteocina crithodes Melvill & Standen, 1907 Indo-Pacific/W Indian cas Acteocina mucronata (Philippi, 1849) Red Sea ques est Alvania dorbignyi (Audouin, 1826) Cosmopolitan cry cry cry Amathina tricarinata (Linnaeus, 1767) Red Sea est Angiola punctostriata (Smith E.A., 1872) Red Sea ques Anteaeolidiella foulisi** (Angas, 1864) Circumtropical cas cas Aplysia dactylomela Rang, 1828 Circumtropical est est est Aplysia parvula Guilding in Mörch, 1863 Circumtropical cry cry cry Atys angustatus Smith, 1872 Red Sea cry Atys cylindricus (Helbling, 1779) Indo-Pacific/Red Sea cas Bostrycapulus odites** Collin, 2005 Tropical Atlantic est Bulla arabica ** Malaquias & Reid, 2008 Indo-Pacific est Bursatella leachii De Blainville, 1817 Circumtropical est est est est Caloria indica (Bergh, 1896) Indo-Pacific cas Canarium mutabile** (Swainson, 1821) Indo-Pacific/Red Sea ques Cantharus tranquebaricus (Gmelin, 1791) Indian cas Cellana rota (Gmelin, 1791) Indian/Red Sea cas cas est Cerithidium diplax** (Watson, 1886) Persian Gulf est Cerithidium perparvulum** (Watson, 1886) Pacific est Cerithiopsis pulvis (Issel, 1869) Red Sea est Cerithiopsis tenthrenois (Melvill, 1896) Indian est Cerithium columna Sowerby 1834 Indo West Pacific ques Cerithium egenum Gould, 1849Indo-Pacific/Red Sea est Cerithium litteratum (Born, 1778) W Atlantic cas Cerithium nesioticum Pilsbry & Vannata, 1906 Indian/Red Sea est Cerithium nodulosum Bruguière, 1792 Indo West Pacific ques Cerithium scabridum Philippi, 1848 Indian/Red Sea est est est Chelidonura fulvipunctata Baba, 1938 Indo-Pacific est est Chromodoris annulata (Eliot, 1904) Indian est Chromodoris quadricolor (Rüppell & Leuckart, 1828) Red Sea cas cas est Chrysallida fischeri Hornung & Mermod, 1925) Red Sea est est Chrysallida maiae (Hornung & Mermod, 1924) Red Sea est
Medit. Mar. Sci., 11/2, 2010, 381-493 403 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Chrysallida micronana Öztürk & van Aartsen, 2006 Red Sea cas Chrysallida pirintella (Melvill, 1910) Red Sea est Cingulina isseli (Tryon, 1886) Subtropical est Clypeomorus bifasciatus (Sowerby G.B. II, 1855) Indo-Pacific/Red Sea cas est Conomurex persicus** (Swainson, 1821) Persian Gulf est cas est Conus arenatus Hwass, 1792 Indo-Pacific/Red Sea cas Conus fumigatus Hwass, 1792 Red Sea cas Conus inscriptus Reeve, 1843 Indian cas Conus rattus Hwass, 1792 Indo-Pacific cas Coralliobia madreporarum (Sowerby, 1832) Pantropical cas cas Crepidula fornicata (Linnaeus, 1758) NW Atlantic est cas est Cuthona perca (Marcus, 1958) Tropical Atlantic cas Cycloscala hyalina (Sowerby, 1844) Indo-Pacific/Red Sea est Cylichnina girardi (Audouin, 1826) Indo-Pacific est Dendrodoris fumata (Rüppell & Leuckart, 1830) Indo Pacific/Red Sea cas Diala semistriata (Philippi, 1849) Indo-Pacific/Red Sea est Diodora funiculata (Reeve, 1850) Indo-Pacific est Diodora ruppellii (Sowerby, 1834) Indo-Pacific/Red Sea cas est Discodoris lilacina (Gould, 1852) Indo-Pacific/Red Sea cry cry Doxander vittatus** (Linnaeus, 1758) Indo-Pacific cas Echinolittorina punctata (Gmelin, 1791) Tropical Atlantic Native est Native Elysia grandifolia Kelaart, 1858 Indian est Elysia tomentosa Jensen, 1997 Indo West Pacific est Engina mendicaria (Linnaeus, 1758) Indo West Pacific cas Ergalatax contracta (Reeve, 1846) Red Sea cas Ergalatax junionae** Houart, 2008 Persian Gulf/Red Sea est Erosaria turdus (Lamarck, 1810) Indian/Red Sea est est Ethminolia hemprichi (Issel, 1869) Red Sea cas Favorinus ghanensis Edmunds, 1968 Tropical Atlantic est Finella pupoides A. Adams, 1860 Indo-Pacific est Flabellina rubrolineata (O’Donoghue, 1929) Indo-Pacific/Red Sea est Fusinus verrucosus (Gmelin, 1791) Indian est Gastrochaena cymbium (Spengler, 1783) Indo-Pacific/Red Sea est Gibborissoa virgata (Philippi, 1849) Indo-Pacific est Gibbula albida (Gmelin, 1791) Adriatic Sea est Native Godiva quadricolor (Barnard, 1927) Eastern Pacific cas Halgerda willeyi Elliot 1904 Indo West Pacific cas Haliotis pustulata cruenta Reeve, 1846 Indian/Red Sea cas cas Haminoea cyanomarginata Heller & Thompson, 1983 Red Sea est est Haminoea japonica** (Pilsbry, 1895) Pacific est Hinemoa cylindrica (de Folin, 1879) Indo-Pacific cas
404 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Hypselodoris infucata (Rüppell & Leuckart, 1828) Indo-Pacific/Red Sea est Iolaea neofelixoides (Nomura, 1936) Pacific cas Leucotina cfr eva Thiele, 1935 Indo-Pacific cas Leucotina natalensis** Smith, 1910 Indo-Pacific/Red Sea est Lienardia mighelsi Iredale & Tomlin, 1917 Pacific cas Littorina saxatilis (Olivi, 1792) Atlantic cry Melibe viridis** (Kelaart, 1858) Indo-Pacific est est est est Metaxia bacillum (Issel, 1869) Red Sea est Mitrella psilla (Duclos, 1846) Tropical Atlantic est Monetaria annulus (Linnaeus, 1758) Indo-Pacific ques Monotigma lauta** (A. Adams, 1853) Indo-Pacific/Red Sea est Murchisonella columna (Hedley, 1907) Indo-Pacific cas Murex forskoehlii Roeding, 1798 Persian Gulf/Red Sea est Nassa situla (Reeve, 1846) Indo-Pacific cas Nassarius arcularia plicatus (Roeding, 1798) Indian/Red Sea cas Nassarius concinnus (Powys, 1835) unknown cas Nassarius stolatus (Gmelin, 1791) Indo-Pacific cas Nerita sanguinolenta Menke, 1829Red Sea cas est Notocochlis gualteriana** (Recluz, 1844) Indo-Pacific/Red Sea cas est Odostomia lorioli (Hornung & Mermod, 1924) Red Sea cas Oscilla jocosa Melvill, 1904 Persian Gulf cas Oxynoe viridis (Pease, 1861) Indo West Pacific est Palmadusta lentiginosa (Gray, 1825) Indian cas lentiginosa Parviturbo dibellai Buzzurro & Celalupo, 2006 unknown cry Patelloida saccharina (Linnaeus, 1758) Pacific cas Philinopsis cyanea (Martens, 1879) Indian/Red Sea cas Planaxis griseus (Brocchi, 1821) Red Sea cas Pleurobranchus forskalii Rüppell & Leuckart, 1828 Red Sea cas Plocamopherus ocellatus Rüppell & Leuckart, 1830 Red Sea est Polycera hedgpethi Marcus Er., 1964 NE Pacific cas cas Polycerella emertoni Verrill, 1881 Pantropical est est Pseudominolia nedyma Melvill, 1897 Persian Gulf/Red Sea est Purpuradusta gracilis notata (Gill, 1858) Indian/Red Sea est Pyrunculus fourierii (Audouin, 1826) Indo-Pacific/Red Sea est Rapana rapiformis (Von Born, 1778) Indo-Pacific/Red Sea ques Rapana venosa (Valenciennes, 1846) Temperate North est est est Pacific Retusa desgenettii (Audouin, 1826) Red Sea cas Rhinoclavis kochi (Philippi, 1848) Indo-Pacific/Red Sea est Rhinoclavis sinensis (Gmelin, 1791) Indo West Pacific cas
Medit. Mar. Sci., 11/2, 2010, 381-493 405 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Rissoina ambigua (Gould, 1849) Indo-Pacific cas Rissoina bertholleti Issel, 1869 Indian/Red Sea est Rissoina spirata Sowerby, 1825 Indo-Pacific cas cas Sabia conica (Schumacher, 1817) Indo-Pacific/Red Sea ques ques Siphonaria crenata de Blainville, 1827 Persian Gulf/ est Arabian Sea Siphonaria pectinata (Linnaeus, 1758) Atlanto-Mediterranean Native est Smaragdia souverbiana (Montrouzier, 1863) Indo-Pacific/Red Sea est Sticteulima cf. lentiginosa (A. Adams, 1861) Indo-Pacific cas Stomatella impertusa (Burrow, 1815) Indo-Pacific/Red Sea cas Syphonota geographica (Adams & Reeve, 1850) Circumtropical est est Syrnola cinctella A. Adams, 1860 Indo-Pacific/Red Sea cas Syrnola fasciata (Jickeli, 1882) Indo-Pacific est Syrnola lendix** (A. Adams, 1863) Red Sea est Thais lacera (von Born, 1778) Indian cas est Thais sacellum (Gmelin, 1791) Indian/Red Sea est Tornus jullieni Adam & Knudsen 1969 W Africa ques Trivirostra triticum Schilder, 1932 Indo-Pacific cas Trochus erithreus Brocchi, 1821 Persian Gulf est Turbonilla edgarii (Melvill, 1896) Indo-Pacific est Vexillum depexum (Deshayes in Laborde, 1834) Indian/Red Sea ques Voorwindia tiberiana (Issel, 1869) Red Sea cas Zafra savignyi (Moazzo, 1939) Red Sea est Zafra selasphora (Melvill & Standen, 1901) Indian/Red Sea est Polyplacophora Acanthopleura gemmata (de Blainville, 1825) Indo-Pacific/Red Sea cas Chiton hululensis (Smith E.E. in Gardiner, 1903) Indo-Pacific/Red Sea ques MISCELLANEA INVERTEBRATA Arthropoda/Pycnogonida Ammothea hilgendorfi (Böhm, 1879) Circumboreal est Anoplodactylus californicus Hall, 1912 Circum(sub)tropical est est Anoplodactylus digitatus (Böhm, 1879) Circum(sub)tropical est Pigrogromitus timsanus Calman, 1927 Circum(sub)tropical ques Bryozoa Aeverrillia setigera (Hincks, 1887) Circumtropical cas Arachnoidea protecta (Harmer, 1915) Indo-Pacific ques Bowerbankia gracillima (Hincks, 1877) E Atlantic cas cas Bugula fulva Ryland, 1960 NW Atlantic est cas cas cas Bugula serrata (Lamarck, 1816) Indo-Pacific est Celleporaria aperta (Hincks, 1882) Circumtropical ques est Celleporaria brunnea (Hincks, 1884) NE Pacific est
406 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Celleporaria pilaefera (Canu & Bassler, 1929) Indo-W Pacific ques Celleporella carolinensis Ryland, 1979 W Atlantic est Crepidacantha poissoni (Audouin, 1826) Circumtropical ques Electra tenella (Hincks, 1880) Circumtropical cas Hippaliosina acutirostris Canu & Bassler, 1929 Indo-Pacific ques Hippopodina feejeensis (Busk, 1884) Indo-Pacific cas Parasmittina egyptiaca (Waters, 1909) Indian est Parasmittina serruloides Harmelin, Bitar & Indo-Pacific est Zibrowius, 2009 Parasmittina spondylicola Harmelin, Bitar & Indo-Pacific cas Zibrowius, 2009 Pherusella brevituba Soule, 1951 NE Pacific cas cas Reteporella jermanensis (Waters, 1909) Red Sea cas Rhynchozoon lareyi (Audouin, 1826) Indo-Pacific est Schizoretepora hassi Harmelin, Bitar & Red Sea? ques Zibrowius, 2007 Scrupocellaria jolloisii (Audouin, 1826) Indo-Pacific est Smittina malleolus (Hincks, 1884) Circumtropical est Tricellaria inopinata d'Hondt & Occhipinti, 1985 Indo-Pacific est est Chaetognatha Aidanosagitta neglecta* Aida, 1897 Indo-Pacific est Ferosagitta galerita* (Dallot, 1971) Indian est Chordata/Ascidiacea Ascidia cannelata Oken, 1820 Indo-Pacific est Botrylloides violaceus Oka, 1927 NW Pacific est Cystodytes philippinensis Herdman, 1886 Circumtropical cry cry Distaplia bermudensis Van Name, 1902 W Atlantic est est Ecteinascidia styeloides (Traustedt, 1882) Circumtropical est est Ecteinascidia thurstoni Herdman, 1890 Indo-Pacific est Herdmania momus (Savigny 1816) Indo-Pacific est Microcosmus exasperatus Heller, 1878 Indo-Pacific est est Microcosmus squamiger Hartmeyer & Michaelsen, 1928 Circumtropical est est Perophora multiclathtrata (Sluiter, 1904) Circumtropical est Phallusia nigra Savigny, 1816 Circumtropical est Polyandrocarpa zorritensis (Van Name, 1931) E Pacific est est Rhodosoma turcicum (Savigny, 1816) Circumtropical cas cas est Styela clava Herdman, 1881 NW Pacific cas Symplegma brakenhielmi (Michaelsen, 1904) Indo-Pacific est Trididemnum cf. savignii (Herdman, 1886) Circum(sub)tropical est cas Cnidaria/Anthozoa Acabaria erythraea (Ehrenberg, 1834) Indo-Pacific est
Medit. Mar. Sci., 11/2, 2010, 381-493 407 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Diadumene cincta Stephenson, 1925 NE Atlantic est Haliplanella lineata (Verrill, 1870) Circumboreal est Oculina patagonica De Angelis, 1908 ?SW Atlantic est est Cnidaria/Hydrozoa Aequorea conica* Browne, 1905 Indo-Pacific cas Amphogona pusilla* Hartlaub, 1909 Indo-Pacific cas Arctapodema australis* Antarctic/Indo-Pacific cas cas Bougainvillia niobe Mayer, 1894 (sub)tropical est W Atlantic Campalecium medusiferum* Torrey, 1902 E Pacific/ Atlantic est cas Cirrholovenia tetranema* Kramp, 1959 Circumtropical est cas Clytia hummelincki* (Leloup, 1935) Circumtropical est est est Clytia linearis** (Thornely, 1900) Circum(sub)tropical est est est est Clytia mccradyi* (Brooks, 1888) Circumtropical est est Cordylophora caspia (Pallas, 1771) Circumglobal cry cry cry cry Coryne eximia* Allman, 1859Circum(sub)tropical ques ques Diphasia digitalis (Busk 1852) Circumtropical cas Dynamena quadridentata (Ellis & Solander, 1786) Circumtropical est Eirene viridula* (Péron & Lesueur, 1810) Circumtropical est est est est Eucheilota paradoxica* Mayer, 1900 Circumtropical cas cas cas est Eudendrium carneum Clarke, 1882 Circumtropical est cas est Eudendrium merulum Watson, 1985 Circumtropical est cas est Euphysora annulata* Kramp, 1928 Indo-Pacific cas cas Euphysora bigelowi* Maas, 1905 Indo-Pacific cas Filellum serratum* (Clarke, 1879) Circum(sub)tropical cas Garveia franciscana (Torrey, 1902) Circum(sub)tropical cas est Gonionemus vertens* A. Agassiz, 1862 Circumboreal est est Kantiella enigmatica* Bouillon, 1978 Indian cas Laodicea fijiana* Agassiz & Mayer, 1899 Indo-Pacific cas Macrorhynchia philippina (Kirchenpauer, 1872) Circumtropical est Moerisia carina* Bouillon, 1978 Indo-Pacific est Moerisia inkermanica* Paltschikowa-Ostroumova, 1925 Circum(sub)tropical cas cas Nubiella mitra* Bouillon, 1980 SW Pacific cas Octotiara russelli* Kramp, 1953 Indo-Pacific cas Olindias singularis* Browne, 1905 Indian est Paracytaeis octona* Bouillon, 1978 W Indian cas Russellia mirabilis Kramp, 1957 Antarctic cas Scolionema suvaensis* (Agassiz & Mayer, 1899) Indo-Pacific est Sertularia marginata (Kirchenpauer, 1864) Circum(sub)tropical est est Sertularia thecocarpa (Jarvis, 1922) Indo-Pacific est Tetrorchis erythrogaster* Bigelow, 1909 Circumtropical est
408 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Trichydra pudica* Wright, 1858 Circumtropical cas cas Cnidaria/Scyphozoa Cassiopea andromeda (Forsska l, 1775) Indo-Pacific est? est Marivagia stellata Galil & Gershwin, 2010 ?Indo-Pacific est Circumtropical Phyllorhiza punctata* von Lendenfeld, 1884 Circumtropical cas cas est Rhopilema nomadica* Galil, Spannier & Red Sea cas est Ferguson, 1990 Stomolophus meleagris* (L. Agassiz, 1862) Atlantic/Pacific cas Ctenophora Beroe ovata* Mayer, 1912 Circum(sub)tropical est est Mnemiopsis leidyi* A. Agassiz, 1865 NW Atlantic est est est est Sulculeolaria angusta* Totton, 1954 Indian est Echinodermata Acanthaster planci (Linnaeus, 1758) Indo-Pacific ques Amphiodia (Amphispina) Mortensen, 1940 Indo-Pacific est obtecta Amphioplus (Lymanella) (Lyman, 1874) Indo-Pacific cas laevis Aquilonastra burtoni** (Gray, 1840) W Indian est Asterias rubens Linnaeus, 1755 NE Atlantic est Diadema setosum (Leske, 1778) Indo West Pacific cas Eucidaris tribuloides (Lamarck, 1816) (sub)tropical Atlantic: est Ophiactis macrolepidota Marktanner-Turneretscher, Circumtropical est 1887 Ophiactis savignyi (Müller & Troschel, 1842) Circumtropical cas est est Protoreaster nodosus (Linnaeus, 1758) Indo-Pacific cas Prionocidaris baculosa (Lamarck, 1816) Indian cas Synaptula reciprocans (Forsska l, 1775) Indo-Pacific est Porifera Cinachyrella australiensis Carter, 1886 Indo-Pacific ques Geodia micropunctata Row, 1911 Indo-Pacific ques Haliclona spinosella (Thiele, 1905) Indo-Pacific ques Haliclona viridis (Keller, 1881) Circum(sub)tropical ques Hyrtios erecta (Keller, 1889) Indo-Pacific ques Lissodendoryx (Ridley, 1844) Indo-Pacific ques schmidti Mycale erythraeana Burton, 1936 Indo-Pacific ques Paraleucilla magna Klautau et al., 2004 SW Atlantic cry cry cry Siphonophora Abyla trigona* Quoy & Gaimard,1827 Indian cas
Medit. Mar. Sci., 11/2, 2010, 381-493 409 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Sipuncula Apionsoma (Apionsoma) (Ikeda, 1904) Circumtropical ques ques misakianum Apionsoma (Apionsoma) (Sluiter, 1902) Circumtropical cas tricocephalus Aspidosiphon (Akrikos) (Murina, 1967) Circumtropical ques cas cas mexicanus Aspidosiphon (Aspidosiphon) (Chamisso & Eysenhardt, Circumtropical est elegans 1821) Phascolion (Isomya) (Sluiter, 1902) Indo-Pacific est cas convestitum Phascolosoma scolops (Selenka & de Man, 1883) Indo-Pacific est est Platyhelminthes Allolepidapedon fistulariae p Yamaguti, 1940 Indo-Pacific cas Glyphidohaptor plectocirrap (Paperna, 1972) Red Sea est Hirudinella ventricosa p (Pallas, 1774) Atlantic cas Hysterolecitha sigani p Manter, 1969 Indo-Pacific ques Lecithochirium Fischthal & Kuntz, 1963 Red Sea cas magnicaudatum Monilicaecum ventricosum p Yamaguti, 1942 Indo-Pacific cas Neothoracocotyle (Meserve, 1938) Hargis, 1956 Atlantic/Pacific cas acanthocybii p Nosema ceratomyxa p Diamant & Paperna, 1985 Red Sea est Polylabris cf. mamaevi p Ogawa & Egusa, 1980 Indo West Pacific est Tetrancistrum polymorphus p (Paperna, 1972) Red Sea cas cas Tetrancistrum Kritsky, Galli & Yang, 2007 Red Sea est strophosolenum p Tetrancistrum suezicus p (Paperna, 1972) Red Sea cas est FISH Elasmobranchii Carcharhinus altimus (Springer, 1950) Indo-Pacific est est Carcharhinus falciformis (Müller & Henle, 1839) Circumtropical est Dasyatis marmorata (Smith, 1828) Tropical Atlantic ques ques Galeocerdo cuvier (Peron & Le Sueur, 1822) Tropical/subtropical cas cas Himantura uarnak (Forsska l, 1775) Indo-Pacific est Isurus paucus Guitart Manday, 1966 Circumtropical cas Rhizoprionodon acutus (Rüppell, 1837) Circumtropical cas Sphyrna mokarran (Rüppell, 1837) Circumtropical cas Torpedo sinuspersici Olfers, 1831 Indo-Pacific ques Actinopterygii Abudefduf vaigiensis (Quoy & Gaimard, 1825) Indo-Pacific cas cas
410 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Acanthurus monroviae Steindachner, 1876 Tropical Atlantic est cas cas Alepes djedaba (Forsska l, 1775) Indo-Pacific est Aluterus monocerus (Linnaeus, 1758) Atlantic cas Anarhichas lupus Linnaeus, 1758 N Atlantic cas Anguilla japonica Temminck & Schlegel, 1847 Pacific ques Apogon fasciatus (White, 1790) Indo-Pacific est Apogon pharaonis Bellotti, 1874 Indo-Pacific est Apogon queketti Gilchrist, 1903 Indian est Apogon smithi (Kotthaus, 1970) Indian est Arius parkii Gunther, 1864 Indo-Pacific cas Atherinomorus forskalii** (Rüppell, 1838) Tropical Atlantic est est Beryx splendens Lowe, 1834 Circumtropical est est Bregmaceros atlanticus Goode & Bean, 1886 Tropical Atlantic cas est Callionymus filamentosus Valenciennes, 1837 Circumtropical est Centrolabrus exoletus (Linnaeus, 1758) Tropical Atlantic cas Cephalopholis taeniops (Valeciennes, 1828) Subtropical Atlantic est cas Champsodon nudivittis (Ogilby, 1895) Tropical Atlantic cas Champsodon vorax Günther, 1867 Indo West Pacific cas Chaunax suttkusi Caruso, 1898 Indo-Pacific cas cas Cheilodipterus (Rüppell, 1838) W Indian cas novemstriatus Cheilopogon furcatus (Mitchill, 1815) Circumtropical cas Chilomycterus reticulatus (Linnaeus, 1758) Circumtropical cas Chirocentrus dorab Forsska l, 1775) Indo-Pacific cas Coryogalops ochetica (Norman, 1927) Red Sea est Crenidens crenidens (Forsska l, 1775) Indian est est Cyclichthys spilostylus (Leis & Randall, 1982) Indo-Pacific est Cyclopterus lumpus Linnaeus, 1758 Atlantic cas Cynoglossus sinusarabici (Chabanaud, 1913) Red Sea est Decapterus russelli (Rüppell, 1830) Indo-Pacific est Diodon hystrix (Linnaeus, 1758) Circumtropical cas ques Diplodus bellottii (Steindachner, 1882) Tropical Atlantic est Dussumieria elopsoides Bleeker, 1849Indo-Pacific est Elates ransonetti (Steindachner, 1876) Tropical Pacific cas cas Enchelycore anatina (Lowe, 1839) Tropical Atlantic est Epinephelus coioides (Hamilton, 1822) Indo-Pacific cas est Epinephelus malabaricus (Bloch & Schneider, 1804) Indo-Pacific est Equulites klunzingeri** (Steindachner, 1898) Indian cas cas est Etrumeus teres (DeKay, 1848) Subtropical est est Favonigobius (Fowler, 1934) Indo West Pacific cas melanobranchus**
Medit. Mar. Sci., 11/2, 2010, 381-493 411 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Fistularia commersonii (Rüppell, 1835) Indo-Pacific est est est est Fistularia petimba Lacepède, 1803 Tropical-Atlantic cas Gephyroberyx darwini (Johnson, 1866) Tropical Atlantic cas ques Glaucostegus halavi (Forsska l, 1775) Indo-Pacific cas ques Gymnammodytes (Jourdain, 1879) N Atlantic est semisquamatus Halosaurus ovenii Johnson, 1863 Tropical Atlantic est Hemiramphus far (Forsska l, 1775) Indo-Pacific est cas est Heniochus intermedius Steindachner, 1893 Indian cas Herklotsichthys punctatus (Rüppell, 1837) Red Sea est Hippocampus fuscus Rueppell, 1838 Indian est Hyporhamphus affinis (Gunther, 1866) Indo-Pacific cas Iniistius pavo Valenciennes, 1840 Indo-Pacific cas Kyphosus incisor (Cuvier, 1831) Tropical Atlantic cas Kyphosus sectator (Linnaeus, 1758) Tropical Atlantic est Lagocephalus sceleratus (Gmelin, 1788) Indo-Pacific cas est Lagocephalus spadiceus (Richardson, 1844) Indo-Pacific est Lagocephalus suezensis Clark & Gohar, 1953 Red Sea est Liza carinata (Valenciennes, 1836) Indian est Liza haematocheila (Temminck & Schlegel, 1845) Indo-Pacific est Lutjanus argentimaculatus (Forsska l, 1775) Indo-Pacific cas Lutjanus jocu (Bloch & Schneider, 1801) Atlantic cas Makaira indica (Cuvier, 1832) Indo-Pacific cas cas Microchirus boscanion Chabanaud, 1926 Tropical Atlantic est cas Microchirus hexophthalmus (Bennet, 1831) Tropical Atlantic est est Monotaxis grandoculis (Forsska l, 1775) Indo-Pacific cas Muraenesox cinereus (Forsska l, 1775) Indo-Pacific cas Mycteroperca fusca (Lowe, 1838) Tropical Atlantic cas Nemipterus randalli Russell, 1986 Indian est Omobranchus punctatus (Valenciennes, 1836) Indo-Pacific cas Oplegnathus fasciatus (Temminck & Schlegel, 1944) Pacific cas Oxyurichthys petersi (Klunzinger, 1871) Red Sea est Pagellus bellottii Steindachner, 1882 Tropical Atlantic est cas Pagrus major (Temminck & Schlegel, 1843) NW Pacific cas Pampus argenteus (Euphrasen, 1788) Indo-Pacific cas Papilloculiceps longiceps (Ehrenberg in Valenciennes, Indian cas 1829) Parexocoetus mento (Valenciennes, 1846) Indo-Pacific est est est Parupeneus forsskali (Fourmanoir & Guézé, 1976) Indian ques cas Pelates quadrilineatus (Bloch, 1790) Indo-Pacific est Pempheris vanicolensis Cuvier, 1831 Indo-Pacific est est
