Additions to the annotated list of marine alien biota in the Mediterranean with special emphasis on Foraminifera and Parasites

Argyro Zenetos

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 origin, 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 insufficient, aliens have increased the total species rich...

More than 60 marine non-indigenous species (NIS) have been removed from previous lists and 84 species have been added, bringing the total to 986 alien species in the Mediterranean [775 in the eastern Mediterranean (EMED), 249 in the central Mediterranean (CMED), 190 in the Adriatic Sea (ADRIA) and 308 in the western Mediterranean (WMED)]. There were 48 new entries since 2011 which can be interpreted as approximately one new entry every two weeks. The number of alien species continues to increase, by 2-3 species per year for macrophytes, molluscs and polychaetes, 3-4 species per year for crustaceans, and 6 species per year for fish. The dominant group among alien species is molluscs (with 215 species), followed by crustaceans (159) and polychaetes (132). Macrophytes are the leading group of NIS in the ADRIA and the WMED, reaching 26-30% of all aliens, whereas in the EMED they barely constitute 10% of the introductions. In the EMED, molluscs are the most species-rich group, followed b...

We measured some nuclear cross sections needed to assess local irradiation models for the early solar system. A one-stage irradiation by protons and alpha particles, either with or without 3He, seems unable to explain meteoritic observations.

Review Article Mediterranean Marine Science Volume 9/1, 2008, 119-165 Additions to the annotated list of marine alien biota in the Mediterranean with special emphasis on Foraminifera and Parasites A. ZENETOS1, E. MER 2, M. VERLAQUE3, P. GALLI4, C.-F. BOUDOURESQUE3, A. GIANGRANDE5, M. E. INAR6 and M. BILECENO LU7 Hellenic Centre for Marine Research, Institute of Marine Biological Resources, P.O. Box 712, 19013, Anavissos, Attica, Hellas 2 Moda, Hüseyin Bey Sokak, 15/4, 34710 Kad köy, stanbul, Turkey 3 UMR 6540, DIMAR, COM, CNRS, Université de la Méditerranée, F13288 Marseille France 4 Wet Lab, Department of Biotechnology and Biosciences, University of Milano Bicocca, Piazza della Scienza 2, 20126 Milano, Italy 5 Department of Biological and Environmental Sciences and Technologies, University of Lecce, Complesso Ecotekne, Via Prov. le Lecce-Monteroni, 73100 Lecce, Italy 6 Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100 Bornova, Izmir, Turkey 7 Adnan Menderes University, Faculty of Arts & Sciences, Department of Biology, 09010 Aydin, Turkey 1 e-mail: zenetos@ath.hcmr.gr Abstract The present work is an update of the annotated list (ZENETOS et al., 2006) based on literature up to April 2008. Emphasis is given to ecofunctional/taxonomic groups poorly addressed in the annotated list, such as the foraminiferan and parasites, while macrophytes are critically reviewed following the CIESM Atlas (VERLAQUE et al., in press). Moreover, in this update the bio-geographic area addressed includes the Sea of Marmara. The update yields a further 175 alien species in the Mediterranean bringing the total to 903. As evidenced by recent findings, more and more previously known ‘casual’ aliens, are becoming established. Approximately 100 more species have become well established in the region, raising the number of established species to 496 versus 385 until 2005. In the period from January 2006 to April 2008 more than 80 published papers have resulted in the recording of 94 new aliens, which is interpreted as a new introduction every 9 days, a rate beyond the worst scenario. Medit. Mar. Sci., 9/1, 2008, 119-165 119 Introduction Biological invasions have become a hot issue in recent decades. An increasing trend towards aquatic invasions has been documented for all European Large Marine Ecosystems but is most pronounced in the Mediterranean Sea (STREFTARIS et al., 2005; EEA, 2007a). A compilation of the available information on the alien marine biota in the Mediterranean resulted in an annotated list that includes 745 alien species (ZENETOS et al., 2006). This covered the groups: fish, zoobenthos (Polychaeta, Crustacea, Mollusca, Echinodermata, Sipuncula, Bryozoa and Ascidiacea) comparatively fully, and less so phytobenthos, phytoplankton and zooplankton. Little information was provided for taxa such as Foraminifera and Isopoda that have not been well studied in the Mediterranean. A gap in information was also noticeable as regards parasites. Since then, new publications have: a) confirmed the establishment and further geographic expansion of rare species; b) reported on the occurrence of new alien species; c) enlightened on the introduction of little studied groups such as parasites, foraminiferans and d) critically reviewed higher taxa such as macrophytes. In parallel, publications based on molecular studies reconsider the status of species excluded in the annotated list, while publications on taxonomy have significantly changed the nomenclature of certain species. The aim of the present work is to update the annotated list by reporting on species presumably omitted in ZENETOS et al. (2006) and on the new species recorded in the 2006-2008 period and possible changes in their establishment success. Emphasis is put on foraminiferan and parasites, through collaboration with regional experts. Methodology The basis for the present work is the annotated list of marine alien species in the Mediterranean (ZENETOS et al., 2006), called here after ‘annotated list’. The list has been updated to incorporate species recorded up to April 2008. The species updates are presented in 10 units, which are ecofunctional/taxonomic groups, namely: 1: Phytoplankton, 2: Zooplankton, 3: Phytobenthos, 4: Zoobenthos/Polychaeta, 5: Zoobenthos/Crustacea, 6: Zoobenthos/Mollusca, 7: Zoobenthos/Foraminifera, 8: Zoobenthos /Miscellanea, 9: Parasites and 10: Fish. Data have been mostly extracted from a simple information system, structured in ACCESS, which has been developed at the Hellenic Centre for Marine Research (internal use only). The database is updated on a monthly basis and results are reported to UNEP/MAP and the European Environment Agency (EEA) (UNEP/MAP, 2004; EEA, 2006). One of the main uses of the aforementioned system is the development of a trend indicator for marine and estuarine species in Europe for the SEBI20101 program (EEA, 2007a, b) and the Mediterranean (UNEP/MAP, 2007). SEBI2010 Streamlining European Biodiverity Indicators: Framework of biodiversity indicators to assess progress towards the 2010 biodiversity target). Expert group on ‘trends in invasive alien species’: http://biodiversity-chm.eea.eu.int/ information/indicator/F1090245995 1 120 Medit. Mar. Sci., 9/1, 2008, 119-165 A major difference with the ‘annotated list’ is the geographic coverage of the Mediterranean. The alien biota of the Sea of Marmara (KAMINSKI et al., 2002; INAR et al., 2006a), which were omitted in the first compilation of data, are included in this work. Concerning their establishment success, alien species are grouped into six broad categories, namely established, casual, questionable, cryptogenic, excluded and invasive, as defined in the ‘annotated list’, For foraminifera a finer distinction of the established category includes the rare species (those observed more than twice in several different localities but always few in number: one or two individuals) and the frequent ones (those species recorded many times in large quantities and showing a wide range of distribution patterns). Besides the presentation of new species (i.e. species missing in ZENETOS et al., 2006 and those recorded in the 2006-2008 period) with notes on the country of their occurrence, special attention is given to those species whose establishment success has altered compared to the ‘annotated list’. For example, in the light of molecular and/or detailed taxonomic studies, some previously assumed questionable or excluded species have been now moved to the cryptogenic or established category. Nomenclature changes are presented under ‘name changes’. These are the result of the latest taxonomic studies (e.g. VAN AARTSEN 2004; 2006); group review papers (GOMEZ, 2008); molecular studies (e.g VERLAQUE et al., 2005) and web databases (GUIRY & GUIRY, 2008). Thanks to collaboration with regional experts, emphasis has been placed on foraminiferan and parasites, two groups Medit. Mar. Sci., 9/1, 2008, 119-165 poorly addressed in the ‘annotated list’, yet with many representatives among aliens. A full list of the experts per taxonomic group who contributed in various ways is provided in the acknowledgements. Results 1. Phytoplankton The validity of categorizing the diatoms and dinoflagellates reported in STREFTARIS et al. (2005) as alien phytoplankton in the European Seas was investigated by GOMEZ (2008) who concluded that the number of alien phytoplankton species in European Seas has been excessively inflated. Most of the discrepancies noted in GOMEZ (2008) had already been rectified in the ‘annotated list’. Exceptions were the cosmopolitan Protoceratium reticulatum reported as Gonyaulax grindleyi, and Gyrodinium falcatum also reported as Gymnodinium fusus/Pseliodinium vaubanii which are now excluded from the Mediterranean aliens. In addition, the benthic/epiphytic dinoflagellates that were included among phytoplankton in the ‘annotated list’ are now moved to phytobenthos. Warm-water species expand their geographical ranges or increase their abundances to detectable levels during warming periods. Alexandrium tamarense, Chaetoceros coarctatus, Proboscia indica and Pyrodinium bahamense are a few examples of marginal dispersal associated with climatic events rather than species introductions from remote areas. Again, these introductions are considered important even if only to document the impact of climatic changes, and are kept on our list. As a result four (4) new phytoplankton species have been added to the list ((Tables 1.1, 1.2). 