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

Review Article

Alien species in the Mediterranean Sea by 2010. A contribution to the applicationof European Union’s Marine Strategy Framework Directive (MSFD).
Part I. Spatial distribution
A. ZENETOS1, S. GOFAS2, M. VERLAQUE3, M.E. INAR4, J.E. GARCI’A RASO5,
C.N. BIANCHI6, C. MORRI6, E. AZZURRO7, M. BILECENOGLU8, C. FROGLIA9,
I. SIOKOU10, D. VIOLANTI11, A. SFRISO12, G. SAN MART N13, A. GIANGRANDE14,
T. KATA AN4, E. BALLESTEROS15, A. RAMOS-ESPLA’ 16, F. MASTROTOTARO17,
O. OCA A18, A. ZINGONE19, M.C. GAMBI19 and N. STREFTARIS10
1

Institute of Marine Biological Resources, Hellenic Centre for Marine Research, P.O. Box 712,
19013 Anavissos, Hellas2
Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Ma’ laga,
E-29071 Ma’ laga, Spain
3
UMR 6540, DIMAR, COM, CNRS, Université de la Méditerranée, France
4
Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100 Bornova, Izmir, Turkey
5
Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Ma’ laga,
E-29071 Ma’ laga, Spain
6
DipTeRis (Dipartimento per lo studio del Territorio e della sue Risorse), University of Genoa,
Corso Europa 26, 16132 Genova, Italy
7
Institut de Ciències del Mar (CSIC) Passeig Mar tim de la Barceloneta, 37-49, E-08003 Barcelona, Spain
8
Adnan Menderes University, Faculty of Arts & Sciences, Department of Biology, 09010 Aydin, Turkey
9c\o CNR-ISMAR, Sede Ancona, Largo Fiera della Pesca, 60125 Ancona, Italy
10
Institute of Oceanography, Hellenic Centre for Marine Research, P.O. Box 712, 19013 Anavissos, Hellas
11
Dipartimento di Scienze della Terra, University of Turin, via Valperga Caluso 35, 10125 Torino, Italy
12
Department of Environmental Sciences, University of Venice, Calle Larga 2137, 30123 Venice, Italy
13
Departamento de Biolog a (Zoolog a), Facultad de Ciencias, Universidad Auto’ noma de Madrid,calle Darwin, 2, 28049 Madrid, Spain
14
Department of Biological and Environmental Sciences and Technologies, University of Lecce,
Complesso Ecotekne, Via Prov. le Lecce-Monteroni, 73100 Lecce, Italy
15
Centre d’Estudis Avanç ats de Blanes (CSIC), Acc. Cala, Sant Francesc 14, 17300 Blanes, Girona, Spain
16
Centro de Investigacio’ n Marina de Santa Pola (CIMAR), Universidad de Alicante, 03130 Santa
Pola, Alicante, Spain
17
Department of Biology, University of Bari, Via Orabona 4, 70125 Bari, Italy
18
Departamento Oceanograf a Biolo’ gica y Biodiversidad, Fundacio’ n Museo del Mar Muelle~Can
onero Dato S/N 51001, Ceuta, Spain
19
Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy
Corresponding author: zenetos@ath.hcmr.gr

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

381

Abstract
The state-of-art on alien species in the Mediterranean Sea is presented, making distinctions
among the four subregions defined in the EU Marine Strategy Framework Directive: (i) the Western
Mediterranean Sea (WMED); (ii) the Central Mediterranean Sea (CMED); (iii) the Adriatic Sea(ADRIA); and (iv) the Eastern Mediterranean Sea (EMED). The updated checklist (December2010) of marine alien species within each subregion, along with their acclimatization status and origin, is provided. A total of 955 alien species is known in the Mediterranean, the vast majority of themhaving being introduced in the EMED (718), less in the WMED (328) and CMED (267) and least in
the Adriatic (171). Of these, 535 species (56%) are established in at least one area.
Despite the collective effort of experts who attempted in this work, the number of introduced
species remains probably underestimated. Excluding microalgae, for which knowledge is still insufficient, aliens have increased the total species richness of the Mediterranean Sea by 5.9%. This figure
should not be directly read as an indication of higher biodiversity, as spreading of so many aliens
within the basin is possibly causing biotic homogenization. Thermophilic species, i.e. Indo-Pacific,
Indian Ocean, Red Sea, Tropical Atlantic, Tropical Pacific, and circum(sub)tropical, account for
88.4% of the introduced species in the EMED, 72.8% in the CMED, 59.3% in the WMED and
56.1% in the Adriatic. Cold water species, i.e. circumboreal, N Atlantic, and N Pacific, make up asmall percentage of the introduced species, ranging between 4.2% and 21.6% and being more
numerous in the Adriatic and less so in the EMED.
Species that are classified as invasive or potentially invasive are 134 in the whole of the Mediterranean: 108 are present in the EMED, 75 in the CMED, 53 in the Adriatic and 64 in the WMED.
The WMED hosts most invasive macrophytes, whereas the EMED has the lion’s share in polychaetes, crustaceans, molluscs and fish.

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

Introduction
The Mediterranean Sea is one of theseas of the world most affected by biological invasions in terms of how long theinvaders have been present (OCCHIPINTI-AMBROGI, 2000; STREFTARISet al., 2005), in number of alien species
detected (COSTELLO et al., 2010) and in
the unprecedented rate of introduction(ZENETOS, 2009; 2010). Triggered bywarming waters and a newly improved
route through the Suez Canal,
tropical/subtropical marine species, somevery invasive, have progressively moved
into the Mediterranean, disrupting
ecosystem stability of the basin.
382

In 1978, there were 128 proven
Lessepsian immigrants and 76 doubtful
cases (POR, 1978). Since the first review
of alien species in the Mediterranean(ZIBROWIUS, 1992), there have been
many papers and reviews dealing with thissubject. More recently, CIESM has published four Atlases of Exotic Species in
the Mediterranean (GALIL et al., 2002;
GOLANI et al., 2002; ZENETOS et al.,
2004; VERLAQUE et al., in press). The
most recent lists show 116 species of fishes, 70 species of decapod and stomatopod
crustaceans, 137 species of molluscs and110 species of macrophytes. This
increased number of records, particularly
over the past few years, has been partly
Medit. Mar. Sci., 11/2, 2010, 381-493

attributed to intensification of research
effort in this topic. ZENETOS &POLYCHRONIDIS (2010), based ondata gathered, analysed for and adopted
in the European Environment Agency(EEA) and UNEP MAP RAC/SPA
reports, estimated that nearly 1000
species entered the Mediterranean during the past century.
A recent synthesis on Mediterraneanmarine biodiversity (COLL et al., 2010)
described the Mediterranean Sea as a
biodiversity hot spot hosting approximately 17,000 marine species, of whichmore than 600 (3.3%) are alien. Thesemost recent detailed biodiversity estimates for alien species show a discrepancy from the figures provided by
ZENETOS et al. (2005; 2008). However,as COLL et al. (2010) have pointed out,
the true numbers of alien species are certainly biased downwards. The data arepresumably accurate for large and conspicuous species that are easily distinguished from the native biota and forspecies that occur along a frequently sampled (or fished) coast and for which taxonomic expertise is readily available, but
are entirely absent for many of the small
members of invertebrates. This underrepresentation is common in many works
and efforts have been made to include
less studied groups in recent reviews (seeZENETOS et al., 2008).
An up-to-date inventory of the alien
species in the Mediterranean, apart fromits scientific merits, can fulfil the needs ofthe regulatory requirements and environmental management options. This is ofparticular importance since the current

