HCMR TEYXOS 6 ergasia 359 - Mediterranean Marine Science
Transcript of HCMR TEYXOS 6 ergasia 359 - Mediterranean Marine Science
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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alien species, technical reports, scientific congresses, academic dissertations,and websites, as well as on unpublished/personal observations.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
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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-
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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-
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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).
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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-
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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-
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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.
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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-
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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-
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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.
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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