121 Table 1.1 New alien phytoplanktonic species and changes in establishment success. *species excluded in the ‘annotated list’ Establishment success: C=casual; E=established; Q=questionable; CR=cryptogenic New species Alexandrium concavum (K.R. Gaarder) E. Balech Alexandrium kutnerae (E. Balech) E. Balech *Proboscia indica (Peragallo) Herna’ ndez-Becerril *1 Pseudosolenia calcar-avis Schultze, 1858 Change in establishment success Karenia mikimotoi (Miyake & Kominami ex Oda) G. Hansen & . Moestrup Prorocentrum mexicanum Osorio-Tafall Establishment success C C CR/E E Source Tunisia: DALY YAHIA-KEFI et al., 2001 Tunisia: DALY YAHIA-KEFI et al., 2001 GOMEZ, 2008 Sea of Marmara: INAR et al., 2006a From Q to CR/E GOMEZ, 2008 From C to CR/E GOMEZ, 2008 Proboscia indica (H. Peragallo) D.U. Hernandez-Becerril (= Rhizosolenia indica) was found previously in the Indian Ocean more than in the Mediterranean or Atlantic. It was historically considered as a variety of the cosmopolitan Rhizolenia alata Brightwell. 1 Table 1.2 Name changes in phytoplankton. NEW NAME Protoceratium reticulatum (Claparède & Lachmann) Butschli Karenia brevis (C.C. Davis) G. Hansen & . Moestrup Karenia mikimotoi (Miyake & Kominami ex Oda) G. Hansen & . Moestrup Gymnodinium aureolum (E. M. Hulburt) G. Hansen Gyrodinium falcatum Kofoid & Swezy Proboscia indica (H. Peragallo) D. U. Herna’ ndez-Becerril OLD NAME Gonyaulax grindleyi Reinecke Source GOMEZ, 2008 Gymnodinium breve C.C. Davis GOMEZ, 2008 Gymnodinium mikimotoi Miyake & Kominami ex Oda Gyrodinium aureolum E.M. Hulburt GOMEZ, 2008 Gymnodinium fusus Schütt Rhizosolenia indica H. Peragallo GOMEZ, 2008 GOMEZ, 2008 2. Zooplankton The number of alien zooplanktonic species in the ‘annotated list’ has been 122 GOMEZ, 2008 reduced by six since the benthic copepod species (Canuellina insignis, Enhydrosoma hopkinsi, Robertsonia salsa, Scottolana longipes, Stenhelia inopinata, Stenhelia minMedit. Mar. Sci., 9/1, 2008, 119-165 uta) reported by POR (1964; 1972) were erroneously placed among zooplankton. Eight (8) new alien species (Table 2) are added to the ‘annotated list’. Five of these species occur on the Mediterranean coast of Egypt: Abyla trigona, Sulculeolaria angusta, Olindias singularis, Rhabdosoma whitei, Aidanosagitta neglecta (ZAKARIA 2004; 2006), one in South Turkey: Ferosagitta galerita (ÜNAL et al., 2002; TERBIYIK et al., 2007) and one in Tunisia: Stomolophus meleagris (DALY YAHIA et al., 2003). It is interesting to note the finding of Beroe ovata, one of the worst Invasive Alien Species (IAS) in the Black Sea, now having been recorded from the South Aegean Sea (SHIGANOVA et al., 2007). The three copepods species, namely Acartia hasanii, Paracartia ioannae and Paracartia janetae (ÜNAL et al., 2002) that were excluded from the ‘annotated list’, as unsupported records, retain their excluded status. After examination of specimens from the area, it was concluded that all specimens were copepodites (future females) of Acartia clausi and Acartia tonsa (A. Gubanova, pers. commun.). The geographic expansion of reported species such as Phyllorhiza punctata (Ionian Sea: NAVANDI & KIKINGER, 2007), and the documented burst of alien zooplanktonic species in general, presum- Table 2 New alien zooplanktonic species in the Mediterranean and changes in establishment success. *species excluded in the ‘annotated list’ Taxon: AMPH=Amphipoda; CHA=Chaetognatha; CTE=Ctenophora; SIPH=Siphonophora; SCY=Scyphozoa Establishment success: C=casual; E=established; Q=questionable Taxon New species SIPH CHA Abyla trigona Quoy & Gaimard, 1827 *Aidanosagitta neglecta1 CTE Beroe ovata Mayer, 1912 CHA Ferosagitta galerita (Dallot, 1971) SCY Olindias singularis Browne, 1905 AMPH Rhabdosoma whitei Bate, 1862 SIPH Stomolophus meleagris (L. Agassiz, 1862) SIPH Sulculeolaria angusta Totton, 1954 Change in establishment success SCY Phyllorhiza punctata von Lendenfeld, 1884 1 Establishment Source success C Egypt: ZAKARIA, 2004 E E Egypt: H. ZAKARIA, pers. commun Greece: SHIGANOVA et al., 2007 Turkey: TERBIYIK et al., 2007 E C C Egypt: ZAKARIA, 2004 Egypt: ZAKARIA, 2006 Tunisia: DALY YAHIA et al., 2003 E Egypt: ZAKARIA, 2004 From C to E Greece: NAVANDI & KIKINGER, 2007 C Reported as Sagitta neglecta. Medit. Mar. Sci., 9/1, 2008, 119-165 123 ably reflect the sensitivity of zooplankton to climatic changes. 3. Phytobenthos A total of 110 species (22 Chromobionta, 71 Rhodobionta, 16 Chlorobionta and 1 Plantae) constitute the main core of the CIESM Atlas of exotic macrophytes of the Mediterranean Sea (VERLAQUE et al., in press). Comparing this information to the ‘annotated list’ has led to the addition to the new list of six cryptogenic species that were reported before the Suez Canal opening and assumed to be native by ZENETOS et al. (2006) (e.g.: Asparagopsis taxiformis, Desmarestia viridis) (Table 3.1); recent rapid changes in their Mediter- ranean distribution give evidence of a recent introduction of either a cryptic species similar to the Mediterranean one or an alien genotype from a remote population. In addition, seven species recently identified as new for the whole or a part of the Mediterranean Sea are also listed in Table 3.1. These include Dictyota ciliolata Sonder ex Kützing (reported from Catalonia: RULL LLUCH et al., 2007); Batophora sp (reported from Italy: BOTTALICO et al., 2006); and 5 species namely, Microspongium tenuissimum (Hauck) A.F. Peters; Lomentaria flaccida Tanaka; Grateloupia minima P.L. Crouan & H.M. Crouan; Polysiphonia stricta (Dillwyn) Greville and Cladophora hutchinsioides Hoek & Womersley, all collected in the Thau Table 3.1 New alien phytobenthic species in the Mediterranean and changes in establishment success. *excluded in the ‘annotated list’. Countries of first observation concerned the populations assumed to be introduced. Taxon: RHO=Rhodobionta, CHL=Chlorobionta, CHR=Chromobionta Establishment success: C=casual; E=established; Q=questionable; CRf=cryptogenic frequent; CRr=cryptogenic rare Taxon New species RHO Acanthophora naydaformis (Delile) Papenfuss Antithamnionella boergesenii (Cormaci & G. Furnari) Athanasiadis *Antithamnionella elegans (Berthold) J.H. Price & D.M. John *Antithamnionella spirographidis (Schiffner) E.M. Wollaston *Asparagopsis taxiformis (Delile) Trevisan de Saint-Léon *Acrochaetium spathoglossi B rgesen Anotrichium okamurae Baldock Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman & Boudouresque RHO RHO RHO RHO RHO RHO CHL Establishment Source success CRf/F Egypt: VERLAQUE et al., 2005 E Algeria: Verlaque et al., in press CRf/Q Italy : VERLAQUE et al., in press CRf/Q Italy : VERLAQUE et al., in press CRf Q E E Egypt: VERLAQUE et al., 2005 Egypt: VERLAQUE et al., in press VERLAQUE et al., in press Libya: VERLAQUE et al., in press (continued) 124 Medit. Mar. Sci., 9/1, 2008, 119-165 Table 3.1 (continued) Taxon New species CHL Caulerpa racemosa var lamourouxi f. requienii (Montagne) Weber-van Bosse Caulerpa racemosa var. turbinata (J. Agardh) Eubank - uvifera (C. Agardh) J. Agardh Chondria coerulescens (J. Agardh) Falkenberg1 *Ceramium bisporum D.L. Ballantine2 Cladophora hutchinsioides Hoek & Womersley3 *Cladosiphon zosterae (J. Agardh) Kylin *Desmarestia viridis (O.F. Müller) J.V. Lamouroux Dictyota ciliolata Sonder ex Kützing Dictyota okamurae (E.Y. Dawson) I. Hörnig, R. Schnetter & W.F. *Ectocarpus siliculosus var. hiemalis (P.L. Crouan & H.M. Crouan) Foslie Feldmannophycus okamurae (Yamada) Mineur, Maggs & Verlaque *Ganonema farinosum (J.V. Lamouroux) K.C. Fan & Yung C. Wang *Gracilaria arcuata Zanardini Grateloupia minima P.L. Crouan & H.M. Crouan4 Lomentaria flaccida Tanaka Microspongium tenuissimum (Hauck) A.F. Peters5 Nemalion vermiculare Suringar *Pilayella littoralis (Linnaeus) Kjellman CHL RHO RHO CHL CHR CHR CHR CHR CHR RHO RHO RHO RHO RHO CHR RHO CHR Establishment Source success E Israel: VERLAQUE et al., in press E Tunisia: VERLAQUE et al., in press E France: VERLAQUE et al., in press Greece: SARTONI & BODDI, 2002 France: VERLAQUE et al., 2007 Q Q E CRf E E CRr/Q E France: VERLAQUE et al., in press France: VERLAQUE et al., in press Spain: RULL LLUCH et al., 2007 France: VERLAQUE et al., in press Italy: VERLAQUE et al., in press CRf France: VERLAQUE et al., in press Egypt: VERLAQUE et al.,in press Q Q Tunisia: TASKIN et al., 2008 France: VERLAQUE et al., 2007 C Q France: VERLAQUE et al., 2007 France: VERLAQUE et al., 2007 E C France: VERLAQUE et al., in press Italy: VERLAQUE et al., in press (continued) Introduced in the Thau Lagoon probably confused with C. codii 3 the taxonomy of the genus Cladophora Kützing is so complicated that the confirmation of the identity of this macrophyte requires further investigation. 4 The determination of the alien or native status of Mediterranean populations requires further investigations. 