412 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Petroscirtes ancylodon Rüppell, 1838 Indian est Pinguipes brasilianus Cuvier & Valenciennes, 1829WAtlantic cas cas Pisodonophis semicinctus (Richardson, 1848) Tropical Atlantic est est cas Platax teira (Forsska l, 1775) Indo-Pacific cas Platycephalus indicus (Linnaeus, 1758) Indo-Pacific cas est Plotosus lineatus (Thunberg, 1787) Indo-Pacific est Pomacanthus imperator (Bloch, 1787) Indo-Pacific cas Pomacanthus maculosus (Forsska l, 1775) Indo-Pacific cas Pomadasys stridens (Forsska l, 1775) Indian est est Priacanthus hamrur (Forsska l, 1775) Indo-Pacific ques ques Priacanthus sagittarius Starnes, 1988 Indo-Pacific cas Psenes pellucidus Lutken, 1880 Tropical Atlantic est cas Pseudupeneus prayensis (Cuvier, 1829) Tropical Atlantic cas Pteragogus pelycus Randall, 1981 Indian est Pterois miles (Bennet, 1803) Indian cas Rachycentron canadum (Linnaeus, 1766) Circumtropical cas cas Rastrelliger kanagurta (Cuvier, 1816) Indo-Pacific cas Rhabdosargus haffara (Forsska l, 1775) Indian est Rhynchoconger trewavasae Ben-Tuvia 1993 Red Sea cas Sargocentron rubrum (Forsska l, 1775) Indo-Pacific cas est Saurida undosquamis (Richardson, 1848) Indo-Pacific est est est Scarus ghobban Forsska l, 1775 Indo-Pacific est Sciaenops ocellatus (Linnaeus, 1766) W Atlantic cas Scomberomorus commerson Lacepède, 1800 Indo-Pacific ? est est Scorpaena stephanica Cadenat, 1943 Tropical Atlantic cas Selene dorsalis Gill, 1862 W Atlantic cas Seriola carpenteri Mather 1971 Tropical Atlantic est Seriola fasciata (Bloch, 1793) Tropical Atlantic est est cas Seriola rivoliana Valenciennes, 1833 Tropical Atlantic cas Siganus javus (Linnaeus, 1766) Indo-Pacific cas Siganus luridus (Rüppell, 1829) Indian cas est est Siganus rivulatus Forsska l, 1775 Red Sea est cas est Silhouetta aegyptia (Chabanaud, 1933) Red Sea est Sillago sihama (Forsska l, 1775) Indo-Pacific est Solea senegalensis Kaup, 1858 Tropical Atlantic est ques Sorsogona prionota (Sauvage, 1873) Indian cas Sphoeroides marmoratus (Lowe, 1838) Atlantic cas cas Sphoeroides pachygaster (Müller & Troschel, 1848) Tropical Atlantic est est est est Sphyraena chrysotaenia Klunzinger, 1884 Indo-Pacific est est est Sphyraena flavicauda Rüppell, 1838 Indian est est Spratelloides delicatulus (Bennett, 1831) Indo-Pacific/Red Sea est
Medit. Mar. Sci., 11/2, 2010, 381-493 413 Table 2 (continued)
Species Author Origin WMED CMED ADRIA EMED Stephanolepis diaspros Fraser-Brunner, 1940 Red Sea est cas est Synagrops japonicus (Doderlein, 1884) Indo-Pacific cas Synaptura lusitanica Capello, 1868) Tropical Atlantic est est Syngnathus rostellatus Nilsson, 1855 N Atlantic cas ques Terapon jarbua (Forsska l, 1775) Indo-Pacific cas Terapon puta (Cuvier, 1929) Indo-Pacific/Red Sea est Terapon theraps Cuvier, 1829Indo-Pacific cas Tetrosomus gibbosus (Linnaeus, 1758) Indo-Pacific est Torquigener flavimaculosus Hardy & Randall, 1983 Indian est Trachurus indicus Necrasov, 1966 Indian cas Trachyscorpia cristulata (Koehler, 1869) Tropical Atlantic est est echinata Tridentiger trigonocephalus (Gill, 1859) Tropical Pacific cas Trypauchen vagina (Bloch & Scheider, 1801) Indo-Pacific cas Tylerius spinosissimus (Regan, 1908) Indo-Pacific est Tylosurus choram (Rüppell, 1837) Indo-Pacific cas Tylosurus crocodilus (Peron & Le Sueur, 1821) Indo West Pacific cas Upeneus moluccensis (Bleeker, 1855) Indo-Pacific est est Upeneus pori Ben-Tuvia & Golani, 1989 Indian est est Vanderhorstia mertensi Klausewitz, 1974 Indo-Pacific est Zenopsis conchifera (Lowe, 1852) Atlantic/Pacific cas
1. THE WESTERN MEDITERRANEAN waters form a permanent clockwise gyre in the Alboran Sea, which is separated from The WMED occupies a key position the remainder of the WMED by a well- because it receives the influx of surface marked hydrographic front between Oran waters from the Atlantic, through the Strait and Almer a. As a consequence of the of Gibraltar. It is further compartmental- admixture of fauna and flora of different ized into fairly isolated sub-basins with dif- origins, the westernmost part of the Albo- ferent climatic and hydrologic conditions. ran Sea, from the Ceuta region to Punta de These sub-basins have a different biogeo- Calaburras on the coast of Malaga, consti- graphic character, which may affect inva- tutes a hot spot of biodiversity. Its enriched sion and settlement of aliens. The Alboran benthic environment includes some species Sea, situated immediately east of Gibraltar, linked with the tropical periods of the exhibits stronger Atlantic affinities, due to Mediterranean Sea (OCAN~A et al., 2009; the continued penetration of Atlantic flora GARC A RASO et al., 2010; URRA et al., and fauna with the incoming influx of water in press). The Tyrrhenian Sea is compara- (HARMELIN & D'HONDT, 1993). In tively isolated from the rest of the WMED return, most Mediterranean endemics are and is surrounded by mountains reducing rare or missing. The incoming Atlantic the impact of the meteorological events
414 Medit. Mar. Sci., 11/2, 2010, 381-493 that strongly influence the internal condi- from the Gulf of Naples, South Tyrrhenian tions of the other Western Mediterranean (SGARRELLA & MONCHARMONT, basins (ASTRALDI et al., 1995). The Gulf 1993), from the Ligurian Sea and north of Lions and the Ligurian Sea are the cold- Tyrrhenian (BANCHETTI et al., 2009) est parts of the WMED. They are charac- and from the coast of Vulcano, Aeolian terized by a severe reduction of ther- Islands (CIMERMAN & LANGER, 1991, mophilic species, while some cold temper- as Conorbella imperatoria). ate species, not found to the south, are present (BIANCHI & MORRI, 1993, 1.2. Alien macrophytes in the Western 1994). Mediterranean A total of 91 alien macrophytes have 1.1. Alien protozoans in the Western been hitherto reported in the WMED, Mediterranean which represent 72% of the alien macro- Four pathogenic protozoans have phytes reported from the Mediterranean been recorded in WMED. Three of them Sea. The major vectors of introduction are are pathogenic to shellfish: Marteilia refrin- the shellfish aquaculture industry and ship gens (RIERA et al., 1993), Bonamia ostreae traffic (fouling, ballast waters). As much as (MONTES & LAMA, 1993), and Perkin- 75 alien macrophyte species are well estab- sus olseni, formerly Perkinsus atlanticus lished, seven are casual, and nine are cryp- (SAGRISTA et al., 1996). The fourth, togenic/questionable. Casual taxa (i.e. Photobacterium damselae, formerly Pas- Cladophora hutchinsioides, Fucus spiralis, teurella piscicida, causes pasteurello- Grateloupia patens, Lomentaria flaccida, sis/photobacteriosis in fish: it was first iso- Rhodophysema georgei, Saccharina japoni- lated in mortalities occurring in natural ca and Sarconema filiforme) might attain populations of white perch and striped established status in the future. The cryp- bass in 1963 in Chesapeake Bay, USA. togenic/questionable category includes From 1990 it has caused economic losses Acanthophora nayadiformis, Anotrichium in different European countries. Cultured okamurae, Antithamnionella boergesenii, A. gilthead seabream (Sparus aurata) and elegans, the Atlantic strain of Asparagopsis seabass (Dicentrarchus labrax) are the most taxiformis, Ceramium bisporum, Ganone- affected species in the WMED (TO- ma farinosum, Osmundea oederi and RANZO et al., 1991). Polysiphonia atlantica. Only four alien foraminiferal species On the 21 invasive or potentially inva- occur in the WMED. Planogypsina acer- sive macrophytes introduced in the valis (BLANC-VERNET, 1969, as Planor- Mediterranean (Table 3), 16 are present in bulina acervalis) and Cymbaloporetta plana the western basin mainly in coastal lagoons. have been previously cited in the other sec- Codium fragile, invasive in the last century, tors of the Mediterranean. Very rare spec- has become less abundant in the open sea. imens of Sorites orbiculus have been The northern coasts are the most reported for the Gulf of Naples impacted region, with a maximum of alien (HOFKER, 1930) and from the Ligurian macrophytes in France (78 taxa out of 91). Sea (BANCHETTI et al., 2009, as S. orbic- On the other hand, the lowest number (23 ularis). Schackoinella imperatoria, probably taxa) was recorded along the northern cryptogenetic/casual, has been recorded Africa coast (Morocco and Algeria). Such
Medit. Mar. Sci., 11/2, 2010, 381-493 415 a difference is probably due to insufficient 1.3. Alien polychaetes in the Western investigations in the south and the huge Mediterranean flood of introductions along with shellfish In the WMED, a total of only 49 alien transfers in the French coastal lagoons polychaete species have been reported to harbouring shellfish aquaculture. date, which is about 39% of the alien poly- VERLAQUE et al. (2007) listed up to 58 chaete species reported from the Mediter- alien macrophytes in the Thau Lagoon ranean Sea. Among them, 26 species are (64% of alien macrophytes of the WMED established in WMED (see Table 2), 12 and 46% of the total of alien macrophytes species are casual, 1 species (Paraprionospio of the Mediterranean), which is the lead- coora) is cryptogenic and 10 species are ing site of shellfish aquaculture in the questionable. Some species seem to have Mediterranean Sea. invasive character, forming dense popula- The majority of alien macrophytes of tions in semi- or heavily polluted areas, the WMED comes from temperate and including harbours (Branchiomma luctuo- cold regions (NE Atlantic and NW Pacific) sum, Hydroides dianthus, H. elegans, Polydo- in relation with the major vector of intro- ra cornuta, and others), brackish waters duction (shellfish transfer). Only 10 puta- (Desdemona ornata, Ficopomatus enigmati- tive Lessepsian immigrants (i.e. Acan- cus, Polydora cornuta, etc.) or shallow-water thophora nayadiformis, Chondria pygmaea, benthic habitats (Eunice antennata, Lum- Ganonema farinosum, Halophila stipu- brineris perkinsi, Notomastus mossambicus, lacea, Hypnea spinella, H. valentiae, Lopho- Prionospio (Aquilaspio) krusadensis, etc.). cladia lallemandii, Padina boergesenii, Sar- The WMED has a relatively low number of conema filiforme and Ulva fasciata) have alien species, in comparison with the been hitherto recorded, mainly out of the EMED, especially the Levantine Sea, which coldest zones (Gulf of Lions and Gulf of is densely colonized by alien polychaetes. Genoa). However, the only WMED report The majority of the established alien of Sarconema filiforme (France) requires species in the WMED are of Pacific, Indo- confirmation. Acanthophora nayadiformis, Pacific, Indian and/or Red Sea origin, Ganonema farinosum, Hypnea spinella, whereas less originate in the tropical Padina boergesenii and Ulva fasciata could Atlantic (six species) or West Atlantic (six alternatively originate from the Atlantic species). Ocean. The strains of Ulva fasciata and A relatively low number of newcomers Hypnea valentiae introduced in the Thau originating in tropical seas and anthro- Lagoon most probably originate from the pogenically introduced via the Gibraltar north-western Pacific (Japan or Korea). Strait are true alien species. Most of them The invasive behaviour of Lophocladia appear to have arrived via shipping, espe- lallemandii in the Balearic Islands con- cially in harbours, as also happens in the trasts with a non-invasive behaviour every- EMED. Although the polychaetes of the where else in the Mediterranean, and a WMED are relatively well known, new different origin for the western strain reports are occurring continuously, mak- deserves to be considered. Thus, there are ing sometimes difficult to elucidate if they hitherto only two unquestionable Lessep- are true invaders, or unrecorded or over- sian macrophytes in the WMED: Chondria looked species. This is especially evident pygmaea and Halophila stipulacea. with small, meiobenthic species; for exam-
416 Medit. Mar. Sci., 11/2, 2010, 381-493 ple, in a recent work (DEL-PILAR-RUSO (ABELL & TORRES, 1998) and Sar- et al., in press) two small syllids (Syllis mau- dinia Channel (MURA et al., 2003), the retanica and Parapionosyllis macaronesien- latter in Alboran Sea and Balearic Island sis; not included in Table) are reported for (CARTES et al., 2000). A similar consider- the first time in the Mediterranean; both ation can be made for Merhippolyte species were described from NE Africa ancistrota, a species occurring in the Albo- and Canary and Madeira islands, relative- ran Sea but also known in the Gulf of ly close to the Strait of Gibraltar. There is Cadiz and Morocco (GARC A RASO, no way to know if these species were acci- 1996; UDEKEM D'ACOZ & URI , dentally introduced, were overlooked in 1996), and Plesionika ensis (A. Milne previous papers, or simply never found. Edwards, 1881), cited in the Alboran Sea (GARC A RASO, 1981) and Gulf of 1.4. Alien crustaceans in the Western Cadiz (GARC A RASO, 1996). However, Mediterranean new references from the island of Crete, In total, 52 alien species of crustaceans Aegean Sea (LABROPOULOU & have been found in the WMED, of which KOSTIKAS, 1999) and its absence in 20 have self-maintaining populations, 20 intermediate Mediterranean areas raise are casual records, eight are questionable doubts about the status of the latter and four are classified as cryptogenic. species in the EMED. There is an old cita- These species belong to amphipods (4), tion of the species Plagusia depressa cirripedes (5), decapods (20), stomatopods (Fabricius, 1775) and Plagusia chabrus (1), cumaceans (1), planktonic and para- (Linnaeus, 1758), now Guinusia chabrus sitic forms of copepods (16), and isopods (Linnaeus, 1758), brought to the port of (5) (Table 2). Marseilles by an iron vessel in 1873 In the decapods, we have not consid- (STEBBING, 1893), but they have not ered alien the Atlantic species found in the been included in Table 2, because they Alboran Sea and whose historical presence have never again been captured (there are is known in Morocco. For example, no later references). Brachynotus atlanticus, a tropical African Half the species originate in the Indo- species known in Morocco (FOREST & Pacific, Pacific, Indian Ocean or Red Sea, GANT S, 1960), was found in Europe for while seven species are known to have a the first time in the littoral of Malaga circumtropical distribution. The remain- (GARC A RASO, 1984a), and later in ders have a tropical/subtropical Atlantic the Bay of Cadiz, in the Atlantic, where it origin, many of them being known in the forms a stable population (GARC A western Atlantic. RASO, 1985). The Atlantic shrimps Two species, Synalpheus tumidomanus Penaeopsis serrata and Hymenopenaeus africanus and Necora puber, not included debilis, known in Moroccan waters (ZA- in earlier Mediterranean alien lists, are RIQUIEY, 1968; PÉREZ FARFANTE, added in Table 2. Synalpheus tumidomanus 1979), were recently found in the western africanus (syn. S. hululensis africanus) was Mediterranean Sea (perhaps due to their first captured in Israel (LEWINSOHN & deep-water habitat); apparently, they are HOLTHUIS, 1964, as Synalpheus hululen- being found in more areas of the Mediter- sis) and considered as alien; later it was ranean; the former in the Alboran Sea found in the Alboran Sea (GARC A
Medit. Mar. Sci., 11/2, 2010, 381-493 417 RASO, 1984b; L PEZ DE LA ROSA & of aliens in the groups of amphipods, cirri- GARC A RASO, 1992; GARC A pedes, cumaceans, isopods and RASO, 1996), a characterization that sub- tanaidaceans and b) a high number of sequently changed following reclassifica- questionable and cryptogenic species. tion of the Mediterranean specimens with- The amphipod Elasmopus pectenicrus in the subspecies S. tumidomanus africanus has a circumtropical distribution: Atlantic (or S. africanus?), known from Casablanca Ocean, Red Sea, Indian Ocean, Pacific and Rabat, Morocco (LAGARD RE, Ocean (described from New Guinea); it 1971). The Alboran populations may rep- has gradually colonized the EMED, the resent a natural expansion of the distribu- Adriatic and the CMED (ZAKHAMA- tion range of the species, which entered SRAIEB & CHARFI-CHEIKHROUHA, through the Strait of Gibraltar and colo- 2010). The latest finding could be clearly nized the southern coast of Spain (perhaps the result of increased scientific effort in with the help of the Atlantic surface water the area. inflow). The references from Greece and In the WMED the only species Turkey (KOUKOURAS & KATTOU- markedly invasive is Percnon gibbesi, which LAS 1974; KOCATA , 1981) could indi- shows a rapid expansion of its geographi- cate a separate advance in the EMED lit- cal distribution range, e.g. in the east Span- toral from the Israeli populations. Recent- ish coast (DEUDERO et al., 2005): first ly it has been captured in Sicily (BACCI et recorded from the Balearic Archipelago in al., 2010). The unresolved question is 1999 (GARCIA & REVIRIEGO, 2000), whether the EMED populations are alien it established populations since 2002 in or not. At this time we have classified it as Barcelona, 2003 in Alicante and Murcia, cryptogenic. 2006 in Almeria. Necora puber is included as ‘alien’ Among alien planktonic copepods species because (in agreement with reported in the WMED Sea, three are ZIBROWIUS, 1992) we believe that its clearly Lessepsian immigrants (Triconia presence in several areas of the Mediter- hawii, T. rufa and T. umerus) and were ranean is due to introductions, most likely found in the Gulf of Naples. Two other related to its widespread food use rather Lessepsian immigrants, Labidocera detrun- than an entry through ballast waters. Only cata and Calanopia elliptica, were reported the specimens collected west of Malaga from the Gulf of Naples (VIVES & (GARC A RASO et al., 2010) could have SHMELEVA, 2007), but were never found entered the Mediterranean through the in the more than 20 years time series of Strait of Gibraltar by their natural means samples collected bi-weekly to weekly in of dispersion. the area (Mazzocchi, pers. comm.): there- With regard to other groups of crus- fore, their presence is considered as ques- taceans we should note that the available tionable. Pseudocalanus elongatus and data are scarce and often difficult to assess. Paracalanus indicus seem to be cryptogenic This is partly due to the smaller number of in the area; according to RAZOULS et al. taxonomic specialists and studies devel- (2005-2010) the latter species could be con- oped on these groups in the different founded with Paracalanus parvus, a very Mediterranean sectors. These limitations common native Mediterranean species. lead to a) a downwards-biased estimation Two copepods parasitic on oyster beds
418 Medit. Mar. Sci., 11/2, 2010, 381-493 (Myicola ostreae and Mytilicola orientalis) actively cultured (for example Crassostrea are well established in the WMED, after gigas and Ruditapes philippinarum, to a their accidental introduction into the lesser extent Mercenaria mercenaria) and French coast with the Pacific oyster Cras- species that were probably introduced sostrea gigas (CLANZIG, 1989). accidentally along with aquaculture (Crepidula fornicata, Gibbula albida, Mus- 1.5. Alien molluscs in the Western Mediter- culista senhousia, Xenostrobus securis), ranean thereby comprising one-fourth of the The WMED has a low incidence of reported aliens. M. senhousia and X. alien Mollusca with 34 recorded species (22 securis are locally invasive in the basin, but established, nine casual, two cryptogenic C. fornicata, which is one of the most inva- and one questionable) in a total of around sive aliens in the NE Atlantic, has difficul- 1500 native species. Two of these (Cerithi- ty in surviving where it has been intro- um scabridum and Brachidontes pharaonis) duced in the Mediterranean. Gibbula albi- are found only very close to the Strait of da was originally an Adriatic endemic, but Messina and are merely outliers from the is now introduced in the Ebro Delta confirmed area where these species have (TRIGO, 1981) and the French Mediter- established in the Ionian Sea. Others, like ranean lagoons (CLANZIG, 1989), as well Chromodoris quadricolor, Pinctada radiata, as on the Atlantic coast of France. Rissoina spirata and Thais lacera, had casu- A unique and certainly recent feature al occurrences that are unlikely to persist, is the occurrence in the vicinity of Tunis although they may be firmly established in (Tunisia) of two alien species (Favorinus the EMED. Leaving aside the outliers and ghanensis, Mitrella psilla) introduced these casual species, the cryptogenic through shipping from tropical West species, and the questionable report of Sac- Africa (BEN SOUISSI et al., 2004; ANTIT costrea cucullata for Tunisia, we are left et al., 2010). More generally, harbours are with not more than 25 alien molluscs defi- a focus for established populations of nitely relevant to the basin. aliens, with random origins. Other exam- The north coast of Tunisia represents ples are the bivalve Theora lubrica in a major step towards the colonization of Leghorn, Italy, and the gastropod the WMED. Two of the hardiest Lessep- Bostrycapulus odites in Alicante, Spain, sian immigrants are present there, having which did not spread out to the neighbour- made their way across the WMED to the ing open sea. coast of Spain and are locally invasive. On the whole, the incidence of alien These are Bursatella leachii mollusc species in the WMED is heavily (WEITZMANN et al., 2009; RAMOS- concentrated in marginal marine environ- ESPL et al., 2010) and Fulvia fragilis ments such as the lagoons of the French (TAMAYO-GOYA, 2008). B. leachii also Mediterranean coast, the Spanish Mar exists in the Atlantic, but the hypothesis Menor and Ebro Delta, and the Italian that WMED populations entered through lagoons of Caprolace and Fusaro on the Gibraltar does not hold since the species Tyrrhenian coast. With the exception of does not exist in Morocco. the recent report of the nudibranch Godi- The largest pool of alien molluscs in va quadricolor from the large harbour of the WMED includes species that are Algeciras (CERVERA et al., 2010), the
