HCMR TEYXOS 6 ergasia 359 - Mediterranean Marine Science

26
Medit. Mar. Sci., 10/2, 2009, 109-133 109 Introduction Biological invasions represent a signif- icant risk for many natural ecosystems and have become an issue of increasing concern worldwide (SCHMITZ & SIMBERLOFF, 1997; EEA, 2009). Invasive alien species are considered to be the second most important cause of global biodiversity change, fol- lowing direct habitat destruction (CBD, 2000). In particular, biological invasions in aquatic habitats represent a recognized glob- Inventory of alien marine species of Cyprus (2009) S. KATSANEVAKIS 1 , K. TSIAMIS 2 , G. IOANNOU 3 , N. MICHAILIDIS 3 and A. ZENETOS 1 1 Institute of Marine Biological Resources, Hellenic Centre for Marine Research, Agios Kosmas, Greece 2 Institute of Oceanography, Hellenic Centre for Marine Research, Anavyssos, Greece 3 Department of Fisheries and Marine Research, Nicosia, Cyprus e-mail: [email protected] Abstract An updated inventory of alien marine species from the coastal and offshore waters of Cyprus is pre- sented. Records were compiled based on the existing scientific and grey literature, including the HCMR database of Mediterranean alien species, technical reports, scientific congresses, academic dissertations, and websites, as well as on unpublished/personal observations. The listed species were classified in one of five categories: established, invasive, casual, cryptogenic, and questionable. The mode of introduction and the year of first sighting were also reported for each species. Eight new records based on personal observa- tions of the authors were reported (Chondria coerulescens, Neosiphonia sphaerocarpa, Enchelycore anati - na, Lagocephalus spadiceus, Lagocephalus suezensis, Scomberomorus commerson, Sillago sihama, and Sphoeroides pachygaster ). Nine species, previously reported as aliens in Cypriot waters, were excluded from the inventory for various reasons. Twelve established species were characterized as invasive (Caulerpa race - mosa var. cylindracea, Cerithium scabridum, Strombus persicus, Trochus erithreus, Brachidontes pharao - nis, Pinctada radiata, Pseudonereis anomala, Charybdis helleri, Fistularia commersonii, Lagocephalus sceleratus, Siganus luridus , and Siganus rivulatus ) as they have a substantial impact on biodiversity and/or local economy. The impact of alien marine species in Cyprus is expected to grow in the near future, and further effort directed towards recording alien invasions and their impact will be needed. Keywords: Alien marine species; Biodiversity; Eastern Mediterranean; Invasive species; Lessepsian immigrants; Cyprus. Review Article Mediterranean Marine Science Volume 10/2, 2009, 109-133

Transcript of HCMR TEYXOS 6 ergasia 359 - Mediterranean Marine Science

Medit. Mar. Sci., 10/2, 2009, 109-133 109

Introduction

Biological invasions represent a signif-icant risk for many natural ecosystems andhave become an issue of increasing concernworldwide (SCHMITZ & SIMBERLOFF,

1997; EEA, 2009). Invasive alien species areconsidered to be the second most importantcause of global biodiversity change, fol-lowing direct habitat destruction (CBD,2000). In particular, biological invasions inaquatic habitats represent a recognized glob-

Inventory of alien marine species of Cyprus (2009)

S. KATSANEVAKIS1, K. TSIAMIS2, G. IOANNOU3, N. MICHAILIDIS3 and A. ZENETOS1

1 Institute of Marine Biological Resources, Hellenic Centre for Marine Research, Agios Kosmas,Greece2 Institute of Oceanography, Hellenic Centre for Marine Research, Anavyssos, Greece3 Department of Fisheries and Marine Research, Nicosia, Cyprus

e-mail: [email protected]

Abstract

An updated inventory of alien marine species from the coastal and offshore waters of Cyprus is pre-sented. Records were compiled based on the existing scientific and grey literature, including the HCMRdatabase of Mediterranean alien species, technical reports, scientific congresses, academic dissertations,and websites, as well as on unpublished/personal observations. The listed species were classified in one offive categories: established, invasive, casual, cryptogenic, and questionable. The mode of introduction andthe year of first sighting were also reported for each species. Eight new records based on personal observa-tions of the authors were reported (Chondria coerulescens, Neosiphonia sphaerocarpa, Enchelycore anati-na, Lagocephalus spadiceus, Lagocephalus suezensis, Scomberomorus commerson, Sillago sihama, andSphoeroides pachygaster). Nine species, previously reported as aliens in Cypriot waters, were excluded fromthe inventory for various reasons. Twelve established species were characterized as invasive (Caulerpa race-mosa var. cylindracea, Cerithium scabridum, Strombus persicus, Trochus erithreus, Brachidontes pharao-nis, Pinctada radiata, Pseudonereis anomala, Charybdis helleri, Fistularia commersonii, Lagocephalussceleratus, Siganus luridus, and Siganus rivulatus) as they have a substantial impact on biodiversity and/orlocal economy. The impact of alien marine species in Cyprus is expected to grow in the near future, andfurther effort directed towards recording alien invasions and their impact will be needed.

Keywords: Alien marine species; Biodiversity; Eastern Mediterranean; Invasive species; Lessepsianimmigrants; Cyprus.

Review Article

Mediterranean Marine ScienceVolume 10/2, 2009, 109-133

al threat with a strong impact on biodiver-sity and local economies (PERRINGS et al.,2002; PIMENTEL et al., 2005; EEA, 2006;STREFTARIS & ZENETOS, 2006).Although natural invasions do occur, nowa-days most biological invasions result fromhuman activities. The Mediterranean Sea isone of the regions most severely affected byalien marine invasions, fostered by the open-ing of the Suez Canal, fouling and ballasttransportation along shipping lines, aqua-culture, and the aquarium trade(STREFTARIS et al., 2005; GALIL, 2009).A recent update of the marine alien specieslist in the Mediterranean Sea reported a to-tal of 925 introduced species listed by Marchof 2009, of which 519 were considered to beestablished and spreading (ZENETOS, 2009).This overall estimation of marine alien speciesin the Mediterranean Sea represents approx-imately 7.5% of the known flora and fauna;however, this percentage is higher in theEastern Mediterranean and may exceed 20%in specific aquatic ecosystems (GALIL &ZENETOS, 2002). The number of recordedalien species in the Mediterranean Sea con-tinues to increase at a rate of one new recordevery nine days (ZENETOS et al., 2008).

In recent years there has been a stronginterest from the scientific community andinternational organisations in monitoringbiological invasions in the MediterraneanSea, assessing their impact on the native flo-ra and fauna and on local economies, in-vestigating the causes and patterns of alienintroductions, and proposing managementmeasures (CBD, 2000; EC, 2006, 2007, 2008;EEA, 2006; IMO, 2009). Within this frame-work, a great effort has been made duringthe last decade to compile fragmentary andsporadic information on the presence anddistribution of alien species in the Mediter-ranean Sea. CIESM has issued a series ofatlases of the alien species in the Mediter-

ranean Sea, covering fish, decapod and stom-atopod crustaceans, molluscs, and macro-phytes (www.ciesm.org). Additionally, sci-entists from several Mediterranean coun-tries have created and continuously updatedatabases of aquatic alien species, and havepublished national inventories usually in-cluding information on the origin of eachspecies, the mode of introduction, and theestablishment success. Such lists for marinealien species have been published for Greece(PANCUCCI-PAPADOPOULOU et al.,2006; ZENETOS et al., in press), Turkey( INAR et al., 2006), Malta (SCIBERRAS& SCHEMBRI, 2007), and Israel (GALIL,2007a). Inventories of marine alien specieson a European level have also been pro-duced in the framework of the DAISIE proj-ect (DAISIE, 2009) but these inventoriesare not regularly updated for all taxonomicgroups.

The Eastern Mediterranean is highlysusceptible to marine biological invasionsbecause of its location at the crossroads be-tween the Ponto-Caspian and the IndianOcean/Red Sea regions, the maritime traf-fic from the Indian Ocean, and a widespreadoccurrence of fish and shellfish farms. Lessep-sian immigrants in particular are mostly con-fined to the eastern part of the Mediter-ranean basin (GALIL & ZENETOS, 2002).Cyprus is the largest, easternmost Mediter-ranean island and the one closest to the SuezCanal. Comparing the list of alien speciesof Cyprus with those of other Eastern Mediter-ranean countries will assist in understand-ing the invasion patterns in the area.

The first systematic effort to record thealien marine species of Cyprus was initiat-ed in 1967 during a joint program (Biota ofthe Red Sea and eastern Mediterranean) bythe Smithsonian Institution, the HebrewUniversity of Jerusalem, and the Sea Fish-eries Research Station in Haifa. This proj-

Medit. Mar. Sci., 10/2, 2009, 109-133110

ect investigated the spread of the Lessep-sian immigrants in the Levant Basin (Egypt,Israel, Cyprus, Rhodes) and listed a total of140 Erythrean and Indo-Pacific species knownto have crossed the Suez Canal into theMediterranean Sea (STEINITZ, 1970; PORet al., 1972). Since then, other studies fo-cusing on specific taxa have appeared in theliterature (e.g. Mollusca: BUZZURRO &GREPPI, 1997; OZTURK et al., 2004; Poly-chaeta: BEN-ELIAHU & PAYIATAS, 1999;

INAR, 2005; Macroalgae: TA KIN et al.,2008) but information remains scattered anda complete updated list of the aliens of Cyprusis still missing. DAISIE (2009) reports a to-tal of 75 alien marine species from Cyprusbut this figure is a marked underestimationof the actual one.

The aim of this work is to present an up-dated list of the marine alien species ofCyprus, based on a thorough compilation ofexisting information, and to provide base-line information on the current situation onthe island. Furthermore, it compares this listwith the inventories of alien species in oth-er eastern Mediterranean countries and com-ments on possible differentiation.

Materials and Methods

Records of marine alien species fromthe coastal and offshore waters of Cypruswere compiled based on the existing scien-tific and grey literature, including the Hel-lenic Center for Marine Research (HCMR)database of Mediterranean alien species(ZENETOS & FRAGGOS, 2008; HCMR,2009), technical reports, scientific congress-es, academic dissertations, and websites, aswell as on unpublished/personal observa-tions. Records from grey literature or per-sonal observations were included as longas they were supported by physical evidencesuch as specimens or photographs. The list

has been updated based on species recordsup to July 2009.