5 The determination of the Atlantic or Mediterranean origin of the Thau populations requires further investigations 1 2 Medit. Mar. Sci., 9/1, 2008, 119-165 125 Table 3.1 (continued) Taxon Establishment Source success RHO *Polysiphonia fucoides (Hudson) E France: VERLAQUE et al., in Greville press Q France: VERLAQUE et al., 2007 RHO Polysiphonia stricta (Dillwyn) Greville6 CHR *Punctaria tenuissima E France: VERLAQUE et al., in (C. Agardh Greville press CHR *Spatoglossum variabile Figari Q Israel: VERLAQUE et al., in press & De Notaris CHL *Ulva fasciata Delile E Egypt: VERLAQUE et al., in press Change in establishment success RHO Antithamnionella sublittoralis From Q to E VERLAQUE et al., in press (Setchell & Gardner) Athanasiadis RHO Antithamnionella ternifolia From C to E VERLAQUE et al., in press (J.D. Hooker & Harvey) Lyle CHL Caulerpa mexicana Sonder ex Kützing From C to E VERLAQUE et al., in press CHL Ceramium strobiliforme G.W. From C to E VERLAQUE et al., in press Lawson & D.M. John CHL Cladophora patentiramea From Q to E VERLAQUE et al., in press (Montagne) Kützing RHO Dasya sessilis Yamada From C to E VERLAQUE et al., in press RHO Dasysiphonia sp. From C to E VERLAQUE et al., in press RHO Goniotrichiopsis sublittoralis From Q to E VERLAQUE et al., in press G.M. Smith RHO Grateloupia patens (Okamura) From E to C VERLAQUE et al., in press Kawaguchi & Wang RHO Laurencia caduciramulosa From Q to E VERLAQUE et al., in press Masuda & Kawaguchi CHR Padina boryana Thivy in W.R. Taylor From C to E VERLAQUE et al., in press RHO Plocamium secundatum (Kützing) From C to E Spain: RODR GUEZ PRIETO, Kützing unpublished RHO Polysiphonia atlantica Kapraun From Q to E VERLAQUE et al., in press & J.N. Norris RHO Porphyra yezoensis Ueda From C to E VERLAQUE et al., in press RHO Solieria dura (Zanardini) F. Schmitz From E to C VERLAQUE et al., in press CHR Sphaerotrichia firma (Gepp) From C to E Occurrence in Levantine, Aegean, A.D.Zinova Marmara, France RHO Symphyocladia marchantioides From C to E VERLAQUE et al., in press (Harvey) Falkenberg 6 New species The determination of the Atlantic or Mediterranean origin of the Thau populations requires further investigation 126 Medit. Mar. Sci., 9/1, 2008, 119-165 Lagoon (VERLAQUE et al., 2007). Ulva ohnoi, recorded in ballast water (Italy: FLAGELLA et al., 2007) is not included in the CIESM atlas. Concerning micro-phytobenthos, there have been few studies of the benthic/epiphytic species such as Ostreopsis lenticularis, O. siamensis, O. ovata in the Mediterranean compared to other regions and these species have simply been recorded when studies were carried out (i.e. MONTI et al., 2007). According to GOMEZ (2008) these new entries should not be considered as recent species introductions in the Mediterranean Sea. However, in the absence of evidence we maintain them as aliens. With the addition of the three afore- mentioned species, thirty four (34) new phytobenthos species have been added to the ‘annotated list’. In the light of new data, changes have occurred in establishment success. Eleven of the previous casual records such as Plocamium secundatum (RODR GUEZ PRIETO, unpublished) are now classified as established (Table 3.1). On the other hand, eight (8) of the questionable species reported in the ‘annotated list’and two of the ‘established’ are excluded in this work (Table 3.2). Additional species that are considered as non-aliens in the CIESM Atlas (VERLAQUE et al., in press) are listed in Table 3.2. Name changes are presented in Table 3.3. Table 3.2 Taxa excluded compared to the ‘annotated list’. + reported as questionable in the ‘annotated list’ reported as established in the ‘annotated list’ ++ Taxon RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO RHO CHL Species Reasoning Acanthophora muscoides (Linnaeus) Bory de Saint-Vincent Unsupported records Acrochaetium balticum (Rosenvinge) Aleem & Schulz Mistake Antithamnion densum (Suhr) M.A. Howe Mistake Antithamnion ogdeniae Abbott Indigenous species (A. decipiens) ++ Chondria collinsiana M.A. Howe Unsupported records ++ Chondria polyrhiza F.S. Collins & Hervey Unsupported records Ceramium giacconei Cormaci & G. Furnari Indigenous species Ceramium graecum Lazaridou & Boudouresque Indigenous species Chondrus crispus Stackhouse Misidentification Gracilaria armata (C. Agardh) Greville Indigenous species + Hypnea variabilis Okamura Unsupported records + Laurencia chondrioides Boergesen Indigenous species + Laurencia intricata J.V. Lamouroux Indigenous species Laurencia japonica Yamada Mistake + Laurencia majuscula (Harvey) A.H.S. Lucas Indigenous species Laurencia microcladia Kützing Indigenous species + Parvocaulis parvulus (Solms-Laubach) S. Berger, Indigenous species U. Fettweiss, S. Gleissberg, L. B. Liddle, U. Richter, H. Sawitsky & G.C. Zuccarello + (continued) Medit. Mar. Sci., 9/1, 2008, 119-165 127 Table 3.2 (continued) Taxon RHO RHO RHO CHR CHR Species + Polysiphonia kampsaxii Boergesen Polystrata fosliei (Weber-van Bosse) Denizot Pterothamnion simile (J.D. Hooker & Harvey) Nägeli Rosenvingea intricata (J. Agardh) B rgesen + Sargassum latifolium (Turner) C. Agardh Reasoning Unsupported records Indigenous species Mistake Indigenous species Unsupported records Table 3.3 Name changes in macroalgae. New name Acrochaetium robustum B rgesen Acrochaetium spathoglossi B rgesen Acrochaetium subseriatum B rgesen Anotrichium okamurae Baldock Old name Audouinella robusta; reported as Acrochaetium sargassicola Audouinella spathoglossi, reported as Kylinia spathoglossi Audouinella subseriata Anotrichium furcellatum (J. Agardh) Baldock) misreported to A. pectinatum Antithamnion nipponicum Yamada & Inagak Botryella parva (Takamatsu) Kim Sorocarpus sp Dasysiphonia sp. reported as Heterosiphonia japonica Feldmannophycus okamurae reported as F. rayssiae (Yamada) Mineur, Maggs & Verlaque Grateloupia subpectinata G. luxurians Hypnea flagelliformis Greville misidentified as Hypnea spicifera ex J. Agardh (Suhr) Harvey Neosiphonia harveyi (J. Bailey) Polysiphonia harveyi J. Bailey M.-S. Kim, H.-G. Choi, Guiry & G.W. Saunders Cladophora herpestica Cladophoropsis javanica (Montagne) Kützing (Kützing) P.C. Silva Ulvaria obscura (Kützing) Gayral Monostroma obscurum (Kützing) J. Agardh Saccharina japonica (Areschoug) Laminaria japonica Areschoug C.E. Lane, C. Mayes, Druehl & G.W. Saunders 128 Source VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., 2005 VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press VERLAQUE et al., in press Medit. Mar. Sci., 9/1, 2008, 119-165 4. Zoobenthos/Polychaeta After the publication by ZENETOS et al. (2006), 10 additional alien polychaete species were reported from the Mediterranean (Table 4). Chaetozone corona, Pseudopolydora paucibranchiata and Timarete punctata were found along the Aegean and Levantine coasts of Turkey ( INAR & ERGEN, 2007; INAR, 2007; DAGLI & INAR, 2008); Syllis bella, Exogone breviantennata and Syllis cf. mayeri from the Lebanon coast (Levantine Sea) (AGUADO & SANMART N, 2007); Novafabricia infratorquata from the Mediterranean coast of Spain and Italy (BICK, 2005; LICCIANO & GIAN- GRANDE 2006); Hesionura serrata and Erinaceusyllis serratosetosa from Spain (CARDELL & MENDEZ, 1996; SAN MART N, 2003) and Megalomma claparedei from Italy (GIANGRANDE & LICCIANO, 2008). Among the species, only T. punctata, E. breviantennata and Megalomma claparedei are Lessepsian immigrants; the others have probably been introduced into the area by shipping. The spionid worm, Pseudopolydora paucibranchiata, which was previously misidentified as P. antennata ( INAR et al., 2006c), invaded the polluted soft bottom of the inner part of Izmir Bay, reaching to a population density of 6180 ind.m-2 in June 2004 (DAGLI & INAR, 2008). Table 4 New alien polychaeta species in the Mediterranean and changes in establishment success. Establishment success: C=casual; E=established; Q=questionable; CRf=cryptogenic frequent New species Establishment success Chaetozone corona Berkeley & Berkeley 1941 CRf Pseudopolydora paucibranchiata Okuda 1937 E Syllis bella (Chamberlin, 1919) E Hesionura serrata (Hartmann-Schröder, 1960) C Timarete punctata (Grube, 1859) Exogone breviantennata HartmannSchröder, 1959 Erinaceusyllis serratosetosa (HartmannSchröder, 1982) Megalomma claparedei Gravier, 1908 E E C Novafabricia infratorquata (Fitzhugh, 1983) Syllis cf. mayeri Musco et Giangrande, 20051 E Q Change in establishment success Epidiopatra hupferiana Augener, 1918 Isolda pulchella Müller, 1858 1 C From Q to C From Q to C Source Turkey: INAR & ERGEN, 2007 Turkey: DAGLI & INAR, 2008 Lebanon: AGUADO & SAN MARTIN, 2007 Spain: CARDELL & MENDEZ, 1996 Turkey: INAR, 2007 Lebanon: AGUADO & SAN MARTIN, 2007 Spain: SAN MART N, 2003 Italy: GIANGRANDE & LICCIANO, 2008 Spain: BICK, 2005 Lebanon: AGUADO & SAN MARTIN, 2007 Italy: Cantone pers. commu Italy: Cantone pers. commu First description of species. Taxonomic posision is questionable Medit. Mar. Sci., 9/1, 2008, 119-165 129 According to INAR (2007), the previous report of Cirriformia semicincta (Ehlers, 1905) from the Levantine coast by LAUBIER (1966), might in fact belong to Timarete punctata. Linopherus incarunculata (Peters, 1858) reported by GALIL, (2007) is excluded as it is considered as misspelling of Linopherus acarunculata, (BEN ELIAHU, 1976). Although there have been many attempts, the Mediterranean polychaete fauna is largely unknown. Therefore, the species that are distributed in the mid-eastern Atlantic cannot be considered as alien species, as they might have been previously overlooked or misidentified in the region. The minute syllid species are a good example. The recent great efforts to understand the syllid fauna in the Mediterranean have doubled the number of species that we had known in the region. For example, Erinaceusyllis belizensis and Syllides bansei, both of Caribbean Atlantic origin, were first reported from the Mediterranean coast of Spain (OLANO et al., 1998; ALOS, 1984) and then subsequently from the other coasts of the Mediterranean, including the Aegean Sea ( INAR & ERGEN, 2002; INAR, 2003). Similarly, representatives of other families of tropical Atlantic origin, such as Neanthes agulhana, were recorded on the Cadiz Coast and Malaga (SARDA, 1985) and recently found along the Italian Coast (Ionian Sea GIANGRANDE unpublished data), and Paranaitis wahlbergi, of north Atlantic origin, were recorded in Ibiza (VIETEZ et al., 2004). 5. Zoobenthos/Crustacea Based on CORBERA (1994), who cast doubt on the origin of Iphinoe crassipes haifae because the species was present off the north-eastern coast of the 130 Iberian Peninsula, this questionable species has been excluded from our list. On the other hand, Penaeopsis serrata, excluded in the ‘annotated list’, is added here. It is argued that it had probably not been discovered earlier due to its deepwater habitat (MURA et al., 2003). Six benthic copepods (Canuellina insignis, Enhydrosoma hopkinsi, Robertsonia salsa, Scottolana longipes, Stenhelia inopinata, Stenhelia minuta) reported in the Bardawil Lagoon (POR, 1964; 1972) that were erroneously placed among zooplankton in the ‘annotated list’ are also added to the zoobenthos. Altogether, research has a) brought to light 10 new alien species (Table 5), belonging mostly to Decapoda and b) expanded the distribution of some species considered as casual so far, thus leading to their classification as established. Two of these new species, namely Eocuma rosea and Urocaridella pulchella are new to science (CORBERA & GALIL, 2007; YOKE & GALIL, 2006b). However, diagnostic characters place them with other species of the same genera known from the Indo-Pacific Ocean and it is thus suggested that they entered the Mediterranean through the Suez Canal (CORBERA & GALIL, 2007; YOKE & GALIL, 2006b). 