Medit. Mar. Sci., 11/2, 2010, 381-493 419 Alboran Sea is free of alien Mollusca, a sit- Alien Ascidiacea are represented by uation which may be related to its excep- eight species (out of 16), most of them tionally high species richness estimated as established. Five species are circumtropi- about 1200 species. More generally, the cal. Trididemnum cf. savignyi was first open sea in the WMED is hardly, if at all, recorded by LAFARGUE (1972) at Port- impacted by alien species of molluscs. Cros, France. LAFARGUE (1974) revised Although a couple of alien molluscs in the Trididemnum tenerum complex, assign- the Mediterranean are of West African ori- ing the record by PÉR S (1954) from gin, there is no species yet that has gradu- southern Tunisia to T. cf. savignyi. Subse- ally entered through the Strait of Gibraltar quent records of this species include Cat- and become newly established in the alonia and Columbretes Islands, Spain Mediterranean. As already noted by (RAMOS-ESPL , 1988). Perophora mul- GOFAS & ZENETOS (2003), all the ticlathrata and Ecteinascia styeloides have prevalently Atlantic species found in the been reported from Corsica (MONNIOT, westernmost Mediterranean have a histor- 1983) and the northern Tyrrhenian Sea ical range since at least the 19th century and (MASTROTOTARO & TURSI, 2010), are, therefore, to be considered native respectively. Microcosmus squamiger, first species. The WMED is nevertheless a recorded in Bizerte (as M. exasperatus) is source area for such species as Siphonaria widespread in the WMED (synthesis in pectinata, native to Algeria and Spain but TUR N et al., 2007). Cystodytes philip- introduced in Greece and recently expand- pinensis, previously observed in southern ing its range towards northern Tunisia. Tunisia (MÉLIANE, 2002), has been Although not qualifying as ‘aliens’ and not recorded recently in the Balearic Islands included in the species counts, one must (D AZ-VALDÉS & RAMOS-ESPL , keep in mind the fact that such species as 2010). This cryptogenic species had proba- Eastonia rugosa (see ALBANO, 2006) are bly been confused in the past with its con- expanding their range. gener C. dellechiajei, widely distributed in the Mediterranean Sea (IZQUIERDO- 1.6. Miscellaneous invertebrates in the MUN~OZ et al., 2009). The Western Western Mediterranean Atlantic species Distaplia bermudensis and Bryozoa include five alien species in the Eastern Pacific species Polyandrocarpa the WMED, out of 23 known from the zorritensis have been introduced by shell- whole basin. Arachnoidea protecta is an fish culture (TUR N & PEREA, 1988; Indo-Pacific species first recorded in the BRUNETTI & MASTROTOTARO, WMED by CHIMENZ GUSSO et al. 2004; MASTROTOTARO & BRU- (1998), further records being provided by NETTI, 2006). Styela clava, originating D'HONDT & CHIMENZ GUSSO from the north-western Pacific, has been (2006). The morphological divergence spreading along the European Atlantic observed between the Indo-Pacific and coast since 1954 and has only recently been Mediterranean specimens led Harmelin recorded in the WMED; however, its (in ZENETOS et al., 2005) to believe that absence from harbours and marinas closed they represent two distinct species. How- to Gibraltar may suggest that this species ever, OCCHIPINTI-AMBROGI et al. has also been accidentally transported into (2010) maintain A. protecta as established. the Mediterranean Sea by shellfish trans-
420 Medit. Mar. Sci., 11/2, 2010, 381-493 fer (DAVIS & DAVIS, 2008). Ophiactis savignyi. The former has been Cnidaria are an exception to most taxa reported from Port Cros, France, in 2000 in that the majority of aliens occur in the (ICES, 2006) with no further details WMED rather than in the EMED: 26 (hence it is classified as questionable), species (out of 46 listed for the whole whereas two individuals of Protoreaster basin). Alien Anthozoa are represented by nodosus, probably released accidentally two species, out of four reported for the from a private aquarium, have been caught whole Mediterranean. Although first by trawling off Majorca, Balearic Islands, recorded in the Ligurian Sea Spain, in 1981 (ALVARADO et al., 1986). (ZIBROWIUS, 1974), Oculina patagonica Casual is considered the presence of Ophi- is abundant in the southern part of the actis savignyi in Banyuls (GUILLE, 1969). WMED, both in Spain (ZIBROWIUS & The only alien Porifera in the WMED RAMOS, 1983; BALLESTEROS, 1998; is Paraleucilla magna, which is proliferat- IZQUIERDO et al., 2007) and along the ing across the Mediterranean North African coast (SARTORETTO et (GUARDIOLA et al., 2010). Platy- al., 2008). Eighteen alien species (out of helminthes are represented by Allolepi- 37) belong to Hydrozoa, a group well rep- dapedon fistulariae, an endoparasite of Fis- resented in the fouling assemblages tularia commersonii reported from Sar- (MORRI & BOERO, 1986) that colonize dinia (PAIS et al., 2007). the large ports of the WMED. Cordy- lophora caspia has been reported only for 1.7. Alien fish in the Western Mediter- the brackish lake of Fondi, Tyrrhenian Sea ranean (MORRI, 1979). Two alien Scyphozoa MASSUTI et al. (2010) reported 38 were found in the WMED: a single speci- new fish in the WMED since the middle men of Phyllorhiza punctata was seen in 20th century. Here we report 45 species (20 October 2009 off NE Sardinia (BOERO et established, 25 casual) inventoried in the al., 2009), while Stomolophus meleagris is 20th century. Most of them are of tropical the only one recorded exclusively in the (subtropical) Atlantic origin. WMED: this may suggest it entered from The entrance of alien fish in the the Atlantic, although the species is also WMED has drawn the attention on the so- known from the Pacific (MORAND & called effect of ‘tropicalization'. This phe- DALLOT, 1985). nomenon has expanded geographically, Both species of alien Ctenophora reaching the EMED. Such is the case for reported for the Mediterranean occur in the Atlantic origin species Carcharhinus the WMED (BOERO et al., 2009; altimus, Acanthurus monroviae, Pisodo- FUENTES et al., 2009; MILLS, 2009), nophis semicinctus, Sphoeroides pachy- while Sipuncula and Pycnogonida have gaster, Pagellus bellottii and Seriola fasciata, one alien species each (PANCUCCI- which have recently entered the Mediter- PAPADOPOULOU et al., 1999; BARTO- ranean through the Strait of Gibraltar, and LINO & CHIMENZ GUSSO, 2010). have already reached the EMED. The only alien echinoderms recorded Documentation on some species is in the WMED are two seastars and one limited so that those here reported as casu- ophiuroid of Indo-Pacific origin: Acan- als could have already established popula- thaster planci, Protoreaster nodosus, and tions. This could be the case of a) benthic
Medit. Mar. Sci., 11/2, 2010, 381-493 421 species with limited swimming capacity where it is exploited in commercial fishery. such as Microchirus boscanion, which has Diplodus bellottii and Pagellus bellottii, been captured at different localities and reported at the Maghrebine and Iberian times (MASSUTI et al., 2010); b) confu- coasts of the Alboran Sea, are occasionally sion with others, such as the soleids captured by commercial fisheries Microchirus hexophthalmus and Synaptura (MASSUTI et al., 2010). lusitanica known for the Iberian Peninsula and the Gulf of Lions (MATALLANAS, 2. CENTRAL MEDITERRANEAN 1984). Some species, such as Anarhichas lupus, Aluterus monocerus, Scorpaena The bulk of the CMED is represented stephanica, Fistularia petimba and Zenopsis by the Ionian Sea, the least known of all conchifera, are known only from single the Mediterranean sub-basins observations and are still limited to the (ZENETOS et al., 1997). Local endemics, WMED. The latest findings include Lut- mostly within molluscs, have been report- janus jocu (VACCHI et al., 2010) and ed for its southern, non-European shores Kyphosus incisor (ORSI-RELINI et al., (SABELLI & TAVIANI, 1980) but also 2010), both in the Ligurian Sea. for Malta (EVANS et al., 2010). The Ion- Within the alien Indo-Pacific species, ian is connected to the WMED through Pomadasys stridens was reported for the the narrow Strait of Messina, a micro-sec- first time in the WMED and then it was tor that harbours a wealth of biogeograph- known in the EMED, where it is common; ic peculiarities, including Pliocene Atlantic Abudefduf vaigiensis, a casual species in the remnants and local endemisms (FREDJ & Levantine basin, has been reported three GIACCONE, 1995), and the larger Strait times in the WMED could be similar. By of Sicily, the meeting point of native contrast, Siganus luridus, and Fistularia Western and Eastern Mediterranean commersonii, which were first reported in species (BIANCHI, 2007), as well as of the EMED, where they are now well aliens of either Atlantic or Indo-Pacific known and even common, have expanded origin (COLL et al., 2010). Reflecting this westwards and are considered established. situation at the crossroads, the composi- Fistularia commersonii has expanded as far tion of the alien faunas is much more bal- north as the Ligurian Sea (OCCHIPINTI- anced between different sources than in AMBROGI & GALIL, 2009) and as far the EMED. Its situation at the transition west as Algeria (HEMIDA & CAPAPÉ, between the eastern and western basins 2009) and Spain (S NCHEZ-TOCINO make it a particularly sensitive place for et al., 2007). monitoring the progression of the much In the WMED, only a few species more numerous aliens already established appear regularly in the catches of some in the EMED. fisheries. That could be the case of Solea senegalensis, whose presence in the 2.1. Alien protozoans in the Central WMED was recorded in 1920 off the Iber- Mediterranean ian coast and has presently extended to Fish pasteurellosis from the protist Algeria, Tunisia and the Gulf of Lions, and Photobacterium damselae was reported Gymnammodytes semisquamatus reported from cultured seabass and seabream only in the north-eastern Iberian coast, installations in the Greek Ionian Sea and
422 Medit. Mar. Sci., 11/2, 2010, 381-493 from Malta (BAKOPOULOS et al., 1995, alien macrophytes respectively, followed 1997). by Tunisia (25) and Libya (12). Such dif- Only 4 alien foraminiferal species are ferences are probably due to a lower inves- reported for the CMED: Amphistegina lob- tigation effort, particularly in Libya. ifera, which probably has established a The alien macrophytes of the CMED population around the island of Malta come from temperate/cold regions (NE (YOKES et al., 2007), A. lessonii and A. Atlantic and NW Pacific) and from tropi- madagascariensis documented along the cal regions in almost equal proportions (51 coast of southern Tunisia (BLANC- and 49%, respectively). A total of 17 puta- VERNET, 1969; HOLLAUS & tive Lessepsian immigrants have been HOTTINGER, 1997). Planogypsina acer- hitherto recorded in the region. valis seems to have been recorded in this area only by BLANC-VERNET (1969). 2.3. Alien polychaetes in the Central Mediterranean 2.2. Alien macrophytes in the Central The CMED includes 36 alien species Mediterranean of polychaetes (17 established, nine casual, In the CMED, 57 alien macrophytes and ten questionable). Aliens already have been hitherto reported, which repre- established in the EMED have colonized sent 46% of the alien macrophytes known this area to subsequently spread to the for the Mediterranean Sea as a whole. The WMED: examples are provided by the major vectors of introduction are the ship three Indo-Pacific species, and likely traffic (fouling, ballast waters), the Suez Lessepsian immigrants, Eunice antennata, Canal, and, in the south Italian coastal Pseudonereis anomala (both established) lagoons, the shellfish aquaculture. Among and Protodorvillea egena (questionable). them, 35 alien macrophytes are well estab- By contrast, Linopherus canariensis and lished, four are casual, 16 are crypto- Ophryotrocha japonica, both established, genic/questionable, and two require con- could be indicative of a passage from west firmation. Casual taxa (i.e. Grateloupia to east. The former is very abundant in the lanceolata, Hypnea anastomosans, Polysi- Faro Lake, on the Messina Strait phonia paniculata and Undaria pinnatifida) (COSENTINO et al., 2009), together with might turn into the established status in another alien species: Syllis hyllebergi, the future. The occurrence of Batophora which is exclusively found in this area sp. and Gracilaria arcuata in the CMED (COSENTINO, in press). Other exclusive requires confirmation. species of the CMED are Eunice floridi- Of the 21 invasive or potentially inva- ana, Epidiopatra hupferiana hupferiana, sive macrophytes introduced in the Epidiopatra hupferiana monroi and Isolda Mediterranean (Table 3), 13 have been pulchella, mostly collected in the Gulf of reported in the CMED. Undaria pinnatifi- Noto (Sicily), all considered casual, non da that was only reported from the Mar established taxa (OCCHIPINTI- Piccolo of Taranto (south Italy), seems to AMBROGI et al., 2010). On the contrary, have disappeared from the region Ophryotrocha japonica is very abundant in (GRAVILI et al., 2010). the Mar Piccolo of Taranto, an area con- The Ionian coasts and Sicily are the sidered a hot spot of alien species diversity most impacted regions, with 35 and 36 within the Italian coasts (OCCHIPINTI-
Medit. Mar. Sci., 11/2, 2010, 381-493 423 AMBROGI et al., 2010). In this locality along the Libyan shores may well be con- one of the most conspicuous populations sequence of a lower research effort, but of Branchiomma luctuosum is present they are also absent from Malta and the (LICCIANO et al., 2002), and it seems southern Italian shores. Another Indo- here to compete with the native species Pacific species is Plagusia squamosa. Sabella spallanzanii (Giangrande, pers. The following species were recorded observ.). Most of the other established only once and with a single specimen: species in this area are the same as those Alpheus inopinatus, Dromia spinirostris, Sir- that are common in the whole basin such pus monodi, Callinectes sapidus, Grapsus as Ficopomatus enigmaticus, Hydroides granulosus. The presence of one adult dianthus, H. diramphus, H. elegans, specimen of Paralithoides camtschaticus, a Lysidice collaris, Metasychis gotoi, Neo- strictly boreal species, in this sector of the pseudocapitella brasiliensis, and Notomas- Mediterranean (FACCIA et al., 2009) is tus aberans. really puzzling. The number of alien copepods is low 2.4. Alien crustaceans in the Central (five) in the CMED and only two are Mediterranean Lessepsian immigrants (Euchaeta concin- A total of 46 alien crustacean are na, Triconia umerus). The presence of the reported in the CMED (26 established, 13 Antarctic to Subantarctic species casual, four questionable, three crypto- Spinocalanus terranovae in the Malta area genic). is questionable (SCIBERRAS & Three West Atlantic species are well SCHEMBRI, 2007). established at least in some regions of the The bentho-planktonic calanoid cope- CMED: Percnon gibbesi, Libinia dubia, pod Pseudocyclops xiphophorus was previ- and Rimapenaeus similis, which was discov- ously recorded only in coastal waters of ered on the trawling grounds of the Gulf of Mozambique. The Mediterranean P. Gabès (BEN HADJ HAMIDA-BEN xiphophorus specimens were collected ABDALLAH et al., 2010). Percnon gibbesi from fouling attached to submerged moor- was first observed in 1999 at Linosa Island ing posts and ropes in the brackish Lake (RELINI et al., 2000). It rapidly spread in Faro, eastern Sicily. It is classified as cryp- other localities of the CMED: Pantelleria togenic because according to ZAGAMI et (GALIL et al., 2002), Malta (BORG & al. ( 2005) it could represent a relict popu- ATTARD-MONTALTO, 2002), the lation of Tethyan origin. The genus Pseu- shores of southern and north-western Sici- docyclops has a worldwide distribution ly (MORI & VACCHI, 2002), the Strait of from temperate to tropical shallow coastal Messina (BELLANTONI & CORAZZA, and brackish waters. 2003) and the Gulf of Taranto (FACCIA & BIANCHI, 2007). 2.5. Alien molluscs in the Central Mediter- Several Lessepsian immigrants, such ranean as Trachysalambria palaestinensis, Eucrate The CMED has reports for 38 alien crenata, Metapenaeus monoceros and species of molluscs (19 established, 13 casu- Metapenaeus stebbingi, established in the al, two cryptogenic and four questionable) EMED have spread westward as far as in a total of 1000 to 1500 native species. Tunisian waters. Their apparent absence Among the species of Indo-Pacific origin,
424 Medit. Mar. Sci., 11/2, 2010, 381-493 there are 12 species that qualify as Lessep- species. The Ionian coast of Greece has sian (e.g. Pinctada radiata, the earliest only nine species reported, of which two reported Lessepsian mollusc) and six that (Bursatella leachii and Pinctada radiata) are definitely non-Lessepsian (e.g. Melibe are among the most widespread Lessep- viridis which was first detected on the Ionian sian immigrants and five are non-Lessep- coast of Greece), keeping the same propor- sian species of tropical Indo-Pacific origin. tion as in SE Turkey but far from the 90% The Ionian coast of Italy holds 14 species. of Lessepsians along the Levantine coast. These three areas share very much of their There are scattered reports of species aliens including the most widespread from the temperate Atlantic (Crepidula Lessepsian species (Bursatella leachii and fornicata, Polycerella emertoni, Anadara Pinctada radiata, Cerithium scabridum and transversa, Zygochlamys patagonica) and Fulvia fragilis in Italy and Malta) which North Pacific (Crassostrea gigas purposely here are secondary introductions from the introduced for farming, Musculista senhou- EMED. Also shared are some of the trop- sia, Ruditapes philippinarum) but most are ical Indo-Pacific species which started far less successful in this particular context, their spread in this area (Aplysia dacty- than in the northern part of the Mediter- lomela, Haminoea cyanomarginata, Melibe ranean (Ligurian Sea, Adriatic Sea) where viridis). Some species, like Anadara trans- some of them are invasive (see Table 3). versa or Musculista senhousia, which are Considering the proximity of the EMED, invasive in the Adriatic, have spread the total numbers are nevertheless towards the Italian Ionian coast. remarkably low. Few species are invasive, among them could be considered the 2.6. Miscellaneous invertebrates in the bivalves Pinctada radiata and Fulvia fragilis Central Mediterranean and the opisthobranch Bursatella leachii in Bryozoa are comprised of eight the Gulf of Gabès. species (out of 23 known for the whole Tunisia holds altogether 24 alien Mediterranean), of various origins (three species (13 established, three casual, three circumtropical, three Pacific, two cryptogenic, five questionable) but a dis- Atlantic). The records of Celleporaria aper- tinction must be made between the eastern ta and C. pilaefera at Malta have been con- and southern coasts bordering the Gulf of sidered questionable by SCIBERRAS & Gabès, which are a continuation of the vir- SCHEMBRI (2007). tually unknown Libyan coast, and the Nearly half of the alien species of north coast west of Cap Bon, which is con- Ascidiacea inventoried in the Mediter- sidered as part of the WMED. A cowry, ranean (seven out of 16) are known for the Erosaria turdus, had a spectacular onset in CMED. These aliens are of various origins recent years and is so invasive that it will (Western Atlantic, circumtropical, Indo- probably displace the native Zonaria Pacific, Eastern Pacific) and both the alien pyrum. The limpet Cellana rota may be the species of Microcosmus, the Indo-Pacific next spectacular invader in this part of the M. exasperatus and the circumtropical M. Mediterranean. squamiger, occur in the CMED Malta has possibly one of the best (IZQUIERDO-MUN~OZ et al., 2009). studied molluscan faunas in the southern Most records of alien ascidians come from part of the EU, and has reports for 17 Taranto harbour (BRUNETTI &
Medit. Mar. Sci., 11/2, 2010, 381-493 425 MASTROTOTARO, 2004; MASTRO- therein), especially from Pelagie Islands and TOTARO et al., 2004; MASTROTO- Malta but also from Tunisia (BEN TARO & BRUNETTI, 2006). SOUISSI et al., 2006a,b) and Libya The CMED hosts nine out of 46 alien (SHAKMAN & KINZELBACH, 2006; species of Cnidaria known for the whole BEN ABDALLAH et al., 2007). A few basin. Hydrozoa include six species, all with records are also available from the northern a wide distribution in warm waters. The Ionian Sea (TORCHIO, 1963; MASTRO- cryptogenic species Cordylophora caspia TOTARO et al., 2007) and the Messina was found abundant in the low salinity Strait, as in the case of Galeocerdo cuvier waters of the ‘Palude del Capitano’, Gulf of (CELONA, 2000), Pinguipes brasilianus Taranto (BIANCHI et al., 1994). Scypho- (ORSI-RELINI, 2002), Platycephalus indi- zoa are represented by the three species cus (CASTRIOTA et al., 2009) and Psenes Cassiopea andromeda (SCHEMBRI et al., pellucidus (NAVARRA et al., 2008). 2010a), Phyllorhiza punctata (ABED- Evidence of established populations in NAVANDI & KIKINGER, 2007) and the CMED are mostly related to the occur- Rhopilema nomadica (SIOKOU- rence of multiple records of the same FRANGOU et al., 2006). species, such as for Cephalopholis taeniops The only alien species of Ctenophora (GUIDETTI et al., 2010) and Pisodonophis is Mnemiopsis leidyi, recorded from Isola di semicinctus (RAGONESE & GIUSTO, Capo Rizzuto, Gulf of Taranto (BOERO 2000) while a few species such as Siganus et al., 2009), whereas the only alien species luridus, Fistularia commersonii, Seriola car- of Porifera is Paraleucilla magna, originat- penteri, Stephanolepis diaspros, Sphoeroides ing from the SW Atlantic (ZAMMIT et al., pachygaster have been collected or observed 2009). In the case of Sipuncula, there are in large numbers (RAGONESE et al., 1997; questionable records of the circumtropical PIZZICORI et al., 2000; AZZURRO & Aspidoshiphon mexicanus from Malta and ANDALORO, 2004; BRADAI et al., 2004; Lampedusa (PANCUCCI-PAPADO- AZZURRO et al., 2007). Latest records POULOU et al., 1999; SCIBERRAS & include Selene dorsalis (VELLA & SCHEMBRI, 2007). DEIDUN, 2009) and Opleognathus fascia- Alien Echinodermata are represented tus (SCHEMBRI et al., 2010b). by three species (out of 12 for the whole The geographical partitioning of Mediterranean). The record of Eucidaris CMED alien species was: 19 of Atlantic tribuloides from Malta was also the first of origin, 25 of Indo-Pacific or Pacific origin, this species for the Mediterranean and three circumtropical. If we look only (TANTI & SCHEMBRI, 2006). at the established species, the number of Indo-Pacific or Pacific fish (15) is almost 2.7. Alien fish in the Central Mediter- double than the number of Atlantic new- ranean comers (eight) indicating unbalanced ratio In the CMED, there are records of 50 towards fish coming from the east. Indeed, alien fish. Of them, 25 seem to be estab- the progressive penetration westwards of lished, 19 are casual, whilst the presence of Lessepsian immigrants is a continuous six species is questionable. The bulk of these phenomenon, which has accelerated dur- records comes from the Sicily Strait (see ing the last decades, as exemplified by GUIDETTI et al., 2010 and references Upeneus pori, Pempheris vanicolensis,