The following definition of aliens wasadopted: A species, subspecies or lower tax-on, introduced outside its natural past orpresent distribution; it includes any part:gametes, seeds, eggs, or propagules of suchspecies that might survive and subsequent-ly reproduce (http://www.cbd.int/inva-sive/terms.shtml). Synonyms are non-native,non-indigenous, allochthonous, and exotic.

The dates of collection of the new records,rather than dates of publication, are reported,since a gap of many years may exist betweenthe two dates. When the date of collectionwas not reported in a publication, we triedto obtain this information by direct com-munication with the author(s). In a few cas-es where the date of collection remained un-known, the date of publication was used in-stead.

The establishment success of each specieswas assessed and is given according to thefollowing terminology, as proposed inZENETOS et al. (2006):

Established: Introduced or feral popu-lation of species settled in the wild with free-living, self maintaining and self-perpetuat-ing populations unsupported by and inde-pendent of humans. Species with at least tworecords in Cyprus spread over time and space(at least three records for fishes) are alsoclassified as established, in the sense of theCIESM atlas series.

Casual: Casual species are those havingbeen recorded only once (no more than twicefor fish) in the scientific and grey litera-ture and are presumed to be non-establishedin Cyprus. In this paper ‘casual’ is used inthe same sense as ‘alien’ in the CIESM at-las series.

Questionable: Species with insufficientinformation – ‘suspects’. Also new entriesnot verified by experts or species with taxo-

Medit. Mar. Sci., 10/2, 2009, 109-133 111

nomic status unresolved.Cryptogenic: Species with no definite ev-

idence of their native or introduced statusaccording to CARLTON (1996) and specieswhose probable introduction occurred in‘early times’ and not been witnessed, e.g. pri-or to 1800.

Excluded: Species previously reportedas aliens in Cyprus but excluded from ourlist due to one of the following criteria:misidentification, native species falsely pre-viously identified as alien, or spurious records(e.g., shells of molluscs transported by manfor food or ornament and left in places wherethey are normally absent). Excluded speciesare not presented in the main list but are re-ported separately.

Invasive species were also indicated. In-vasive were defined as those established aliensthat have overcome biotic and abiotic bar-riers and are able to disseminate away fromtheir area of initial introduction through theproduction of fertile offspring with notice-able impact, such as threat to the diversityor abundance of native species, the ecolog-ical stability of infested ecosystems, eco-nomic activities dependent on these ecosys-tems, and human health.

The records of marine alien species ofCyprus were compared with those of the oth-er countries of the Levantine basin (Egypt,Israel, Lebanon, Syria, southern Turkey)(Fig. 1). Such records were taken from theHCMR database of Mediterranean alienspecies (HCMR, 2009).

Results

Up to July 2009, a total of 126 alien ma-rine species has been reported in Cyprus,among which are 42 molluscs, 28 fish, 19polychaetes, 15 phytobenthic species, 12crustaceans, and 10 species from other taxa(ANNEX 1). Most of the species (101) have

an Indo-Pacific or Indian Ocean origin. Thedominant mode of introduction (104 species)was via the Suez Canal.

Among the 126 reported alien marinespecies, 80 are established (including 12 in-vasive), 31 are casual, 9 are cryptogenic, and6 are questionable. The species classified asinvasive were the green alga Caulerpa race-mosa var. cylindracea, the gastropod mol-luscs Cerithium scabridum, Strombus persi-cus, and Trochus erithreus, the bivalve mol-luscs Brachidontes pharaonis and Pinctadaradiata, the polychaete Pseudonereis anom-ala, the crustacean Charybdis helleri, and thefish Fistularia commersonni, Lagocephalussceleratus, Siganus luridus, and Siganus rivu-latus. These species have overcome bioticand abiotic barriers, have high abundanceand extensive spatial distribution in the coastalwaters of Cyprus, and have evident eco-logical and/or economic impacts (DFMR,2008; STREFTARIS & ZENETOS, 2006).

Six species were characterized as ques-tionable, for various reasons:1. Polysiphonia atlantica is easily confused

with other Mediterranean species of thegenus Polysiphonia (BEN MAIZ et al.,1986) and its occurrence in the Mediter-ranean Sea needs further study.

2. Laodicea fijiana has been reported onlyonce by SCHMIDT (1973) (as cited inPOR, 1978) and could be a misidentifi-cation of L. undulata (Forbes & Good-sir, 1853) (DE VITO et al., 2006).

3. Hydroides dianthus was found only oncein a state of decomposition, and takenfrom a ship’s propeller. There was noknowledge of the history of the ship, thusit was uncertain whether settlement tookplace in Cypriot waters (BEN-ELIAHU& PAYIATAS, 1999).

4. Lysidice collaris was probably confusedwith the native species L. margaritaceaClaparède, 1868 (ZENETOS et al., 2006).

Medit. Mar. Sci., 10/2 2009, 109-133112

5. For Terebella ehrenbergi, insufficient da-ta were provided in the initial recordand identification was judged uncertainby ZENETOS et al. (2006).

6. In the distribution map of Himanturauarnak, provided by LAST & STEVENS(1994), Cyprus is included; however noother evidence is provided that the specieswas actually found in Cypriot waters.

Nine species previously reported as aliensin Cypriot waters were excluded from the in-ventory for various reasons (Table 1).

In the light of present knowledge, theprevious reports of Linopherus acarunculataand Branchiomma boholense ( INAR, 2005)actually refer to L. canariensis and B. bairdirespectively ( INAR, 2009).

A comparison of the records of the alienmarine species of Cyprus with those of theother neighboring areas of the Levantine re-gion revealed a relatively low number ofrecorded species (Fig. 2). With the exceptionof Syria, where only 119 alien marine specieshave been recorded, in all the other Levan-tine countries much higher numbers of record-ed species appear (Egypt: 212, Israel: 389,Lebanon: 262, South Turkey: 257) (ZENETOS,2009). The relative difference of recordedalien species varies greatly among taxonom-ic groups. The reported number of alien poly-chaetes in Cyprus is similar to that of mostother Levantine areas (Fig. 2). For all oth-er major taxa there is a deficit in reportedspecies in Cyprus when compared with theother Levantine countries, except for mol-

Medit. Mar. Sci., 10/2, 2009, 109-133 113

Fig. 1: Map of the Levantine basin and surrounding countries.

luscs, where Syria had fewer records (Fig. 2).However, there is an increasing trend inthe reporting of alien marine species in Cyprus(Fig. 3), and thus their total number is ex-pected to increase in the near future.

A total of 352 alien marine species havebeen established in the Levantine Sea(HCMR, 2009). The established alien ma-rine species of Cyprus represent 22% ofthe Levantine established alien marinespecies, while an extra 8% have been re-ported as casual or questionable. The vastmajority of established Levantine alien ma-

rine species (70%) have not been report-ed from Cyprus.

Discussion

The present compilation of the alienmarine species of Cyprus revealed a total of126 species, which is much higher than thepreviously reported figures. GALIL (2006)reported 78 alien marine species from Cyprus,while 75 species are reported in DAISIE(2009). Both these studies are based in themain on 2005 data and the DAISIE data-

Medit. Mar. Sci., 10/2, 2009, 109-133114

Table 1Species excluded from the inventory of alien marine species of Cyprus.

Species Reported by Reasoning for exclusionLeonnates persicus Wesenberg-Lund, 1949 GALIL, 2006 Absent from Cyprus

(M.E. INAR, pers. comm.).Rhodine loveni Malmgren, 1865 BEN-ELIAHU, 1972 Atlanto-Mediterranean species

(ZENETOS et al., 2006).Branchiosyllis exilis (Gravier, 1900), INAR 2003 A widespread species

(Atlantic, Indo-Pacific), including eastAtlantic (ZENETOS et al., 2006).

Monticellina dorsobranchialis INAR, 2005 Atlanto-Mediterranean species (Kirkegaard, 1959) (ZENETOS et al., 2006).Umbonium vestiarium (Melvill, 1896) DELONGUEVILLE & ZENETOS et al. (2004)

SCAILLET, 2007 consider it to be a spurious record.DELONGUEVILLE & SCAILLET(2007) state also that the identificationis not verified.

Sebastapistes nuchalis (Günther, 1874) FRÖILAND, 1972 The initial record was based upon amisidentification of a Scorpaenamaderensis Valenciennes, 1833 specimen (GOLANI et al., 2002).

Radicilingua thysanorhizans (Holmes) CIRIK et al., 2000 It was considered a native Papenfuss 1956, species (VERLAQUE et al., in press).Thais sacellum (Gmelin, 1791) GALIL, 2006. Not present in CyprusPseudochama corbieri (Jonas, 1846) GALIL, 2006 DEMETROPOULOS (1971) refers

to Pseudochama ferruginea (Reeve,1846), a distinct species.

Medit. Mar. Sci., 10/2, 2009, 109-133 115

Fig. 2: Comparison of the number of reported alien marine species in the surrounding countries of theLevantine basin. For Turkey only the Levantine part (south Turkey) was considered. Non-establishedare the casual and questionable.

Fig. 3: Cumulative number of alien marine species in Cyprus per decade, based on the reported yearof first sighting.

base seems not to have been updated sincethen. In the present study the number ofalien marine species from Cyprus has in-creased by 51 species (~68%) comparedwith the DAISIE (2009) inventory. This in-crease is mainly due to: 1. The introduction of new alien species in

Cyprus. Twenty-three alien marine specieshave been reported since 2005 (althoughthis may be partly due to point (2) be-low). The rate of new biological inva-sions in the Mediterranean Sea is highand has been estimated at 1 new speciesevery 9 days (ZENETOS et al., 2008).These species are mostly introduced viathe Suez Canal and their successful es-tablishment is believed to have been as-sisted in recent years by climate change(OCCHIPINTI-AMBROGI, 2007).

2. More intense research in the field in re-cent years and rapid publication. Theawareness and response of the scientif-ic community to the issue of alien marineinvasions has resulted in taxonomic com-petence and multinational collaborationin related projects/publications. Databas-es of alien marine species have been cre-ated (e.g., Si.Di.Mar, 2008; HCMR, 2009;DAISIE, 2009; ELNAIS, 2009) and elec-tronic scientific journals promote promptpublication of findings related to alienmarine species (e.g., Aquatic Invasions,Journal of the Marine Biological Asso-ciation of the UK – Biodiversity Records,Mediterranean Marine Science)(GKOUVOUSI & ZENETOS, 2009).

3. A more thorough investigation of the lit-erature. Scientific publications and greyliterature that were not taken into ac-count in the previous inventories wereincluded in the present review.