6. Zoobenthos /Mollusca Mollusca is one of the best studied groups (ZENETOS et al., 2004) with significant contribution among alien biota in the Mediterranean (ZENETOS, 2008). The number of molluscan alien species recorded by December 2005 was estimated to be 196 (ZENETOS et al., 2006). The rate of increase, compared to the 139 reported by GOFAS & ZENETOS (2003), was attributed to the increased Medit. Mar. Sci., 9/1, 2008, 119-165 Table 5 New alien crustacean species in the Mediterranean and changes in establishment success. * species excluded in the ‘annotated list’. Taxon: AMP=Amphipoda; CIR=Cirripedia; CUM=Cumacea; DEC=Decapoda; ISO=Isopoda; STO=Stomatopoda. Establishment success: C=casual; E=established; Q=questionable Taxon New species DEC Charybdis feriata (Linnaeus, 1758) STO Clorida albolitura Ahyong & Naiyanetr, 2000 Eocuma rosae Corbera & Galil, 20071 Fenneropenaeus merguiensis (De Man, 1888) Grapsus granulosus H. MilneEdwards, 1853 *Penaeopsis serrata Bate, 1881 Scherocumella gurneyi (Calman, 1927) Sirpus monodi Gordon, 1953 CUM DEC DEC DEC CUM DEC Establishment Source success C Spain: ABELL & HISPANO, 2006 C Israel: AHYONG & GALIL, 2006 CIR DEC 2 Israel: CORBERA & GALIL, 2007 C Turkey: ÖZCAN et al., 2006 C Tunisia: ZAOUALI et al., 2007a Q C Sardinian Channel: MURA et al., 2003 Israel: CORBERA & GALIL, 2007 C Greece: PANCUCCI-PAPADOPOULOU & NALETAKI, 2007 Tunisia: ZAOUALI et al., 2007b Turkey: YOKE & GALIL, 2006b Tetraclita squamosa Pilsbry, 1916 Urocaridella pulchella Yoke & Galil, 20062 Change in establishment success ISO Apanthura sandalensis Stebbing, 1900 DEC Dorippe quadridens (Fabricius, 1793) From C to E From E to C AMP From C to E Israel: NEGOESCU, 1981 Record based on a single animal: GALIL, 2005. Turkey: BAKIR et al., 2007 From C to E From C to E Turkey: INAR et al., 2008 Tunisia: ZAOUALI et al., 2007a ISO DEC 1 Q Gammaropsis togoensis (Schellenberg, 1925) Paradella dianae Menzies, 1962 Plagusia squamosa (Herbst, 1790) C E First description of the species First description of the species interest of malacologists and the relatively easy collection/identification of mollusca. However, 31 species were classified as questionable. The questionable taxa were reported mostly from the Israeli coast; the records were based on a single specimen or a few empty shells. In the 2006-2007 Medit. Mar. Sci., 9/1, 2008, 119-165 period, nine of them (Table 6.1) were reported as casual in other areas or even as having become established, such as Chama aspera (MIFSUD & OVALIS, 2007; OVALIS & ZENETOS, 2007) or Nanostrea exigua (MIENIS, 2008). The fact is that some of these species are present in collec131 Table 6.1 New alien molluscan species in the Mediterranean and changes in establishment success. *Species excluded in the ‘annotated list’ Establishment success: C=casual; E=established; Q=questionable; New species Acanthopleura gemmata (de Blainville, 1825) Anadara granosa (Linnaeus, 1758) Atys angustatus Smith, 1872 Establishment success Q C E Source Bractechlamys vexillum (Reeve, 1853) Cerithidium perparvulum (Watson 1886) Chrysallida micronana Öztürk & van Aartsen, 2006 Circe scripta (Linnaeus, 1758) *Conus arenatus arenatus Hwass, 1792 Conus inscriptus Reeve, 1843 Conus rattus Hwass, 1792 Mytilopsis sallei Recluz, 1849 Nassa situla (Reeve, 1846) Nassarius concinnus (Powys, 1835) Nassarius stolatus (Gmelin, 1791) Parviturbo dibellai Buzzurro & Celalupo, 2006 C C C C C C C C C *Petricola hemprichi (Issel, 1869) C Sepia gibba Ehrenberg, 1831 Sepia pharaonis Ehrenberg, 1831 C Q Strombus vittatus vittatus Linnaeus, 1758 Trivirostra triticum Schilder, 1932 Change in establishment success Acteocina crithodes Melvill & Standen, 1907 Angulus flacca (Roemer, 1871) Atys cylindricus (Helbling, 1779) Chama aspera Reeve, 1846 Chama asperella Lamarck, 1819 Chromodoris quadricolor (Rüppell & Leuckart, 1828) Ergalatax contracta (Reeve, 1846) Ethminolia hemprichi (Issel, 1869) C C Libya: ZAOUALI et al., 2007b Greece: YOUNG et al., 2007 Turkey: VAN AARTSEN & GOUD, 2006a France: WGITMO, 2006 Israel: BOGI & GALIL, 2006 Turkey: ÖZTÜRK & VAN AARTSEN, 2006 Greece: YOUNG et al., 2007 Israel: MIENIS,2008 Greece: YOUNG et al., 2007 Israel: MIENIS,2008 Egypt: HOFFMAN et al., 2006 Israel: MIENIS,2008 Israel: MIENIS,2008 Greece: YOUNG et al., 2007 Turkey: BUZZURRO & CELALUPO, 2006 Turkey: CEVIKER & ALBAYRAK S., 2006 Israel: MIENIS, 2008 Known only from cuttlebones: MIENIS 2003a Turkey: KABASAKAL et al., 2005 Israel: MIENIS, 2008 From Q to C From Q to C From Q to C From C to E From Q to E From C to E Turkey: MIENIS, 2004 Israel: MIENIS, 2004 Israel: MIENIS, 2004 Turkey: MIFSUD & OVALIS, 2007 Greece: OVALIS & ZENETOS, 2007 Turkey: ÖZTÜRK & CAN, 2006. From E to C From Q to C R. HOUART, pers. Commun. MIENIS, 2004 Great Bitter Lake, HOFFMAN et al., 2006 C E C (continued) 132 Medit. Mar. Sci., 9/1, 2008, 119-165 Table 6.1 (continued) New species Nanostrea exigua Harry, 1985 Nerita sanguinolenta Menke, 1829 Pinctada margaritifera (Linnaeus, 1758) Rhinoclavis sinensis (Gmelin, 1791) Rissoina ambigua (Gould, 1849) Septifer bilocularis (Linnaeus, 1758) Establishment success From Q to E From C to E From E to C From Q to C From Q to C From Q to C tions from the Mediterranean and are frequently reported in periodicals of Malacological Societies (ARION, BASTERIA, GLORIA MARIS, HALIOTIS, LEVANTINA, NAUTILUS, NEPTUNEA, NOVAPEX, SPIRULA, TRITON, etc.). The ‘annotated list’ has been updated based on contributions that include summaries of findings of invasive molluscs in the Mediterranean, such as MIENIS (2004; 2008) and DELONGUEVILLE & SCAILLET (2007), or just sparse records of new alien species (MIENIS, 2003a, 2003b, 2005). From the latest publications it appears that the centre of bio-invasions, which used to be the Israeli coast, is spreading northward. Most of the new records refer to the Levantine coast of Turkey (Table 6.1) where more alien mollusca are met, including species excluded in the CIESM Atlas (ZENETOS et al., 2004) such as Petricola hemprichi (CEVIKER & ALBAYRAK, 2006) and Conus arenatus arenatus (MIENIS, 2008). Newly-established species in areas along the coast of Turkey include Chromodoris quadricolor (CAN, 2006) and Nerita sanguinolenta (MUTAF et al., 2007). Eight new species are reported as occasional new visitors along the Mediterranean coast of Israel (Table 6.1) including a variation of the well established gastropod Medit. Mar. Sci., 9/1, 2008, 119-165 Source Israel: MIENIS,2008 Turkey: MUTAF et al., 2007 MIENIS, 2004 MIENIS, 2004 MIENIS, 2004 Turkey: ALBAYRAK & CALGAR, 2006 Rhinoclavis kochi var. recurva and the rare Planaxis griseum (MIENIS, 2008). Another hotspot area for alien biota is the Saronikos Gulf. The presence of four alien molluscan species (shells only), new to the Mediterranean, is attributed to shipping in combination with climatic change (YOUNG et al., 2007). Meanwhile taxonomic studies have led to nomenclature changes of several species (Table 6.2). There are now approximately 216 alien mollusca in the Mediterranean twenty (20) new molluscs since the ‘annotated list’. 7. Zoobenthos /Miscellanea Twenty (20) new species are added to the ‘annotated list’; the main invading groups are echinodermata, ascidia and bryozoa (Table 7). Most of the new species are fouling organisms reported from port areas. Echinodermata: The number of alien echinodermata has doubled with the present records from 5 to 11. Four of the newcomers were reported in the 2006-2007 period while Prionocidaris babulosa and Asterias rubens were reported earlier (Table 7). Asterias rubens is a well-established species in the Sea of Marmara, known since 1990 (YUCE & SADLER, 133 Table 6.2 Name changes in mollusca. New name Cerithidium diplax (Watson, 1886) Cyrnola lendix (A. Adams 1863) Old name Clathrofenella ferruginea (A. Adams, 1860) Styloptygma beatrix Melvill, 1911 VAN AARTSEN & GOUD, 2006b Diplodonta subrotunda Issel, 1869 VAN AARTSEN, 2004 Diplodonta bogii Van Aartsen, 2004 Ergalatax junionae Houart, 2008 Ergalatax obscura Houart, 1996 Melibe viridis (Kelaart, 1858) Melibe fimbriata Alder & Hancock, 1864 Monotigma lauta Adelacteon fulvus (A. Adams, 1853) (A. Adams, 1851) Leucotina natalensis Smith, 1910 Adelacteon amoenus (A. Adams, 1851) Timoclea marica Timoclea maurica (Linnaeus, 1758) 2000). MORTENSEN (1927) reported that it was found accidentally in the Mediterranean but did not specify a locality. ALBAYRAK, (1996) reports it from the Bosphorus Straits. The suggestion that A. rubens was introduced by shipping (ZIBROWIUS, 2002) is reasonable. Its planktotrophic larvae may have been transferred in the ballast waters on ships. Other ship-mediated introductions include Eucidaris tribuloides (TANTI & SCHEMBRI, 2006), Prionocidaris babulosa (SCHEMBRI, 1978), while there remain the introductions of Diadema setosum, and Ophiocoma scolopendrina (YOKE & GALIL, 2006a; ZAOUALI et al., 2007b) presumably via the Suez Canal. Bryozoa: Three new bryozoan species are reported, all three presumably shiptransfered and reported close to harbour areas. Bowerbankia gracillima of Atlantic origin, reported from Lazio, Italy (D’HONDT & CHIMENZ GUSSO, 2006); Celleporaria brunnea and Cellepo134 Source VAN AARTSEN, 2006 HOUART, 2008 GOSLINER & SMITH, 2003 VAN AARTSEN & HORI, 2006 VAN AARTSEN & HORI, 2006 Spelling mistake in ZENETOS et al., 2006 raria pilaefera, of Indo-Pacific