426 Medit. Mar. Sci., 11/2, 2010, 381-493 Sphyraena chrysotaenia and Siganus rivula- 2004). The Central Adriatic is character- tus (BRADAI et al., 2004; BEN SOUISSI ized by the lack of both Northern-Adriatic et al., 2006a,b). endemics and Atlantic-Adriatic disjunc- tions. Mediterranean endemics and ther- 3. THE ADRIATIC SEA mophilic species are still scarce, so that this sector is less diverse than the Gulf of Lions The Adriatic Sea is a rather unique and Ligurian Sea, to which it resembles. and differentiated area within the However, it is significantly richer in species Mediterranean, with a strong contrast than the Northern Adriatic. Finally, the between the predominantly linear sandy Southern Adriatic exhibits a transitional shores along the western (Italian) side, and character between the Adriatic and the the opposite complex coasts of the eastern Ionian Sea, with which it communicates side (Slovenia, Croatia, Montenegro and through the Otranto Strait. Affinities with Albania) forming a maze of islands and the WMED are strongly reduced. inlets with rocky shores. The hydrographic conditions are also peculiar, with very low 3.1. Alien protozoans in the Adriatic Sea winter temperatures in the northern part, Three protozoans pathogenic to shell- which is also quite shallow (40 m depth), fish, namely Marteilia refringens, Bonamia and very hot summers in the southern part, ostrea and Perkinsus olseni (formerly which is much deeper. All these features Perkinsus atlanticus) have been reported lead to differentiation between the north- from shellfish cultures in the Adriatic Sea. ern and southern Adriatic areas. Serious outbreaks of pasteurellosis caused From a biogeographic standpoint, the by Photobacterium damselae (formerly Adriatic Sea is divided in three sectors, Pasteurella piscicida) in the Adriatic are arranged more or less latitudinally. The reported by CESCHIA et al. (1991). Northern Adriatic Sea is perhaps the most Foraminiferal aliens from the Adriatic peculiar sector of the whole Mediter- are few (six species). Moreover, all of them ranean. Strong winter cooling, low salinity have been reported based on single speci- due to significant river input, and compar- mens, in only one publication or site and atively great tidal range make it more sim- could be considered as cryptogenic/casual. ilar to the Northern Atlantic than to the This is the case for warm water alien rest of the Mediterranean. This is reflected species such as Archais angulatus in the biota, which includes disjunct (LANGER & HOTTINGER, 2000), Atlantic-Adriatic species whose occur- Clavulina angularis (BANCHETTI et al., rence contributes to what has been called 2009), Coscinospira hemprichii, Cymbalo- the Northern-Adriatic ‘sub-Atlanticism’ poretta plana (as Trethomphalus bulloides) (SACCHI et al., 1985). Mediterranean and Sorites orbiculus (CIMERMAN & endemics are scarce and the overall diver- LANGER, 1991). A. angulatus is only sity is the lowest of the Mediterranean reported from the Adriatic (LANGER & basin, so that the phrase ‘North-Adriatic HOTTINGER, 2000). Schackoinella gap’ has also been coined (SACCHI, imperatoria, described from the coast of 1983). In addition, peculiar to this sector central Adriatic (CIMERMAN & are some taxa with Black Sea affinity LANGER, 1991), could be cautiously con- (SACCHI et al., 1985; BIANCHI et al., sidered a cryptogenic/casual form.
Medit. Mar. Sci., 11/2, 2010, 381-493 427 3.2. Alien macrophytes in the Adriatic Sea listed up to 33 aliens (67% of alien macro- A total of 49 alien macrophytes has phytes of the Adriatic) in the Lagoon of been hitherto reported from the Adriatic Venice. Such a difference is probably due Sea, which represent 39% of the alien to a flood of introductions along with shell- macrophytes reported from the Mediter- fish transfers (oysters, mussels, Manila ranean Sea. The major vectors of intro- clams) on the north-Italian coasts where duction are the shellfish aquaculture aquaculture is extensively developed. industry in northern coastal lagoons and The majority of alien macrophytes of the ship traffic (fouling, ballast waters). the Adriatic basin (35 taxa) come from Among them, 35 alien macrophytes are temperate and cold regions (NE Atlantic well established, three are casual, and 11 and NW Pacific) in relation with the major are cryptogenic/questionable. Casual taxa vector of introduction (shellfish transfer). (i.e. Acrothamnion preissii, Hypnea valenti- Only seven putative Lessepsian immi- ae and Polysiphonia paniculata) might turn grants (i.e. Acanthophora nayadiformis, into the established status in the future. Chondria pygmaea, Halophila stipulacea, The cryptogenic/questionable category Hypnea spinella, Hypnea valentiae, Lopho- includes 11 species (Table 2). cladia lallemandii and Ulva fasciata) have Of the 21 invasive or potentially inva- been hitherto registered, mainly from the sive macrophytes introduced into the northern coasts. However, Acanthophora Mediterranean (Table 3), 14 are present in nayadiformis and Hypnea spinella could the Adriatic Sea. Among them, cold tem- alternatively originate from the Atlantic perate taxa (i.e. Codium fragile, Gracilaria Ocean. The strains of Hypnea valentiae and vermiculophylla, Grateloupia turuturu, Ulva fasciata found in the Lagoon of Dasysiphonia sp., Sargassum muticum and Venice may originate from the north-west- Undaria pinnatifida) grow in north-western ern Pacific as in the Thau Lagoon. coastal lagoons, especially in the Lagoon of Venice and the lagoons of the Po Delta, 3.3. Alien polychaetes in the Adriatic Sea while the warm temperate and tropical The Adriatic hosts only 22 alien poly- taxa (i.e. Asparagopsis spp., Caulerpa race- chaete species (11 established, six casual and mosa var. cylindracea, Caulerpa taxifolia, five questionable), which represent 17% of Halophila stipulacea, Lophocladia lalle- the Mediterranean polychaete alien species. mandii, and Womersleyella setacea) invade At least four species appeared exclusive of the marine habitats of southern Italy and the Adriatic: Fabriciola ghardaqa, Megalom- the east coasts of the Adriatic (Croatia, ma claparedei, Platynereis australis and Syllis Albania), with the exception of Acrotham- alose, the latter two being considered as nion preissii. The invasive Gracilaria ver- questionable taxa. miculophylla is hitherto known only in the The established species are among lagoons of the Po Delta. those well known and settled in the whole Northern Adriatic coasts are the most Mediterranean area, and most of them are impacted, with 34 aliens out of a total of 49 also invasive, such as Branchiomma luctuo- (69% of alien macrophytes of the Adriat- sum, Ficopomatus enigmaticus, Hydroides ic), while only 15 aliens were reported from dianthus, H. elegans, but also Lysidice col- the Central and Southern Adriatic. laris, Metasychis gotoi, Neopseudocapitella SFRISO et al. (2009 and unpublished data) brasiliensis, and Notomastus aberans. The
428 Medit. Mar. Sci., 11/2, 2010, 381-493 most recently introduced species is B. luc- species are excluded from our list follow- tuosum, present with very large populations ing GALIL et al. (2002), whereas P. along the Mediterranean coasts. At present squamosa is kept as questionable because this species seems to have reached also the thriving populations were recently report- Brazilian Coast of Sa~o Paulo, in the Bay of ed from elsewhere in the Mediterranean: Santos, in the vicinity of the largest seaport Tunisia and Libya (ZAOUALI et al., in Latin America (DE MATOS 2007). NOGUEIRA et al., 2006), underlining its Only three crabs (Callinectes sapidus, ability to invade new areas, possibly trans- Dyspanopeus sayi and Rhithropanopeus ported by ships. An introduction via the harrisii) are definitely established in the Suez Canal and possible transfer via ship- area and all three originate from the ping, can be postulated not only for B. luc- Atlantic coast of USA. The spreading of tuosum, but also for the other two Red Sea the two panopeid crabs in the Adriatic Sea sabellid species, exclusive of this area and in recent years (FROGLIA & SPE- collected on the hard bottoms of the South- RANZA, 1993; MIZZAN & ZANELLA, ern Adriatic coast: M. claparedei and F. 1996; ONOFRI et al., 2008) has been facil- ghardaqa that were, however, found only itated by the development of mussel aqua- once with very few individuals and consid- culture both in lagoons and open sea, with ered therefore casual (GIANGRANDE & transfer of mussels seed and half grown MONTANARO, 1999; GIANGRANDE mussels among aquaculture plants. & LICCIANO, 2008). By contrast, the The latest record is that of the Indo- other sabellid Novafabricia infratorquata is Pacific crab Charybdis lucifera, caught six a Caribbean species that must have been miles off the Venetian coast (MIZZAN & introduced from the Strait of Gibraltar, VIANELLO, 2009). appearing established in the WMED and The following species were recorded casual in the Southern Adriatic along the only once and with single specimens: Mar- Apulian coast (LICCIANO & supenaeus japonicus, Scyllarus caparti, GIANGRANDE, 2006). Callinectes danae, Charybdis lucifera, Eri- ocheir sinensis, Hemigrapsus sanguineus, 3.4. Alien crustaceans in the Adriatic Sea and Charybdis japonica (FROGLIA, in A total of 24 alien crustacean are press). reported from the Adriatic (11 established, Only three planktonic copepod species 11 casual, two cryptogenic). were recorded as aliens in the Adriatic The first records of alien species intro- Sea, originating from the Atlantic and duced in the Northern Adriatic by mar- Indo-Pacific oceans. itime traffic date back to the years before World War I, when STIASNY (1908) 3.5. Alien molluscs in the Adriatic Sea recorded the capture of Plagusia squamosa The Adriatic Sea has possibly the old- (Herbst, 1790), Thenus orientalis (Lund, est alien (treated as cryptogenic) marine 1793) and Portunus sanguinolentus (Herb- species in the Mediterranean Sea, viz. Lit- st, 1796), respectively in the harbours of torina saxatilis, originally described from Trieste and Fiume (Rijeka), at that time Venice (OLIVI, 1792) before it could be the main harbours of the Austro-Hungari- suspected of being introduced from the an Empire (BABIC, 1913). The latter two NE Atlantic (JANSON, 1985). The Adri-
Medit. Mar. Sci., 11/2, 2010, 381-493 429 atic holds a few endemic species, one of 3.6. Miscellaneous invertebrates in the which (Gibbula albida) has been intro- Adriatic Sea duced into the WMED and elsewhere. Three alien species of Bryozoa (out of The Adriatic holds only 27 alien 23 known for the whole Mediterranean) species (15 established, nine casual and have been recorded in the Adriatic. The three cryptogenic) but the striking charac- Indo-Pacific species Tricellaria inopinata teristic is the high proportion of them was originally described on material com- which have become invasive. Together ing from the Lagoon of Venice by with the Levantine basin, the Adriatic may D'HONDT & OCCHIPINTI-AMBROGI be the part of the Mediterranean which (1985). It has been subsequently found in has been most transformed by the onset of many European seas (DYRYNDA et al., alien species. The most invasive species 2000). The alien status of Bugula fulva is include Anadara kagoshimensis (formerly supposed based on its patchy occurrence known as A. inaequivalvis), Musculista sen- nearly restricted to harbours housia, Rapana venosa and Ruditapes (HAYWARD & MCKINNEY, 2002). philippinarum, all originating from the The only alien species of Ascidiacea is temperate North Pacific and therefore Botrylloides violaceus, introduced in the comfortable despite seasonal lows in sea Lagoon of Venice by the north-western water temperature. Anadara kagoshimen- Pacific by shellfish culture (ZANIOLO et sis, now coupled to the possibly Atlantic al., 1998). Anadara transversa, has formed spectacu- Alien Anthozoa are represented only lar accumulations on the NE Italian coast by the north-eastern Atlantic species Diad- (RINALDI, 1985) and has profoundly umene cincta (BIRKEMEYER, 1996), impacted the sandy infralittoral bottoms which in a sense reinforces the Northern- which are a habitat for commercial species Adriatic ‘sub-Atlanticism’. such as the venerid Chamelea gallina. Porifera are represented by Paraleucil- Farmed species include Crassostrea gigas la magna only (LONGO et al., 2007). and Ruditapes philippinarum, and the activ- Fourteen alien species of Cnidaria ity related to farming may be responsible (out of 46 for the whole Mediterranean) for an important part of the introductions are known for the Adriatic. Alien Hydro- to and from the Adriatic; the latter species zoa are comprised of 13 species (out of is invasive and likely to displace or to drive 37). Most of them have circum(sub)tropi- locally extinct the native clam Ruditapes cal origin, but two estuarine species decussatus. The Sidney rock oyster Sac- (Cordylophora caspia and Gonionemus costrea commercialis was introduced to the vertens) prefer temperate waters. The first Venice Lagoon in the 1980s but seems to Mediterranean record of Garveia francis- have dwindled and has not been reported cana comes from the Venice Lagoon recently (MIZZAN, 1999). (MORRI, 1982). It is debatable whether Twenty-one of the alien species known Coryne eximia can be considered as alien from the Adriatic are recorded in Italy, species since the Mediterranean occur- and only the few tropical species Brachi- rence of this species seems to need re-con- dontes pharaonis, Cellana rota, Melibe firmation (BOUILLON et al., 2004). viridis and Halgerda wileyi have records Two of the three alien species of exclusively along the eastern coast. Ctenophora known for the Mediterranean
430 Medit. Mar. Sci., 11/2, 2010, 381-493 are established in the Adriatic: Beroe ovata Sea, together with the smaller Sea of Mar- and Mnemiopsis leidyi (SHIGANOVA & mara, which connects it to the Black Sea. MALEJ, 2009). These authors refer earli- The Levant Sea is warmer than the rest of er citations of Beroe ovata in the Mediter- the Mediterranean and harbours a signifi- ranean to the native species B. cucumis, cant number of circumtropical species. and report the real B. ovata as a Western Atlantic-Mediterranean elements and Atlantic species first introduced to the Mediterranean endemics are comparative- Black Sea and possibly from there to the ly scarce (MORRI et al., 2009). Adriatic. Since the construction of the Suez Sipuncula number two alien species: Canal, the Levant Sea is experiencing an both are circum(sub)tropical (PANCUCCI- important influx of Red Sea species. POR PAPADOPOULOU et al., 1999). The (1990) defined the geographical limits to only alien pycnogonid is Ammothea hilgen- the expansion of Red Sea immigrants in dorfi, first reported for the Mediterranean the Mediterranean as the ‘Anti-Psara line’ from the Lagoon of Venice (KRAPP & to the north (Anti-Psara being an island in SCONFIETTI, 1983). the Aegean) and the Strait of Sicily to the west: these boundaries match the 15 ÆC 3.7. Alien fish in the Adriatic Sea surface isotherm for February (BIANCHI, Adding Fistularia commersonii 2007). The Aegean Sea has local (DUL IC’ et al., 2008), 11 Lessepsian fish endemics, but the typical Mediterranean species had reached the Adriatic Sea by biota is impoverished with respect to the 2009 (DRAGI EVIC’ & DUL IC’, WMED (BIANCHI & MORRI, 1983); 2010). The occurrence of Terapon theraps this, however, may be partly due to insuffi- in 2007 (LIPEJ et al., 2008) is of particular cient inventory effort (MORRI et al., 1999; interest since its record was the first for the KOUKOURAS et al., 2001). The Sea of Mediterranean; furthermore, it was found Marmara exhibits peculiar hydrological at great distance from its usual distribution conditions, with low salinity waters coming area. Similar are the cases of Pampus from the Black Sea stratifying over saline argenteus and Epinephelus coioides. waters of Mediterranean origin on the bot- The catch of Elates ransonnetii in tom (ÜNLÜATA et al., 1990). This hydro- March 2010 in the eastern Adriatic logical regime should facilitate the diffu- (DUL IC’ et al., 2010) brought the num- sion of Black Sea species into the Northern ber of Lessepsian fish immigrants that Aegean rather than vice-versa, but our were recorded in the Adriatic Sea to 12. knowledge on the exchanges between the Sphoeroides pachygaster, of tropical two areas is limited and their biotic affini- Atlantic origin, has undoubtedly formed a ty is low (KOUKOURAS et al., 2001). In well established population in the Adriat- recent times, climatic change favoured an ic (Albania, Slovenia, Montenegro, Croat- increase of biotic penetration from the Sea ia, Italy). of Marmara into the Black Sea, which therefore has been undergoing a process 4. EASTERN MEDITERRANEAN of ‘mediterranization’ (TOKAREV & SHULMAN, 2007). The EMED includes two major bodies Some scientists (e.g., TORTONESE, of water: the Levant Sea and the Aegean 1973; OLIVERIO & TAVIANI, 2003;
Medit. Mar. Sci., 11/2, 2010, 381-493 431 POR, 2009) have argued that this basin, Pegidia lacunata and Pseudolachlanella particularly in its eastern stretches, suf- slitella (as Quinqueloculina eburnea) from fered from an ‘ecological vacuum’ that is north-eastern Africa (BLANC-VERNET now being filled by Red Sea immigrants. et al., 1979) and Cyprus (ABU TAIR & This idea has been criticized by GALIL LANGER, 2010). (2007). The new recovery of alien foraminifers mainly comes from recent researches car- 4.1. Alien protozoans in the Eastern ried out on living coastal assemblages of Mediterranean the EMED, which testify to the increasing Fish pasteurellosis caused by Photo- scientific interest in alien and Lessepsian bacterium damselae was reported from cul- immigrants. These studies confirm the tured seabass (Dicentrarchus labrax) in wide distribution of many large endosym- Turkey (CANDAN et al., 1996). biont-bearing taxa, such as Amphiste- Among the total of 50 alien ginids, along the coasts of the Levantine foraminiferal species presently known for basin (LANGER, 2008), Greece the Mediterranean, all except one are (TRIANTAPHYLLOU et al., 2009; present in the EMED (LANGER & KOUKOUSIOURA et al., 2010a), Turkey HOTTINGER, 2000). All of them have (MERI et al., 2010) and Cyprus (ABU tropical affinity. Despite a significant num- TAIR & LANGER, 2010). Amphistegina ber of species with circumtropical distribu- lobifera appears to have established suc- tion (KOUKOUSIOURA et al., 2010b), cessful populations in Greek coastal most have Indo-Pacific origin and their ecosystems (TRIANTAPHYLLOU et al., penetration in the EMED is likely to have 2009; KOUKOUSIOURA et al., 2010a,b). been favoured by the opening of the Suez Taxa of Indo-Pacific origin, probably true Canal (MERI et al., 2007a; LANGER, Lessepsian forms, are represented by 2008). Off the west Turkish coast, alien Borelis sp., Coscinospira hemprichii, Cyclo- foraminifers showed high abundance at forina sp., Heterostegina depressa, Pegidia hot submarine springs that allowed higher lacunata, Pseudolachlanella slitella and bottom water temperatures in winter Sorites orbiculus. (MERI et al., 2010). Similar observa- Taxa undetermined at the specific tions on alien macrophytes in the Aegean level (Borelis sp., Cycloforina sp.) have (DE BIASI & ALIANI, 2003) and in the been included as alien forms only when Tyrrhenian Sea (GAMBI et al., 2008) sug- described and figured in previous work on gest that shallow water hydrothermal vents Red Sea or Indo-Pacific areas in the Mediterranean Sea may act as step- (HOTTINGER et al., 1993; LANGER & ping stones for the penetration of tropical HOTTINGER, 2000; LANGER, 2008 aliens (BIANCHI et al., in press). with references) or (Haddonia sp.) if per- Among the nine new taxa added to tain to endemic Indo-Pacific genera those recorded by ZENETOS et al. (2008), (LOEBLICH & TAPPAN, 1988). eight occur in the EMED (LANGER, At the present state of studies, it seems 2008) and 4 of them have been reported prudent to exclude some very seldom only from single or few localities: Brizalina recorded species cited by ZENETOS et al. simpsoni and Euthymonacha polita from (2008), yet very rare or poorly preserved, the coasts of Turkey (MERI et al., 2010), but also known at various latitudes or as
432 Medit. Mar. Sci., 11/2, 2010, 381-493 fossils in the Mediterranean area. These which represent 50% of the alien macro- species are here represented by Acervulina phytes reported from the Mediterranean inhaerens, Iridia diaphana, Cymbaloporetta Sea. This high percentage is mainly due to squammosa, Pyramidulina perversa and the proximity to the Suez Canal and dense Triloculina affinis. international ship traffic in the area. Schackoinella imperatoria, a small Among them, 35 alien macrophytes are species originally described from fossil well established, nine are casual, 15 are material of the Tertiary Basin of Vienna, cryptogenic/questionable, and four require pertains to a genus now known only in the confirmation. Casual taxa (i.e. Indo-Pacific Ocean (LOEBLICH & Acrochaetium spp., Derbesia boergesenii, TAPPAN, 1988). Rare specimens have Padina antillarum, Rhodophysema georgii, been recorded along the eastern coasts of Rhodymenia erythraea, Solieria dura and North Africa (BLANC-VERNET, 1969) Spatoglossum variabile) might turn into the as Glabratella imperatoria. More data are established status in the future. The cryp- needed to verify its introduction by ship- togenic/questionable category includes ping through the Suez Canal. Its rare both temperate taxa mainly registered in occurrence suggests a cryptogenic/casual the north of the basin, i.e. the Sea of Mar- acclimatization status. mara and the Northern Aegean Sea In addition to all these benthic species, (Anotrichium okamurae, Chondria planktonic foraminifers are represented coerulescens, Cladosiphon zosterae, Ecto- only by Pulleniatina obliquiloculata, a ther- carpus siliculosus var. hiemalis, Halothrix mocline dwelling species characteristic of lumbricalis, Polysiphonia atlantica, P. equatorial Atlantic and Indo-Pacific fucoides and Pylaiella littoralis), and tropi- Oceans (BE & TOLDERLUND, 1971; cal taxa distributed in the Aegean Sea and KENNETT & SRINIVASAN, 1983). Its the Levantine Basin (Acanthophora nayad- occurrence has been reported only by iformis, Antithamnionella elegans, the MERI et al. (2004) along the Turkish Atlantic strain of Asparagopsis taxiformis, coasts and its introduction was inferred Caulerpa racemosa var. turbinata, Cerami- from the Atlantic Ocean via Gibraltar um bisporum, Ganonema farinosum and (ZENETOS et al., 2008). Recently, P. Ulva fasciata). The presence of Antitham- obliquiloculata has been reported from nionella spirographidis, Asparagopsis arma- Canary Islands, Eastern Atlantic (WILKE ta, Gracilaria arcuata and Hypnea flagelli- et al., 2009). It is absent in the Red Sea, but formis in the EMED requires confirma- it has been frequently reported from the tion. The latest record is that of the sea- Arabian Sea and Bay of Bengal (BE & weed Codium parvulum, recent blooms of TOLDERLUND, 1971; CHEN & which were observed on the northern FARRELL, 1991, inter alii). More data are shores of Israel (ISRAEL et al., 2010). required to interpret its occurrence and to On the 21 invasive or potentially inva- exclude its transport within ballast waters. sive macrophytes introduced into the Mediterranean (Table 3), 11 are reported 4.2. Alien macrophytes in the Eastern in marine benthic habitats of the EMED. Mediterranean No data are available for coastal lagoons. A total of 63 alien macrophytes have Cladophora cf. patentiramea and Codium been hitherto reported in the EMED parvulum are hitherto known only for the