4. New records based on personal obser-vations of the authors. Eight species notpreviously reported elsewhere have been

included in this inventory. The new records are the two Rhodophy-

ta Chondria coerulescens and Neosiphoniasphaerocarpa (K. Tsiamis, pers. observ.) andthe fish Enchelycore anatina, Lagocephalusspadiceus, Lagocephalus suezensis, Scombero-morus commerson, Sillago sihama, and Spho-eroides pachygaster (G. Ioannou & N. Michai-lidis, pers. observ.). The two Rhodophytawere characterized as cryptogenic, two ofthe fish were of Atlantic origin (E. anatinaand S. pachygaster) and the rest of Indo-Pa-cific origin. Two of the fish species (E. anati-na and S. sihama) were casual records, whilethe rest are considered established.

The number of alien marine species re-ported in a country is largely dependenton the scientific effort directed towardsrecording alien species, and on the availableexpertise. For example, the markedly lownumber of alien polychaetes from Syria (Fig.2) is attributed rather to the lack of focussedresearch on these taxa than to the absenceof alien polychates from Syrian coastal wa-ters. Similarly, the observed low numbers ofreported alien marine species from Cypruson many taxonomic groups such as crus-taceans, fish, and molluscs, in comparisonto the neighbouring countries of the Le-vantine Sea (Fig. 2), indicates a lack of rel-evant information. A directed survey to recordthe distribution of alien marine species inCypriot waters will surely uncover manymore species than the 126 reported here, es-pecially molluscs, fish, crustaceans, and phy-tobenthic species.

Additionally, the relatively low numberof alien marine species may be partly due tothe insular character of Cyprus. Accordingto the theory of island biogeography(MACARTHUR & WILSON, 2001), amongthe factors that influence species richnesson islands is their location relative to oceancurrents. The general circulation in the Le-

Medit. Mar. Sci., 10/2, 2009, 109-133116

vantine Sea is described as a counter clock-wise flow along the Libyo-Egyptian and Mid-dle East slopes (GERIN et al., 2009). Theisolation of Cyprus, with a minimum dis-tance of 70 km from the continent and thelack of a strong connecting current with theLevant coast, is a barrier for many speciesin successfully colonising the island (BEN-ELIAHU & PAYIATAS, 1999). Thus, trans-ported by the current, marine alien speciesentering through the Suez Canal progres-sively advance along the coastline throughEgypt, Israel, Lebanon, Syria, Turkey, andGreece. The colonization of Cyprus by thosespecies may be hindered by water circula-tion.

The established (and invasive) speciesin Cypriot waters are expected to increasein the future, as they currently represent asmall percentage of the established alienspecies in the Levantine basin. The alienspecies introduced via the Suez Canal arethermophilic species, and it is likely that anincreasing seawater temperature will favourtheir reproduction, growth, survival, and thusestablishment in the area. Such an increaseof established alien marine species is ex-pected to have an adverse impact on the stateof many native populations as well as an un-predictable impact on fisheries and localeconomies.

Twelve established species were char-acterized as invasive. Apart from Trochuserithreus, the other eleven species (Caulerparacemosa var. cylindracea, Cerithium scabridum,Strombus persicus, Brachidontes pharaonis,Pinctada radiata, Pseudonereis anomala,Charybdis helleri, Fistularia commersonii,Lagocephalus sceleratus, Siganus luridus, andSiganus rivulatus) are included in the list ofthe 100 ‘Worst Invasives’ in the Mediter-ranean (STREFTARIS & ZENETOS, 2006).

C. racemosa var. cylindracea exhibitedan excessive rate of proliferation and has in-

vaded the entire basin of the MediterraneanSea within the last 18 years. The rate of ex-pansion for C. racemosa appears to be amongthe most significant ever recorded for an in-vasive species (KLEIN & VERLAQUE,2008). It has stolons that can quickly elon-gate, and easily overgrow other macroal-gal (PIAZZI et al., 1997) or invertebratespecies (KRU IC’ et al., 2008) and poses asubstantial threat to biodiversity. In Cyprus,it has rapidly expanded forming extensivemats and causing changes in macrofaunalassemblages (ARGYROU et al., 1999).

All four invasive fish (F. commersonii,L. sceleratus, S. luridus, and S. rivulatus) arequite abundant and represent a substan-tial portion of the catches of the coastal fleet.F. commersonii is a high order carnivore andhas been nicknamed the "Lessepsian sprint-er" (KARACHLE et al., 2004) due to its veryrapid spread and successful establishmentin the Mediterranean Sea, reaching as faras Spain (SANCHEZ-TOCINO, 2007) with-in less than a decade from its introductioninto the Mediterranean Sea. The invasionof F. commersonii is expected to have asignificant impact on the composition of thenative ichthyofauna.

L. sceleratus, although a relative new in-vader (first record in the Mediterranean Seawas in 2003 by AKYOL et al., 2005), has be-come abundant and well-established in theLevantine basin. In Cyprus this species hasa significant negative impact on the artisanalfisheries, since it often damages both thefishing gear and the catch of the fishermenwith its powerful jaws (DFMR, 2008). Insome areas, many fishermen have even al-tered their fishing methods (gear, depths,time of the day, etc.) in order to avoid in-teraction with this species (G. Ioannou &N. Michailidis, pers. observ.). L. sceleratushas also a potential risk to humans, since itcontains tetrodotoxin, which may cause poi-

Medit. Mar. Sci., 10/2, 2009, 109-133 117

soning and even death (BENTUR et al, 2008;KATIKOU et al., 2009).

The two species of siganid fish, Siganusrivulatus and S. luridus have become dom-inant in many eastern Mediterranean coastalareas. They compete with the two main na-tive herbivores, Sparisoma cretense (Lin-naeus, 1758) and Sarpa salpa (Linnaeus,1758) and it seems that they have replacedthe latter in many areas (BARICHE et al.,2004). These two siganids have altered thecommunity structure and the native foodweb along the Levantine rocky infralit-toral zone (GALIL, 2007b). In 2008, thesetwo species represented 4.6% in weight ofthe total catch of the Cypriot artisanal fish-eries (DFMR, 2008).

Most of the invasive molluscs are ‘oldinvaders’ in Cyprus and have successfullyspread, altering the biocommunity structureand impacting on native species. Some ofthem (Strombus persicus and Pinctada ra-diata) are edible and are being exploited forhuman consumption in certain areas(STREFTARIS & ZENETOS, 2006;KATSANEVAKIS et al., 2008).

The polychaete Pseudonereis anomalaand the crab Charybdis helleri have reachedhigh population densities in many localitiesin Cyprus, impacting native species and af-fecting benthic biodiversity.

It is foreseen that the impact of alienspecies will intensify in the future in allMediterranean countries but especially inthe Levantine basin. The present study servesas a tool in the effort to monitor the expan-sion of alien species in Cyprus and may beutilized as a baseline for future assessments.Although the present inventory has sub-stantially added to our previous knowledgeof the extent of alien invasions in Cyprus, itis probably still an underestimation of theactual situation. Further directed effort isneeded to assess the number of established

alien species and their potential impact onthe island.

Acknowledgements

We gratefully acknowledge the contri-bution of the following colleagues who pro-vided their latest publications, clarifications,and comments at various stages: Murat Bile-cenoglu (Turkey), Melih Ertan inar(Turkey), ükran Cirik (Turkey), AngelosHannides (Cyprus), Roland Houart (Bel-gium), Unsal Karhan (Turkey), Martin Langer(Germany), Henk Mienis (Israel), FrancisDov Por (Israel), Evi Vardala-Theodorou(Greece), George Tornaritis (Cyprus), Ba-ki Yokes (Turkey), Helmut Zibrowius(France). The comments made by threeanonymous reviewers contributed to thequality improvement of the manuscript.

References

A IK, S., MURINA, G.V., INAR, M.E.& ERGEN, Z., 2005. Sipunculans fromthe coast of northern Cyprus (easternMediterranean Sea). Zootaxa, 1077: 1-23.

AKYOL, O., UNAL, V., CEYHAN, T. &BILECENOGLU M., 2005. First con-firmed record of Lagocephalus scelera-tus (Gmelin, 1789) in the MediterraneanSea. Journal of fish biology, 66: 1183-1186.

ARGYROU, M., 2000. The "Cladophora"phenomenon in the coastal waters ofCyprus. Proceedings of the First Mediter-ranean Symposium on Marine Vegeta-tion. Ajaccio, France, 3-4 October 2000,p. 69-73.

ARGYROU, M., DEMETROPOULOS,A. & HADJICHRISTOPHOROU, M.,1999. Expansion of the macroalga Cauler-pa racemosa and changes in softbottommacrofaunal assemblages in Moni Bay,

Medit. Mar. Sci., 10/2, 2009, 109-133118

Cyprus. Oceanologica Acta, 22: 517-528.BARICHE, M., LETOURNEUR, Y. &

HARMELIN-VIVIEN, M., 2004. Tem-poral fluctuations and settlement pat-terns of native and Lessepsian herbivo-rous fishes on the Lebanese coast (east-ern Mediterranean). Environmental Bi-ology of Fishes, 70: 81-90.

BAZZOCCHI, P., 1985. Prima segnalazionedi Strombus (Conomurex) decorus ray-baudii Nicolay & Romagna-Manoja,1983 per l’Isola di Cipro. Bolletino Mala-cologico, 21: 64.

BEN-ELIAHU, M.N., 1972. Littoral poly-chaeta from Cyprus. Tethys, 4: 85–94.

BEN-ELIAHU, M.N., 1995. A list of Poly-chaeta along the Levant coast. Haasiana,1: 78-93.

BEN-ELIAHU, M.N. & PAYIATAS, G.,1999. Searching for lessepsian migrantserpulids (Annelida: Polychaeta) on Cyprus– some results of a recent expedition.Israel Journal of Zoology, 45: 101-119.

BEN-MAIZ, N., BOUDOURESQUE, C.F.& GERBAL, M., 1986. Flore algale del’ étang de Thau: Grateloupia dorypho-ra (Montagne) Howe et Grateloupia fil-icina (Wulfen) C. Agardh. Thalasso-graphica, 9: 39-49.

BENTUR Y, ASHKAR J, LURIE Y, LEVYY, AZZAM ZS, LITMANOVICH M,GOLIK M, GUREVYCH B, GOLANID, EISENMAN A, 2008. Lessepsian mi-gration and tetrodotoxin poisoning dueto Lagocephalus sceleratus in the easternMediterranean. Toxicon, 52(8): 964-8.

BEN-TUVIA, A., 1962. Collection of fish-es from Cyprus. Bulletin of the ResearchCouncil of Israel, 11B: 132-145.