emergence of the new generation of EU
political actions covering major maritime
strategic objectives, such as the Marine
Strategy Framework Directive (MSFD)(2008/56/EC), the European Strategy for
Marine and Maritime Research (COM(2008) 534) encompassing the Marine
Spatial Planning and the Ecosystem
Approach (ECAP) within the Barcelona
Convention by UNEP/MAP1, as well as
many initiatives of the UNEP RAC/SPA.
Alien species regulations are of major
importance in those policies. In the current MSFD the descriptor: "Non-indigenous species introduced by human activities are at levels that do not adversely
alter the ecosystems" is actually one of theeleven qualitative descriptors for determining Good Environmental Status(GES). It is also closely related to six
other GES descriptors because of thegreat variety of impacts that Invasive
Alien Species (IAS) may exert on native
biodiversity, ecosystem functioning, andseabed habitats as well as commercial
marine resources (OLENIN et al., 2010).
This territorial and institutional scenario raises the need for a multi-levelgovernance system with a quite complex
coordination system in which non-EU
Member States can also participate. Thisis particularly important in the Mediterranean, where the southern and eastern
coast belong to non-EU Member States
or countries with accession status (e.g.
Turkey and Israel). Furthermore, theBarcelona Convention of the UNEP/MAP
supports the introduction of MSFD
(2009) concepts that can be usefulthroughout the basin despite the handi-

1

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

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

383

cap that only seven of 21 Mediterraneancountries are Member States of
UNEP/MAP.
Considering a) the high introduction
rate, b) recent publications and reviewson little studied taxa and c) the need toassess the extent of the phenomenon atsubregional level and inform the EU,
UNEP/MED and other stakeholders
accordingly, this work aims to present the2010 state-of-art on the presence andacclimatization status of alien species at
each MSFD Mediterranean subregion,
namely: (i) the Western Mediterranean
Sea (WMED); (ii) the Central Mediterranean Sea (CMED); (iii) the AdriaticSea (ADRIA); and (iv) the Eastern
Mediterranean Sea (EMED). The latteris also commonly referred to as the
Aegean-Levantine basin, but in this work
the Marmara Sea, bearing more similarities to the Aegean than to the neighbouring Black Sea, is also included.

Methodology
Biological invasions have become ahot issue at a broad geographical scale
such as that of Large Marine Ecosystems(LME). Among European LMEs, biological invasions are most pronounced in the
Mediterranean (EEA, 2007a; ZENETOS
et al., 2009).
On the European scale, over a dozen
partitions can be found dividing up thewaters of the NE Atlantic, the Mediterranean Sea and the Black Sea on oceanographic or purely geometric bases, or a
combination of both, as well as divisions
based on national jurisdictions2 (Fig. 1).
In this work the Mediterranean is being
2

studied along the four subregions as
adopted and described under the MSFD.
The coastal areas of the countries and theregions included in these subregions arelisted in Table 1. This division imposessome restrictions/difficulties in the case ofcountries whose waters lie within theboundaries of these subregions as in thecase of Albania, Tunisia and Italy.
The borders of the CMED are hereby
defined as the Kythira-Anti-Kythira
Straits (Greece) and Libya-Egypt borders
to the east, Otranto Strait (Italy, Albania)
to the north, Cap Bon (Tunisia) andsouth-west Sicily to the west.
The list is updated based on valid
species records up to December 2010.
Alien species are investigated within all
four regions in terms of their establishment success (casual, established, questionable, cryptogenic), as well as invasive
success, their native range and geographical expansion. The species lists are presented for seven units which are systematic groups namely: 1) Protozoa (including
Foraminifera), 2) Macrophytes, 3) Polychaeta, 4) Crustacea, 5) Mollusca, 6) Miscellaneous Invertebrata (Arthropoda
Pycnogonida, Bryozoa, Chaetognatha,
Chordata
Ascidiacea,
Cnidaria,
Ctenophora, Echinodermata, Porifera,
Sipuncula, Nematoda and Platyhelminthes), and 7) Fish, in each part.
The introduction of alien species in the
Mediterranean Sea is hard to demonstrate
for marine microalgae as for microrganisms in general (WYATT & CARLTON,
2002). Bearing in mind that it can rarely be
excluded that a suspected microalgal invader was already present as part of the rare,hidden and unsampled phytoplankton, we
refrain from citing a detailed list of plank-

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

384

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

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

Table 1
Subregions, areas and coast of the Mediterranean studied in this work.
Western
Mediterranean
Tyrrhenian Sea
Ligurian Sea
Sardinia
CorsicaFranceMonacoSpain
Baleares
Gibraltar
MoroccoAlgeria
North Tunisia
West Sicily

Central Mediterranean
including
the Ionian Sea
Greek Ionian Sea
Italian Ionian Sea
Albanian Ionian Sea
Apulian Sea
South East Sicily
Malta
South Tunisia
Libya

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

Adriatic Sea

Italian Adriatic Sea
Slovenia
Croatia
Montenegro
Albanian Adriatic Sea

Eastern Mediterranean(the Aegean-Levantine Marmara Seas)
Greek Aegean Sea
Turkish Aegean Sea
Sea of Marmara
South Turkey
Cyprus
Syria
Lebanon
Palestine Authority
IsraelEgypt

385

tonic and benthic microalgae as in
ZENETOS et al. (2005, 2008).
The revised checklist has been as wide
as possible. Contrary to the CIESM Atlas,
we have taken into account alien species
introduced from elsewhere within the
Mediterranean when the introduction
event was evident (e.g. the mollusc
Siphonaria pectinata introduced from the
WMED to Greece, or the algae Cladosiphon zosterae, Desmarestia viridis,
Ectocarpus siliculosus var. hiemalis andPylaiella littoralis from the Adriatic to theThau Lagoon, WMED).
Freshwater species occurring in estuarine waters such as Acipenser gueldenstaedtii,
Acipenser baeri and Huso huso in the North
Aegean estuaries and Micropterus salmoides
in Ionian Sea estuarine waters are not included in our list. Similarly, the freshwater crayfish Procambarus clarki, though it can standslightly brackish waters and has been reported in the inner part of the Varano Lagoon,
Adriatic Sea (FLORIO et al., 2008) and in
the Palude di Torre Flavia, Tyrrhenian Sea(SCALICI et al., 2010), is not included.
Alien [synonyms: Non-indigenous
species (NIS), exotic, non-native,
allochthonous] are species, subspecies orlower taxa, present in the wild, introduced
outside of their natural range (past or
present) and beyond their natural dispersal potential. This includes any part,
gamete or propagule of such species thatmight survive and subsequently reproduce. Their presence in the given regionis due to intentional or unintentional
introduction resulting from human activities. Natural shifts in distribution ranges(e.g. due to climate change or dispersal byocean currents) do not qualify a species as
a NIS (OLENIN et al., 2010). Specimens
kept in captivity and specimens still
attached to a ship’s hull or other man386