Medit. Mar. Sci., 11/2, 2010, 381-493 433 EMED, for Cyprus and Israel, respectively. prionospio coora) are cryptogenic and 28 The majority of alien macrophytes of species are questionable. 22 species seem the EMED (63%) come from tropical to have invasive character (see Table 3), regions, mainly the Red Sea and the Indo- forming dense populations in semi- or Pacific Ocean. The major vector of intro- heavily polluted areas (including har- duction is the Suez Canal. Hitherto 35 bours) or shallow water benthic habitats. putative Lessepsian immigrants (56% of There is a relatively high number of ques- the total) have been registered, of which tionable species in the area. They include Cladophora cf. patentiramea, Codium the species only listed in ecological papers parvulum, Halophila stipulacea, Lophocla- (i.e. Cossura coasta, Loimia medusa), those dia lallemandii and Stypopodium schimperi identified as ‘cf.’ in the area (i.e. Oenone cf. are considered invasive. Acanthophora fulgida, Syllis cf. mayeri) or whose descrip- nayadiformis, Ganonema farinosum and tions based on the Mediterranean speci- Ulva fasciata were found in the Mediter- mens were insufficient (i.e. Lepidonotus ranean before the opening of the Suez carinulatus), hindering us in deciding their Canal in 1869, but the occurrence, at least real taxonomic positions. Future studies to in the EMED, of exotic strains coming be performed in the region would enable from the Red Sea and the Indian Ocean us to understand if these species really via the Canal is highly probable today. The exist in the Mediterranean. A total of ten other alien macrophytes reported from the casual species (i.e. Laonome elegans, EMED come from cold temperate regions Nereis gilchristi) might turn into the estab- (NE Atlantic and NW Pacific) and were lished status if data regarding them are recorded along the north coasts of Greece accumulated. The native species, Chaeto- and Turkey (Sea of Marmara and North- zone setosa (in part) and Paraprionospio ern Aegean Sea). Some of these last pinnata, previously widely reported from macrophytes, i.e. Ectocarpus siliculosus the Mediterranean Sea in polluted and var. hiemalis, Polysiphonia fucoides and deep waters, have been recently re-identi- Pylaiella littoralis, also occur in the Black fied as C. corona and P. coora, respectively Sea (TA KIN et al., 2008): so, a native sta- ( INAR & ERGEN, 2007; YOKOYA- tus cannot be excluded in the northern MA et al., 2010). These species were part of the EMED. regarded as cryptogenic because the data on their distributions are very limited; they 4.3. Alien polychaetes in the Eastern are relatively newly described species and Mediterranean could have been in existence in the area for A total of 98 alien polychaete species many years. have been reported to date from the The Levantine Sea is densely colo- EMED, almost 75% of the alien poly- nized by alien polychaete species. In the chaete species reported from the Mediter- region, a total of 77 species (79% of the ranean Sea, mainly due to the proximity to alien polychaetes from the EMED) have the Suez Canal and dense international been hitherto reported. A total of 11 ship traffics in the area. Among them, 52 species are categorized as questionable. species have been well established in habi- The highest number of alien species (58 tats of the EMED, 16 species are casual, species) are known from the Levantine two species (Chaetozone corona and Para- coast of Turkey, followed by the Israeli (29
434 Medit. Mar. Sci., 11/2, 2010, 381-493 species) and Egyptian (28 species) coasts. species of alien status were reported from On the Syrian coast, only two alien species the region. However, three species were (Spirobranchus tetraceros and Spirorbis excluded from the list, eight species are marioni) were encountered. The majority questionable and six species (Nereis persi- of species are Lessepsian immigrants (47 ca, Paraprionospio coora, Polydora cornuta, species, 61% of total species), the others Streblospio gynobranchiata, Desdemona were introduced to the area via ships, ornata and Ficopomatus enigmatus) have except for Chaetozone corona which is a been established. The report of the cryptogenic species. A total of 66 species Lessepsian species N. persica by (86%) could have been originated in the RULLIER (1963) needs to be confirmed. Red Sea and Indo-Pacific areas. The oth- The other species could have been intro- ers (11 species) were introduced to the duced to the area by ships. area from the Atlantic Ocean (mainly from the western part). 4.4. Alien crustaceans in the Eastern The Aegean Sea is on the route of the Mediterranean Lessepsian species that have become well A total of 119 alien crustaceans have acclimatized to the Mediterranean envi- been reported in the EMED among which ronment and have a large dispersal capac- six are cryptogenic species (Cymadusa ity. Up to date, 39 alien polychaetes were filosa, Gammaropsis togoensis, Mono- reported from the Aegean Sea, of which 16 corophium sextonae, Pseudocalanus elonga- species are Lessepsian immigrants. In the tus, Synalpheus tumidomanus africanus and area, 20 species have been well estab- Thalamita poissonii). Introduced crus- lished, one species, Prionospio (Minuspio) taceans belong to Amphipoda (10), Cirri- pulchra, is casual, 16 species are question- pedia (5), Copepoda (33), Cumacea (3), able and two species, Paraprionospio coora Decapoda (58), Isopoda (7), Tanaidacea and Chaetozone corona, are cryptogenic. (1) and Stomatopoda (2). These numbers Ship-transported species dominate semi- include both benthic and pelagic forms of or highly polluted areas, especially near copepods as well as parasites among cirri- the large international harbours. The pedes, copepods and isopods. The vast opportunistic species Malacoceros fuligi- majority of them occur in the Levantine nosus and Capitella spp. seem to have been Sea (e.g. 68% of the benthic forms, all replaced by the North Atlantic species planktonic taxa and all parasites). Two Polydora cornuta and Streblospio gyno- thirds of the species have colonized the branchiata and the Pacific species Pseu- Levantine and are spreading to the dopolydora paucibranchiata ( INAR et Aegean, while almost one third is known al., 2005, 2006a). In the eastern part of the from single records. Very few species, such Aegean Sea, 19 alien polychaete species as the decapods Calappa pelii, Metape- were found, of which two species are ques- naeus affinis and Sirpus monodi, are limit- tionable. In the western part, 31 species ed to the Aegean Sea. Reasoning for the were reported, of which 14 species are relatively high number of questionable questionable. copepod species (ca. 10%) is provided in Polychaetes from the Sea of Marmara ZENETOS et al. (2005, 2008). have been recently reviewed by INAR With the exception of a few species (2010). He postulated that a total of 17 originating in the Atlantic (Calappa pelii,
Medit. Mar. Sci., 11/2, 2010, 381-493 435 Callinectes sapidus, Farfantepenaeus information is available almost exclusively aztecus, Necora puber, Percnon gibbesi, Pro- for copepods, since it is the most studied cessa macrodactyla, Sirpus monodi, group (as in the entire Mediterranean Synalpheus tumidomanus africanus and Sea). Most of the newcomers are Lessep- Thalamita indistincta), the native range of sian immigrants, two originating in the four-fifths of the alien crustacean in the Atlantic (Calanopia biloba, Paracartia Mediterranean is in the Indo-Pacific grani), while the origin of Acartia tonsa, Ocean, Indian Ocean and Red Sea. The Arietellus pavoninus, Centropages furcatus pathway of introduction of decapod and Parvocalanus crassirostris is question- species is the Suez Canal, but penetration able, since they are distributed in tropical has been either unintentional (Lessepsian to subtropical waters both of the Atlantic migration) or ship-mediated. Shipping and Indian oceans and even of the Pacific. appears to be also responsible for the The species Pontellina plumata, Corycaeus immigration of species of Atlantic origin. speciosus and Subeucalanus crassus were However, spreading of self maintaining excluded from the list of alien species populations from the CMED appears to given by ZENETOS et al. (2005) after be the mode of introduction of the west reconsideration of the available informa- Atlantic crab, Percnon gibbesi now present tion on their distribution (RAZOULS et in Greece, Turkey, Syria, Cyprus, Israel al., 2005-2010). The consideration of (KATSANEVAKIS et al., in press and ref- Subeucalanus subcrassus as alien is ques- erences therein) and Egypt (AZZURRO tionable because it is present in most areas et al., in press). Thalamita poissonii could of the Mediterranean Sea as well as in the be a Lessepsian species, but there are temperate to tropical waters of the records of it before the Suez Canal open- Atlantic and Pacific oceans. The presence ing (UDEKEM D’ACOZ, 1999), which of Spinocalanus terranovae in the EMED could support a pre-Messinian Mediter- seems questionable, given its restricted ranean presence. If so, the alien status distribution in Antarctic and subantarctic should be downgraded to ‘questionable’. areas. Regarding the records of Triconia Perhaps at this moment it is best to con- rufa (originating from the Red Sea) in the sider it as ‘crypogenic’. EMED (Lebanon) by MALT et al. (1989), The latest additions to the inventory of BÖTTGER-SCHNACK & SCHNACK the marine decapod species in the EMED (2009) assumed that the previous authors are the pilumnid crab Eurycarcinus inte- may have confounded the specimens with grifrons (ÖZCAN et al., 2010), the red Triconia sp. 8. Finally, Pseudocalanus elon- swimming crab Gonioinfradens pauciden- gatus should be considered as cryptogenic tata (CORSINI-FOKA et al., 2010), both in the Mediterranean Sea: it has been of Indo-Pacific origin, and the western interpreted as a boreal relict species Atlantic shrimp Farfantepenaeus aztecus (FURNESTIN, 1979) or an immigrant, (CENZIG et al., 2010). either from the Black Sea (SIOKOU- The Indo-Pacific stomatopod Eru- FRANGOU, 1985) or the north-eastern gosquilla massavensis thrives along the Atlantic (VIVES et al., 1981). Levantine coast and is expanding in the Two introduced cymothoid species of Aegean (ÖZCAN et al., 2009). Indo-Pacific origin, Anilocra pilchardi and Among the planktonic crustaceans, Cymothoa indica, typically parasitic of
436 Medit. Mar. Sci., 11/2, 2010, 381-493 teleost fish, have been reported from species are of Indo-Pacific origin and their Lebanon (BARICHE & TRILLES, 2006; presence is related to the proximity of the TRILLES & BARICHE, 2006) parasitiz- Red Sea and to the Suez Canal connec- ing mainly barracudas (Sphyraenidae). tion. Only six species (Siphonaria pectinata, Three parasitic copepods new to the Mya arenaria, Cerithium litteratum, Mediterranean fauna were reported in Anadara transversa, Mytilopsis sallei, and waters off the Egyptian coast near Alexan- Petricola pholadiformis) are Atlantic or dria by EL-RASHIDY & BOXSHALL Atlanto-Mediterranean in origin. Exam- (2009). Two of them, Mitrapus oblongus ples of Indo-Pacific species yet non- and Clavellisa ilishae, are of Indo-Pacific Lessepsians, among the now well-estab- origin and are considered to have co- lished species, are the strombid Cono- invaded the Mediterranean through the murex persicus, first recorded in Turkey Suez Canal on Red Sea immigrant hosts. and only later arrived on the Levantine The third parasite, Nothobomolochus coast, or the opisthobranch Melibe viridis, fradei, was previously known from the Gulf first recorded in Greek waters. Some of of Guinea and the Arabian Gulf. these species, like Conomurex persicus, are A rhizocephalan barnacle, Heterosac- not even known from the Red Sea and cus dollfusi, followed its portunid host their introduction must therefore have crab, Charybdis longicollis, from the Red involved maritime traffic at some stage. Sea through the Suez Canal to the POR (2009, 2010) expressed the view Mediterranean Sea. that the thermophilic species of Indo- Pacific origin should not be viewed as 4.5. Alien molluscs in the Eastern Mediter- aliens but rather as a restoration of the ranean Tethyan faunal realm as it existed previ- The EMED has the lion’s share of ously to the closure of the communication alien molluscs, building up impressive with the Indian Ocean in the late Miocene. numbers and by far the highest proportion Another quite distinct pool is made up worldwide. To date there are 190 alien by species which originate from the tem- molluscs reported (105 established, 64 perate North Pacific, rather than from the casual, 15 questionable and six crypto- tropical areas, and whose introduction is genic), to be added to around one thou- linked with transfers of living molluscs for sand native species. This represents more aquaculture and/or maritime traffic. than 90% of all alien mollusc species The EMED has comparatively limited reported in the Mediterranean as a whole, aquaculture activity and this category a percentage that remains remarkably sta- would hold, apart from the cultured ble if we consider established species only. species proper Crassostrea gigas and Rudi- Six species are classified as cryptogenic tapes philippinarum, very few species such (Alvania dorbignyi, Aplysia parvula, Atys as Rapana venosa, Anadara kagoshimensis angustatus, Discodoris lilacina, Parviturbo or Musculista senhousia. dibellai, Teredo navalis), as there are A particular aspect regarding molluscs doubts on whether they are native or intro- is the high incidence of unsupported duced or perhaps represent very old intro- records, often based on single shells and ductions. carried perpetuated in subsequent check- Predictably, the vast majority of these lists. The rationale for their exclusion has
Medit. Mar. Sci., 11/2, 2010, 381-493 437 been detailed in GOFAS & ZENETOS Suez Canal and along the Levantine coast, (2003). We have not considered in this all established except the casual Psam- report the records for 32 additional motreta praerupta versus 20 species of the species, including those found on an oil rig tropical Indo-Pacific which are definitely (MIENIS, 2004), among them oysters and not Lessepsian and started their Mediter- other bivalves firmly attached to their sub- ranean settlement in the area. strate, considering that these have not yet The Aegean Sea is the natural route of been found in the wild. progression for the most successful Indo- The Levantine Sea is the area which Pacific invaders, both Lessepsian and holds the highest number of reports, with those introduced first in the southern 149 species (83 established, 45 casual, 5 Turkey/Cyprus area. Nevertheless it has cryptogenic, 16 questionable); historically also its own pools of species introduced this is the area where pioneer reports were through other pathways. Contrary to the made on alien species of Indo-Pacific ori- south coast of Turkey, there are several gin (e.g. GRUVEL & MOAZZO, 1931; species that originate from the Atlantic HAAS, 1937). (Anadara transversa, Crepidula fornicata, There are 48 of the species recorded Cerithium litteratum, Petricola pholadi- from Israel, Lebanon and Syria, which formis, Mya arenaria) or the temperate have a record in the Suez Canal and may North Pacific (Ruditapes philippinarum, be safely qualified as Lessepsian, against Rapana venosa) and are involved neither only seven (Melibe viridis, Murchisonella in the Lessepsian migration nor in the SE columna, Sticteulima cf. lentiginosa, Turkey focus of introductions. Siphonaria Smaragdia souverbiana, Conomurex persi- pectinata, introduced in the Gulf of cus, Zafra savignyi, Zafra selasphora) Saronikos, is a clear example of intra- species of tropical Indo-Pacific origin Mediterranean introduction. which are proven non-Lessepsians (whose As may be derived from this, the progression started elsewhere than on the importance of Lessepsian species is mod- Levantine coast and have no Suez Canal erate, and even the proven Lessepsian record). The remainder cannot be posi- species found there may be secondary tively assessed but the ratio of Lessepsian introductions from populations settled to non-Lessepsian among them should elsewhere in the EMED. probably remain high. Among the 18 species which could The coasts of southern Turkey and hold as Lessepsian because they were Cyprus are the next largest centres where recorded in the Suez Canal, species like alien species have been reported, but there Cellana rota, Fulvia fragilis, Murex the exploration is much more recent and forskoehlii, first showed up in the the fauna had remained virtually unknown Saronikos Gulf heavily impacted by mar- until the 1980s. Here the totality of the itime traffic: Cerithium scabridum species is of tropical Indo-Pacific origin appeared in Greece (ZENETOS et al., (save the cryptogenic species) but the 2009) much later than in Sicily, where it share is more balanced between Lessep- was obviously a secondary introduction. sian and non-Lessepsian. For the coast of Only a few Lessepsian species such as southern Turkey, we have 32 proven Pinctada radiata or Trochus erithreus may Lessepsians previously recorded in the have arrived in the Aegean as a result of
438 Medit. Mar. Sci., 11/2, 2010, 381-493 regular progression inside the Mediter- 2006b; SALOMIDI et al., 2006). Paradoxi- ranean. cally no Red Sea coral species has yet pen- etrated in the EMED (BIANCHI et al., 4.6. Miscellaneous invertebrates in the 2010). In the case of the Hydrozoa, the Eastern Mediterranean EMED harbours two thirds of the alien Fourteen alien species of Bryozoa, out species hitherto known for the Mediter- of the 23 recorded in the whole Mediter- ranean (26 out of 37) in accordance with ranean, are known for the EMED. Most of what is known for virtually all the other them are of Indo-Pacific origin, and most groups of organisms, which typically show are established. Three recently described a preponderance of aliens in that basin. species from Lebanon have presumably Among the 26 alien planktonic hydrozoa, penetrated from the Red Sea: this seems 17 were recorded in the EMED. Clytia lin- sufficiently documented for Parasmittina earis and Eirene viridula are the only estab- serruloides and P. spondylicola lished alien species in the entire Mediter- (HARMELIN et al., 2009) but the status ranean Sea. Most of the alien Hydrozoa in as a Levantine endemic cannot be exclud- the EMED have a circumtropical distribu- ed for Schizoretepora hassi (HARMELIN tion, the remainders have Indo-Pacific ori- et al., 2007). gin. Macrorhynchia philippina is the most Seven of the 16 alien species of Ascidi- widespread and abundant (MORRI, acea known for the Mediterranean are 2008). Four Scyphozoa aliens known for established in the EMED. They are in the Mediterranean supposedly came in majority Indo-Pacific, or alternatively cir- through the Suez Canal and are now estab- cumtropical. Rhodosoma turcicum, lished along the coast of Israel (GALIL et Ecteinascidia thurstoni, Phallusia nigra, al., 1990, 2010). Cassiopea andromeda has Ascidia cannellata, Symplegma brakenhiel- extended its penetration range to Turkey mi, Herdmania momus, and probably (ÖZGÜR & ÖZTÜRK, 2008) and Microcosmus exasperatus are regarded as Greece (ZENETOS et al., in press). Abyla Red Sea immigrants ( INAR et al., trigona is the single alien siphonophore in 2006b; BITAR et al., 2007; IZQUIERDO- the Mediterranean Sea, found in Egypt MUN~OZ et al., 2009; KATSANEVAKIS (ZAKARIA, 2006). et al., 2009; SHENKAR & LOYA, 2009; After having successfully invaded the KONDILATOS et al., 2010). Black Sea, with severe impact on both the Cnidaria are represented by 32 alien native ecosystem-functioning and the fish- species out of 46 known for the whole ery yield, the Western Atlantic combjelly Mediterranean. Anthozoans are repre- Mnemiopsis leidyi started to spread in the sented by two species: the Indo-Pacific Marmara and North Aegean Sea and gorgonian Acabaria erythraea has been reached Israel in 2009 (GALIL et al., found in the harbour of Hadera, Israel 2009). Its predator Beroe ovata followed (FINE et al., 2004), whereas the scleractin- the same path and arrived in the Aegean ian coral Oculina patagonica, possibly orig- Sea (SHIGANOVA et al., 2007). Among inating from the SW Atlantic, has been the three alien ctenophore species in the recorded from many sites through the entire Mediterranean Sea, Sulculeolaria whole basin (BITAR & ZIBROWIUS, angusta was found only in the Egyptian 1997; FINE et al., 2001; INAR et al., coasts (ZAKARIA, 2006).