BOGI, C., CIANFANELLI, S. & TALENTI,E., 1989. La malacofauna dell’ Isola diCipro. p. 187–214. In: Atti Prima Gior-nata di Studi Malacologici C.I.S.MA. Cen-tro Italiano di Studi Malacologi, I. Nofroni

(Ed.), Roma, Italy.BUZZURRO, G. & GREPPI, E., 1997.

Notes on the molluscs of Cyprus, withspecial attention to the allochthone species.La Conchiglia, 283: 21-31 and 61-62.

CARLTON, J.T., 1996. Biological invasionsand cryptogenic species. Ecology, 77:1653-1655.

CBD, 2000. Global strategy on invasive alienspecies. Convention on Biological Diver-sity, UNEP/CBD/SBSTTA/6/INF/9: 1-2.

CECALUPO, A. & QUADRI, P. 1994. Con-tributo alla conoscenza malacologicaper il nord dell’ isola di Cipro (parte I).Bollettino Malacologico, 30(1-4): 5-16.

CECALUPO, A. & QUADRI, P., 1995.Contributo alla conoscenza malacolog-ica per il nord dell’ isola di Cipro (parteII). Bollettino Malacologico, 30(10-12):269-276.

CECALUPO, A. & QUADRI, P., 1996.Contributo alla conoscenza malacolog-ica per il nord dell’ isola di Cipro (parteIII). Bollettino Malacologico, 31(5-8):95-118.

CHERBONNIER, G., 1986. Holothuriensde la Méditerranée et du nord de la merRouge. Bulletin du Muséum Nationald'Historie Naturelle, 8: 43-46.

INAR, M.E., 2003. Ecology of Syllidae (An-nelida: Polychaeta) from northern Cyprus(eastern Mediterranean Sea). Bulletin ofMarine Science, 72(3): 795-811.

INAR, M.E., 2005. Polychaetes fromthe coast of northern Cyprus (easternMediterranean Sea), with two new recordsfor the Mediterranean Sea. Cahiers deBiologie Marine, 46: 143-159.

INAR, M.E., 2009. Alien polychaete species(Annelida: Polychaeta) on the southerncoast of Turkey (Levantine Sea, easternMediterranean), with 13 new records forthe Mediterranean Sea. Journal of Nat-ural History, 43: 2283-2328.

Medit. Mar. Sci., 10/2, 2009, 109-133 119

INAR, M.E. & ERGEN, Z., 2003. Eu-syllinae and Syllidae (Annelida: Poly-chaeta) from northern Cyprus (easternMediterranean Sea) with a checklistof species reported from the Levant Sea.Bulletin of Marine Science, 72(3): 769-793.

INAR, M.E., ERGEN, Z. & BENLI, H.A.,2003. Autolytinae and Exogoninae (Poly-chaeta: Syllidae) from northern Cyprus(eastern Mediterranean Sea) with achecklist of species reported from theLevant Sea. Bulletin of Marine Science,72(3): 741-767.

INAR, M.E., BILECENOGLU, M.,ÖZTÜRK, B., KATAGAN, T. &AYSEL, V., 2006. Alien species on thecoasts of Turkey. Mediterranean MarineScience, 6(2): 119-146 [2005].

CIRIK, ., AYSEL, V., BENLI, H.A.,CIHANGIR, B. & ÜNLÜO LOU, A.,2000. Preliminary studies on the marinevegetation of northern Cyprus. TurkishJournal of Marine Sciences, 6: 31-40.

DAISIE, 2009. Delivering Alien InvasiveSpecies Inventories for Europe. Availableon line: http://www.europe-aliens.org/.Accessed on 24/6/09.

DELONGUEVILLE, C. & SCAILLET, R.,2007. Les espèces de mollusques inva-sives de Méditerranée. Novapex, 8(3):47-70.

DEMETROPOULOS, A., 1971. Marinemolluscs of Cyprus. Part B: Bivalvia(Lamellibranchiata). Fisheries Bulletin,Ministry of Agriculture and Natural Re-sources, Fisheries Department, 3: 1-24.

DEMETROPOULOS, A. & HADJICHRI-STOPHOROU, M., 1976. Some addi-tions to the knowledge of the malaco-fauna of Cyprus. Fisheries Bulletin, Min-istry of Agriculture and Natural Resources,Fisheries Department, 4: 75-83.

DEMETROPOULOS, A. & NEOCLEOUS,

D., 1969. The fishes and crustaceans ofCyprus. Fisheries Bulletin, Ministry ofAgriculture and Natural Resources, Fish-eries Department, 1: 1-21.

DE VITO, D., PIRAINO, S., SCHMICH,J., BOUILLON, J. & BOERO, F., 2006.Evidence of reverse development inLeptomedusae (Cnidaria, Hydrozoa):the case of Laodicea undulata (Forbesand Goodsir 1851). Marine Biology, 149:339-346.

DFMR, 2006. Toxic Fish. Technical Report427/2006, Department of Fisheries andMarine Research, Ministry of Agricul-ture, Natural Resources and Environ-ment, Nicosia, Cyprus.

DFMR, 2008. Annual report on the Cyprusfisheries for the year 2008. Departmentof Fisheries and Marine Research. Min-istry of Agriculture, Natural Resourcesand Environment, Nicosia, Cyprus.

EC, 2006. Halting the loss of biodiversity by2010 – and beyond. Sustaining ecosystemservices for human well-being. COM (2006)216. Brussels, Commission of the Euro-pean Communities.

EC, 2007. Council Regulation concerning useof alien and locally absent species in aqua-culture. Regulation 708/2007, OJ L 168/1.

EC, 2008. Marine strategy framework direc-tive. Directive 2008/56/EC, OJ L 164.

EEA, 2006. Priority issues in the Mediter-ranean environment. EEA Report 4/2006.European Environmental Agency, UNEP.ISSN 1725-9177.

EEA, 2009. Progress towards the European2010 biodiversity target. EEA Report4/2009. European Environmental Agency,UNEP. ISSN 1725-9177.

ELNAIS, 2009. Hellenic Network on Aquat-ic Invasive Species. Available on line:http://elnais.ath.hcmr.gr. Accessed on24/6/09.

ELTON, CS, 1958. The ecology of invasions

Medit. Mar. Sci., 10/2, 2009, 109-133120

by animals and plants. Methuen & Co.London. 181 pp.

FISCHER, W., 1993. Beitraege zur Kennt-nis der rezenten und fossilen marinenMolluskenfauna Zyperns (II). Die Mol-lusken des Kap Drepanum (Peyia,Paphos). Club Conchylia Information,15(2): 147-152.

FODER , V., 1961. Report to the gov-ernment of Cyprus on fishery develop-ment possibilities. Expanded Programof Technical Assistance, FAO, ReportNo 1436.

FRÖILAND, O., 1972. Fishes of the fami-ly Scorpaenidae from Cyprus, includingthree new records. Bulletin Sea FisheriesResearch Station, Haifai, 59: 5–16.

GALIL, B.S., 2006. The marine caravan –the Suez Canal and the Erythrean in-vasion. p. 207-300. In: Bridging divides:maritime canals as invasion corridors, S.Gollasch, B.S. Galil, & A.N. Cohen (Eds)(Monographiae Biologicae 83), Berlin,Springer Verlag.

GALIL, B.S., 2007a. Seeing Red: Alienspecies along the Mediterranean coastof Israel. Aquatic Invasions, 2: 281-312.

GALIL, B.S., 2007b. Loss or gain? Invasivealiens and biodiversity in the Mediter-ranean Sea. Marine Pollution Bulletin,55: 314-322.

GALIL, B.S., 2009. Taking stock: invento-ry of alien species in the MediterraneanSea. Biological Invasions, 11: 359-372.

GALIL, B.S. & ZENETOS, A., 2002. A seachange – Exotics in the Eastern Mediter-ranean. p. 325-336. In: Invasive Aquat-ic Species of Europe: Distributions, Im-pacts and Management, E. Leppakoski,S. Gollasch & S Olenin (Eds.), Dor-drecht, Boston, London, Kluwer Acad-emic Publishers.

GALIL, B., FROGLIA, C. & NOEL, P.Y.,2002. CIESM Atlas of exotic species in

the Mediterranean. Vol. 2 Crustaceans:decapods and stomatopods. Monaco,CIESM Publishers, 192 pp.

GERIN, R., POULAIN, P.-M., TAUPIER-LETAGE, I., MILLOT, C., BENISMAIL, S. & SAMMARI, C., 2009.Surface circulation in the Eastern Mediter-ranean using drifters (2005–2007). OceanScience Discussions, 6, 525-555.

GITARAKOS, G., KALLIANIOTIS, A. &HADJISTEFANOU, N., 2007. Dem-ersal fish and megafaunal assemblageson the Cyprus continental shelf and slope(Eastern Mediterranean). Rapports etProcès-verbaux des Réunions, Commi-sion Internationale pour l’Exploration Sci-entifique de la Mer Méditerranée, 38: 488.

GKOUVOUSI, A. & ZENETOS, A., 2009.Alien species: new trends of scientificjournals for the record of a global phe-nomenon. p. 681-686. In: Proceedings ofthe 9th Panhellenic Symposium onOceanography and Fisheries, Patra, 13-16 May 2009 [in Greek].

GOLANI, D., 2000. The Lessepsian migrant,the Red-eye round herring Etrumeusteres (DeKay, 1842), a new record fromCyprus. Zoology in the Middle East, 20:61-64.

GOLANI, D., ORSI-RELINI, L., MASSUTI,E. & QUINGNARD, J.P., 2002. CIESMAtlas of exotic species in the Mediterranean.Vol. 1 Fishes. Monaco, CIESM Pub-lishers, 256 pp.

HADJICHRISTOPHOROU, M., ARGY-ROU, M., DEMETROPOULOS, A. &BIANCHI, T.S., 1997. A species list ofthe sublittoral soft-bottom macrobenthosof Cyprus. Acta Adriatica, 38(1): 3-32.

HCMR, 2009. European Database on alienspecies. Developed for EEA by the Hel-lenic Centre for Marine Research. Up-date of June 2009.

HORNEL, J., 1935. Report on the fisheries

A

Medit. Mar. Sci., 10/2, 2009, 109-133 121

of Palestine. London, Government ofPalestine, Crown Agent for the Colonies.

HOUART, R., 2001. A review of the recentMediterranean and Northeastern Atlanticspecies of Muricidae. Roma, EdizioniEvolver.