made crafts are not considered.
The acclimatization status of eachspecies was assessed, and is given hereaccording to the following terminology.
Established: Introduced or feral population of species settled in the wild with
free-living, self-maintaining and self-perpetuating populations unsupported by andindependent of humans. Species with atleast two records in the area spread overtime and space (at least three records forfishes) are also classified as established, inthe sense of the CIESM Atlas series.
Casual: Casual species are those having been recorded only once (no morethan twice for fish) in the scientific andgrey literature and are presumed to be
non-established in the area. In this paper‘casual’ is used in the same sense as ‘alien’in the CIESM Atlas series.
Questionable: Species with insufficient information–‘suspect’. This includes
old casual records that have not been subsequently found despite appropriate
investigation, and also new entries notverified by experts or species with taxonomic status unresolved.
Cryptogenic: Species with no definite
evidence of their native or introduced status according to CARLTON (1996) andspecies whose probable introduction
occurred in ‘early times’ and has not been
witnessed (e.g. prior to 1800).
Invasive: Species defined as establishedaliens that have overcome biotic and abiotic barriers and are able to disseminate awayfrom their area of initial introduction
through the production of fertile offspringwith noticeable impact, such as threat to thediversity or abundance of native species, theecological stability of infested ecosystems,
economic activities dependent on theseecosystems, and human health.
True aliens need to be separated from
Medit. Mar. Sci., 11/2, 2010, 381-493

species with seemingly isolated records,
which have naturally spread to regions
beyond their usual range. BOUCHET &TAVIANI (1992) illustrated this scenario
with the case of deep-sea molluscs in the
Mediterranean, brought into the basin aslarvae carried by the inflowing surface
waters, but unable to establish fertile populations. This is also the case of someEastern Atlantic crustacean species seldom recorded in the West Mediterranean
but occurring in the Alboran sea (Penaeopsis serrata, Merhippolyte ancistrota, etc.),
and of the big-eye thresher shark, Alopias
superciliosus, which is distributed worldwide in tropical and temperate seas andwas formerly classified as alien in the
Mediterranean (ZENETOS et al., 2008).
Occurrences in this kind of context havebeen filtered out of alien species lists.
Regarding the native range, caution has
been taken in establishing the origin of alien
species. The true origin for many species ismuddled after becoming widely dispersed
over a long time. Precise localities will beknown for some, but for others only a general region may be known. This is particularly important in the case of the Mediterranean where the true origin of populationsof a species widely distributed in the IndoPacific Ocean may be its population in the
Red Sea, or much further afield. The
expression "Lessepsian migration" wascoined by POR (1978) for those species that
inched their way through the Suez Canal
into the Mediterranean. However it is far
from straightforward that all the Indo-Pacific species documented in the Mediterranean
are Lessepsian immigrants, and at least two
more pools of species can be individualized.
One such pool would include tropical IndoPacific species that did not first show up on
the Levantine or Egyptian coasts, but further away from the Suez Canal.
Medit. Mar. Sci., 11/2, 2010, 381-493

Finally, attention was paid to recent
nomenclatural updates. These are theresult of the latest taxonomic and/or
molecular studies, such those of LAI et al.
(2010) on the Portunus pelagicus species
complex; of HUBER (2010) on Anadarakagoshimensis, formerly known as Anadarainaequivalvis; and of MALAQUIAS &REID (2008) on Bulla arabica, formerly
known as Bulla ampulla. A list of all
species catalogued under their currently
valid names along with the old names isprovided in the ANNEX.
Nomenclature adopted in this paper
follows the World Register of Marine
Species (WoRMS <www.marinespecies.
org>) and contributing databases(AlgaeBase, CLEMAM, FishBase…).
The time boundary is generally set fromthe date of the opening of the Suez Canal(1869): species established earlier are treated as cryptogenic (e.g. Teredo navalis).
Results and Discussion
The core of this work is Table 2, which
provides a full list of alien species withineach Mediterranean sub-basin, along with
their acclimatization status and origin. Furthermore, this part is built around eightsections. The first four deal in detail withthe distribution of the seven systematicgroups within each MSFD area. They are
structured in a way so as to be autonomousand therefore understood by the independent taxonomist reader. The fifth section
describes the state of art of phytoplankton
within the Mediterranean LME. The last
three sections are more general andattempt to discuss some trends from theelaboration of Table 2. A list of invasive or
potentially invasive species for each basin is
included. The possible role of warming in
the observed trends is also addressed.
387

Table 2
List of species with origin and establishment success in all Mediterranean MSFD areas(WMED=Western Mediterranean, CMED=Central Mediterranean, ADRIA=Adriatic Sea,
EMED=Eastern Mediterranean). Species in alphabetic order within each taxon.
Establishment success abbreviated as est=established, cas=casual, cry=cryptogenic, que=questionable.
One asterisk denotes planktonic form. Two asterisks denote recent name changes. P= Parasite.
Species

Author

WMED CMED ADRIA EMED

OriginPROTOZOA
Miscellaneous Protozoa
Bonamia ostreae(Pichot et al., 1979)
Circumboreal
Marteilia refringens
Cavalier-Smith, 2002
unknown
Perkinsus olseni**
Lester & Davis, 1981
Cosmopolitan
Photobacterium damselae NE Atlantic
Foraminifera
Agglutinella arenata(Said, 1949)
Indo-PacificAgglutinella compressa
El-Nakhal, 1983
Indo-PacificAgglutinella robusta
El-Nakhal, 1983
Indo-PacificAgglutinella soriformis
El-Nakhal, 1983
Indo-Pacific/Atlantic
Amphisorus hemprichii
Ehrenberg, 1840
CircumtropicalAmphistegina lessoniid'Orbigny, 1826
CircumtropicalAmphistegina lobifera
Larsen, 1976
CircumtropicalAmphistegina
d’Orbigny, 1826
Indo-Pacificmadagascariensis
Archais angulatus(Fichtel & Moll, 1798)
W Atlantic
Articulina alticostata
Cushman, 1922
Indo-PacificAstacolus insolitus(Schwager, 1866)
Indo-PacificAstacolus sublegumen(Parr, 1950)
Indo-PacificBorelis sp.
Circumtropical
Brizalina simpsoni(Heron-Allen & Earland, 1915) Indo-Pacific
Clavulina angularisd' Orbigny, 1826
Circumtropical
Clavulina cf. multicamerata Chapman, 1907
Indo-PacificCoscinospira hemprichii
Ehrenberg, 1839
Indo-PacificCushmanina striatopunctata (Parker & Jones, 1865)
Circumtropical
Cycloforina sp.
Indian/Red Sea
Cyclorbiculina compressa (d'Orbigny, 1839 )Circumtropical
Cymbaloporetta plana(Cushman, 1924)
Indo-Pacific
Edentostomina cultrata(Brady, 1881)
Indo-Pacific
Elphidium cf. charlottensis (Vella, 1957)
Indo-Pacific
Elphidium striatopunctatum (Fichtel & Moll, 1798)
Indo-PacificEuthymonacha polita(Chapman, 1904)
Indo-PacificHaddonia sp.
Chapman, 1898
Indo-Pacific
Hauerina diversa
Cushman, 1946
Circumtropical
Heterocyclina tuberculata (Moebius, 1880)
Indian
388

cas
cas
cry
est

?

cas
cas
cry
?