Medit. Mar. Sci., 11/2, 2010, 381-493 439 Seven species of alien Porifera with VAFIDIS, 2009) might exert a consider- Indo-Pacific origin have been traditionally able impact on Mediterranean ecosystems. considered as Red Sea immigrants The EMED hosts the vast majority of (TSURNAMAL, 1969). More recently, the alien platyhelminthes, all reported as this has been questioned by VACELET et fish parasites, all with an Indo-Pacific al. (2007), who evidenced taxonomic prob- and/or Red Sea origin. Of the nine mono- lems with all of them except perhaps Geo- geneids introduced into this basin, five are dia micropunctata. According to considered established. Both host and par- VACELET et al. (2007), sponges with asite are Lessepsian immigrants that have tropical affinities hitherto known only for co-invaded the Mediterranean Sea via the the EMED are more likely to be remnants Suez Canal. The first documented case of of the thermophilic biota that thrived in a monogeneid invading a new biogeo- the Mediterranean during warmer periods graphical region by ‘natural’ extension of of the late Pliocene or Pleistocene and that its host range is that of the gill ectoparasite disappeared from the rest of the basin dur- Polylabris cf. mamaevi infecting the rabbit- ing the Würmian cooling (BIANCHI et al., fish Siganus rivulatus. The greater abun- 2010). dance of P. cf. mamaevi in the invading All six alien species of Sipuncula (Mediterranean) populations is probably occurring in the Mediterranean have been due to the changed, new environment, recorded for the EMED. Phascolion possibly impacting on host resistance to (Isomya) convestitum and Phascolosoma the parasite and encouraging heavier scolop are of Indo-Pacific origin, the infections (PASTERNAK et al., 2007). A remaining four are circumtropical possible explanation for the unexpected (PANCUCCI-PAPADOPOULOU et al., colonization success of immigrant fish par- 1999). All are known from a single or few asite species belonging to the class Myx- localities, where they can however be osporea, despite their putative complex abundant (A IK, 2007; 2008). life cycles, is discussed in DIAMANT KRAPP et al. (2008) reviewed the Pyc- (2010). nogonida of the EMED: three species are alien, although the status of Pigrogromitus 4.7. Alien fish in the Eastern Mediter- timsanus is questionable. Anoplodactylus ranean californicus and A. digitatus are estab- A total of 106 alien fish (nearly 70% of lished. the alien fish known in the Mediterranean) The echinoderm fauna of the EMED has been reported from the EMED, the includes eight alien species (out of twelve majority of which originate from the Indo- for the whole Mediterranean). Most of Pacific and Red Sea regions. Details on them are likely to have penetrated through their distribution per country, origin and the Suez Canal, but Asterias rubens origi- mode of introduction are provided in nated from the North Atlantic and was GOLANI & APPELBAUM-GOLANI first introduced in the Black Sea (2010). New data extending the distribu- (KARHAN et al., 2008). Two conspicuous tion of already reported alien species in Indo-Pacific species, Diadema setosum the Levantine basin include: Pisodonophis (YOKES & GALIL, 2006) and Synaptula semicinctus and Pomadasys stridens reciprocans (ANTONIADOU & (BILECENOGLU et al., 2009); Apogon
440 Medit. Mar. Sci., 11/2, 2010, 381-493 smithi and A. queketti (GOREN et al., overlooked or confused with Mycteroperca 2008); Enchelycore anatina, Lagocephalus rubra. Alternatively, it was introduced in spadiceus and Lagocephalus suezensis the ballast waters of a ship (HEEMSTRA (KATSANEVAKIS et al., 2009). et al., 2010). The complicated flood of confamilial species is prominent and deserves special 5. PHYTOPLANKTON interest, as it is unlikely to be coincidental. Alien cardinal fishes represent a good The introduction of marine microal- example of this, since only a single species gae in the Mediterranean Sea is hard to was known for over 60 years and four document. Theoretically, these microscop- species have been added to the fauna with- ic organisms can easily be transported by in the last five years (GOREN et al., 2009a; currents, but also by the feet of migratory GOREN et al., 2010; TURAN et al., 2010). birds, or introduced through human com- Two species of the family Champsodonti- merce of marine animals, ship’s ballast dae were observed from Turkey (Champ- waters or fouling organisms. The conse- sodon nudivittis) and Lebanon (Champ- quences of a microalgal invasion are only sodon vorax), interestingly both without evident if there are conspicuous changes in records from the Red Sea ( I EK & an area, e.g. water discoloration or BILECENOGLU, 2009; BARICHE, mucilage, or where the species is implied 2010a). Among recent alien gobies, Van- in toxicity events or other harmful impacts derhorstia mertensi was found off southern (ZINGONE & WYATT, 2005), while Turkish coasts and became quite common invasions go totally unobserved in case of shortly after its first observation rare species, which are a conspicuous part (BILECENOGLU et al., 2008; YOKES et of the phytoplankton biodiversity in all al., 2009), while the far-east chameleon seas. In addition, to prove that a species is goby Tridentiger trigonocephalus is report- an alien requires very sound background ed from Israel (GOREN et al., 2009b). knowledge of the species of a given area. Sudden occurrence of two alien angelfish- Unfortunately, the diversity of marine es, Pomacanthus imperator and Pomacan- microalgae is scarcely known in wide areas thus maculosus, from the Israeli and of the Mediterranean Sea, e.g. the south- Lebanese coasts, respectively, is also note- ern shores, where only a few sites have worthy (BARICHE, 2010b; GOLANI et been investigated, or the offshore waters, al., 2010). Among the latest records is that where studies are limited to occasional of the Indo-Pacific goby Trypauchen vagina sampling during cruises. Even in the north- in Israel (SALAMEH et al., 2010). ern Mediterranean waters the knowledge The appearance of Mycteroperca fusca, of the distribution of these unicellular a large Atlantic fish, in Israel in 2010, and organisms in a given area is far from being without having been spotted on the way, exhaustive, for several reasons. First, phy- raises the question of the route of its toplankton is difficult to sample properly, arrival. A reasonable possibility is that M. as most species have an ephemeral occur- fusca entered the Mediterranean through rence and a discontinuous spatial distribu- the Strait of Gibraltar, as many Atlantic tion. The resolution of sampling programs species do, then expanded its distribution rarely matches the space and time scale of along the North African coast but was occurrence of microalgae, and new species
Medit. Mar. Sci., 11/2, 2010, 381-493 441 show up when more detailed or integrative For the reasons stated above, it can sampling approaches are applied. For rarely be excluded that a suspected example, many cysts of the toxic dinofla- microalgal invader was already present as gellate Alexandrium andersonii were found part of the rare, hidden and unsampled in sediment traps deployed in the Gulf of phytoplankton. These factors are probably Naples (MONTRESOR et al., 1998). The at the origin of the extremely low number species was not known from the Mediter- of proven invaders among phytoplankton ranean Sea before, and has never been species (WYATT & CARLTON, 2002). recovered in any other plankton study in Nonetheless, a number of cases of pos- the area since. Probably Alexandrium sibly introduced microalgal species in the andersonii appears in the plankton for very Mediterranean Sea have been brought to short periods, or it is restricted to specific the general attention over the last few depths of the water column that are not years. One of the first cases was provided covered during investigations. Another by the toxic planktonic dinoflagellate major problem concerns all organisms that Alexandrium catenella, appearing in high are difficult to preserve and either break concentrations in a French coastal lagoon, or loose their peculiar characteristics with the Thau Lagoon, in 1998 (LILLY et al., common fixatives such as formalin and 2002). Based on the molecular similarity of lugol. Micromonas pusilla, a 2-3 Ìm naked the species with strains from Japan, it was prasinophyte, is easily identifiable when it proposed that the species had been trans- grows in cultures due to a very typical ported recently to the Mediterranean Sea. swimming pattern. It can reach concentra- The species had been seen before in the tions of up to 106 cells l-3 in coastal Balearic Sea in 1983 (MARGALEF & Mediterranean waters in winter ESTRADA, 1987), along the Spanish coast (THRONDSEN & ZINGONE 1994; since 1996 (VILA et al., 2001) and along ZINGONE et al., 1999). However, its cells the Sardinian coast in 1999 (LUGLI et generally burst upon preservation. This is al., 2003). Apparently its range is expand- also the case for much larger flagellates, ing across the Mediterranean Sea, as it has e.g. the raphidophyte Chattonella subsalsa been found recently on the Tunisian coast (30-40 Ìm) or naked dinoflagellates, which (TURKI & BALTI, 2007). On the other explode or lose their shape when fixed. hand, more recent studies using alternative Finally, many species require particular molecular tools have questioned the close skills or methods to be identified. This is relationships between Japanese and the case for cryptomonads, generally very Mediterranean populations, and hence the abundant in inshore waters, which require identification of Japan as the source area of cultivation, electron microscopy and at A. catenella (MASSERET et al., 2009). times molecular techniques to be identi- While the knowledge of planktonic fied at the species level. While their abun- species distribution poses many problems dance as a group can be traced with pig- in terms of sampling and detection, benth- ment analyses (HPLC) that allow alloxan- ic and epiphytic microalgae should be easi- thin to be detected, or by flow-citometry, er to collect. Yet these organisms have individual species are only identified by been studied very rarely in the Mediter- combining a number of different tech- ranean Sea until the end of the last centu- niques (CERINO & ZINGONE, 2006). ry, when they became popular mainly due
442 Medit. Mar. Sci., 11/2, 2010, 381-493 to health and environmental problems microphytobenthos sampling carried out caused by a toxic species Ostreopsis cf. to study O. ovata at several sites over the ovata, which forms conspicuous blooms in last decade. In this case, an introduction many coastal areas (e.g., TOTTI et al., from the Indian Ocean could be hypothe- 2010; MANGIALAJO et al., in press). Also sised, but the source population would be in this case, the sudden emergence of a missing for the time being, as the taxo- conspicuous problem was initially attrib- nomic identity of the species is still uncer- uted to a new invasion, probably in relation tain. On the other hand, data on benthic to climate change, as Ostreopsis species microalgae are now more complete than were previously known mainly from tropi- 10 years ago, which would allow one to test cal areas. However, an Ostreospis species a predictable range expansion westward. did form a conspicuous bloom in Ville- Indeed, marginal range expansion have franche Bay in 1972, where macroalgae been observed for other species, such as appeared to be covered by a brown mucous Alexandrium catenella (LUGLI et al., layer (Taylor, pers. comm.), but it was just 2003), G. catenatum (G MEZ, 2003), and recorded due to the curiosity of an expert a number of diatom species (G MEZ, taxonomist swimming in the area. The 2008), which could be related to the cli- species was reported as O. siamensis, the matic changes observed in the Mediter- only species known at the time (TAYLOR, ranean Sea or to other unexplained long- 1979), but it was likely to be the same as the term fluctuations in species abundance. one reported in more recent years in the A sounder background knowledge on same area, i.e. O. cf. ovata. From the phytoplankton distribution is available for molecular point of view, the Mediter- places where time series exist and accurate ranean populations of this species were identification is performed over a long- known to be similar only to Brazilian pop- term period. At the LTER-MC station in ulations (PENNA et al., 2010), but compa- the Gulf of Naples, two species never rable ribotypes have very recently been recorded despite their relatively easy iden- found also in Japanese waters (SATO et al., tification, have suddenly appeared in 2010). Molecular analyses on other mark- recent years. These were Skeletonema trop- ers are required to establish where, among icum (ZINGONE et al., 2003; KOOISTRA the three sites, the molecular diversity of et al., 2008), which is easily identifiable as Ostreopsis cf. ovata is the highest, which it is the only species in the genus having would indicate the source site for these dif- many chloroplasts, and Pseudo-nitzschia ferent populations around the world. multistriata (ZINGONE et al., 2003), Species of the benthic dinoflagellate which has a typical sigmoid shape. The two genus Gambierdiscus, which produce tox- species had never been recorded before ins that are responsible for the neurologi- 1995 and 2002, respectively, and have cal syndrome ciguatera, had never been shown increasing concentrations since recorded in the Mediterranean Sea until their first finding. In both cases, an intro- 2003, when specimens were found in duction could be hypothesised but it can- Greek waters off Crete (ALIGIZAKI & not be excluded that these are other exam- NIKOLAIDIS, 2008). No other record is ples of marginal range expansion from the confirmed from anywhere else in the southern Mediterranean Sea shore north- Mediterranean Sea, despite the intensive ward, or that these species were extremely
Medit. Mar. Sci., 11/2, 2010, 381-493 443 rare for many years, suddenly finding decades and continue to do so, forming appropriate environmental conditions to conspicuous blooms in places where they increase their abundance. The latter were not recorded at all before. However, hypothesis takes into consideration long- whether these species are really alien term fluctuations in species abundance should be tested effectively, considering at and is partially supported by the absence least two alternative hypotheses: 1) that of discoveries since 1992 of another they belong to the rare component of the Pseudonitzschia species, P. subpacifica. plankton and that they were missed in pre- This species was relatively common in the vious studies; 2) that their range expanded 1984-1987 period in the Gulf of Naples, from unexplored southern or eastern then it became rarer and eventually disap- Mediterranean areas. peared. Although P. subpacifica could still be in the Gulf of Naples, this case indicates 6. INVASIVE SPECIES that there is always the possibility of unex- plained local extinctions, and more gener- The ongoing incoming of immigrants ally points to important changes in species and their increasing success in the abundance over the long term. Mediterranean basin imply a profound Another relevant case is that of species and continuous alteration in the species showing discontinuous geographical distribution patterns that seems to have ranges. The dinoflagellate Alexandrium accelerated in the last decade. Up to balechii is only found in the Gulf of Mexico recent times the influence of these immi- and in Tyrrhenian waters, suggesting that grants was geographically limited to the the species originated in one of the two areas close to the source of introduction, places and was transported to the other, but today the idea of biogeographical although there are no data to establish boundaries to Lessepsian invasions (POR, where the species should be considered an 1981; QUIGNARD & TOMASSINI, alien. Several other dinoflagellates are only 2000) seems to be overcome. found in the Indo-Pacific area and in the The most invasive species per MSFD Mediterranean Sea so far (G MEZ, are listed in Table 3; their share per taxo- 2006), but the suspicion that they were nomic group is depicted in Figure 2. introduced through the Suez Canal should A total of 134 species are classified as be substantiated when their absence corre- invasive or potentially invasive in the sponds to their actual absence in the Mediterranean. The present list is based on Atlantic Ocean and not to our limited data on worst invasive marine species col- knowledge of protist biogeography. For lated from various sources for the Mediter- similar reasons, the high number of flagel- ranean (STREFTARIS & ZENETOS, late species that have been described in the 2006; GALIL, 2007; STREFTARIS et al., Mediterranean Sea and never recorded 2008), European Seas (EEA, 2007b; elsewhere cannot be considered as proof of DAISIE: VILA et al., 2008), and globally their endemism in the Mediterranean Sea (Global Invasive Species Database), man- (THRONDSEN & ZINGONE, 1994). aged by the Invasive Specialist Species In synthesis, in the Mediterranean Sea Group (ISSG: www.issg.org). The list was a number of planktonic and benthic enriched with species such as the bigfin reef- microalgae has appeared over the last squid Sepioteuthis lessoniana, an invader
444 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 3 Distribution of invasive (++) or potentially invasive alien species (+) in the Mediterranean MSFD areas. (-) Species absent from the area, *marketable species.
SPECIES WMED CMED ADRIA EMED Foraminifera Amphistegina lobifera + + + ++ Macrophyta 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 ++ ++ + + Polychaeta Branchiomma 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 + - - ++
Medit. Mar. Sci., 11/2, 2010, 381-493 445 Table 3 (continued)
SPECIES WMED CMED ADRIA EMED Polydora cornuta + - - ++ Pomatoleios kraussii - - - ++ Prionospio (Aquilaspio) krusadensis - - - ++ Prionospio (Prionospio) saccifera - - - ++ Pseudonereis anomala - + - ++ Pseudopolydora paucibranchiata - - - + Spirorbis (Dexiospira) marioni ++ - - ++ Streblospio gynobranchiata - - - ++ Crustacea Callinectes 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 + + ++ - Mollusca/Bivalvia Anadara kagoshimensis + - ++ + Anadara transversa + + ++ ++ Brachidontes pharaonis + + + ++ Chama pacifica - - - ++ Dendrostrea frons - - - ++ Fulvia fragilis ++ ++ ++ Musculista senhousia ++ + ++ + Pinctada radiata* + ++ + ++ Ruditapes philippinarum* + + ++ + Septifer forskali - - - ++ Spondylus spinosus - - - ++ Xenostrobus securis ++ - ++ - Mollusca/Cephalopoda Sepioeuthis lessoniana* - - - ++ Mollusca/Gastropoda Aplysia dactylomela ++ + ++ Bursatella leachii ++ ++ ++ ++
446 Medit. Mar. Sci., 11/2, 2010, 381-493 Table 3 (continued)
SPECIES WMED CMED ADRIA EMED Cellana rota + + ++ Cerithium scabridum + + - ++ Conomurex persicus* + + ++ Crepidula fornicata + + - + Ergalatax junionae - - - ++ Erosaria turdus - ++ - + Melibe viridis + ++ ++ ++ Pseudominolia nedyma - - - ++ Rapana venosa +? - + + Rhinoclavis kochi - - - ++ Porifera Paraleucilla magna ++ + + - Cnidaria/Anthozoa Oculina patagonica ++ + Cnidaria/Hydrozoa Clytia hummelincki ++ ++ ++ - Clytia linearis ++ ++ ++ ++ Garveia franciscana + - ++ - Macrorhynchia philippina - - - ++ Cnidaria/Scyphozoa Cassiopea andromeda - + - + Phyllorhiza punctata - + - + Rhopilema nomadica - + - ++ Ctenophora Mnemiopsis leidyi + + + ++ Bryozoa Bugula fulva + + + + Tricellaria inopinata - + ++ - Echinodermata Aquilonastra burtoni - - - ++ Asterias rubens - - - + Synaptula reciprocans - - - ++ Ascidiacea Botrylloides violaceus - - + - Distaplia bermudensis + + - - Microcosmus exasperatus - + - + Microcosmus squamiger ++ + - - Phallusia nigra - - - ++ Polyandrocarpa zorritensis + + - - Rhodosoma turcicum + + + Styela clava + - - -
Medit. Mar. Sci., 11/2, 2010, 381-493 447 Table 3 (continued)
SPECIES WMED CMED ADRIA EMED Symplegma brakenhielmi - - - + Fish Apogon 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- The share of invasive species does not tion explosion within 2010 (ZENETOS et support POR’s (2009, 2010) statement that al., in press). species from the tropical Indo-Pacific A total of 108 species are locally inva- should behave more like one more element sive or merely present in the EMED, 75 in of the native fauna and be less aggressive in the CMED, 53 in the Adriatic and 64 in their onset than real aliens, e.g. those orig- the WMED (Fig. 2). The WMED hosts inating from the temperate Atlantic or most macrophytes, whereas the EMED Pacific. Rather, it would conform to the has the lion’s share in polychaetes, crus- rule of thumb that one out of every ten taceans, molluscs and fish. established alien species becomes invasive.