IMO, 2009. International Conventionfor the Control and Management ofShips' Ballast Water and Sediments.http://www.imo.org/

INGLE, R.W., 1963. Crustacea Stomatopo-da from the Red Sea and Gulf of Aden.Bulletin Sea Fisheries Research Station,Haifa, 33: 1-69.

KARACHLE, P.K., TRIANTAPHYLLIDIS,C. & STERGIOU, K. I., 2004: Bluespot-ted cornetfish, Fistularia commersoniiRüppell, 1838: a lessepsian sprinter. Ac-ta Ichthyologica et Piscatoria, 34: 103-108.

KATIKOU, P., GEORGANTELIS, D.,SINOURIS, N., PETSI, A. & FOTARAS,T., 2009. First report on toxicity assess-ment of the Lessepsian migrant puffer-fish Lagocephalus sceleratus (Gmelin,1789) from European waters (AegeanSea, Greece). Toxicon 54(1): 50-55.

KATSANEVAKIS S, LEFKADITOU E,GALINOU-MITSOUDI S, KOU-TSOUBAS D, ZENETOS A, 2008. Mol-luscan species of minor commercial in-terest in Hellenic Seas: Distribution, ex-ploitation and conservation status. Mediter-ranean Marine Science, 9(1): 77-118.

KAYA, M., BILECENOGLU, M. &GOLANI, D., 2000. New record of aLessepsian migrant Pteragogus pelycusRandall, 1981 (Teleostei: Labridae) fornorthern Cyprus. Zoology in the MiddleEast, 20: 65-68.

KIMOR, B. & BERDUGO, V., 1967. Cruiseto the eastern Mediterranean Cyprus03, Plankton Reports. Sea Fisheries Re-search Station Haifa, Bulletin No: 45.

KIRMITZOGLOU, I., KITSOS, M.S.,

THESSALOU-LEGAKI, M., TSELE-PIDES, A. & KOUKOURAS, A., 2006.Investigation of the progress and possi-ble expansion of the limits of the lessep-sian migratory current regarding De-capoda (Crustacea). Proceedings of the10th International Congress on the Zoo-geography and Ecology of Greece and ad-jacent regions, Patra, Greece, 26-30 June2006: 51.

KLEIN, J. & VERLAQUE, M., 2008. TheCaulerpa racemosa invasion: A criticalreview. Marine Pollution Bulletin, 56:205–225.

KOCATA , A., KATA AN, T. & BENLI,H.A., 2001. Contribution to the knowl-edge of the Crustacean fauna of Cyprus.Israel Journal of Zoology, 47: 147-160.

KRU IC’, P., ULJEVIC’, A. & NIKOLIC’,V., 2008. The highly invasive alga Cauler-pa racemosa var. cylindracea poses a newthreat to the banks of the coral Clado-cora caespitosa in the Adriatic Sea.Coral Reefs, 27: 441-441.

LANGER, M.R. & HOTTINGER, L., 2000.Biogeography of selected "larger"foraminifera. Micropaleontology, 46(1):105-126.

LAST, P.R. & STEVENS, J.D., 1994. Sharksand rays of Australia. Australia, CSIRO,513 pp.

LEWINSOHN, C.H. & HOLTHUIS, L.B.,1986. The Crustacea Decapoda of Cyprus.Zoologische Verhandelingen, Leiden, 230:1-64.

LIPKIN, Y., 1975. Halophila stipulacea inCyprus and Rhodes, 1967-1970. Aquat-ic Botany, 1: 309-320.

MAAS, O., 1903. Die Scyphomedusen derSiboga Expedition. Siboga Expedition1901, 11(9): 1-91.

MACARTHUR, R.H. & WILSON, E.O.,2001. The theory of island biogeogra-phy. Princeton University Press, 224 pp.

Medit. Mar. Sci., 10/2, 2009, 109-133122

MONTEROSATO, T., 1899. Coquillesmarines de Chypre. Journal de Conchyli-ologie, Paris, 47(4): 392-401.

NISHIKAWA, T., 2002. Revision of thegenus Herdmania (Urochordata:As-cidiacea) inhabiting Japanese waters.Species Diversity, 7: 217-250.

NOFRONI, I. & TRINGALI, L., 1995. Ran-dom notes on eastern Atlantic, Mediter-ranean and lessepsian Pyramidellidae(Gastropoda: Heterobranchia: Pyramidel-loidea). Notiziario del CISMA, 17: 21-49.

NORMAN, J.R., 1929. Notes on the fish-es of the Suez Canal. Proceedings of theZoological Society, London, 2: 616.

OCCHIPINTI-AMBROGI, A., 2007. Glob-al change and marine communities: Alienspecies and climate change. Marine Pol-lution Bulletin 55: 342-352.

OZTURK, B., BUZZURRO, G. & AVNIBENLI, H., 2004. Marine molluscs fromCyprus: new data and checklist. Bollet-tino Malacologico, 39 (5-8): 49-78.

PANCUCCI-PAPADOPOULOU, M.A.,ZENETOS, A., CORSINI-FOKA, M.& POLITOU, C.Y., 2006. Update ofmarine alien species in Hellenic waters.Mediterranean Marine Science, 6(2): 147-158 [2005].

PERRINGS, C., WILLIAMSON, M.,BARBIER, E.B., DELFINO, D.,DALMAZZONE, S., SHOGREN, J.,SIMMONS, P. & WATKINSON, A.,2002. Biological invasion risks and thepublic good: an economic perspective.Conservation Ecology, 6:1-7.

PIAZZI, L., BALESTRI, E., MAGRI, M. &CINELLI, F., 1997. Expansion de l'alguetropicale Caulerpa racemosa (Forsskal)J. Agardh (Bryopsidophyceae, Chloro-phyta) le long de la côte Toscane (Italie).Cryptogamie Algologie, 18: 343-350.

PIMENTEL, D., ZUNIGA, R. &MORRISON, D., 2005. Update on the

environmental and economic costs as-sociated with alien-invasive species inthe United States. Ecological Econom-ics, 52: 273-288.

POR, FD., 1978. Lessepsian migration. Theinflux of Red Sea biota into Mediter-ranean by way of the Suez Canal. In:WD Billings et al (Eds.) Ecological Stud-ies. Analysis and Synthesis, vol. 23, Springer-Verlag, Heidelberg, UNESCO 228 p.

POR, F.D., STEINITZ, H., FERBER, I. &ARON, W., 1972. The biota of the RedSea and the Eastern Mediterranean(1967–1972). A survey of the marine lifeof Israel and surroundings. Israel Jour-nal of Zoology, 21: 459-523.

SANCHEZ-TOCINO, L., HIDALGOPUERTAS, F. & PONTES, M., 2007.First record of Fistularia commersoniiRuppell 1838 (Osteichthyes, Fistularri-dae) in Mediterranean waters of theIberian Peninsula. Zoologica Baetica,18: 79-84.

SANCHEZ VILLAREJO, F., 2007. Melibe'fimbriata' from Cyprus. [Message in] SeaSlug Forum. Australian Museum, Syd-ney. Available from http://www.seaslug-forum.net/find.cfm?id=20207

SCHMIDT, H.E., 1973. Hydromedusae fromthe eastern Mediterranean Sea. IsraelJournal of Zoology, 22: 151-167

SCHMITZ, D.C. & SIMBERLOFF, D.,1997. Biological invasions: a growingthreat. Issues in Science and Technolo-gy, 13(4): 33-40.

SCIBERRAS, M. & SCHEMBRI, P.J., 2007.A critical review of records of alien ma-rine species from the Maltese Islandsand surrounding waters (Central Mediter-ranean). Mediterranean Marine Science,8(1): 41-66.

Si.Di.Mar 2008. La banca dati del SistemaDifesa Mare. http://www.sidimar.tute-lamare.it/distribuzione_alieni.jsp

Medit. Mar. Sci., 10/2, 2009, 109-133 123

STEINITZ, H., 1970. A critical list of im-migrants via the Suez Canal. Biota ofthe Red Sea and Eastern Mediterranean,pp. 59-63.

STEINITZ, H., 1970. A critical list of im-migrants via the Suez Canal. p. 64-74.In: The Hebrew University-SmithsonianInstitution Joint Program: Biota of theRed Sea and Eastern Mediterranean,Progress Report. Dept. Zool., HebrewUniv. Jerusalem [mimeo].

STREFTARIS, N. & ZENETOS, A., 2006.Alien marine species in the Mediter-ranean - the 100 ‘worst invasives’ andtheir impact. Mediterranean MarineScience, 7: 87-118.

STREFTARIS, N., ZENETOS, A. &PAPATHANASSIOU, E., 2005. Glob-alisation in marine ecosystems – Thestory of non-indigenous marine speciesacross European Seas. Oceanographyand Marine Biology: An annual Review,43: 419–453.

TA KIN, E., ÖZTÜRK, M. & KURT, O.,2008. Alien and invasive marine plants ofNorth Cyprus. In: Cicek BA, Oniz H, eds.Proceedings of the 2nd international sym-posium on Underwater Research. EasternMediterranean University, Famagusta,TRN Cyprus, March 2008, pp. 20.

TORCU, H., AKA, Z. & I BILIR, A., 2001.An investigation on fishes of northernCyprus. Turkish Journal of Veterinary andAnimal Sciences, 25: 155-159.

TORNARITIS, G., 1987. Mediterranean SeaShells. Nicosia, Cyprus, published by theauthor.

TZOMOS, T., KITSOS, S., CHRISTO-DOULOU, M., KOUKOURAS, A.& ELEFTHERIOU, A., 2007. Investi-gation of the evolution and dispersionrates of the lessepsian Pisces in theMediterranean. 1st Panhellenic meetingfor the invasion of alien aquatic species

in the eastern Mediterranean, Heraklion,Crete, 5-6 November 2007.

VERLAQUE, M. & BOUDOURESQUE,C.F., 1991. Stypopodium schimperi(Buchinger ex Kützing) Verlaque etBoudouresque comb. nov. (Dictyotales,Fucophyceae), algue de mer Rougerécemment apparue en Méditerranée.Cryptogamie Algologie, 12: 195-211.

VERLAQUE, M., BOUDOURESQUE,C.F., MEINESZ, A. & GRAVEZ, V.,2000. The Caulerpa racemosa complex(Caulerpales, Ulvophyceae) in theMediterranean Sea. Botanica Marina,43: 49-68.

VERLAQUE, M., RUITTON, S., MINEUR,F. & BOUDOURESQUE, C.F., in press.CIESM atlas of exotic species in the Mediter-ranean. Vol. 4 Macrophytes. Monaco,CIESM Publishers.