?
cas
cas
cas
cas
est
est
est
est

est?
est?
est
cry/cascry/cascry/cascry/estcry/cas

est
est
est
est
cas
cry/estest
cry/estcas
est
est
cry/est
est
est
est
cas
est
est
est

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

Table 2 (continued)
Species
Heterostegina depressa
Miliolinella cf. hybrida
Nodophthalmidium
antillarum
Operculina ammonoides
Pegidia lacunata
Peneroplis antillarum

Authord’Orbigny, 1826
(Terquem, 1878)(Cushman, 1922)

OriginCircumtropicalIndo-PacificCircumtropical

Red Sea
Indo-Pacific
Caribbean or SW
Atlantic
Planogypsina acervalis(Brady, 1884)
Circumtropical
Planogypsina squamiformis (Chapman, 1901)
Circumtropical
Planorbulinella larvata(Parker & Jones, 1865)
Indo-PacificPseudolachlanella slitella Langer, 1992
Indo-PacificPseudomassilina reticulata (Heron-Allen & Earland, 1915) Indo-PacificPulleniatina obliquiloculata* (Parker & Jones, 1865)
Circumtropical
Pyramidulina catesbyi(d'Orbigny, 1839)
Indo-Pacific/Atlantic
Pyrgo denticulata(Brady, 1844)
Indo-Pacific
Quinqueloculina cf.
Said, 1949
Indo-Pacificmosharrafai
Schackoinella imperatoria d’Orbigny, 1846
Indo-Pacific
Schlumbergerina(Brady, 1879)
Circumtropicalalveoliniformis
Sorites orbiculus
Forssk a l, 1775
Circumtropical
Sorites variabilis
Lacroix, 1941
Indo-PacificSpiroloculina antillarumd'Orbigny, 1839
Circumtropical
Spiroloculina cf. angulata Cushman, 1917
Indo-Pacific
Triloculina fichtelianad'Orbigny, 1839
Circumtropical

WMED CMED ADRIA EMED
est
est
est

(Gronovius, 1781)
McCulloch, 1977
d’Orbigny, 1839

cas
est
cas
cry

cry

cry
est
cas
est
est
cas
est
est
est

cry/cascry/cascry/casest

cry/estcry/cascry/est
est
est
cas
est

MACROPHYTESChlorophyta
Batophora sp.
Atlantic
Caulerpa distichophylla
SonderIndo-PacificCaulerpa mexicana
Sonder ex Kützing
Indo-PacificCaulerpa racemosa var.
(Sonder) Verlaque, Huisman Indo-Pacificcylindracea& BoudouresqueCaulerpa racemosa var.
(Montagne) Weber-van
Indo-Pacificlamourouxii f. requienii
Bosse
Caulerpa racemosa var.
(J. Agardh) Eubank/Indo-Pacificturbinata /uvifera(C. Agardh) J. Agardh
Caulerpa scalpelliformis(Brown ex Turner) C. Agardh Indo-PacificCaulerpa taxifolia(Vahl) C. Agardh
Indo-PacificCladophora cf. patentiramea (Montagne) Kützing
Indo-PacificCladophora herpestica(Montagne) Kützing
Indo-Pacific
Medit. Mar. Sci., 11/2, 2010, 381-493

ques
est
est

est

est

est
est
est

est

est

cry/ques

cry/ques
est

est

est

est
est
est
389

Table 2 (continued)
Species
Cladophora hutchinsioides
Codium fragile subsp.
fragile**Codium parvulum
Codium taylorii
Derbesia boergesenii

Author
Hoek & Womersley(Suringar) Hariot

OriginPacific
NW Pacific(Bory ex Audouin) P.C. Silva
P.C. Silva(Lyengar & Ramanathan)
Mayhoub
YamadaDickieDelileKjellman(Kützing) Gayral

Indo-PacificIndo-PacificIndo West Pacific

Derbesia rhizophora
NW Pacific
Neomeris annulata
Indo-PacificUlva fasciata
Cosmopolitan
Ulva pertusa
Indo-PacificUlvaria obscura
NW Pacific
Fucophycae
Acrothrix gracilis
Kylin
Circumboreal
Botryella cf. parva(Takamatsu) H.-S. Kim
Pacific
Chorda filum(Linnaeus) Stackhouse
N Atlantic/N PacificCladosiphon zosterae(J. Agardh) Kylin
Atlantic
Colpomenia peregrina
Sauvageau
Indo-PacificDesmarestia viridis(O.F. Müller) J.V. Lamouroux Atlantic/Pacific
Ectocarpus siliculosus(P.L. Crouan & H.M.
Atlantic
var. hiemalis
Crouan) Foslie
Fucus spiralis
LinnaeusAtlantic
Halothrix lumbricalis(Kützing) Reinke
N Atlantic/N Pacific
Leathesia marina(Lyngbye) Decaisne
Cosmopolitan
Microspongium tenuissimum (Hauck) A.F. Peters
Atlantic
Padina antillarum(Kützing) Piccone
Indo-Pacific
Padina boergesenii
Allender & Kraft
Indo-Pacific
Padina boryana
Thivy in W.R. Taylor
Indo-PacificPunctaria tenuissima(C. Agardh) Greville
NE Atlantic
Pylaiella littoralis(Linnaeus) Kjellman
N Atlantic/N Pacific
Rugulopteryx okamurae(Dawson) I.K. Hwang,
Pacific
W.J. Lee & H.S. Kim
Saccharina japonica**(Areschoug) C.E. Lane,NW Pacific
C. Mayes, Druehl &G.W. Saunders
Sargassum muticum(Yendo) Fensholt
NW Pacific
Scytosiphon dotyi
Wynne
NE PacificSpatoglossum variabile
Figari & De Notaris
Indo-PacificSphaerotrichia firma(Gepp) A.D.Zinova
NW PacificStypopodium schimperi(Kützing) VerlaqueIndo West Pacific& Boudouresque390