448 Medit. Mar. Sci., 11/2, 2010, 381-493 Fig. 2: Breakdown of invasive or potentially invasive species per taxonomic group.
In addition to documented impacts on Little has been done to evaluate the biodiversity, some of the invasive species, risk posed by the introduction and spread among decapods, molluscs and fish, have of invasive species, or to minimize the risk turned to be beneficial to man and be com- of introducing additional alien species. mercially exploited in several countries Effective legislation on alien species exists (Table 3). Though no extinction of a native only in few countries and for selective vec- species is known, sudden decline in abun- tors e.g. in EU countries for aquaculture dance, and even local extirpations, concur- introductions; Egypt, France, Spain and rent with proliferation of aliens, has been Syria for introductions in ballast waters recorded (GALIL, 2007). (IMO/ BWC, Ballast Water Convention). Yet, there is no way to predict invasi- bility of ecosystems. However, community 6.1. Macrophytes ‘invasibility' appears to be the result of the Most spectacular has been the arrival relationship between native species rich- of several invasive exotic varieties of green ness and alien species ability to colonize and red algae. These algae have flourished new habitats (BULLERI et al., 2008). This at the expense of benthic Mediterranean concept implies that habitats with high lev- organisms. Their dense and creeping els of diversity are difficult to invade. In meadows - which feature up to 27,000 contrast, species-poor communities, or erected blades per square meter in Cauler- stressed ecosystems, are arguably more pa racemosa var. cylindracea - overwhelm prone to invasion, primarily due to lack of the benthic assemblages. Their success is biotic resistance (OCCHIPINTI- usually explained by a fast growth, perma- AMBROGI & SAVINI, 2003). At nent meadows and synthesis of secondary Mediterranean basin scale, this appears to metabolites active against natural enemies be true (see chapter 7). (herbivorous, epiphytes, pathogens, etc.).
Medit. Mar. Sci., 11/2, 2010, 381-493 449 Once they have spread, species like this molluscs, dock’s pilings, ropes and tires) are impossible to eliminate. along the Levantine coast of Turkey. Among the 125 alien macrophytes, 21 Pomatoleios kraussii formed a densely pop- are invasive or potentially invasive (17% of ulated belt in shallow-water areas in total taxa) (Table 3). The total number of Mersin Bay ( INAR, 2006). The invasive invasive or potentially invasive alien nereid species Pseudonereis anomala is a macrophytes decreases eastwards: WMED dominant component of the algal commu- (16 taxa), ADRIA (14 taxa), CMED (13 nities in the Levant Sea, attaining a popu- taxa), and EMED (11 taxa). Only six inva- lation density of 2475 ind.m-2 on the alga sive alien macrophytes have colonized the Jania rubens in Iskenderun Bay ( INAR four basins: the Indo-Pacific strain of & ALTUN, 2007). BEN-ELIAHU (1989) Asparagopsis taxiformis, Caulerpa racemosa reported that a native nereidid species of var. cylindracea, Codium fragile, Halophila the Mediterranean, Perinereis cultrifera stipulacea, Lophocladia lallemandii and (Grube), was excluded from the habitats of Womersleyella setacea. the Levant Sea by P. anomala. This species expanded its distributional range to Izmir 6.2. Polychaetes Bay, North Aegean Sea ( INAR & A total of 22 species (17% of the total ERGEN, 2005) and the coasts of Greece alien polychaetes) seem to have invasive (KAMBOUROGLOU & NICOLAI- character, dominating shallow-water ben- DOU, 2006). The eunicid worm Eunice thic habitats, estuarine areas, polluted antennata was found to be a common areas and harbour environments. All of species on the southern coast of Turkey them are invasive in the EMED, five are (KURT SAHIN & INAR, 2009) and present in the CMED, six in the Adriatic was also previously reported from the and nine in the WMED. coasts of Italy (CANTONE, 1993). Polluted soft bottoms especially near harbours greatly facilitate the establish- 6.3. Crustacea ment of ship-mediated species such as The Indo-Pacific prawns, in particular Polydora cornuta, Streblospio gynobranchi- Marsupenaeus japonicus, Metapenaeus ata and Pseudopolydora paucibranchiata monoceros, and Penaeus semisulcatus, are ( INAR et al., 2005, 2006a; DAGLI & highly prized and are considered a boon to INAR, 2008). Polluted environments, the Levantine fisheries (DURUER et al., where competition among species is low 2008). Portunus segnis, formerly known as and food is ample, provided these species P. pelagicus, is commercially exploited in with an excellent opportunity to build up Egypt (ABDEL-RAZEK, 1987). It is also dense populations. The maximum density commercially important but caught in of P. cornuta was calculated as 3170 ind.m-2, small quantities in Turkey (DURUER et that of S. gynobranchiata as 60480 ind.m-2 al., 2008). The latest record of P. segnis is and that of P. paucibranchiata as 6180 from the northern Tyrrhenian Sea ind.m-2 in Izmir Bay ( INAR et al., 2005; (CROCETTA, 2006). DAGLI & INAR, 2008). The invasive crab Percnon gibbesi is In a study by INAR (2006), alien one of the most recent and successful species comprised more than 95% of the invaders in the Mediterranean Sea. Its dis- serpulids found on hard substrata (rocks, tribution has been recently updated by
450 Medit. Mar. Sci., 11/2, 2010, 381-493 KATSANEVAKIS et al. (in press) who Rapana venosa, one of the most inva- report that the relative abundance of the sive species worldwide, has been rapidly species in the Kas -Kekova Marine Pro- increasing in the Adriatic Sea, thus tected Area (SE Turkey) reached 112 indi- enhancing the risk of invasion and further viduals in 2010, from two individuals in spreading. Maximum estimated popula- 2006. tion densities of > 500 ind..km-2 were The blue crab Callinectes sapidus was reached near the shores of Cesenatico transported into the Mediterranean in bal- (SAVINI et al., 2007). last tanks from the north-east coast of the The spread of the alien species across USA. After the first Mediterranean record the Mediterranean is not easy. Only six of (Venice, Italy, 1949), this species has been the ca. 50 Lessepsian species have crossed widely recorded in different Mediter- the Sicily Strait westwards and only two ranean regions, especially in the EMED (Bursatella leachii and Fulvia fragilis) have (Turkey, Greece, Syria, Lebanon, Israel reached the Spanish coast so far. and Egypt) where it became invasive and is None of the other species of Indo- commercially exploited (STREFTARIS & Pacific origin, so successful in the EMED, ZENETOS, 2006; DURUER et al., 2008). have a real invasion success in the western A decade after its introduction, basin. Some of the tropical Indo-Pacific Heterosaccus dollfusi, a parasite on Charyb- species which started in the CMED dis longicollis, became invasive in the south (Aplysia dactylomela, Haminoea cyanomar- Levantine coast. Despite the high preva- ginata, Melibe viridis) are now spreading to lence of the parasite and its injurious Malta (BORG et al., 2009; SCHEMBRI, impact on the host reproduction, the inva- 2009) or have even reached the western sive host-parasite pair reached a modus Mediterranean. Several individuals of vivendi with no noticeable reduction in the Melibe viridis were recently observed in host population (INNOCENTI & GALIL, Gulf Aranci, NE Sardinia (DONEDDU & 2007). TRAINITO, 2008). The case history of the forerunners 6.4. Mollusca indicates that the spread across the The gastropods Cerithium scabridum, Mediterranean involves a complex history Rhinoclavis kochi, Conomurex persicus and of natural spread with larvae and transport Bursatella leachi and the bivalves Pinctada via shipping from the original point of radiata, Brachidontes pharaonis, Fulvia introduction. In any case the distribution fragilis, Chama pacifica and Spondylus of these aliens in the western and central spinosus have been reported as locally parts of the Mediterranean is patchy and invasive in the EMED by ZENETOS et al. does not suggest a continuous progression. (2005, 2008). Conomurex persicus has been The limiting factors may be on the one reported to achieve densities of tens of hand, the natural species richness of the specimens per m2 at some sites western basin; on the other, the severe dif- (FISHELSON, 2000) on the Israeli coast. ference in sea surface temperatures (POR, To these may be added the limpet Cellana 2009, 2010) which may make it inhos- rota, reported as thriving to the point of pitable to thermophilic species such as displacing the native limpet Patella Pinctada radiata. caerulea (MIENIS, 2002, 2003). The impact of invasive species is com-
Medit. Mar. Sci., 11/2, 2010, 381-493 451 parable within these two main poles of marine invader with a high invasive alien molluscs in the Mediterranean, but potential. the pool of invasive species is not the same. A recent immigrant, Styela clava, To the south, an aggressive onset is seen among the worst invasives in Europe, has for some tropical Indo-Pacific species such been accidentally transported into the as Conomurex persicus, Cellana rota or (at Western Mediterranean by shellfish trans- the beginning of the invasion) Pinctada fer (DAVIS & DAVIS, 2008). Styela clava, radiata, which fail to become massively a solitary, hermaphroditic ascidian, is con- established in other parts of the basin. One sidered to be an aggressive invader of the latest introductions is Sepioteuthis throughout the world; this species is a lessoniana, a commercially important major pest to the mussel farming industry squid species for inshore fisheries through- of Canada (ARSENAULT et al., 2009). Its out its distributional range, which was recent discovery in the Mediterranean Sea observed in Iskenderun Bay (SALMAN, caused a considerable alarm to mussel 2002) and has rapidly spread in the Levant farms. and Aegean seas (KATSANEVAKIS et Phallusia nigra is an old invader in the al., 2009; LEFKADITOU et al., 2009; Mediterranean Sea. The quite sudden ZENETOS et al., in press). In the northern appearance of the species in the Aegean parts (Northern Aegean, Adriatic, and Sea and the extended distribution that it French Mediterranean lagoons) invasive demonstrated within a short time along species like Anadara kagoshimensis, Rudi- the coasts of Rodos and Kriti tapes philippinarum or Rapana venosa orig- (KONDILATOS et al., 2010; ZENETOS inate from temperate areas, and none of et al., in press) support the invasive charac- these are reported as invasive in the south- ter of the species, as already pointed out by ern parts of the Mediterranean. Therefore IZQUIERDO-MUN~ OZ et al. (2009). the climatic gradient in the Mediterranean Other species such as Botrylloides vio- is likely to promote natural boundaries for laceus, Symplegma brakenhemi and possi- the alien species, which should reach a sta- bly Polyandrocarpa zorritensis and Distaplia ble range at some time. bermudensis are locally invasive in altered habitats (harbours, aquaculture rafts, Po 6.5. Miscellaneous invertebrates and Ebro deltas). The ascidian Microcosmus squamiger, The scleractinian coral Oculina patag- first recorded at 1963 in Tunisia, is widely onica, a species of uncertain, but probably spread in the Western Mediterranean Atlantic origin, has taken advantage of the (France, Spain, Italy, North Africa: see warming sea and has explosively expanded synthesis in TUR N et al., 2007), and around the southern Mediterranean dur- locally in the CMED (Malta: ing the last years, building coral pavements IZQUIERDO-MUN~ OZ et al., 2009). (see latest updates in SARTORETTO et This species forms dense populations al., 2008). reaching up to 2300 ind..m-2 and its pres- Of the 37 alien hydrozoan species ence has strong implications in the struc- recorded for the Mediterranean, four can ture and functioning of the native com- be considered invasive. The occurrence of munities (RIUS et al., 2009). Microcos- Clytia hummelincki in the Mediterranean mus squamiger is considered a global Sea was only discovered in 1996 in the
452 Medit. Mar. Sci., 11/2, 2010, 381-493 northern Ionian (BOERO et al., 1997), BITAR-KOULI, 1995). It is now wide- and within the subsequent decade further spread and frequent in the whole Levant records came from the Southern Adriatic Sea ( INAR et al., 2006b). Its diffusion and the Tyrrhenian seas (BOERO et al., on coastal rocks from just below the sur- 2005; GRAVILI et al., 2008). At present, face to more than 40 m and the frequent C. hummelincki is extremely abundant in reproductive status (MORRI et al., 2009) the summer in shallow rocks and is rapid- suggest a high invasive potential. Its ly expanding its range thanks to the dis- colonies, up to 30 cm tall and stinging to persal capacity with currents of the the touch, are easily recognizable, so that medusa stage. Clytia linearis was recorded its further expansion in the Mediter- in the Suez Canal as early as in the 1930s ranean should be easily tracked (MORRI, (BILLARD, 1938) and rapidly spread 2008). through the whole Mediterranean to get One of the top 100 worldwide invasive its north-western reaches by the 1950s species, the comb jelly, Mnemiopsis leidyi, (PICARD, 1951): thus, it is probably one was first seen in the Saronikos Gulf of the first and most successful Lessepsian (EMED) in 1990 (SHIGANOVA et al., immigrants. The medusae of the two 2001). Sporadic sightings ever since from species of Clytia are relatively large and the Turkish and Greek coasts raised no are produced in great number: their vora- concern about its notorious impacts on cious predatory behaviour on coastal zoo- the ecosystem and the fisheries, until 2009 plankton may imply a significant impact when swarms if it spread across the on native Mediterranean ecosystems by Mediterranean, from Israel to Spain these two invasive species (GRAVILI et (FUENTES et al., 2010). Each summer al., 2008). since the mid-1980s swarms of the invad- The first Mediterranean record of ing jellyfish, Rhopilema nomadica, have Garveia franciscana came from the Lagoon appeared along the Levantine coast. of Venice in 1978, where it showed among These swarms of voracious planktotrophs the most abundant hydroids (MORRI, can stretch up to 100 km long. As they 1981). In subsequent years it was also draw nearer to shore, they adversely affect found in other estuarine habitats of the tourism, fisheries, and coastal installa- North Adriatic and Catalonia, NE Spain tions. (MORRI, 1982; GILI, 1986). Its spread in many brackish environments world-wide 6.6. Fish and its occurrence in ship-hull fouling Classically, Lessepsian species were (MORRI & BOERO, 1986), suggest high expected to be limited to the EMED, due invasive potential for this species: lack of to hydrological conditions, but the spread recent studies on Mediterranean brackish- to the western basin is now a reality. So water hydroids, however, prevents any far, only a few Lessepsian fish species have solid evaluation of its present degree of been observed in the Western Mediter- success. ranean: Abudefduf vaigiensis, Siganus Macrorhynchia philippina has luridus, and Fistularia commersonii. This occurred in the EMED since at least the latter is probably the fastest-spreading early 1990s, when has been found in pol- species in the Mediterranean: since the luted waters near Beirut (BITAR & first record in Mediterranean ten years
Medit. Mar. Sci., 11/2, 2010, 381-493 453 ago, F. commersonii is now reported more striped catfish, Plotosus lineatus, and the or less in the whole Mediterranean (Table pufferfish Lagocephalus sceleratus are a 2). The very recent records of F. commer- threat to human health. sonii and S. luridus in the north-western Mediterranean (S NCHEZ-TOCINO et 7. SYNTHESIS: STATUS AND al., 2007; DANIEL et al., 2009) could then GEOGRAPHICAL ORIGIN be the hint of future extensive spreading of Lessepsian species. The opposite direc- A total of 955 alien species occur in tion was followed by Atlantic invaders the Mediterranean, the vast majority of such as Seriola fasciata (SONIN et al., them having being introduced in the 2009), Pagellus bellottii (SAAD & EMED (718), fewer in the WMED (328) SBAIHI, 1995) and Sphoeroides pachy- and CMED (267), and least in the Adriat- gaster (GOLANI, 1996) that have reached ic (171). Of these, 535 species (56%) have the Levantine basin. been established in at least one area. We Today, some of these species consti- calculated that the established species tute an important part of local fishery accounted for 57.6% of the total number catches; 16 such species are listed (Table of alien species in the WMED, 51.5%, in 3) among the most invasive species in the the CMED, 52.6% in the Adriatic Sea and Mediterranean, Siganus luridus and 56.3% in the EMED. The casual records Siganus rivulatus though marketable, they constitute 27-29% of the alien species at are classified as venomous species in each basin. On the contrary, the number of FishBase. Lessepsian immigrants consti- questionable species in the EMED is three tuted 19% of the total abundance of fish times as many as those of the WMED. captured at the Lebanese rocky coast. In Macrophytes, polychaetes and crustaceans decreasing order of importance, were have the highest number of questionable Atherinomorus lacunosus (11.5%), Siganus species. The questionable status for luridus (2.3%), S. rivulatus (2.3%), Pem- macrophytes is generally due to the uncer- pheris vanicolensis (1.6%), Sargocentron tainty regarding their origin. For poly- rubrum (1.0%), Stephanolepis diaspros chaetes and crustaceans, the reason for the (0.2%), and Apogon nigripinnis (presently high percentage is the lack of precise taxo- Apogon pharaonis) (0.02%) (HARME- nomic works in the area and the corollary LIN-VIVIEN et al., 2005). Siganus luridus that these species were largely reported and S. rivulatus contributed 23% to the from ecological studies and seldom docu- species composition captured in purse mented by museum specimens or illustra- seine operating during the daytime in tions. Abu-Qir and El-Mex Bays, Alexandria The number of species per establish- (Egypt) (AKEL, 2005). ment success category along with the con- Invasive species such as: Apogon tribution of alien taxa distributed at each pharaonis, Fistularia commersonii, Gym- MSFD area is illustrated in Figure 3. Fur- nammodytes semisquamatus, Parexocoetus ther details per taxonomic group are pro- mento, Pempheris vanicolensis, Pteragogus vided below. pelycus, Sphoeroides pachygaster, Stephano- Figure 4 shows the origin of alien lepis diaspros, are just a nuisance to fish- species for each sub-basin. Species origi- eries. Others as the highly venomous nally distributed either in the Indo-Pacific
454 Medit. Mar. Sci., 11/2, 2010, 381-493 Fig. 3: a: Establishment success per basin (all taxa); b, c, d: Partitioning of alien taxa per establishment success.
Medit. Mar. Sci., 11/2, 2010, 381-493 455 Fig. 4: Origin of species per basin. IP-IO-RS: Indo-Pacific, Indian Ocean, Red Sea.
or Indian or Red Sea are clustered togeth- (2.7%) (Fig. 4). Cold water species, cir- er. Similarly, species originating in either cumboreal, N Atlantic, N Pacific, make up the east or west of the tropical Atlantic are a small percentage ranging between 4.2% grouped together, and so are those origi- and 21.6%, being more pronounced in the nating from the North Atlantic or the Adriatic and less so in the EMED. North Pacific. While maritime traffic and other It is estimated that almost 60% of the human activities such as aquaculture are alien species in the Mediterranean Sea important vectors for the introduction of come from the Suez Canal. By adding alien species worldwide (RUIZ et al., those species introduced via shipping 2000), in the Mediterranean they are not and/or aquaculture and trade, it is quite the main responsibles for the large differ- clear that most of the alien species in the ences observed among the four basins. In Mediterranean are warm water species, the EMED the human intervention originating in (sub)tropical seas. In partic- responsible for most of the aliens is the re- ular, thermophilic species, (Indo-Pacific, establishment of the connection with the Indian Ocean, Red Sea, tropical Atlantic, Indo-Pacific through the Suez Canal tropical Pacific, circum(sub)tropical), (1869), rather than the actual transfer of account for 88.4% of the introduced the invaders. In addition, with the present species in the EMED, 72.8% in the climate change (BELKIN, 2009), the trop- CMED, 59.3% in the WMED and 56.1% ical features and temperature of the waters in the Adriatic. The respective figures for are increasing more quickly in the EMED, Indo-Pacific species only are 47.5% for implying dramatic modifications of the EMED, 24.9% for CMED, 22.8% for the biota (POR 2009, 2010). As a consequence, Adriatic and 20% for WMED, whereas Indo-Pacific species (regardless of the tropical Atlantic species show a sharp mode introduction) have found optimal decrease from WMED (10.1%) to EMED environment for settlement in the EMED.