WHITEHEAD, P.J.P., 1985. FAO speciescatalogue Vol. 7. Clupeoid fishes of theworld (suborder Clupeioidei). An anno-tated and illustrated catalogue of the her-rings, sardines, pilchards, sprats, shads,anchovies and wolf-herrings. Part 1 - Chiro-centridae, Clupeidae and Pristigasteri-dae. FAO Fish. Synop. 125(7/1):1-303.

WIRTZ, P. & DEBELIUS, H., 2003. Mediter-ranean and Atlantic invertebrate guide.Hackenheim, Conchbooks.

YOKES, M.B., 2006. Aplysia dactylomela:an alien opisthobranch in the Mediter-ranean. Journal of the Marine BiologicalAssociation of the UK – Biodiversity Records5299, published online.

ZENETOS, A., 2009. Marine biolo-gical invasions. Technical Report20/2009/RAC/SPA. UNEP/MAP/RAC/SPA, Tunis.

ZENETOS, A. & FRAGGOS, G., 2008.Marine Alien Species: HCMR databasesfor the needs of EEA and UNEP/MAP.International Conference On Marine

Medit. Mar. Sci., 10/2, 2009, 109-133124

Data and Information Systems - IMDIS2008 -Athens (Greece) March 31- April2, 2008, http://hnodc.hcmr.gr/imdis-2008/IMDIS_Presentations/imdis_POSTERS /Session1/IMG_7444.JPG

ZENETOS, A., GOFAS, S., RUSSO, G. &TEMPLADO, J., 2004. CIESM Atlas ofexotic species in the Mediterranean. Vol.3 Molluscs. Monaco, CIESM Publishers.

ZENETOS, A., INAR, M.E., PANCUCCI-PAPADOPOULOU, M.A., HARME-LIN, J.G., FURNARI, G., ANDALORO,F., BELLOU, N., STREFTARIS, N. &ZIBROWIUS, H., 2006. Annotated listof marine alien species in the Mediter-ranean with records of the worst inva-sive species. Mediterranean Marine Science,6(2): 63-118 [2005].

ZENETOS, A., MERI , M., VERLAQUE,M., GALLI, P., BOUDOURESQUE,C.F., GIANGRANDE, A., INAR,M.E. & BILECENO LU, M., 2008.Additions to the annotated list of ma-rine alien biota in the Mediterraneanwith special emphasis on Foraminiferaand Parasites. Mediterranean Marine

Science, 9: 119-165.ZENETOS, A., PANCUCCI-PAPADO-

POULOU, M.A., ZOGARIS, S.,PAPASTERGIADOU, E., VARDA-KAS, L. & ECONOMOU, A.N., in press.Aquatic alien species in Greece: Track-ing sources, patterns and effects on theecosystem. Journal of Biological Research-Thessaloniki: 10.1017/ S1755267209990832;Vol. 2; e156; 2009 Published on line.

ZENETOS, A., KONSTANTINOU, F.,KONSTANTINOU, G., 2009.Towards homogenization of theLevantine alien biota: Additions to thealien molluscan fauna along the Cypri-ot coast. Marine Biodiversit Records:10.1017/S175 526 720 999 0832.

ZIBROWIUS, H., 2002. Assessing scaleand impact of ship-transported alienfauna in the Mediterranean? p. 63-68.In: Alien marine organisms intro-duced by ships in the Mediterraneanand Black Seas, F. Briand (Ed.),Monaco, CIESM Workshop Mono-graphs 20.

Medit. Mar. Sci., 10/2, 2009, 109-133 125

Submitted: July 2009Accepted: August 2009

Published on line: October 2009

Medit. Mar. Sci., 10/2, 2009, 109-133126

AN

NE

X 1

:A

lien

mar

ine

spec

ies

repo

rted

from

Cyp

rus.

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

PHYT

OBE

NTH

OS

Rhod

ophy

taAc

anth

opho

ra n

ayad

iform

is(D

elile)

Pap

enfu

ss19

97In

dian

Oce

anSu

ezCr

ypto

geni

cCI

RIK

et al.

,200

0As

para

gops

is ar

mat

a H

arve

y19

98Au

strali

ash

ippi

ng (f

oulin

g)Ca

sual

CIRI

K et

al., 2

000

Chon

dria

coeru

lesce

ns (J

. Aga

rdh)

Falk

enbe

rg20

08At

lantic

aqua

cultu

reCr

ypto

geni

cK.

TSI

AMIS

, per

s. ob

serv

.G

anon

ema

farin

osum

(J.V

. Lam

ouro

ux) K

.C. F

an

1997

Indi

an O

cean

Suez

Cryp

toge

nic

CIRI

K et

al.,2

000

& Y

ung C

. Wan

gLo

phoc

ladia

lalle

man

dii (

Mon

tagn

e) F

. Sch

mitz

1997

Indo

-Pac

ific

Suez

/ Sh

ippi

ng

Esta

blish

edCI

RIK

et al.

, 200

0(fo

ulin

g)Ne

osiph

onia

spha

eroca

rpa

(B_r

gese

n) M

.S. K

im

2008

Atlan

tic

ship

ping

(fo

ulin

g)Cr

ypto

geni

cK.

TSI

AMIS

, per

s. ob

serv

.&

I.K.

Lee

Polys

iphon

ia a

tlant

ica K

apra

un &

J.N.

Nor

ris20

08N.

Atla

ntic/

Indo

-Pac

ific

ship

ping

Que

stion

able

TASK

IN et

al.,

2008

Polys

iphon

ia fu

coid

es (H

udso

n) G

revil

le20

08N.

Atla

ntic

fishi

ng n

ets

Cryp

toge

nic

TASK

IN et

al.,

2008

Wom

ersley

ella

seta

cea

(Hol

lenbe

rg)

2008

Indo

-Pac

ific

ship

ping

(fou

ling)

Casu

alTA

SKIN

et a

l.,20

08Ch

loro

phyt

aCa

ulerp

a rac

emos

a var

. cyli

ndra

cea (

Sond

er) V

erlaq

ue,

1991

SW A

ustra

liaaq

uariu

mIn

vasiv

eH

ADJI

CHRI

STO

-H

uism

an &

Bou

dour

esqu

ePH

ORO

U et

al.,

1997

Caul

erpa

race

mos

a va

r.lam

ouro

uxii

f.req

uien

ii 19

97In

do-P

acifi

cSu

ezCa

sual

VERL

AQU

E et

al.,2

000

(Mon

tagn

e) W

eber

-van

Bosse

Caul

erpa

race

mos

a va

r. tu

rbin

ata

(J. A

gard

h)

1992

Indo

-Pac

ific

Suez

Cryp

toge

nic

VERL

AQU

E et

al., 2

000

Euba

nk -

uvife

ra (C

. Aga

rdh)

J. A

gard

hCl

adop

hora

cfpa

tentir

amea

(Mon

tagn

e) K

ützin

g 19

91In

do-P

acifi

cSu

ezCa

sual

ARG

YRO

U, 2

000

(con

tinue

d)

Medit. Mar. Sci., 10/2, 2009, 109-133 127

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Phae

ophy

taSt

ypop

odiu

m sc

him

peri

(Buc

hing

er ex

Küt

zing)

19

90Re

d Se

aSu

ezEs

tabl

ished

VERL

AQU

E &

Verla

que &

Bou

dour

esqu

eBO

UD

OU

RESQ

UE,

1991

Sper

mat

ophy

taH

aloph

ila st

ipulac

ea(F

orssk

l) As

cher

son

1967

Red

Sea

Suez

Esta

blish

edLI

PKIN

, 197

5PH

YTO

PLAN

KTO

NPs

eudo

solen

ia ca

lvar-a

vis S

chul

tze, 1

858

1964

Blac

k Se

ash

ippi

ngEs

tabl

ished

KIM

OR

& B

ERD

UG

O, 1

967

FORA

MIN

IFER

AAm

phist

egin

a lo

bifera

Lar

sen,

1976

1976

Indo

-Pac

ific/A

tlant

icSu

ezCr

ypto

geni

cLA

NGER

& H

OTT

ING

ER,

2000

; M. L

ange

r, pe

rs. co

mm

un.

CNID

ARIA

Scyp

hozo

aCa

ssiop

eia a

ndro

med

a (F

orssk

l, 17

75)

1903

Indo

-Pac

ific

Suez

Esta

blish

edM

AAS,

1903

Rhop

ilem

a no

mad

ica G

alil,

1990

1995

Red

Sea

Suez

Esta

blish

edG

ALIL

, 200

6H

ydro

zoa

Laod

icea

fijia

na A

gassi

z & M

ayer

, 189

919

72In

do-P

acifi

cSu

ezQ

uesti

onab

leSC

HM

IDT,

1973

, cite

d in P

OR,

1978

ANNE

LIDA

Bran

chio

mm

a ba

irdi (

McI

ntso

sh, 1

885)

1998

Pacif

ic/At

lantic

Suez

/ship

ping

Esta

blish

edIN

AR, 2

009

Bran

chio

mm

a lu

ctuos

um (G

rube

, 186

9)19

98In

do-P

acifi

cSu

ez/sh

ippi

ngEs

tabl

ished

INAR

, 200

5Ce

rato

nerei

s mira

bilis

Kinb

erg,

1866

1997

Indo

-Pac

ific

Suez

Esta

blish

edIN

AR, 2

005

Eusy

llis k

upffe

riLa

nger

hans

, 187

919

98At

lantic

ship

ping

Esta

blish

edIN

AR &

ERG

EN, 2

003

Hyd

roid

es d

iant

hus(

Verri

ll, 18

73)

1997

North

Wes

t Atla

ntic

ship

ping

Que

stion

able

BEN-

ELIA

HU

& P

AYIA

TAS,

1999

Hyd

roid

es el

egan

s (H

aswe

ll, 18

83)

1996

circu

mtro

pica

lsh

ippi

ngEs

tabl

ished

BEN-

ELIA

HU

& P

AYIA

TAS,

1999

Hyd

roid

es h

etero

ceru

s (G

rube

, 186

8)19

98In

dian

Oce

anSu

ez/sh

ippi

ngEs

tabl

ished

BEN-

ELIA

HU

& P

AYIA

TAS,

1999

Lysid

ice co

llaris

Gru

be, 1

870

1968

Pacif

ic O

cean

Suez

Que

stion

able

BEN-

ELIA

HU

, 197

2

a

a

(con

tinue

d)

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Meta

sych

is go

toi(

Izuk

a, 19

02)