origin, reported near Alsancak Harbour, Aegean Turkey and Rinella, Malta, respectively (KOCAK, 2007; AGIUS et al., 1977). Hydrozoa: Two species are added to the ‘annotated list’ namely: Eudendrium carneum, which has been recorded from the western Mediterranean in recent years (GILI, 1986; BAVESTRELLO & PIRAINO, 1991) and Sertularia marginata found in the eastern Mediterranean (PICARD, 1958). Ascidiacea: Six new ascidians are reported. Of them, Styela clava is classified among the top worst invasive IAS in Europe (EEA, 2007b). The proximity of commercial shell fisheries to the discovered populations in S. France, and the absence of S. clava from other harbours and marinas along the coast, suggests that the species may have been introduced by shellfish transfer (DAVIS & DAVIS, 2008). Several colonies of Distaplia bermudensis were observed from the Taranto Seas (Mar PicMedit. Mar. Sci., 9/1, 2008, 119-165 Table 7 New alien zoobenthic species (Miscellanea taxa) in the Mediterranean and changes in establishment success. Taxon: ECH=Echinodermata; BRY=Bryozoa; HYD=Hydrozoa; ASC=Ascidiacea; SIP=Sipuncula; PLA=Platyelminthes. Establishment success: C=casual; E=established; Q=questionable; CR=Cryptogenic Taxon ECH ECH ECH ECH ECH ECH BRY BRY BRY HYD HYD ASC ASC ASC ASC ASC ASC ASC POR SIP PLA New species Establishment success Acanthaster planci (Linnaeus, 1758) C Asterias rubens Linnaeus, 1755 E Diadema setosum (Leske, 1778) C Eucidaris tribuloides (Lamarck, 1816) E Ophiocoma scolopendrina (Lamarck, 1816) Prionocidaris babulosa (Lamarck, 1816) Bowerbankia gracillima Hinks, 1880 Celleporaria brunnea (Hincks, 1884) Celleporaria pilaefera (Canu & Bassler, 1929) Eudendrium carneum Clarke, 1882 Sertularia marginata (Kirchenpauer, 1864) Botrylloides violaceus Oka,1927 Cystodytes philippinensis Herdman 1886 Distaplia bermudensis Van Name, 1902 Ecteinscidia styeloides (Traustedt, 1882) Perophora multiclathrata (Sluiter,1904) Styela clava Herdman, 1882 Botrylloides violaceus Oka, 1927 Paraleucilla magna Klautau et al., 2004 Apionsoma misakianum Ikeda, 19041 Boninia neotethydis Curini-Galletti & Campus, 2007 Source C France: WGITMO, 2006 Turkey: YUCE & SADLER, 2000 Turkey: YOKE_ & GALIL, 2006a Malta:TANTI & SCHEMBRI, 2006 Libya: ZAOUALI et al., 2007b C Malta: SCHEMBRI, 1978 C C Q Italy: D’HONDT & CHIMENZ GUSSO, 2006 Turkey: KOCAK, 2007 Malta: AGIUS et al., 1977 E C Spain: GILI, 1986 Lebanon: PICARD, 1958 E CR E E E E E Italy: ZANIOLO et al .,1998 MELIANE, 2002 Spain: PERES, 1957 Italy: MASTROTOTARO & BRUNETTI, 2006 MASTROTOTARO & TURSI, 2006 MONNIOT, 1983 France: DAVIS & DAVIS, 2008 Italy: ZANIOLO et al., 1998 Italy: LONGO et al., 2007 Q Turkey: A IK, 2007 E Israel: CURINI-GALLETTI & CAMPUS, 2007 E (continued) 1 Origin uncertain Medit. Mar. Sci., 9/1, 2008, 119-165 135 Table 7 (continued) Taxon New species Establishment Source success Change in establishment success SIP Aspidosiphon elegans (Chamisso & Eysenhardt, 1821) ASC Microcosmus squamifer Michaelsen, 1927 ASC Eusynstyela hartmeyeri Michaelsen, 1904 ASC Ascidia cannelata Oken,1820 colo and Mar Grande) since 2000 (MASTROTOTARO & BRUNETTI, 2006). Botryllus schlosseri, a cosmopolitan species normally present in the Mediterranean Sea, was erroneously cited as established in the ‘annotated list’. In contrast, Botrylloides violaceus, native to Northwest Pacific, is recorded from the Venice Lagoon in Italy (ZANIOLO et al., 1998). Perophora multiclathrata and Ecteinascidia styeloides have only been found in harbours (MONNIOT, 1983, MASTROTOTARO & TURSI 2006) while Cystodytes philippinensis is a cryptogenic species probably confused with C. dellechiajei (MELIANE, 2002). Symplegma viride, a casual species in the ‘annotated list’ is excluded as synonym of Symplegma brakenhielmi (IZQUIERDO-MUNOZ et al., in press). Ascidia savigny is also excluded as a misidentification of Ascidia cannelata (A. Ramos-Espla, pers. commun.). Porifera: Current research on the Lebanese coast has brought to life five species that are new for the Mediterranean but which are widely distributed in the tropics. However, these species do not appear to be Lessepsian immigrants, but are interpreted as remnants of an ancient 136 From Q to E Turkey: A IK, 2008 From Q to E TURON et al., 2007 From C to Q IZQUIERDO-MUNOZ et al., in press IZQUIERDO-MUNOZ et al., in press From C to E thermophilous fauna that survived in the easternmost part of the Mediterranean (VACELET et al., 2007). On the other hand, the calcareous sponge Paraleucilla magna has been detected at different Mediterranean sites (Taranto, Porto Cesareo, Brindisi and Naples). Its record in well-studied areas where several benthic surveys have previously been carried out suggests a recent introduction of the species into the Mediterranean Sea (LONGO et al., 2007). Sipuncula: Aspidosiphon elegans, a rare species to date, has recently established viable populations on the Levantine coast of Turkey (A IK, 2008). The status of a second Apionsoma species A. (A.) misakianum, though widely distributed in the Aegean Sea (not confined to a recipient area such as harbours or near canals) was most probably previously confused with the other Apionsoma species in the Mediterranean Sea. The previous reports of Apionsoma species should be re-examined to determine the real distributional boundary of A. (A.) misakianum within the Mediterranean basins (A IK, 2007). Platyelminthes: Based on specimens Medit. Mar. Sci., 9/1, 2008, 119-165 collected in the Mediterranean and the Red Sea coasts of Israel, a new Lessepsian flatworm (Boninia neotethydis) was described (CURINI-GALLETTI & CAMPUS, 2007). An Indo-Pacific origin of the species is supported by the history of its records. In 1998 the species was common in the Mediterranean, particularly in shelly, very coarse sediments at the bottom of rocky pools, and among clumps of the mussel Brachidontes pharaonis. 8. Zoobenthos /Foraminifera In the Mediterranean, very little attention was paid to alien foraminifera until the early 1990s. Since then, they have been reported from many localities such as the Adriatic Sea (CIMERMAN & LANGER, 1991), Italy: the Tyrrhenian Sea (SGARRELLA & MONCHARMONTZEI, 1993), Israel (HYAMS et al., 2002), Egypt (SAMIR et al., 2003), Greece (DEBENAY et al., 2005; TRIANTAPHYLLOU et al., 2005) and the Maltese Islands (YOKE et al., 2007). The greatest amount of information however, comes from the Turkish coast. Recent studies have revealed the presence of many alien benthic foraminifera species on the southern and western Mediterranean coasts of Turkey (MERI & AV AR, 2001; MERI et al., 2002 a, b, c; 2003 a, b; 2004 a, b), and in the Dardanelles (MERI et al., 2008a) and the Sea of Marmara (KAMINSKI et al., 2002; MERI et al., 2005, 2007). In the ‘annotated list’ only seven species have been reported with the authors acknowledging the difficulty in properly documenting the records of foraminiferans in the area and identifying the need for an update. Table 8 summarizes information on alien foraminifera Medit. Mar. Sci., 9/1, 2008, 119-165 recorded in the Mediterranean, also providing the geographic range they have been reported from at country level. A summary of the Turkish alien foraminifera is provided in MERI et al, 2008d. Most of the alien foraminifer species observed in the Mediterranean Sea are of Indo-Pacific origin presumably introduced via the Suez Canal. The most abundant species are Sorites orbiculus and Amphistegina lobifera. Some researchers reported the presence of Amphistegina in Haifa-Israel, Greece and the Gulf of Gabes without indicating a specific species (LANGER & HOTTINGER, 2000). Iridia diaphana, which is rarely observed in the Northern Aegean Sea, is suggested as being of Atlantic origin and widely distributed in the Mediterranean (LOEBLICH & TAPAN, 1988). The species Pulleniatina obliquiloculata occurs both in the Atlantic and IndoPacific; however, its absence from the Red Sea fauna suggests that it has been introduced and has spread into the Mediterranean from the Atlantic via Gibraltar. In contrast, the species Cushmanina striatopunctata, also known to have a wide distribution in the Atlantic and in the Indo-Pacific (HOTTINGER et al., 1993; LOEBLICH & TAPAN, 1994), is absent from the western Mediterranean, which suggests that the eastern Mediterranean population has an Indo-Pacific origin. Thus, it might have been introduced from the Red Sea and settled in the Dardanelles Straits. Like Agglutinella and Amphistegina spp., it may also establish a population in the Sea of Marmara. In total, 45 alien foraminiferan species (49 including 4 identified at the genera level) are found in the Mediterranean. 32 genera and 42 species are suggested to have an Indo-Pacific origin, whereas, only 2 genera and 2 species are of Atlantic origin. 