WMED CMED ADRIA EMED
cas
est
est
est
est
est
est
cas

est

est
est
est
est

est
cry/ques cry/ques cry/ques
est

est

est
est
est
est
est
est

cry/ques
est

cas
est
est
est

cry/ques cry/ques cry/ques
est
est

est
est
est
est

est

est
cry/ques
est

cry/ques
est cry/ques

cas
est
est

est
est

est
cry/ques cry/ques cry/ques

cas

est
est

est
est

est
est

cas
est
est

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

Table 2 (continued)
Species
AuthorOriginUndaria pinnatifida(Harvey) Suringar
Pacific
Magnoliophyta
Halophila stipulacea(Forssk a l) Ascherson
Red SeaPelagophyceae
Chrysonephos lewisii(W.R.Taylor) W.R.Taylor W Atlantic
Rhodophyta
Acanthophora nayadiformis (Delile) Papenfuss
Indo-PacificAcrochaetium codicola
B rgesen
Atlantic/PacificAcrochaetium robustum
B rgesenIndo-PacificAcrochaetium spathoglossi B rgesen
Indo-PacificAcrochaetium subseriatum B rgesen
Indo-PacificAcrothamnion preissii(Sonder) E.M. Wollaston
Indo-PacificAgardhiella subulata(C. Agardh) Kraft &Atlantic/Pacific
M.J. Wynne
Aglaothamnion feldmanniae Halos
N Atlantic
Ahnfeltiopsis flabelliformis (Harvey) Masuda
Pacific
Anotrichium okamurae
Baldock
NW Pacific
Antithamnion amphigeneum A. Millar
SW Pacific
Antithamnion hubbsii
DawsonIndo-Pacific
Antithamnionella boergesenii (Cormaci & G. Furnari)
Indo-PacificAthanasiadis
Antithamnionella elegans (Berthold) J.H. Price &unknown
D.M. John
Antithamnionella(Schiffner) E.M. Wollaston Indo-Pacificspirographidis
Antithamnionella sublittoralis (Setchell & Gardner)
Pacific
Athanasiadis
Antithamnionella ternifolia (J.D. Hooker & Harvey) Lyle unknown
Apoglossum gregarium(E.Y. Dawson) M.J. Wynne PacificAsparagopsis armata
HarveySW Pacific

WMED CMED ADRIA EMED
est
cas
est

Asparagopsis taxiformis
DelileAsparagopsis taxiformis
invasive strain
Bonnemaisonia hamifera Hariot
Botryocladia
Feldmann-Mazoyer
madagascariensis
Caulacanthus okamurae** Yamada
Ceramium bisporum
D.L. Ballantine
Ceramium strobiliforme
G.W. Lawson & D.M. John
Chondria coerulescens(J. Agardh) Falkenberg
Chondria curvilineata
F.S. Collins & Hervey

Atlantic
Indo-PacificIndo-PacificIndiancry/ques cry/ques
est
est
est
est
est
est

NW Pacific
W Atlantic
N Atlantic
E Atlantic
Atlantic

est
cry/ques

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

est

est

est

est

est
cry/ques cry/ques cry/ques cry/ques
est
est
est
est
cas
cas
cas
est
est
cas
est
est
est
est
est
est
cry/ques cry/ques cry/ques cry/ques
est
est
est
cry/ques cry/quescry/ques cry/ques cry/ques cry/ques
est

est
est
est

est
est

est

est

est

est

est
est

est

est
est
est

Requiresconfirmation

Requiresconfirmation
cry/ques
est
est
est

cry/ques
cry/ques cry/quescry/ques cry/ques cry/quesest391

Table 2 (continued)
Species
AuthorChondria pygmaea
Garbary & Vandermeulen
Chondrus giganteus f.
Mikamiflabellatus
Chrysymenia wrightii(Harvey) Yamada
Dasya sessilis
YamadaDasysiphonia sp.
Yendo(= Heterosiphonia japonica)
Galaxaura rugosa(J. Ellis & Solander)
J.V. Lamouroux
Ganonema farinosum(J.V. Lamouroux) K.C. Fan& Yung C. Wang
Goniotrichiopsis sublittoralis G.M. Smith
Gracilaria arcuata
Zanardini

OriginIndo-PacificPacific

Gracilaria vermiculophylla
Grateloupia asiatica
Grateloupia lanceolata
Grateloupia minima

NW Pacific
NW PacificPacific
Pacific

est
est
est

Pacific

cas

Pacific
NW PacificAtlantic/PacificIndo-Pacific
Red Sea
Cosmopolitan
Indo-Pacific

est
est
est
est

Grateloupia patens
Grateloupia subpectinata
Grateloupia turuturu
Griffithsia corallinoides
Herposiphonia parca
Hypnea anastomosans**
Hypnea cornuta
Hypnea flagelliformis
Hypnea flexicaulis
Hypnea spinella
Hypnea valentiae
Laurencia caduciramulosa
Laurencia okamurae
Lithophyllum yessoense
Lomentaria flaccida
Lomentaria hakodatensis
Lophocladia lallemandii
Nemalion vermiculare
Neosiphonia harveyi

392(Ohmi) Papenfuss
Kawaguchi & Wang
(Okamura) Kawaguchi
P.L. Crouan & H.M.
Crouan(Okamura) S. Kawaguchi& H.W. Wang
HolmesYamada(Linnaeus) Batters
SetchellPapenfuss, Lipkin & Silva(Kützing) J.Agardh
Greville ex J. Agardh

WMED CMED ADRIA EMED
est
est
est
est

Pacific
Pacific
NW Pacific

est
est
est

est

Red Sea

est

Indiancry/ques cry/ques

NW PacificIndo-Pacific/Red Sea

Yamagishi & Masuda
Indo-Pacific(C. Agardh) Kützing
Pantropical(Turner) Montagne
Red Sea
Masuda & Kawaguchi
SW PacificYamadaPacific
FosliePacific
TanakaPacific
Yendo
Pacific(Montagne) F. Schmitz
Indo-Pacific
Suringar
NW Pacific(J. Bailey) M.-S. Kim, H.-G. NW PacificChoi, Guiry & G.W. Saunders

cry/ques

est
Requiresconfirmation

Requiresconfirmation
est

cas

est
est

est
est

cas
est

est
est
est
est
est
cas
est
est
est
est

est
est
Requiresconfirmation
est
est
cas

est
est

est

est
est

est

est

est

est

est
est
est

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

Table 2 (continued)
Species
AuthorNitophyllum
Okamurastellato-corticatum
Osmundea oederi(Gunnerus) G. Furnari
Plocamium secundatum(Kützing) Kützing
Polysiphonia atlantica
Kapraun & J.N. Norris
Polysiphonia fucoides(Hudson) Greville
Polysiphonia morrowii
HarveyPolysiphonia paniculata
Montagne
Polysiphonia stricta(Dillwyn) Greville
Porphyra yezoensis
Ueda
Pterosiphonia tanakae
Uwai & Masuda
Rhodophysema georgei
Batters
Rhodymenia erythraea
Zanardini
Sarconema filiforme(Sonder) Kylin
Sarconema scinaioides
B rgesen
Solieria dura
(Zanardini) F. Schmitz
Solieria filiformis(Kützing) Gabrielson
Spongoclonium caribaeum** (B rgesen) M.J. Wynne
Symphyocladia(Harvey) Falkenberg
marchantioides
Womersleyella setacea(Hollenberg) R.E. Norris