1997

Indo

-Pac

ific

Suez

/ship

ping

Esta

blish

edIN

AR, 2

005

Noto

mas

tus a

bera

ns D

ay, 1

957

1997

Indo

-Pac

ific

Suez

Esta

blish

edIN

AR, 2

005

Noto

mas

tus m

ossa

mbic

us(T

hom

assin

, 197

0)19

97Pa

cific

Oce

anSu

ezEs

tabl

ished

INAR

, 200

5O

enon

e cf.

fulgi

da (S

avign

y, 18

18)

1997

Indo

-Pac

ific

Suez

Casu

alIN

AR, 2

005

Pista

uni

bran

chia

taD

ay, 1

963

1997

Indo

-Pac

ific

Suez

Esta

blish

edIN

AR, 2

005

Pseu

done

reis a

nom

ala(G

ravie

r, 19

00)

1969

Indo

-Pac

ific

Suez

/ship

ping

Inva

sive

BEN-

ELIA

HU

, 197

2Li

noph

erus c

anar

iensis

Lan

gerh

ans,

1881

1997

Atlan

ticSh

ippi

ngEs

tabl

ished

INAR

, 200

9Sp

haero

syllis

long

ipapil

lata

Har

tman

n-Sc

hröd

er &

19

97Pa

cific

Oce

anSu

ez/sh

ippi

ngCa

sual

INAR

et a

l.,20

03H

artm

ann,

1979

Spiro

bran

chus

tetra

cero

s(Sc

hmar

da, 1

861)

1996

circu

mtro

pica

lSu

ez/sh

ippi

ngEs

tabl

ished

BEN-

ELIA

HU

& P

AYIA

TAS,

1999

Spiro

rbis

mar

ioni

Cau

llery

and

Mes

nil,

1997

1998

East

Pacif

icsh

ippi

ngCa

sual

ZIBR

OW

IUS,

2002

Tereb

ella

ehren

bergi

Gru

be, 1

870

1969

Indo

-Pac

ific

Suez

/ship

ping

Que

stion

able

BEN-

ELIA

HU

, 197

2CR

USTA

CEA

Isop

oda

Apan

thur

a sa

ndale

nsis

Steb

bing

, 190

019

98So

uth

Afric

aSu

ezEs

tabl

ished

H. Z

IBRO

WIU

S, p

ers c

omm

.Am

phip

oda

Mae

ra h

amige

ra H

aswe

ll, 18

7919

97In

do-P

acifi

cSu

ezEs

tabl

ished

KOCA

TAet

al.,2

001

Deca

poda

Callin

ectes

sapid

us R

athb

un, 1

896

1964

Wes

t Atla

ntic

Gib

ralta

r/shi

ppin

gEs

tabl

ished

DEM

ETRO

POU

LOS

&

NEO

CLEO

US,

1969

Char

ybdi

s hell

eri (A

. Miln

e-Ed

ward

s, 18

67)

1998

Indo

-Wes

t Pac

ific

Suez

/ship

ping

Inva

sive

GAL

IL et

al.,

2002

Char

ybdi

s lon

gicol

lisLe

ene,

1938

1986

Indi

an O

cean

Suez

Esta

blish

edLE

WIN

SOH

N &

HO

LTH

UIS

, 198

6M

arsu

pena

eus j

apon

icus (

Bate

, 188

8)19

61In

do-P

acifi

cSu

ez/aq

uacu

lture

Esta

blish

edD

EMET

ROPO

ULO

S &

NEO

CLEO

US,

1969

Meta

pena

eops

is ae

gypt

ia G

alil &

Gol

ani,

1990

2004

Indo

Pac

ific

Suez

Esta

blish

edKI

RMIR

TZO

GLO

U et

al.,

2006

Medit. Mar. Sci., 10/2, 2009, 109-133128

(con

tinue

d)

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Meta

pena

eus m

onoc

eros (

Fabr

icius

, 179

8)19

61In

do-W

est P

acifi

cSu

ezEs

tabl

ished

DEM

ETRO

POU

LOS

&NE

OCL

EOU

S, 19

69Pi

lum

nope

us va

uque

lini (

Audo

uin,

1826

)19

63In

dian

Oce

anSu

ezEs

tabl

ished

GAL

IL, 2

006

Portu

nus

pelag

icus (

Linn

aeus

, 175

8)19

58In

do P

acifi

cSu

ez/sh

ippi

ngEs

tabl

ished

ELTO

N, 19

58Th

alam

ita p

oisso

nii(

Audo

uin,

1826

)19

69In

do-W

est P

acifi

cSu

ezCr

ypto

geni

cG

ILAT

E in

PO

R, 19

78;

GIT

ARAK

OS

et al.

,200

7St

omat

opod

aEr

ugos

quilla

mas

save

nsis

(Kos

sman

n, 18

80)

1963

Indi

an O

cean

Suez

Esta

blish

edIN

GLE

, 196

3M

OLL

USCA

Gas

trop

oda

Acteo

cina

muc

rona

ta(P

hilip

pi, 1

849)

1992

Red

Sea

Suez

Casu

alCE

CALU

PO &

QU

ADRI

, 199

4Al

vani

a do

rbign

yi (A

udou

in, 1

826)

1985

Cosm

opol

itan

Suez

?Cr

ypto

geni

cBO

GI e

t al.,

1989

Aplys

ia d

actyl

omela

Ran

g, 18

2820

04Ci

rcum

tropi

cal

unkn

own

Esta

blish

edYO

KES,

2006

Bulla

am

pulla

Lin

naeu

s, 17

5820

00In

do-P

acifi

cSu

ez/sh

ippi

ngCa

sual

ZENE

TOS

et al.

, 200

4Cl

athr

ofen

ella

perp

arvu

la (W

atso

n, 18

86)

1995

Indo

-Pac

ific

Suez

Casu

alCE

CALU

PO &

QU

ADRI

, 199

5Ce

rithi

opsis

pul

vis (M

elvill

, 189

6)19

85Re

d Se

aSu

ezCa

sual

BOG

I et a

l.,19

89Ce

rithi

opsis

tent

hren

ois P

ilsbr

y & V

anna

ta, 1

906

1985

Indi

an O

cean

Suez

?Es

tabl

ished

TORN

ARIT

IS, 1

987

Cerit

hium

nes

iotic

um P

hilip

pi, 1

848

1985

Indi

an O

cean

Suez

?Ca

sual

BOG

I et a

l.,19

89Ce

rithi

um sc

abrid

um (W

atso

n 18

86)

1983

Indi

an O

cean

Suez

/ship

ping

Inva

sive

FISC

HER

, 199

3Ch

rysall

ida

mai

ae (H

ornu

ng &

Mer

mod

, 192

4)19

95Re

d Se

aSu

ezCa

sual

BUZZ

URO

& G

REPP

I, 19

97Ci

ngul

ina

isseli

(Try

on, 1

886)

1998

Subt

ropi

cal

Suez

?Ca

sual

ZENE

TOS

et al.

, 200

4Cy

closc

ala h

yalin

a (S

ower

by, 1

844)

1992

Indo

Pac

ific

unkn

own

Casu

alCE

CALU

PO &

QU

ADRI

, 199

4Cy

lichn

ina

girar

di (A

udou

in, 1

826)

1996

Indo

-Pac

ific

Suez

Casu

alCE

CALU

PO &

QU

ADRI

, 199

6Er

galat

ax ju

nion

ae H

ouar

t, 20

0819

93In

dian

Oce

anSu

ezEs

tabl

ished

BUZZ

URO

& G

REPP

I, 19

97Fi

nella

pup

oide

s A. A

dam

s, 18

6019

96In

do-P

acifi

cSu

ezEs

tabl

ished

CECA

LUPO

& Q

UAD

RI, 1

996

Medit. Mar. Sci., 10/2, 2009, 109-133 129

(con

tinue

d)

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Leuc

otin

a na

talen

sis S

mith

, 191

019

96In

do P

acifi

cSu

ezCa

sual

CECA

LUPO

& Q

UAD

RI, 1

996

Meli

be vi

ridis

(Kela

art,

1858

)20

07In

do-P

acifi

csh

ippi

ngCa

sual

SANC

HEZ

VIL

LARE

JO, 2

007

Meta

xia b

acillu

m (I

ssel,

1869

)<

1995

Red

Sea

Suez

?Ca

sual

CECA

LUPO

& Q

UAD

RI, 1

995

Purp

urad

usta

grac

ilis n

otat

a (G

ill, 1

858)

2000

Indi

an O

cean

Suez

?Es

tabl

ished

ZENE

TOS

et al.

,200

4Py

runc

ulus

four

ierii

(Aud

ouin

, 182

6)19

95In

do P

acifi

cSu

ezCa

sual

BUZZ

URO

& G

REPP

I, 19

97Rh

inoc

lavis

koch

i (Ph

ilipp

i, 18

48)

1976

Indo

Pac

ific

Suez

Esta

blish

edD

EMET

ROPO

ULO

S &

HAD

JICH

RIST

OPH

ORO

U, 1

976

Risso

ina

berth

ollet

i Isse

l, 18

6919

85In

dian

Oce

anSu

ezCa

sual

BOG

I et a

l.,19

89Sm

arag

dia

souv

erbia

na (M

ontro

uzier

, 186

3)19

95In

do P

acifi

cSu

ezCa

sual

BUZZ

URO

& G

REPP

I, 19

97St

icteu

lima

cf. le

ntigi

nosa

(A. A

dam

s, 18

61)

1995

Indo

-Pac

ific

Suez

Casu

alBU

ZZU

RO &

GRE

PPI,

1997

Stro

mbu

s pers

icus S

wain

son,

1821

(Roe

ding

, 179

8)19

85In

dian

Oce

anSu

ez/sh

ippi

ngIn

vasiv

eBA

ZZO

CCH

I, 19

85Sy

rnol

a fa

scia

ta (J

ickeli

, 188

2)19

95In

do-P

acifi

cSu

ez?

Casu

alNO

FRO

NI &

TRI

NGAL

I, 19

95Th

ais l

acera

(von

Bor

n, 17

78)

1988

Indi

an O

cean

Suez

/ship

ping

Casu

alH

OU

ART,

2001

Troc

hus e

rithr

eus B

rocc

hi, 1

821

1985

Indi

an O

cean

Suez

Inva

sive

TORN

ARIT

IS, 1

987

Turb

onilla

edga

rii B

rocc

hi, 1

821

1996

Indo

-Pac

ific

Suez

?Ca

sual

CECA

LUPO

& Q

UAD

RI, 1

996

Zafra

savig

nyi (

Moa

zzo,

1939

)19

95Re

d Se

aSu

ezEs

tabl

ished

BUZZ

URO

& G

REPP

I, 19

97Za

fra se

lasph

ora

(Melv

ill &

Sta

nden

, 190

1)19

95In

dian

Oce

anSu

ez?