137 Table 8 Alien Foraminiferan species in the Mediterranean. + species reported in the ‘annotated list’. Establishment success: C=casual; R=Established/Rare; F=Established/Frequent Species Acervulina inhaerens Schultze Agglutinella arenata (Said) Agglutinella compressa El-Nakhal Agglutinella robusta El-Nakhal Agglutinella soriformis El-Nakhal + Amphisorus hemprichii Ehrenberg Establishment success R C R R R F Amphistegina lessonii D’Orbigny R + Amphistegina lobifera Larsen F Amphistegina madagascariensis (D’Orbigny) Articulina alticostata Cushman + Astacolus insolithus (Schwager) + Astacolus sublegumen (Parr) Borelis sp Clavulina angularis D’Orbigny C C C C R R Clavulina cf. multicamerata Chapman Cushmanina striatopunctata (Parker & Jones) Cyclorbiculina compressa (D’Orbigny) Cymbaloporetta plana (Cushman) Cymbaloporetta squammosa (D’Orbigny) Edentostomina cultrata (Brady) R R C R R R Elphidium cf. charlottense (Vella) Elphidium striatopunctatum (Fichtel & Moll) R R Entosigmomorphina sp Euuvigerina sp Haddonia sp Hauerina diversa Cushman R R F F Heterocyclina tuberculata (Mobius) R Source Turkey: MERI et al., 2004 a Turkey: KAMINSKI et al., 2002 Egypt: SAMIR et al., 2003 Egypt: SAMIR et al., 2003 Egypt: SAMIR et al., 2003 YANKO, 1995; Egypt: SAMIR et al., 2003; Turkey: MERI et al., 2008c Greece: HOLLAUS & HOTTINGER, 1997; Israel: HYAMS et al., 2002 Israel: YANKO et al., 1993; Turkey: MERI & AV AR, 2001; Malta: YOKE et al., 2007 Egypt: SAMIR et al., 2003 Turkey: MERI et al., 2004a Turkey: MERI et al., 2004a Turkey: MERI et al., 2004a Israel: HYAMS et al., 2002 Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001 Turkey: MERI et al., 2008c Turkey: MERI et al., 2008b Turkey: MERI et al., 2008c Turkey: MERI et al., 2008c Turkey: MERI et al., 2008c Israel: YANKO et al., 1993; Turkey: MERI et al., 2004a Turkey: MERI et al., 2008c Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001 Turkey: MERI et al., 2008c Turkey: MERI et al., 2004b Turkey: MERI et al., 2008c Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001; Israel: YANKO, 1995; Turkey: AV AR et al., 2001 (continued) 138 Medit. Mar. Sci., 9/1, 2008, 119-165 Table 8 (continued) Species Establishment Source success + Heterostegina depressa D’Orbigny F Italy: MONCHARMONT-ZE , 1968; Greece: MORARIU & HOTTINGER, 1988; Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001 Iridia diaphana Heron-Allen & Earland F Turkey: MERI et al., 2008a Miliolinella cf. hybrida (Terquem) R Turkey: MERI et al., 2008c Nodopthalmidium antillarum (Cushman) F Israel: YANKO, 1995; Turkey: MERI et al., 2008c Operculina ammonoides (Gronovius) C Israel: YANKO, 1995 Peneroplis antillarum D’Orbigny C Israel: HYAMS et al., 2002 Peneroplis arietinus (Batsch) F Turkey: MERI et al., 2008c + Planogypsina acervalis (Brady) R Turkey: MERI et al., 2004a + Planogypsina squamiformis (Chapman) R Turkey: MERI et al., 2004a Planorbulinella larvata (Parker & Jones) C Israel: YANKO, 1995 Pseudomassilina reticulata R Israel: YANKO et al., 1993; (Heron-Allen & Earland) Turkey: AV AR et al., 2001 Pulleniatina obliquiloculata (Parker & Jones) R Turkey: MERI et al., 2004b Pyramidulina catesbyi (D’Orbigny) R Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001 Pyramidulina perversa (Schwager) R Turkey: MERI et al., 2008c Pyrgo denticulata (Brady) R Israel : YANKO et al., 1993; Egypt: SAMIR et al., 2003; Turkey: MERI et al., 2008c Quinqueloculina cf. mosharrafai Said R Turkey: MERI et al., 2008c Schlumbergerina alveoliniformis (Brady) R Turkey: MERI et al., 2008c Sorites orbiculus Ehrenberg F Italy: HOFKER, 1930, Greece: CHERIF, 1970; France: BLANC VERNET et al., 1979; Italy: CRAPON-DE CAPRONA & BENIER, 1985; Adriatic: CIMERMAN & LANGER, 1991; Israel: YANKO, 1995; Turkey: AV AR et al., 2001; Egypt: SAMIR et al., 2003 Sorites variabilis Lacroix F Israel: YANKO et al., 1993; Turkey: MERI et al., 2008c Spiroloculina angulata Cushman F Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001 Spiroloculina antillarum D’Orbigny F Israel: YANKO et al., 1993; Turkey: AV AR et al., 2001; Egypt: SAMIR et al., 2003; Greece: DEBENAY et al., 2005 Triloculina cf. fichteliana D’Orbigny R Turkey: MERI et al., 2004a Medit. Mar. Sci., 9/1, 2008, 119-165 139 9. Parasites Unlike the recent increase in reports of introduced free-living marine species, reports of parasites invading marine environments are relatively uncommon. When species are introduced into a new territory their parasites may also follow, forming a biotic unit called ‘symbiota’ (GALLI et al., 2005). Parasites are potentially able to control host populations and reduce host density by affecting their growth, reproduction, and survival (TORCHIN & MITCHELL, 2004). They can also affect community structure by indirectly acting on predation or competition (MOURITSEN & POULIN, 2006). A recent study examining all parasites from different groups of animals (molluscs, crustaceans, fish, birds, mammals, amphibians and reptiles) showed that the number of parasite species found in exotic populations is roughly half the number found in native populations (TORCHIN et al. 2003). In the same study, it was shown that introduced populations were less heavily parasitized in terms of average prevalence (percentage of hosts infected with one or more individuals of a parasite species (BUSH et al., 1997). The parasites with a lower probability of colonizing new areas tended to be those that were less prevalent in native populations. The success of establishment of parasites (both in new geographical areas and new hosts) has been described in detail by PASTERNAK et al. (2007) and PAIS et al. (2007) who illustrated that it may depend on: 1) how often the host population has been introduced into the new area; 2) the complexity of the parasite life-cycle; 3) the age of the host (according to BAUER (1991) younger fish, fry or fingerlings, are less infected than adults) and 4) global climatic change. 140 Few scientists have been only looking for parasitological aspects in the wild, while investigations have focused mostly on the micro- and macro- parasites of Indo-Pacific immigrants, the most popular hosts being Siganus luridus and S. rivulatus and more recently Fistularia commersoni. As knowledge concerning the original distribution of parasites is not complete, we propose cross-comparison of parasitological data regarding both native and alien hosts. We consider as alien the parasites detected only on alien hosts, or recognized as aliens from the authors of the research. Parasites recorded from both native and alien fish are regarded as of unknown origin. In particular, we consider of unknown origin the genera Ceratomyxa, Entamoeba, Hexamita (syn Octomitus) and Ortholinea, which in fact were found on both Red Sea and Mediterranean hosts. To date, 14 species of parasites have been recoqnized as Lessepsian (BARICHE & TRILLES, 2006; TRILLES & BARICHE, 2006; GALIL & LÜTZEN 1995; PAIS et al., 2007; PASTERNAK et al., 2007; PAPERNA, 1972; DIAMANT et al., 1999). The most representative group belongs to the class Monogenoidea (Table 9.1). In the Red Sea, monogenoids have been studied principally by PAPERNA (1965, 1972a,b,c), PAPERNA et al. (1984), DIAMANT & PAPERNA (1986), and DIAMANT (1989), and more recently by KRITSKY et al. (2007) and KRITSKY & GALLI (2007); In Mediterranean Sea Monogenoidea were investigated by DIAMANT (1989) and PASTERNAK et al. (2007). The first documented case of a monogenoidean invading a new biogeographical region by ‘natural’ extension of its host range is that of the gill ectoparasite Polylabris cf. mamaevi infecting rabbitfish, Medit. Mar. Sci., 9/1, 2008, 119-165 Table 9.1 Alien parasite species in the Mediterranean. + species reported in the ‘annotated list’. Taxon: CRU=Crustacea; MON=Monogenoidea/Platyelminthes; DIG= Digenea/ Platyelminthes; PRO=Protozoa. Establishment success: C=casual; E=established; Q=questionable Taxon CRU CRU CRU CRU CRU DIG DIG MON MON MON MON MON MON MON MON MON MON PRO PRO PRO PRO PRO PRO Species Establishment Source success Anilocra pilchardi Bariche C Lebanon: BARICHE & & Trilles, 2006 TRILLES, 2006 Cymothoa indica (Schiodte. C Lebanon: TRILLES & & Meinert, 1884) BARICHE, 2006 + Heterosaccus dollfusi Boschma, 1960 E Israel: GALIL & LÜTZEN 1995 + Mytilicola orientalis Mori, 1935 E France: HIS, 1977 + Myicola ostrea Hoshina & Sigiura, 1953 E France: POLLIO, 1981 Allolepidapedon fistulariae C Italy: PAIS et al., 2007 Yamaguti, 1940 + Hysterolecitha sigani Manter, 1969 Q Israel: FISCHTHAL 1980 Polylabris cf mamaevi Ogawa E Israel: DIAMANT, 1989 & Egusa, 1980 Paperna, 1972 Tetrancistrum polymorphus E Israel: PAPERNA, 1972d (Paperna, 1972) Tetrancistrum suezicum (Paperna, 1972) E Israel: PAPERNA, 1972d Tetrancistrum strophosolenum E Israel: DIAMANT et al., 1999 Kritsky, Galli & Tingbao 2007 Lecithochirium magnicaudatum C Israel: FISCHTHAL 1980 Fischthal & Kuntz, 1963 Monilicaecum ventricosum C Israel: FISCHTHAL 1980 Yamaguti, 1942 Glyphidohaptor plectocirra E Israel: PAPERNA, 1972d (Paperna, 1972) Lecithochirium magnicaudatum C Israel: FISCHTHAL 1980 Fischthal &Kuntz,1963 Monilicaecum ventricosum C Israel: FISCHTHAL 1980 Yamaguti,1942 + Neothoracocotyle acanthocybii C Italy: ROMEO et al., 2005 (Meserve, 1938) + Hirudinella ventricosa (Pallas, 1774) C Italy: ROMEO et al., 2005 Balantidium sigani Diamant Q Israel: DIAMANT et al., 1999 & Wilbert, 1985 Nosema ceratomyxa Diamant Q Israel: DIAMANT et al., 1999 & Paperna, 1985 + Bonamia ostrea (Pichot et al., 1979) C Spain: MONTES & LAMA, 1993 Perkinsus atlanticus (Levine 1978) E Spain: SANTMART et al., 1995 Marteilia refringens Cavalier-Smith, 2002 C Spain: RIERA et al., 1995 Medit. Mar. Sci., 9/1, 2008, 119-165 141 Siganus rivulatus. The prevalence levels of P. cf. mamaevi in the Israeli Mediterranean rabbitfish populations were very high, approximately three times greater than those found in the native populations of the northern Red Sea. The subfamily Protomicrocotylinae, to which Polylabris belongs, is almost entirely restricted to warmer waters of the Indo-west Pacific Ocean. Both host and parasite are Lessepsian immigrants that have co-invaded the Mediterranean Sea via the Suez Canal. The greater abundance of P. cf. mamaevi in the invading (Mediterranean) populations is probably due to the changed, new environment, possibly impacting on host resistance to the parasite and encouraging heavier infections (PASTERNAK et al., 2007). One Red Sea Digenea, Allolepidapedon fistulariae, was found by PAIS et al. (2007). A. fistulariae was reported for the first time from the Mediterranean Sea, parasitizing the bluespotted cornetfish Fistularia commersonii. The finding of this Indo-Pacific digenean, acquired by juvenile or adult fish predating on second intermediate hosts, would suggest that F. commersonii colonized the Mediterranean by actively swimming individuals and not by passive planktonic larvae (PAIS et al., 2007). Cymothoids (Isopoda) are a group of crustaceans typically parasitic of teleost fish. However, they are poorly studied animals and some groups remain completely undescribed. Studies of parasitic isopods from Lebanon have revealed the occurrence of two introduced species Anilocra pilchardi and Cymothoa indica (BARICHE & TRILLES, 2006; TRILLES & BARICHE, 2006) from the Indo-Pacific region parasitizing mainly barracudas (Sphyraenidae). 