OriginPacific

WMED CMED ADRIA EMED
est

Atlantic
cry/ques cry/questropical/subtropicalest
est
N Atlantic/N Pacific cry/ques cry/ques cry/ques cry/ques
N Atlantic
est cry/ques cry/ques cry/ques
NW Pacific
est
est
est
E Pacific
est
cas
cas
NE Atlantic
estPacific
est
Pacific
est
Atlantic/Pacificcas
casIndo West PacificcasIndo West PacificcasestIndianest
Indo-Pacificcas
N Atlantic
est
est
est
W Atlantic
est
Indo-Pacificest
Indo-Pacific

est

est

cas
cas

cas

est
est

est
est

est

est

POLYCHAETA
Amphicorina pectinata
Apoprionospio pygmaea**
Branchiomma bairdi
Branchiomma boholensis
Branchiomma luctuosum
Capitellethus dispar
Ceratonereis mirabilis
Chaetozone corona
Cirriformia semicincta
Cossura coasta
Dasybranchus carneus
Desdemona ornata
Dispio magnus
Dispio uncinata
Dodecaceria capensis
Dorvillea similis
Eunice antennata
Eunice cf. cariboea
Eunice floridana(Banse, 1957)(Hartman, 1955)(McIntosh, 1885)(Grube, 1878)(Grube, 1869)(Ehlers, 1907)
Kinberg, 1866
Berkeley & Berkeley, 1941
(Ehlers, 1905)
Litamori, 1960
Grube, 1870
Banse, 1957
(Day, 1955)
Hartman, 1951
Day, 1961(Crossland, 1924)(Savigny, 1820)
Grube, 1856
(Pourtalès, 1867)

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

Pacific
Pacific
Atlantic/PacificIndo-PacificIndo-PacificIndo-Pacific/Red Sea
Indo-Pacificunknown
Red Seaunknown
Red Sea
Indo-PacificIndianW Atlantic
IndianIndo-PacificIndo-Pacific
W Atlantic
W Atlantic

est

cas

est
cas

est
ques

est
ques
cas
ques
est

est
est
est
ques
est
cry
ques
cry/ques
ques
est

ques
est
est

ques
393

Table 2 (continued)
Species
AuthorEpidiopatra hupferiana
Augener, 1918
hupferina
Epidiopatra hupferiana
Day, 1957
monroi
Eunice indica
Kinberg, 1865
Erinaceusyllis serratosetosa (Hartmann-Schroder, 1982)
Euniphysa tubifex(Crossland, 1904)
Eurythoe complanata(Pallas, 1766)
Eusyllis kupfferi
Langerhans, 1879
Exogone africana
Hartmann-Schröder, 1974
Exogone breviantennata
Hartmann-Schröder, 1959
Fabriciola qhardaqa
Banse, 1959
Ficopomatus enigmaticus (Fauvel, 1923)
Glycinde bonhourei
Gravier, 1904
Haploscoloplos kerguelensis Mc Intosh, 1885
Hesionura serrata(Hartmann-Schroder, 1960)
Hyboscolex longiseta
Schmarda, 1861
Hydroides albiceps
Grube, 1870
Hydroides branchyacanthus Rioja, 1941
Hydroides dianthus(Verrill, 1873)
Hydroides diramphus
Mörch, 1863
Hydroides elegans(Haswell, 1883)
Hydroides heterocerus(Grube, 1868)
Hydroides homocerus
Pixell, 1913
Hydroides minax(Grube, 1878)
Hydroides operculatus(Treadwell, 1929)
Hydroides steinitzi
Ben-Eliahu, 1972
Janua (Dexiospira) steueri (Sterzinger, 1909)
Isolda pulchella
Muller, 1858
Laonome elegans
Gravier, 1906
Laonome triangularis
Hutchings & Murray, 1984
Leiochrides australis
Augener, 1914
Leocrates chinensis
Kinberg, 1866
Leonnates decipiens
Fauvel, 1929
Leonnates indicus
Kinberg, 1966
Leonnates persicus
Wesenberg-Lund, 1949
Lepidonotus carinulatus(Grube, 1870)
Lepidonotus tenuisetosus (Gravier, 1902)
Linopherus canariensis** Langerhans, 1881
Longibranchium atlanticum (Day, 1973)
Loimia medusa(Savigny, 1818)
394

OriginTropical Atlantic

WMED CMED ADRIA EMED
cas

Indiancas

Indo-PacificPacific
IndianAtlantic/PacificAtlantic
Indo-PacificCircumtropical
Red Sea
SubtropicalIndo-Pacific
Antarctic
Red Sea
Cosmopolitan
Indo Pacific/Red Sea
Indo-Pacific
NW Atlantic
Circumtropical
CircumtropicalIndian/Red Sea
IndianIndo-Pacific/Red Sea
IndianRed Sea
Indo-PacificCircumtropical
Red Sea
SW PacificPacific
Pacific
Indo-PacificIndo-PacificIndo-PacificIndo-Pacific/Red Sea
Indo-Pacific/Red Sea
Atlantic
Atlantic
Cosmopolitan

ques
est
ques
ques
est
cas
est

ques

est
ques
cas
ques
cas
est
est
est

est

cas
est

ques

est
est
est

est
est

ques

est
est

cas

est
est
est
est
est
est
est
est
cas
cas

cas
cas
est
cas
ques

cas
ques

cas

est
cas
ques

cas
ques

ques
est
est
est
ques
cas
est
ques

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

Table 2 (continued)
Species
AuthorOriginWMED CMED ADRIA EMED
Lumbrineris acutifrons(Gallardo, 1967)
Pacific
ques
cas
Lumbrineris neogesae
Miura, 1980
S Africa
cas
cas
Lumbrineris perkinsi**
Carrera-Parra, 2001
Indo-Pacific
est
est
est
Lysidice collaris
Grube, 1870
Pacific/Red Sea
est
est
est
est
Lysidice natalensis(Kinberg, 1865)
Indo-Pacificques
Marphysa disjuncta
Hartman, 1961
Pacific
cas
Mediomastus capensis
Day, 1961
Indian
ques
ques
ques
Megalomma claparedei
Gravier, 1908
Red Seacas
Metasychis gotoi(Izuka, 1902)
Indo-Pacific
est
est
est
est
Naineris quadraticeps
Day, 1965
Red Seaques
Neopseudocapitella
Rullier & Amoureux, 1979 W Atlantic/
est
est
est
est
brasiliensis
Red Sea
Neanthes agulhana(Day, 1963)
S Africa
est
est
Neanthes willeyi(Day, 1934)
Indo-Pacificcas
Nereis gilchristi
Day, 1960
Red Seacas
Nereis jacksoni
Kinberg, 1866
Indo-Pacificest
est
Nereis persica
Fauvel, 1911
Indo-Pacificques
estNotomastus aberansDay, 1957
Indian/Red Sea
est
est
est
est
Notomastus mossambicus (Thomassin, 1970)
Indianest
est
Notopygos crinita
Grube, 1855
W Atlanticques
Novafabricia infratorquata Fitzhugh, 1983
W Atlantic
est
est
A
Ophryotrocha diadema
kesson, 1976
Pacific
est
Ophryotrocha japonica
Claparède & Mecznikow, 1968 Pacific
est
est
est
Oenone cf. fulgida(Savigny, 1818)
Indo-Pacific/Red Sea ques
ques
Onuphis eremita oculata Hartman, 1951
W Atlantic
estPalola valida(Gravier, 1900)
Red Sea
est
Paradyte cf. crinoidicola(Potts, 1910)
Indo-Pacificques
Paraehlersia weissmaniodes (Augener,1913)
Indo-Pacificcas
Paraprionospio coora
Wilson, 1990
PacificcrycryPerinereis nuntia(Savigny, 1818)
Indiancas
Pherusa parmata(Grube, 1878)
Indo-PacificcasPherusa saldanhaDay, 1961
Indiancas
Pileolaria berkeleyana(Rioja, 1942)
E Pacific
estTropical Atlantic
cas
Pisione guanche
San Mart›n, Lo’ pez &Nu’ ~nez, 1999
Pista unibranchia
Day, 1963
Indo-Pacificest
estPlatynereis australis(Schmarda, 1861)
Pacific
ques
Podarkeopsis capensis(Day, 1963)
Indo-Pacificcasques
Polycirrus twisti
Potts, 1928
Suez Canal
est
Polydora colonia
Moore 1907
W Atlanticcascas
Polydora cornuta
Bosc, 1802
CosmopolitanestestMedit. Mar. Sci., 11/2, 2010, 381-493