Casu

alBU

ZZU

RO &

GRE

PPI,

1997

Biva

lvia

Brac

hido

ntes

pha

raon

is (F

ische

r P., 1

870)

1960

Indi

an O

cean

Suez

/ship

ping

Inva

sive

H. M

IENI

S, p

ers.

com

m.;

TORN

ARIT

IS, 1

987

Cham

a pa

cifica

Bro

derip

, 183

419

98In

do-P

acifi

cSu

ezEs

tabl

ished

ZENE

TOS

et al.

, 200

4De

ndro

strea

fron

s (Li

nnae

us, 1

758)

2008

Indo

-Pac

ific

ship

ping

Esta

blish

edZE

NETO

S et

al.,2

009

Fulvi

a fra

gilis

(For

sskl i

n Ni

ebuh

r, 17

75)

1983

Indi

an O

cean

Suez

/ship

ping

Esta

blish

edFI

SCH

ER, 1

993

Gaf

rariu

m p

ectin

atum

(Lin

naeu

s, 17

58)

2005

Indo

-Pac

ific

Suez

Esta

blish

edZE

NETO

S et

al.,2

009

Malv

ufun

dus r

egul

us (F

orssk

l, 17

75)

1970

Indo

-Pac

ific

Suez

Esta

blish

edD

EMET

ROPO

ULO

S, 19

71a

a

Medit. Mar. Sci., 10/2, 2009, 109-133130

(con

tinue

d)

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Paph

ia te

xtile

(Gm

elin,

1791

)20

04In

do-P

acifi

cSu

ezEs

tabl

ished

ZENE

TOS

et al.

,200

9Pi

ncta

da ra

diat

a (L

each

, 181

4)18

99In

do-P

acifi

cSu

ez/aq

uacu

lture

Inva

sive

MO

NTER

OSA

TO, 1

899

Psam

mot

reta

prae

rupt

a (S

alisb

ury,

1934

)20

09In

do-P

acifi

cSu

ez?

Casu

alZE

NETO

S et

al.,2

009

Sept

ifer f

orsk

ali (D

unke

r, 18

55)

2005

Indi

an O

cean

Suez

/ship

ping

Esta

blish

edZE

NETO

S et

al.,2

009

Spon

dylu

s spin

osus

Sch

reib

ers,

1793

2001

Indo

-Pac

ific

Suez

/ship

ping

Esta

blish

edZE

NETO

S et

al.,2

009

ECH

INO

DERM

ATA

Hol

othu

roid

eaSy

napt

ula

recipr

ocan

s (Fo

rssk

l, 17

75)

1986

Indo

-Pac

ific

Suez

Esta

blish

edCH

ERBO

NNIE

R, 19

86O

phiu

roid

eaO

phia

ctis p

arva

Mor

tens

en, 1

926

<20

02Re

d Se

aSu

ezEs

tabl

ished

ZIBR

OW

IUS,

2002

Oph

iacti

s sav

ignyi

Mül

ler &

<

2002

Circ

umtro

pica

lSu

ezEs

tabl

ished

ZIBR

OW

IUS,

2002

Tros

chel,

1842

SIPU

NCUL

APh

asco

loso

ma

scol

ops (

Selen

ka &

de M

an, 1

883)

1998

Trop

ical/s

ubtro

pica

lSu

ez/sh

ippi

ngEs

tabl

ished

ACIK

et a

l.,20

05CH

ORD

ATA

Ascid

iace

aH

erdm

ania

mom

us (S

avign

y 181

6)19

98In

do-P

acifi

cSu

ez/sh

ippi

ngEs

tabl

ished

NISH

IKAW

A, 20

02

PISC

ESCh

ondr

ichth

yes

Him

antu

ra u

arna

k (F

orssk

l, 17

75)

1994

Indo

-Wes

t Pac

ific

Suez

Que

stion

able

LAST

& S

TEVE

NS, 1

994

Ost

eicht

hyes

Alep

es d

jedab

a (F

orssk

l, 17

75)

1964

Indo

-Pac

ific

Suez

/ship

ping

Esta

blish

edD

EMET

ROPO

ULO

S &

NEO

CLEO

US,

1969

Apog

on p

hara

onis

Bello

tti, 1

874

1964

Indo

-Pac

ific

Suez

Esta

blish

edD

EMET

ROPO

ULO

S &

NEO

CLEO

US,

1969

a

a

a

Medit. Mar. Sci., 10/2, 2009, 109-133 131

(con

tinue

d)

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Athe

rinom

orus

lacu

nosu

s (Fo

rster

in B

loch

19

29In

do-P

acifi

cSu

ezEs

tabl

ished

NORM

AN, 1

929;

and

Schn

eider

, 180

1)D

EMET

ROPO

ULO

S &

NE

OCL

EOU

S, 19

69Du

ssum

ieria

elop

soid

es B

leeke

r, 18

49

1985

Indo

-Pac

ific

Suez

Esta

blish

edW

HIT

EHEA

D, 1

985;

GIT

ARAK

OS

et al.

2007

Ench

elyco

re an

atin

a (L

owe,

1838

)20

08At

lantic

Gib

ralta

rCa

sual

G. I

OAN

NOU

& N

. M

ICH

AILI

DIS

, per

s. ob

serv

.Et

rum

eus t

eres (

DeK

ay, 1

848)

1999

Indo

-Pac

ific

Suez

Esta

blish

edG

OLA

NI, 2

000

Fistu

laria

com

mers

onii

(Rüp

pell,

1835

)19

99In

do-P

acifi

cSu

ezIn

vasiv

eW

IRTZ

& D

EBEL

IUS,

2003

Hem

iram

phus

far (

Forss

kl,

1775

)19

64In

do-P

acifi

cSu

ezEs

tabl

ished

DEM

ETRO

POU

LOS

&NE

OCL

EOU

S, 19

69H

erklo

tsich

thys

pun

ctatu

s(Rü

ppell

, 183

7)19

85Re

d Se

aSu

ezCa

sual

WH

ITEH

EAD

, 198

5La

goce

phalu

s sce

lerat

us (G

meli

n 17

89)

2004

Indo

-Pac

ific

Suez

Inva

sive

DFM

R (2

006)

Lago

ceph

alus s

padi

ceus

(Rich

ards

on, 1

844)

2006

Indo

-Pac

ific

Suez

Esta

blish

edG

. IO

ANNO

U &

N.

MIC

HAI

LID

IS, p

ers.

obse

rv.

Lago

ceph

alus s

ueze

nsis

Clar

k &

Goh

ar, 1

953

2007

Red

Sea

Suez

Esta

blish

edG

. IO

ANNO

U &

N.

MIC

HAI

LID

IS, p

ers.

obse

rv.

Leio

gnat

hus k

lunz

inge

ri (S

tein

dach

ner,

1898

)19

61Re

d Se

aSu

ezEs

tabl

ished

FOD

ERA,

1961

Parex

ocoe

tus m

ento

(Vale

ncien

nes,

1846

)<

2002

Indo

-Pac

ific

Suez

Esta

blish

edG

OLA

NI et

al.,

2002

Pem

pheri

s van

icolen

sis (C

uvier

1831

)19

95In

do-P

acifi

cEs

tabl

ished

TORC

U et

al.,

2001

Ptera

gogu

s pely

cus R

anda

ll, 19

8119

97In

do-P

acifi

cSu

ezEs

tabl

ished

KAYA

et a

l.,20

00Sa

rgoce

ntro

n ru

brum

(For

sskl,

1775

)19

61In

do-P

acifi

cSu

ezEs

tabl

ished

FOD

ERA,

1961

Saur

ida

undo

squa

mis

(Rich

ards

on, 1

848)

1960

Indo

-Pac

ific

Suez

Esta

blish

edBE

N TU

VIA,

1962

Scom

bero

mor

us co

mm

erson

(Lac

epèd

e, 18

00)

2008

Indo

-Pac

ific

Suez

Esta

blish

edG

. IO

ANNO

U &

N.

MIC

HAI

LID

IS, p

ers.

obse

rv.

a

a

Medit. Mar. Sci., 10/2, 2009, 109-133132

(con

tinue

d)

Medit. Mar. Sci., 10/2, 2009, 109-133 133

Year

of

MAR

INE

SPEC

IES

first

Orig

inM

ade o

fEs

tabl

ishm

ent

Refe

renc

esig

htin

gIn

trod

uctio

nSu

cces

s

Siga

nus l

urid

us (R

üppe

ll, 18

29)

1964

Indo

-Pac

ific

Suez

Inva

sive

DEM

ETRO

POU

LOS

&NE

OCL

EOU

S, 19

69Si

ganu

s rivu

latus

For

sskl,

1775

1928

Indo

-Pac

ific

Suez

Inva

sive

NORM

AN, 1

929

Silla

go si

ham

a (F

orssk

l, 17

75)

2009

Indo

-Pac

ific

Suez

Casu

alG

. IO

ANNO

U &

N.

MIC

HAI

LID

IS, p

ers.

obse

rv.

Spho

eroid

es p

achy

gaste

r(M

üller

& T

rosc

hel,

1848

)20

05Ci

rcum

globa

l G

ibra

ltar

Esta

blish

edG

. IO

ANNO

U &

N.

(tem

pera

te an

d M

ICH

AILI

DIS

, per

s. ob

serv

.tro

pica

l sea

s)Sp

hyra

ena

chrys

otae

nia

Klun

zinge

r, 18

8419

64In

do-P

acifi

cSu

ezEs

tabl

ished

DEM

ETRO

POU

LOS

&NE

OCL

EOU

S, 19

69St

epha

nolep

is di

aspr

osFr

aser

-Bru

nner

, 194

019

35In

do-P

acifi

cSu

ez/sh

ippi

ngEs

tabl

ished

HO

RNEL

L, 19

35; G

ITAR

AKO

Set

al.,2

007

Upen

eus m

oluc

cens

is(B

leeke

r, 18

55)

1964

Indo

-Pac

ific

Suez

Esta

blish

edD

EMET

ROPO

ULO

S &

NEO

CLEO

US,

1969

Upen

eus p

oriB

en-T

uvia

& G

olan

i, 19

8920

04In

do-P

acifi

cSu

ezEs

tabl

ished

TZO

MO

S et

al.,2

007

aa