142 The global transport of marine bivalves for aquaculture has lead to widespread introductions of parasitic copepods. Some parasitic copepods that infect shellfish have been widely introduced with the transport and culture of bivalves. Mytilicola orientalis and Myicola ostrae are both parasitic copepods of the Pacific oyster, Crassostrea gigas in Asia, where they are native. Both species infect native bivalves and M. orientalis is considered a serious pest (STREFTARIS & ZENETOS, 2006). Protistan parasites have often caused significant and widespread effects in marine systems and aquaculture operations. Some of these pathogens are likely to be introduced species. However, for other pathogens there is limited evidence of an exotic source. The protistan reported as aliens in the Mediterranean are the species Bonamia ostrea, Perkinsus atlanticus and Marteilia refringens, all introduced accidentally with aquaculture. Bonamia ostreae is an intracellular haplosporidian parasite in European flat oysters Ostrea edulis that occurs on both coasts of the United States and causes significant mortality in Europe (MARTY et al., 2006); since 1990 Perkinsus atlanticus has been responsible for abnormal mortalities of Ruditapes decussatus in Spain (SANTMART et al., 1995; SAGRIST A et al, 1996); Marteilia refringens is a haplosporidium protozoan parasite affecting the digestive system of the flat oyster. In Europe this disease is commonly known as Aber disease, digestive gland disease of the European oyster, or marteiliosis. Infection with Marteilia refringens produces high mortality, associated with sporulation in the epithelial cells of the digestive tubules. An intermediate host or a free-living stage is thought to be required in the lifecycle of this parasite. Marteilia refrinMedit. Mar. Sci., 9/1, 2008, 119-165 gens can occur in some oysters without causing disease. Table 9.1 summarizes the information known to the authors where fourteen (14) additions have been made to the ‘annotated list’. In the absence of further evidence, one of the crustaceans rated as casual in the ‘annotated list’, namely the cirripedia Loxothylacus texanus known to parasitise on Callinectes sapidus is excluded in this work as an unsupported record. This brings the number total of known alien parasites in the Mediterranean to 21. Undoubtedly, more introduced marine parasites exist than are reported (TORCHIN & KURIS, 2005). Partly, this may be because, unlike their introduced hosts, parasites are often difficult to observe and thus some alien parasites go unreported. The hosts analysed are illustrated in Table 9.2 while the most recent nomenclature changes are presented in Table 9.3. Table 9.2 Alien parasites with their hosts. Taxon: CRU=Crustacea; MON=Monogenoidea/Platyelminthes; DIG= Digenea /Platyelminthes; PRO=Protozoa Taxon CRU Species Anilocra pilchardi CRU CRU CRU CRU DIG MON MON Cymothoa indica Heterosaccus dollfusi Mytilicola orientalis Myicola ostrea Allolepidapedon fistulariae Polylabris cf mamaevi Tetrancistrum polymorphus MON MON MON Tetrancistrum suezicum Tetrancistrum strophosolenum Glyphidohaptor plectocirra MON PRO PRO PRO PRO Hysterolecitha sigani Balantidium sigani Nosema ceratomyxae Bonamia ostrea Perkinsus atlanticus PRO Marteilia refringens Medit. Mar. Sci., 9/1, 2008, 119-165 host Sardina pilchardus Boops boops Pagellus acarne Pagellus erythrinus Engraulis encrasicolus Sphyraena chrysotaenia Charybdis longicollis Oyster beds Oyster beds Fistularia commersonii Siganus rivulatus Siganus rivulatus Siganus luridus Siganus rivulatus Siganus rivulatus Siganus rivulatus Siganus luridus Siganidae Siganus rivulatus Siganus rivulatus Oyster beds Ruditapes philippinarum Tapes decussatus Tapes semidecussatus Oyster beds 143 Table 9.3 Name changes in parasites. New name Tetrancistrum polymorphus (Paperna, 1972) Old name Pseudohaliotrematoides polymorphus eilaticus: Paperna, 1972 Pseudohaliotrematodides polymorphus eilaticus: KTARI & KTARI (1974) Tetrancistrum suezicum Pseudohaliotrematoides polymorphus (Paperna, 1972) suezicus: Paperna, 1972 Pseudohaliotrematodides polymorphus suezicus: KTARI & KTARI (1974) (misspelling); Pseudohaliotrematoides suezicus Paperna, 1972 Tetrancistrum strophosolenum Pseudohaliotrematoides nagatyi Diamant, Kritsky, Galli & Tingbao, 2007 1985 (nomen nudum) Pseudohaliotrematoides polymorphus ssp.: DIAMANT & PAPERNA (1986); P. nagatyi: Diamant (1989) (nomen nudum) Pseudohaliotrematoides polymorphus "nagatyi" of DIAMANT et al. (1999) Glyphidohaptor plectocirra Synonyms. Pseudohaliotrema plectocirra (Paperna, 1972) Paperna, 1972; Tetrancistrum plectocirra: Lim, 2002 10. Fish To the 110 species in the ‘annotated list’ sixteen (16) records have been added in the 2006-April 2008 period (Table 10) and one is excluded in this work. The established aliens increased from 63 to 69, the casual aliens from 42 to 50 and the number of questionable aliens (5) remaining the same, thus bringing the total number of alien fish to 125 species. Glaucostegus halavi, which was previously excluded from the list of alien fish by GOLANI et al. (2002 and whose vector of introduction was stated to be evidently from the Red Sea (BEN SOUISSI et al., 2007), has recently been found off southern Tunisia. Among questionable species in the 144 Source KRITSKY et al. 2007 KRITSKY et al. 2007 KRITSKY et al. 2007 KRITSKY & GALLI 2007 ‘annotated list’, Torpedo sinuspersici was recently evaluated by some authors as a casual Lessepsian species (i.e. SERENA, 2005; GALIL, 2008). However, the single specimen reported from Syria (SAAD et al., 2004) was not supported by any taxonomical information to confirm species identification, thus retaining its questionable occurrence until substantiated by further studies. The status of the bigeye thresher shark (Alopias superciliosus), that was classified as questionable due to uncertainty regarding its origin, is still not clear despite recent records from the Mediterranean Sea. Some morphological characters of the species (i.e. very large eyes reaching the dorsal surface of head; the v-shaped ridge on the head; long snout etc.) clearly distinMedit. Mar. Sci., 9/1, 2008, 119-165 Table 10 New alien fish species and changes in establishment success. * species excluded in the ‘annotated list’ Establishment success: C=casual, E=established, Q= questionable New species Apogon queketti Gilchrist, 1903 Apogon smithi (Kotthaus, 1970) Bregmaceros atlanticus Goode & Bean, 1886 1 2 Establishment success C E E Source Turkey: ERYILMAZ & DALYAN, 2006 Israel: GOLANI et al., 2008 Turkey: YILMAZ, et al., 2004; FILIZ et al., 2007 Israel: GOREN & GALIL, 2006 Libya: BEN ABDALLAH et al., 2007 Croatia: DUL I & GOLANI, 2006 Israel: GOLANI & CAPAPÉ, 2004 Israel: GOLANI, 2006 Tunisia: BEN SOUSSI et al., 2007 Turkey: BILECENO LU, 2007 Israel: GOLANI & SONIN, 2006 Turkey: BILECENO LU & RUSSELL, in press Croatia: DUL I & KRALJEVIC, 2007, Greece: CORSINI-FOKA & ECONOMIDIS, 2007 Egypt: KOVA I & GOLANI, 2007 Cephalopolis taeniops (Valenciennes, 1828) Cyclopterus lumpus Linnaeus, 1758 Dasyatis marmorata (Steindachner, 1892) Decapterus russelli (Rüppell, 1830) *Glaucostegus halavi1 Monotaxis grandoculis (ForsskaÆl, 1775) Nemipterus randalii Russell, 19862 C C Q E C C E Pagrus major (Temminck & Schlegel, 1843) C Papillogobius mel·nobranchus (Fowler, 1934) Parupeneus forsskali (Fourmanoir and Guézé, 1976) C Platax teira (ForsskaÆl, 1775) C Sciaenops ocellatus (Linnaeus, 1766) Zenopsis conchifera (Lowe, 1852) Change in establishment success Coryogalops ochetica (Norman, 1927) Hippocampus fuscus Rüppell, 1838 Torquigener flavimaculosus Hardy & Randall, 1983 C C Turkey: INAR et al., 2006b Malta (?): SCIBERRAS & SCHEMBRI, 2007 Turkey: BILECENO LU & KAYA, 2006 Israel: GOLANI & MIRES, 2000 Tunisia: RAGONESE & GIUSTO, 2007 From C to E From C to E From C to E Egypt: KOVA I & GOLANI, 2007 Turkey: BILECENO LU, pers.obs. Greece: CORSINI-FOKA et al., 2006 C Recorded as Rhinobatos halavi by BEN SOUISSI et al .(2007) Recorded as N.japonicus by GOLANI &SONIN (2006) guish A. superciliosus from its Mediterranean congeneric A. vulpinus, so the probability that it was previously overlooked is Medit. Mar. Sci., 9/1, 2008, 119-165 not justified. TORTONESE (1956) stated that the bigeye thresher shark is a typical species of the tropical Atlantic, and its 145 absence in the Mediterranean (at least until the 1980’s) was indicated in two monumental works (HUREAU & MONOD, 1973; WHITEHEAD et al., 1984). Although a range expansion of A. superciliosus during the last two decades from western and central basin to eastwards is evident (see CLO et al., 2008), a probable introduction via the Gibraltar Strait requires verification. Tortonese’s stingray (Dasyatis tortonesei) was listed among questionable fish in the ‘annotated list’, but should be excluded due to its distribution being restricted to the Mediterranean (WHITEHEAD et al., 1984). The validity of the species is still a matter of dispute (COMPAGNO, 1999). On the other hand, the eastern Atlantic originated Dasyatis marmorata is now included on our list as a questionable alien species. Its first record from Gulf of Gabès in southern Tunisia (MAURIN & BONNET, 1970) was followed by specimens captured along Israeli coasts (GOLANI & CAPAPÉ, 2004). Although D.marmorata has never been evaluated as an alien fish (GOLANI et al., 2002; SERENA, 2005), its pattern of zoogeographical distribution is similar to other tropical Atlantic originated fishes such as Enchelycore anatina, Arius parkii and Acanthurus monroviae (GOLANI & CAPAPÉ, 2004). The taxonomy of the species, which was previously mentioned under the names of Dasyatis pastinaca marmorata and D. chrysonota, is currently not clear. The antenna codlet (Bregmaceros atlanticus) finding from Turkey (YILMAZ et al., 2004) did not put forward the possible origin of the species, but GOREN & GALIL (2006) evaluated the species as alien, which is likely