456 Medit. Mar. Sci., 11/2, 2010, 381-493 On the contrary, the Adriatic Sea, blages, almost totally made up by taxa of which is topographically a dead end, is the Atlantic affinity, more useful for the bios- area with the lowest number of aliens, tratigraphical and paleobathymetrical receiving them among those already applications that make foraminifers so established in the EMED and CMED important for micropaleontologists. As a that spread northwards, or among those consequence, it is very difficult to recog- introduced via shipping or aquaculture in nize new Atlantic immigrants entering hot spot areas such as the Venice Lagoon the Mediterranean. On the basis of the (OCCHIPINTI-AMBROGI et al., 2010). (scarce) geological data the assumption A significant number of Indo-Pacific that some more tolerant large fora- species reaches the WMED, which is minifers, such as amphisteginids, could enriched by ship-transferred species of have survived during the Pleistocene in Pacific origin mostly among macrophytes. the southern Mediterranean cannot be The Strait of Gibraltar is essentially dif- ruled out. ferent from the Suez Canal as a potential pathway for alien species. First of all, it is 7.2. Alien macrophytes in the Mediter- an ancient waterway, believed to have ranean originated 5.33 million years ago Compared to previous lists of alien (HILGEN & LANGEREIS, 1993), com- macrophytes in the Mediterranean Sea pared to the 142 years of the Suez Canal. (RIBERA & BOUDOURESQUE, 1995; Therefore, the current distribution of BOUDOURESQUE & VERLAQUE, Atlantic species, tropical or not, with part 2002; RIBERA SIGU N, 2002; of their range in the Mediterranean is the CORMACI et al., 2004; ZENETOS et al., result of a natural process over a long 2008), the present revised checklist has time; these species do not in any case taken into account all the possible intro- qualify as aliens, even if their discovery in ductions of exotic genotypes. Several the Alboran Sea comes later than their species already represented in the first description in the Atlantic. Secondly, Mediterranean Sea by putative native the Atlantic coast of Morocco is swept by populations, such as Cladosiphon zosterae, a prevalently southward oceanic circula- Desmarestia viridis, Ectocarpus siliculosus tion that prevents many potential new- var. hiemalis, Pylaiella littoralis, have been comers to approach the Strait of Gibral- only considered here if introduction tar. events are locally evident (e.g. Thau Lagoon, Lagoon of Venice). A total of 7.1. Alien foraminiferans in the Mediter- 125 taxa (79 Rhodophyta, 24 Fuco- ranean phyceae, 20 Chlorophyta, 1 Pelago- A total of 50 foraminiferan species phyceae and 1 Magnoliophyta) has been are reported in this work, all of them listed on the basis of recent works being warm water species. The high diver- (FURNARI et al., 2010; SFRISO et al., sity of Indo-Pacific species in the EMED 2010; TSIAMIS et al., 2010; CECERE et probably implies that many of them are al., in press; VERLAQUE et al., in press) Lessepsian immigrants. Data on shallow and on-going researches. Among them, water foraminifers are very few and scat- 97 taxa (78% of alien macrophytes) are tered. Most works concern deeper assem- established in at least one of the four
Medit. Mar. Sci., 11/2, 2010, 381-493 457 Mediterranean basins, the remainders able ( INAR, 2010). (22%) belong to casual and crypto- The Mediterranean Sea is largely genic/questionable categories. under the siege of polychaete species, As far as the major vectors of intro- mainly introduced from the Red duction are concerned, the transfer of Sea/Indo-Pacific areas (99 species, 77% of shellfish stands the first (46% of alien total species) and Atlantic Ocean (24 macrophytes), followed by the shipping species). The Suez Canal is one of the and the Suez Canal (29% and 27%, main vectors for the introduction of the respectively each). Some species have species. The EMED habitats are largely been probably introduced several times by dominated by the Red Sea immigrants. different vectors. The predominantly Polychaete species that, most probably, donor regions are temperate and cold entered the Mediterranean Sea via the regions (54% of alien macrophytes) vs Suez Canal and expanded their distribu- tropical regions (46%). Such a result may tion to the WMED are Nereis jacksoni, be an artefact due to the unequal effort of Lysidice collaris, Lumbrineris perkinsi, investigation between the WMED and the Notomastus aberans, N. mossambicus, EMED. Introductions along with shellfish Metasychis gotoi, Pista unibranchiata, Stre- transfers from north-eastern Atlantic and blosoma comatus and Branchiomma luctu- north-western Pacific have been thor- osum. oughly studied in the north-western basin The knowledge about alien poly- and the Adriatic, while in the EMED chaete species in the Mediterranean Sea smallest alien macrophytes remain proba- has greatly increased recently. A total of bly insufficiently studied (most of report- 59 species were reported after 2005. This ed Lessepsian migrants are large-sized number also includes some species (i.e. species). Sigambra constricta, S. parva, Dasy- branchus carneus) that were not classified 7.3. Alien polychaetes in the Mediterranean as alien species by ZENETOS et al. (2005, Compiling data on alien polychaete 2008). Due to the lack of precise taxo- species reported from the Mediter- nomic works on polychaetes, there is a ranean Sea revealed a total of 129 high number of questionable species. species belonging to 26 families, of These species were largely reported from which 59 species have become estab- ecological studies. The future taxonomical lished in the region, 31 species are casu- studies to be held in the whole basin al, 37 species are questionable and 2 would enable us to realize the real number species (Chaetozone corona and Parapri- of alien polychaete species in the Mediter- onospio coora) are cryptogenic. The ranean Sea. majority of alien polychaete species have been found in the EMED (97 species), 7.4. Alien crustaceans in the Mediterranean followed by the WMED (49 species), the A total of 153 alien crustacean species CMED (35 species) and the Adriatic Sea have entered the Mediterranean. The (22 species). In the Sea of Marmara, 14 majority are decapods (78 species), fol- alien polychaete species are known, of lowed by copepods (42), amphipod (11), which 6 species have become established isopods (8), cirripedes (7), cumaceans (3), in the area and 8 species are question- stomatopods (3) and tanaids (1).
458 Medit. Mar. Sci., 11/2, 2010, 381-493 Approximately 77% of all alien Parhyale explorator, a NE Atlantic species Mediterranean decapod species have an found in 2005 on the coast of Iskenderun Indo-Pacific/Indian/Red Sea origin, while Bay (BAKIR et al., 2008). Yet, the lack of only 23% are from the Atlantic. The dom- studies to determine the actual original inance of these ‘eastern alien species’ distribution of species such as Cymadusa over the total is obvious; it mainly reflects filosa, Gammaropsis togoensis and Mono- the situation (environment characteristics corophium sextonae has led to the assign- mainly) in the EMED and CMED where ment of the cryptogenic status to almost 82% of all Mediterranean alien species 30% of the species. could be found (only the following alien species are not present: Actumnus globu- 7.5. Alien molluscs in the Mediterranean lus, Charybdis feriata, Charybdis japonica, Molluscs are one of the major groups Charybdis lucifera, Cryptosoma cristatum, in the marine fauna worldwide and con- Dyspanopeus sayi, Eriocheir sinensis, Hem- tinue as the first contributors to the alien igrapsus sanguineus, Herbstia nitida, fauna in the Mediterranean Sea. The total Menaethius monoceros, Scyllarus caparti, number of reported alien species amounts Scyllarus posteli and Thalamita gloriensis). to 212, a small but steady increase of In the WMED, the situation is more bal- about 8% since the recent review of anced, where only 52.6% of the alien ZENETOS et al. (2005). More than half decapods are Indo-Pacific and the of the species (115 in total) are established remainders are Atlantic. The Adriatic is a in at least one of the four basins, 74 are peculiar sector with 13 alien species, six of casual and the remaining 24 are question- which (Callinectes danae, Charybdis able or cryptogenic. japonica, Charybdis lucifera, Dyspanopeus There is a clear decrease in the impor- sayi, Herbstia nitida and Scyllarus caparti) tance of aliens from the Levantine coast are exclusive to this sector. The EMED where extra-Mediterranean species may and WMED share only 10 species represent 10 to 20% of the local fauna, (12.8%), three of which inhabit estuarine towards the Alboran Sea where only one environments or are related to aquacul- species (Godiva quadricolor) is reported, ture or food (Rhithropanopeus harrisii, i.e. less than 1% of the total. Nevertheless Marsupenaeus japonicus and Necora the numbers drop dramatically if we disre- puber) and other two are invasive gard as aliens the Lessepsian immigrants, (Callinectes sapidus and especially Perc- following POR’s (2009, 2010) view that non gibbesi). these species have made their way by their The contribution of Indo-Pacific own means following the restoration of an species reaches 86% in copepods vs 11.6% ancient waterway, and therefore should of Atlantic origin and one debatable case: not be regarded as aliens. In this perspec- the Antarctic visitor (Spinocalanus terra- tive, there are hardly some thirty aliens in novae). Recent taxonomic studies have the Mediterranean which originate from shed some light on alien microcopepods donor regions other than the tropical in the Mediterranean (BÖTTGER- Indo-Pacific. These have arrived mainly as SCHNACK & SCHNACK, 2009). The a consequence of shipping and transfers of contribution of warm water species is also living spat for aquaculture, and the most high among alien amphipods, excepting heavily impacted areas are not the Levan-
Medit. Mar. Sci., 11/2, 2010, 381-493 459 tine basin, but the marginal marine envi- four species, the most widespread of ronments of the northern part of the which is Oculina patagonica. It is a species basins, including the Adriatic and the of presumably temperate SW Atlantic lagoons of the WMED. origin, probably introduced into the It is noteworthy that the Adriatic Sea Mediterranean by shipping. Invasive in shares more established species of mol- natural and artificial habitats, especially luscs (12 out of 16) with the WMED than along the southern WMED and EMED with any of the other parts of the Mediter- coasts (SARTORETTO et al., 2008). Five ranean, even the neighbouring Ionian Sea species coming from the subtropical east- (8 shared established species and 2 casu- ern Atlantic which have recently expand- al). This reflects not only the climatic sim- ed their range into the Mediterranean ilarity, with colder sea water temperature and are presently restricted to the Albo- in the northern parts, but also the fact that ran Sea (OCAN~ A et al., 2007, 2009) are most of these aliens are estuarine species excluded from our list; yet we keep some having some connections with aquacul- reservations on their mode of introduc- ture, therefore likely to be transported tion. Dendrophyllia laboreli is a coral in from the Venice area to the French expansion at the Canaries and also the Mediterranean lagoons or vice versa. continental Atlantic coast of Africa. Para- muricea grayi, Eunicella gazella and Euni- 7.6. Alien miscellaneous invertebrates in cella labiata are mainly known from the the Mediterranean Atlantic coast of Africa including a wide The prevailing taxa among alien tropical area (GRASSHOFF, 1992), but invertebrates other than polychaetes, have not been recognized as part of the crustaceans and molluscs, are the Mediterranean fauna by SHUCHERT et cnidarians (46 species: 37 hydrozoans, 4 al. (2003). According to recent observa- anthozoans, 5 scyphozoans) followed by tions, boats have borne larvae of Eunicel- bryozoans (23), ascidians (16), echino- la spp. from the Atlantic coast of Africa in derms (12), flatworms (12), sponges (8), their ballast waters. Antipathella wollas- sipunculans (6), pycnogonids (4), toni is a typical Macaronesian faunal ele- ctenophores (3), chaetognaths (2) and ment with clear tropical affinities siphonophores (1). With the exception (BRITO & OCAN~ A, 2004). Its recent of ascidians, the remaining taxa were settlement in the Alboran Sea may have poorly treated by COLL et al. (2010), if been favoured by the present climate addressed at all. warming. The current information is always The 16 alien ascidian species hitherto proportional to the degree of expertise in recorded in the Mediterranean basin rep- the area. Along this line, cnidarians and resent about 7% of the known species in bryozoans appear to be the best studied the whole basin. In many cases (13 out of taxa represented by many alien species, 16 species) the introduced ascidian species while sponges, although well studied in form established populations in antago- the region, have relatively few represen- nism with the endemic species of the tatives, and the reported ones are either Mediterranean benthos (ascidian or not). questionable or cryptogenic forms. The species Phallusia nigra, Herdmania Alien Anthozoa are represented by momus, Symplegma brakenhielmi, Rho-
460 Medit. Mar. Sci., 11/2, 2010, 381-493 dosoma turcicum, Ascidia cannellata and form a significant portion of the Mediter- probably Microcosmus exasperatus are con- ranean marine alien species list. The rate sidered immigrants via the Suez Canal of introduction is significantly high, where (IZQUIERDO-MUN~ OZ et al., 2009), 25 species were added since April 2008 while species such as Botrylloides violaceus, (ZENETOS et al., 2008), bringing the Polyandrocarpa zorritensis, Distaplia total number of alien fish species to 149. bermudensis and Styela clava have been This figure reflects the relevance of recently introduced by shellfish culture Mediterranean invasion that has no equiv- (BRUNETTI & MASTROTOTARO, alents on other seas. The richness of 2004; MASTROTOTARO & BRUNETTI, Mediterranean alien fish species is 2006; DAVIS & DAVIS, 2008; impressive if compared with native ones. IZQUIERDO-MUN~OZ et al., 2009). In Even if there is no general consensus on many cases the first record has been the total number of Mediterranean fish - reported when the species was already for example FREDJ & MAURIN (1987) present with dense populations often in listed 612 species while QUIGNARD & antagonism with native species (e.g. TOMASINI (2000) numbered 664 species BRUNETTI & MASTROTOTARO, – this review can give a more precise idea 2004; MASTROTOTARO & BRU- of how the original identity of Mediter- NETTI, 2006; DAVIS & DAVIS, 2008). ranean fish fauna has changed following The majority of echinoderms origi- the arrival of these immigrants. nate in the Indo-Pacific or are circum- So far, no species extirpation has been tropical species and their distribution is documented. Therefore the ingression of limited to the EMED; only Ophiactis savi- new species generates an increase of gnyi reaches the WMED. Asterias rubens, species richness, the main routes con- originating in the N Atlantic, is limited to tributing to Mediterranean fish diversity the Marmara Sea where it was introduced being the Suez Canal and the Strait of from an established population in the Gibraltar. Almost half of the alien fish Black Sea (KARHAN et al., 2008). species that have been recorded in the Chaetognaths are represented by two Mediterranean have given origin to per- species, both of Indo-Pacific origin, limit- manent populations. The magnitude of ed to the innermost part of the EMED. the phenomenon of fish invasion in the The presence of the parasitic nema- Mediterranean Sea raises concern on the tode Anisakis simplex s.str., in some indi- ecological and economic consequences vidual hosts (Gaidropsarus granti) from and solicits the urgent need of ongoing the WMED, which seemed to be the monitoring, especially over the southern result of immigration of these fish from rim of the basin were research effort is less the Atlantic into the WMED intense. (MATTIUCCI & NASCETTI, 2006), has been excluded, because, according to 8. THE STATE OF THE ART IN MANFREDI et al. (2000), it is a common MEDITERRANEAN MARINE ALIEN parasite in many Mediterranean fishes. SPECIES
7.7. Alien fish in the Mediterranean Species of tropical/subtropical affini- In terms of number of species, fish ty, favoured by climate warming, are intro-
Medit. Mar. Sci., 11/2, 2010, 381-493 461 duced and colonize the Mediterranean at treated by COLL et al. (2010). In contrast a fast rate, while cold water species are to ZENETOS et al. (2008), we do not settling at a lower rate. The net outcome is address microalgal species (either pelagic a jump in species richness. or benthic) for the reasons described in Compared to the latest lists of alien chapter 5. This implies that despite the biota (COLL et al., 2010; COSTELLO et collective effort of experts attempted in al., 2010) the present work includes a larg- this work, the number of introduced er number (954) of species (Table 4). The species remains underestimated. It is clear divergence in numbers is more obvious in that the introduced species have increased fish (149 this work vs. 116 in COLL et al., the biodiversity of the Mediterranean Sea, 2010), polychaetes (129 this work vs. 75 in as an entity, by 5.9% excluding phyto- COLL et al., 2010), bryozoans (23 this plankton and microzooplankton (Table work vs. one in COLL et al., 2010), and 4). The figure is even higher reaching cnidarians (46 this work vs three in COLL 27.9% in fish and 9% on average for the et al., 2010) (Table 4). Here, we also studied taxa. While species richness is include aliens among Porifera, Platy- increasing at whole basin scale, leading to helminthes and Foraminifera, taxa not a higher Á diversity, cases of local replace-
Table 4 The number of alien species in the Mediterranean by taxonomic group.
COLL et al., 2010 This work Taxon all aliens native aliens % aliens Protozoa (excluding Foraminifera) 0 0 4 Foraminifera >600 0 600 50 8.3 Rhodophyta 657 73 584 7913.5 Phaeophyta & Pelagophyceae 277 23 254 24+1 9.8 Chlorophyta 190 17 173 20 11.6 Magnoliophyta 7 1 6 1 16.7 Polychaeta 1172 75 1097 129 11.8 Crustacea 2239106 2133 153 7.2 Mollusca 2113 200 1913 212 11.1 Cnidaria 757 3 754 46 6.1 Bryozoa 388 1 387 23 5.9 Ascidiacea 22915 214 16 7.5 Echinodermata 154 5 14912 8.1 Porifera 681 0 681 8 1.2 Platyhelminthes 1000 0 1000 12 1.2 Other Invertebrates 2168 2 2166 16 0.7 Fish 650 116 534 14927.9 Total 637* 954* Average % 3.3% 5.9%
*excluding monocellular algae
462 Medit. Mar. Sci., 11/2, 2010, 381-493 ment have been reported (GALIL, 2007), lished, aliens if undertaken seriously. which may imply an alteration of · diver- The application of molecular meth- sity. In addition, the spatial overlap ods recently developed will be of great between alien and native species causes help in the definition of the relation- biotic homogenization (BEN RAIS ships between populations from differ- LASRAM & MOUILLOT, 2009), and ent areas, and in the assessment of pos- hence a depression in ‚ diversity. Thus, sible routes of introduction and immi- the relationship between the entering of gration. In addition, metagenomic tech- alien species and threats to biodiversity is niques will presumably allow a much not straightforward (BIANCHI et al., more complete if not exhaustive knowl- 2010). edge of the microbial species in given While biodiversity is currently a key areas through their molecular signa- indicator of an ecosystem's health, statis- tures. In the meantime, classical taxo- tics tell too simple a story in the Mediter- nomic studies should be intensified to ranean Sea. In a recent study (2008) investigate the still wide proportion of assessing fish assemblages associated with unreported alien species of small-sized Posidonia oceanica meadows in the taxa in the Mediterranean Sea including Rodos Island (EMED), the silverver- microalgae. stripe blaasop, Lagocephalus sceleratus, ranked among the 10 most abundant Acknowledgements species, totalling 2% of total biomass, while bluespotted cornetfish, Fistularia The work was initiated under the aus- commersonii, and reticulated leatherjack- pices of the SEBI2010 - Streamlining et, Stephanolepis diaspros contributed European 2010 Biodiversity Indicators - 0.6% and 0.3%, respectively Expert Group 5: Numbers and costs of (KALOGIROU et al., 2010). Of the near- invasive alien species. ly 30000 specimens collected by commer- Further research on the distribution cial benthic trawler off the central Israeli of alien species in the Mediterranean Sea coast at depths between 15 and 30 m, in has been carried out in the frame of the October 2008 (OCCHIPINTI- Integrated Project ‘SESAME’ (Southern AMBROGI & GALIL, 2010), only 9% European Seas: Assessing and Modelling were native Mediterranean species, the Ecosystem changes: Project no: 036949) rest consisted of Erythrean aliens, such as funded by the EU [A. Zenetos, I. Siokou the highly venomous striped catfish, Plo- and N. Streftaris] and in the frame of the tosus lineatus (11437 specimens), the research project ‘The impacts of biologi- blotchfin dragonet, Callionymus filamen- cal invasions and climate change on the tosus (5745), the silver sillago, Sillago biodiversity of the Mediterranean Sea’ sihama (1423), the kuruma prawn, Marsu- (C.N. Bianchi and C. Morri), an Italy- penaeus japonicus (1154) and the velvet Israel co-operation funded by the Italian shrimp Metapenaeopsis consobrina Ministry for the Environment. Special (1138). thanks are due to C. Salas Casanova and The faunal survey of large harbours A. Logan whose constructive criticism has been largely neglected and is likely to and suggestions have improved the man- yield many more localized, but estab- uscript.
Medit. Mar. Sci., 11/2, 2010, 381-493 463 ANNEX: Name changes
Current name Old /others name(s) used Protozoa Perkinsus olseni Lester & Davis Perkinsus atlanticus Macrophytes Caulacanthus okamurae Yamada Feldmannophycus okamurae Codium fragile subsp. fragile (Suringar) Hariot Codium fragile subsp. tomentosoides Hypnea anastomosans Papenfuss, Lipkin & Silva Hypnea esperi Saccharina japonica (Areschoug) C.E. Lane, C. Mayes, Laminaria japonica Druehl & G.W. Saunders Spongoclonium caribaeum (B rgesen) M.J. Wynne Pleonosporium caribaeum Polychaeta Apoprionospio pygmaea (Hartman, 1955) Prionospio pygmaea Linopherus canariensis Langerhans, 1881 Pseudeurythoe acarunculata Lumbrineris perkinsi Carrera-Parra, 2001 Lumbrineris inflata Prosphaerosyllis longipapillata (Hartmann-Schroeder, 1979) Sphaerosyllis longipapillata Streblosoma comatus (Grube, 1856) Streblosoma hesslei Crustacea Austrominius modestus Darwin, 1854 Elminius modestus Amphibalanus eburneus (Gould, 1841) Balanus eburneus Glabropilumnus laevis (Dana, 1852) Heteropanope laevis Linguimaera caesaris Krapp-Schickel, 2003 Maera hamigera Parvocalanus crassirostris Dahl, 1894 Paracalanus crassirostris Pilumnus minutus De Haan, 1835 Pilumnus hirsutus Portunus segnis (Forsska l, 1775) Portunus pelagicus Subeucalanus subcrassus Giesbrecht, 1888 Eucalanus subcrassus Sternodromia spinirostris (Miers, 1881) Dromia spinirostris Synalpheus tumidomanus africanus Crosnier & Forest, 1966 Synalpheus hululensis Mollusca Anadara kagoshimensis (Tokunaga, 1906) Anadara inaequivalvis / Arca/Scapharca Anadara transversa (Say, 1822) Anadara demiri Anteaeolidiella foulisi (Angas, 1864) Aeolidiella indica Bostrycapulus odites Collin, 2005 Crepidula calyptraeiformis Bostrycapulus aculeatus Bulla arabica Malaquias & Reid, 2008 Bulla ampulla Diplodonta bogii Van Aartsen, 2004 Diplodonta cf. subrotunda Canarium mutabile (Swainson, 1821) Strombus mutabilis Cerithidium diplax (Watson, 1886) Clathrofenella fusca Clathrofenella ferruginea Cerithidium perparvulum (Watson, 1886) Cerithiopsis tenthrenois Conomurex persicus (Swainson, 1821) Strombus persicus Strombus decorus raybaudii
464 Medit. Mar. Sci., 11/2, 2010, 381-493 ANNEX (continued)
Current name Old /others name(s) used Doxander vittatus (Linnaeus, 1758) Strombus vittatus vittatus Ergalatax junionae Houart, 2008 Ergalatax obscura/ Morula martensi Haminoea japonica (Pilsbry, 1895) Haminoea callidegenita Leucotina natalensis Smith, 1910 Adelactaeon amoenus/ Monotygma amoena Melibe viridis (Kelaart, 1858) Melibe fimbriata Monotigma lauta (A. Adams, 1853) Adelactaeon fulvus/ Monotygma fulva Notocochlis gualteriana (Recluz, 1844) Natica gualteriana Syrnola lendix (A. Adams, 1863) Styloptygma beatrix Zygochlamys patagonica (King & Broderip, 1832) Chlamys lischkei Miscellaneous invertebrates Clytia linearis (Thornely, 1899) Clytia gravieri Aquilonastra burtoni (Gray, 1840) Asterina burtoni Fish Atherinomorus forskalii (Ruppell, 1838) Atherinomorus lacunosus Equulites klunzingeri (Steindachner, 1898) Leiognathus kluzingeri Favonigobius melanobranchus (Fowler, 1934) Papillogobius melanobranchus
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