395

Table 2 (continued)
Species
Polydora spongicola
Pomatoleios kraussii
Prionospio (Aquilaspio)
krusadensis
Prionospio (Aquilaspio)
sexoculata
Prionospio (Prionospio)
depauperata
Prionospio (Prionospio)
saccifera
Prionospio (Prionospio)
paucipinnulata
Prionospio (Minuspio)
pulchraProtodorvillea egena
Protodorvillea biarticulata
Prosphaerosyllis
longipapillata**
Pseudonereis anomala
Pseudopolydora
paucibranchiata
Scoletoma debilis
Sigambra constricta
Sigambra parva
Spirobranchus tetraceros
Spirorbis marioni
Streblosoma comatus**
Streblospio
gynobranchiata
Syllis alosae
Syllis bella
Syllis hyllebergi
Syllis cf. mayeri
Syllis pectinans
Synelmis rigida
Syllis schulzi
Terebella ehrenbergi
Timarete anchylochaeta
Timarete caribous
Timarete dasylophius
Timarete punctata
396

AuthorBerkeley & Berkeley, 1950
(Baird, 1865)
Fauvel, 1929

OriginWMED CMED ADRIA EMED
Pacific
ques
Indo-Pacific/Red Sea
est
Indo West Pacificest

Augener, 1918

Atlantic/PacificestImajima, 1990

NE Pacific

est

Mackie & Hartley, 1990

Indo-Pacificest

Blake & Kudenov, 1978

Pacific

est

Imajima, 1990

Atlantic/Pacificcas(Ehlers, 1913)
Indian/Red Sea
Day, 1963
Indian(Hartmann-Schröder, 1979) SW Pacific

ques
ques

(Gravier, 1900)
Okuda, 1937

Indo-PacificPacific

est(Grube, 1878)(Southern, 1921)(Day, 1963)(Schmarda, 1861)
Caullery & Mesnil, 1897
(Grube, 1856)
Rice & Levin, 1998

Indo-PacificIndo-Pacific/Red Sea
IndianCircumtropicalAtlantic/PacificIndo-Pacific
W Atlantic

San Mart›n, 1992
(Chamberlin, 1919)(Licher, 1999)
Musco & Giangrande, 2005
Haswell, 1920
(Fauvel, 1919)(Hartmann-Schröder, 1962)
Grube, 1870
(Schmarda, 1861)(Grube, 1859)(Marenzeller, 1879)(Grube, 1859)

W Atlantic
Pacific
Red Sea
W Atlantic
Pacific
Indo-PacificIndian/Red Sea
Indo-Pacific/Red Sea
Pacific
W Atlantic
Indo-PacificIndo-Pacific

ques
cas
est
est
ques
ques

cas
est
est

cas

est
est
est
est

ques
est
est
est
ques
ques

ques
est
ques
ques
ques
ques
cas
ques
est

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

Table 2 (continued)
Species

AuthorAmphipoda
Bemlos leptocheirus(Walker, 1909)
Caprella scaura
Templeton, 1836
Cymadusa filosa(Savigny, 1816)
Elasmopus pectenicrus(Bate, 1862)
Gammaropsis togoensis(Schellenberg, 1925)
Linguimaera caesaris**
Krapp-Schickel, 2003
Monocorophium sextonae (Crawford, 1937)
Parhyale explorator
Arresti, 1989
Photis lamellifera
Schellenberg, 1928
Rhabdosoma whitei
Bate, 1862
Stenothoe gallensis
Walker, 1904
Cirripedia
Amphibalanus eburneus** (Gould, 1841)
Austrominius modestus** Darwin, 1854
Balanus reticulatus
Utinomi, 1967
Balanus trigonus
Darwin, 1854
Boschma, 1960
Heterosaccus dollfusi p
Megabalanus tintinnabulum (Linnaeus, 1758)
Tetraclita squamosa
Pilsbry, 1916
rufotinta
Copepoda
Acartia centrura
Giesbrecht, 1889
Acartia fossae
Gurney, 1927
Acartia tonsa
Dana, 1849
Arietellus pavoninus
Calanopia biloba
Calanopia elliptica
Calanopia media
Calanopia minor
Canuellina insignis
Centropages furcatus
Clavellisa ilishaep
Enhydrosoma vicinum
Euchaeta concinna
Labidocera agilis
Labidocera detruncata
Labidocera madurae
Labidocera orsinii(G. O. Sars, 1905)
Bowman, 1957
(Dana, 1846)
Gurney, 1927
A. Scott, 1902
Gurney, 1927
(Dana, 1852)
Pillai, 1962
Por, 1967
Dana, 1849
(Dana, 1849)(Dana, 1849)(A. Scott, 1909)
Giesbrecht, 1889

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

OriginCRUSTACEA
IndianIndianIndo-PacificCircumtropicalCosmopolitan
Indo-Pacificunknown
NE Atlantic
IndianRed Sea
CircumtropicalW Atlantic
Tropical PacificCircumtropical
Circumtropical
Red Sea
CircumtropicalIndo-PacificIndianIndo-Pacific
W Atlantic /Indo-PacificTropical-subtropicalW subtropical Atlantic
Indo-PacificIndo-PacificIndo-PacificIndo-PacificIndo-Pacific/Atlantic
IndianIndo-PacificIndo-PacificIndo-PacificIndo-PacificIndo-Pacific
Red Sea

WMED CMED ADRIA EMED

cas
est
cas?