to have arrived in the Mediterranean with discharged ballast water. The antenna codlet is currently quite abundant at the northeastern Mediter146 ranean and several specimens may be observed during bottom trawl catch compositions. The species has recently been found in the Aegean Sea (FILIZ et al., 2007). The record of Nemipterus japonicus (Bloch, 1791) of GOLANI & SONIN (2006) is invalid as it is considered a misidentification of N. randalli Russell, 1986 (for a full account, see BILECENO LU & RUSSELL, in press; LELLI et al., in press). Even though Decapturus russelli and Apogon smithi are known by single records from the Mediterranean Sea, they were rated as established following GOLANI (2006) and GOLANI et al. (2008), respectively. According to a recent review of alien fish from the Maltese islands, SCIBERRAS & SCHEMBRI (2007) noted the possible occurrence of Parupeneus forsskali, which was hitherto known by a single report from Turkey ( INAR et al., 2006b). However, the relevant record from Malta should be evaluated as questionable due to lack of preserved material and information. Aphanius dispar was considered as a Lessepsian immigrant for many years. It is one of the worst alien fish in the eastern Mediterranean according to some authors (i.e. GOREN & GALIL, 2005; GALIL, 2008), since the species and its hybrids have replaced the native killifish, A. fasciatus, along the Mediterranean coast of Israel. On the other hand, GOLANI et al. (2002) excluded A. dispar from the list of exotic species, based on a comparative electrophoretic study by KORNFIELD & NEVO (1976) who suggested that the species was a permanent Mediterranean resident. Based on evidence presented by the latter authors that the occurrence of A. dispar in the Mediterranean precedes the opening of Suez Canal, we are retaining the species within excluded taxa in the ‘annotated list’. Medit. Mar. Sci., 9/1, 2008, 119-165 Discussion Biological invasions have been recognised as one of the priority issues of concern to the health of marine ecosystems in the Mediterranean (EEA, 2006). ZENETOS et al. (2006) reported approximately 745 alien species in an annotated list, up to December 2005. The figures presented in ZENETOS et al. (2006) were criticised by Prof D.F. Por (pers. Commun.) who argues that they present an artificial inflation because the Lessepsian immigrants are mixed with aliens introduced in other ways. In his opinion, the tropical immigrants fit in well in the impoverished and sub-tropical Levantine basin. GALIL (2008) claims a total of 558 alien metazoan species in the Mediterranean Sea, ignoring the 745 that were already assembled in the ‘annotated list’. The latest figures documented in this work reveal an even more spectacular increase in the number of marine alien species in the Mediterranean. In addition to the 94 new species, reported in the period 2006-2008, the ‘annotated list’ is further enriched with 81 records reported before 2005, which had either escaped the author’s attention, and thus are missing from the ‘annotated list’: or had been rated as excluded but have to be reinstated as aliens according to recent taxonomic/molecular studies. On the other hand, a few species (17 in all) included in the ‘annotated list’are excluded in this work as cosmopolitan taxa/ unsupported records/ indigenous species. To sum up, a total of 903 alien species are to date present in the Mediterranean; their breakdown per ecofunctional/taxonomic group is presented in Figure 1. To these, we could add a few fresh/brackish water species that occasionally occur in estuarine/coastal areas. In the latter category, some mollusca and fish are Medit. Mar. Sci., 9/1, 2008, 119-165 included. Among mollusca, we refer to the bivalvia Dreissena polymorpha and Anodonta woodiana and the gastropoda Potamopyrgus antipodarum, Congeria leucophaeta and Ferrissia wautieri, all cryptogenic species well established in the Mediterranean. Among fish, Micropterus salmoides, Acipenser baeri, Acipenser gueldenstaedtii have been reported from coastal areas in Greece (CORSINI-FOKA & ECONOMIDIS, 2007) and France (WGITMO, 2006). Worth noting is that the planktonic alien species appear to be underestimated. According to Prof. S. Lakkis (Lebanon) approximately two hundred alien phytoand zooplanktonic species have been collected from the Lebanese coast between 1970 and 2005. However, a critical evaluation is needed based on voucher specimens before these species can be rated as aliens in the Mediterranean. Excluding the parasites and foraminiferans that are treated in detail in this work, hence the increase is 3- fold to 6fold respectively, an increase of approximately 10-25% is observed in the other groups. Thus, the rate of biological invasions is highest in phytobenthos (24%) followed by zooplankton (16%), fish (13%) and zoobenthos (13%). These newcomers are mostly of tropical and subtropical origin, which presumably indicates the ‘tropicalization’ of the Mediterranean. Indeed, it has been argued that present-day warming ultimately favours the spread of warmwater species through direct and indirect effects, and especially by changing water circulation. It is impossible at present to foresee to what extent the exuberance of warm-water species will affect the trophic web and the functioning of marine ecosystems in the Mediterranean Sea of tomorrow (BIANCHI, 2007). The eastern Mediterranean is still the favourite destina147 tion for alien species, but the centre of introduction seems to have spread from Israel to Turkey. This observation suggests a shift from Lessepsian migration to shipmediated transfer, a further argument towards the ‘tropicalisation’ hypothesis for the Mediterranean. Besides the new species presented in this work, changes in their establishment success are also indicated. The state of establishment success, as in April 2008, is presented in Figure 1. On considering the overall increase, the percentage of the established species to the total alien species has remained stable (about 55% vs 52% reported by December 2005. However, the fact that more than 30 species have changed their establishment status to ‘established’ indicates that alien species are becoming all the more permanent inhabitants in the Mediterranean. In con- trast to the ‘annotated list’ (385 established species), there are to date 496 established alien species, an increase of 29%. GALIL (2008) reports approximately 80 species in the 2000-2007 period. This seems an underestimate. Based on the publications of the recent 28 months (Jan 2006-April 2008), it is here demonstrated that there are 94 new records (14 fish, 1 phytoplankton, 3 zooplankton, 59 zoobenthos, 13 macrophytes and 4 parasites), indicating a rate of 1 new entry every 9 days (1.3 weeks). This rate of introduction is in agreement with STREFTARIS et al. (2005) who noticed an increasing pattern in the rate of introduction over the years 1998-2003. In particular, they calculated the time span for alien species’ introduction in the Mediterranean to be 5.2 weeks in 1998 vs 2.9 weeks in 2003. The increasing trend in bio-invasions Fig. 1: Number of marine alien biota in the Mediterranean and their establishment success . Numbers indicate totals per group. 148 Medit. Mar. Sci., 9/1, 2008, 119-165 that has been mostly attributed to climatic change worldwide is partly due to the awareness and response of the scientific community on the issue, resulting in: a) taxonomic competence and multinational collaboration on projects and/or publications - tracing/identifying aliens has never been an easy task; and b) the development of electronic scientific journals that promote prompt publication of findings related to alien species. One good example is the new electronic journal Aquatic Invasions AI: an important part of the developing. European early warning system on aquatic invasive species. Fourteen out of the 94 new marine alien species in the Mediterranean were first recorded in the 2006-2007 volumes of AI. Conclusions ñ A total of 903 alien species by April 2008 in the Mediterranean. ñ The rate of biological invasions in the Mediterranean not only has not paused but is increases astronomically. 94 species within 28 months means 1 new species entry every 9 days! ñ The newcomers are becoming all the more permanent inhabitants. 496 established species in 2008 versus 385 in 2005 means a 29% increase in the number of permanent visitors! Acknowledgements We gratefully acknowledge the following individuals for generous contributions to this project, including the provision of the latest publications and comments at various stages. Fish: Jamila BEN SOUISSI (Tunisia); Jakov DUL I (Croatia); Daniel GOLANI (Israel) Mollusca: Serhat ALBAYRAK (Turkey); Medit. Mar. Sci., 9/1, 2008, 119-165 Ghazi BITAR (Lebanon); Fabio CROCETTA (Italy); Henk DEKKER (The Netherlands); Jeroen GOUD (The Netherlands); Christiane DELONGUEVILLE (Belgium); Alper DOGAN (Turkey); Roland HOUART (Belgium); Henk MIENIS (Israel); Jose TEMPLADO (Spain); Jacobus J. VAN AARTSEN (The Netherlands) Polychaeta: Guillermo SAN MARTIN (Spain); Maria Teresa AGUADO (Spain) Crustacea: Cedric D’ UDEKEM D’ ACOZ (Belgium); Gianna INNOCENTI (Italy) Echinodermata: Patrick SCHEMBRI (Malta) Foraminifera: Baki YOKE (Turkey) Ascidia: Francesco MASTROTOTARO (Italy), Alfonso RAMOS-ESPLA (Spain) Bryozoa: Jean-Loup D’HONDT (France) Zooplankton: Alexandra GUBANOVA (Russia); Sami LAKKIS (Lebanon); Ioanna SIOKOU-FRANGOU (Greece); Nejib DALY YAHIA (Tunisia); Howaida ZAKARIA (Egypt) Phytobenthos: Enric BALLESTEROS (Spain) This work was supported by the SESAME project, EC Contract No GOCE-036949, funded by the European Commission's Sixth Framework Programme under the priority 'Sustainable Development, Global Change and Ecosystems'. The publication reflects only the views of the authors and the EC is not liable for any use that may be made of the information contained herein. References ABELL , P. & HISPANO, C., 2006. 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