est
cry
est

est
est

est
cry

cry

ques

est

est
est
ques
est

est

est
est

est

est

ques

est

ques
cas

est

est
cas

ques

cas
ques

cry
est
cry/est
est
cry
cas
cas
cas
est
est
est
est
est
est

cas
est
est
est
cas
est
est
cas
ques
est
est
ques
cas
cas
cas
est
cas
397

Table 2 (continued)
Species
AuthorLabidocera pavo
Giesbrecht, 1889
Metacalanus acutioperculum Ohtsuka, 1984
(Pillai, 1964)
Mitrapus oblongusp
Hoshina & Sigiura, 1953
Myicola ostreaep
Mori, 1935
Mytilicola orientalisppNothobomolochus fradei Marques, 1965
Paracalanus indicus
Wolfenden, 1905
Paracartia grani
G. O. Sars, 1904
Parvocalanus crassirostris** Dahl, 1894
Parvocalanus elegans
Andronov, 1972
Parvocalanus latus
Andronov, 1972
Pseudocyclops xiphophorusp Wells, 1967
Pseudocalanus elongatus (Boeck, 1865)
Robertsonia salsa
Gurney, 1927
Scaphocalanus amplius
Park, 1970
Scaphocalanus brevirostris Park, 1970

OriginWMED CMED ADRIA EMED
Indo-Pacificest
Pacific
est
est
Indianest
Pacific
est
Pacific
est
Atlantic/Indian
est
Subtropical Atlantic, cry/ques
Red SeaAtlantic
est
est
est
est
Indo-Pacific/Atlanticcas
est
Indo-PacificcasIndiancas
Indiancry
E Atlantic
cry
cry
cry
Indo-Pacificques
Subtropical Atlanticcas
Subtropical Atlantic/
cas
IndianSubtropical AtlanticcasIndo-Pacificques
Antarctic
ques
ques
Indo-Pacificques
Indo-Pacificques
Atlantic/Pacific
ques
ques
ques
Red Sea
est

Scolecithrix valens
Farran, 1926
Scottolana longipes(Thompson & Scott, 1903)
Spinocalanus terranovae Damkaer, 1975
Stenhelia inopinata(A. Scott, 1902)
Stenhelia minuta(A. Scott, 1902)
Subeucalanus subcrassus** Giesbrecht, 1888
Triconia hawii(Böttger-Schnack& Boxshall, 1990)
Triconia rufa
(Boxshall & Böttger, 1987) Indian/Red Sea
Triconia umerus(Böttger-Schnack
Red Sea& Boxshall, 1990)
Cumacea
Eocuma rosae
Corbera & Galil, 2007
Indo-Pacific/Red Sea
Eocuma sarsii(Kossmann, 1880)
Indo-Pacific
Scherocumella gurneyi(Calman, 1927)
Red Sea
Decapoda
Actumnus globulus
Heller, 1861
Red Sea
Alpheus audouini
Coutiere, 1905
Indo West Pacific
Alpheus inopinatus
Holthuis & Gottlieb, 1958 Indian/Red Sea
Alpheus migrans
Lewinsohn & Holthuis, 1978 Red Sea
Alpheus rapacida
de Man, 1908
Indo West PacificAshtoret lunaris(Forssk a l, 1775)
Indo West PacificAtergatis roseus(Rüppell, 1830)
Indo-Pacific
398

est
est

cas

ques
cas

cas
est
cas

est

cas
cas

est
est
est
est
cas
est

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

Table 2 (continued)
Species
AuthorCalappa hepatica(Linnaeus, 1758)
Calappa pelii
Herklots, 1851
Callinectes danae
Smith, 1869
Callinectes sapidus
Rathbun, 1896
Carupa tenuipes
Dana, 1851
Charybdis feriata(Linnaeus, 1758)
Charybdis japonica(A. Milne-Edwards, 1861)
Charybdis helleri(A. Milne-Edwards, 1867)
Charybdis longicollis
Leene, 1938
Charybdis lucifera(Fabricius, 1798)
Coleusia signata(Paulson, 1875)
Cryptosoma cristatum
Brulle, 1837
Daira perlata(Herbst, 1790)
Dorippe quadridens(Fabricius, 1793)
Dyspanopeus sayi
(Smith, 1869)
Eriocheir sinensis
H. Milne Edwards, 1853
Eucrate crenata
de Haan, 1835
Eurycarcinus integrifrons de Man, 1879
Farfantepenaeus aztecus(Ives, 1891)
Fenneropenaeus merguiensis (De Man, 1888)
Glabropilumnus laevis** (Dana, 1852)
Gonioinfradens paucidentata (A. Milne Edwards, 1861)
Grapsus granulosus
H. Milne-Edwards, 1853
Halimede tyche(Herbst, 1801)
Hemigrapsus sanguineus (de Haan, 1835)
Herbstia nitida
Manning & Holthuis, 1981
Hyastenus hilgendorfi
de Man, 1887
Ixa monodi
Holthuis & Gottlieb, 1956
Leptochela aculeocaudata Paulson, 1875
Leptochela pugnax
de Man, 1916
Libinia dubia
H. Milne Edwards, 1834
Lucifer hanseni
Nobili, 1905
Macrophthalmus graeffei A. Milne Ewdards, 1873
Marsupenaeus japonicus (Bate, 1888)
Melicertus hathor(Burkenroad, 1959)
Menaethius monoceros(Latreille, 1825)
Metapenaeopsis aegyptia Galil & Golani, 1990
Metapenaeopsis mogiensis (Nobili, 1904)
consobrina
Metapenaeus affinis(H. Milne Edwards, 1837)
Metapenaeus monoceros (Fabricius, 1798)
Medit. Mar. Sci., 11/2, 2010, 381-493

OriginWMED CMED ADRIA EMED
Indo-Pacificcas
E Atlantic
cas
cas
cas
W Atlanticcas
W Atlantic
cas
cas
est
est
Indo-Pacificest
Indo-PacificcasIndo-PacificcasIndo West PacificestIndian/Red Sea
est
Indo-PacificcasIndo West Pacificest
Tropical East Atlantic cas
Indo West PacificcasIndo West Pacificest
N East Atlantic
estPacificcas
cas
Indo-PacificestestIndian/Red Sea
cas
W Atlantic
est
Indo West PacificcasIndiancas
cas
Indo-Pacificest?
Red Seacas
Indo West PacificcasPacific
cas
cas
Tropical East AtlanticcasIndo West Pacificest
Red Sea
est
Indo West PacificestIndo West Pacificest
W Atlantic
est
Indo West Pacificques
Indo West Pacificest
Indo-Pacific
cas
cas
cas
est
Indianest
Indo-PacificcasIndo-PacificestIndo West PacificestIndo West PacificIndo West Pacific

est

est
est
399

Table 2 (continued)
Species
AuthorMetapenaeus stebbingi(Nobili, 1904)
Micippa thalia(Herbst, 1803)
Myra subgranulata
Kossmann, 1877
Necora puber(Linnaeus, 1767)
Notopus dorsipes(Linnaeus, 1758)
Ogyrides mjoebergi(Balss, 1921)
Palaemonella rotumana(Borradaile, 1898)
Palinurus ornatus(Fabricius, 1798)
Paralithodes camtschaticus (Tilesius, 1815)
Penaeus semisulcatus
de Haan, 1844
Percnon gibbesi(H. Milne Edwards, 1853)
Periclimenes calmani
Tattersall, 1921
Pilumnopeus vauquelini(Audouin, 1826)
Pilumnus minutus**
de Haan, 1835
Plagusia squamosa(Herbst, 1790)
Portunus segnis**(Forssk a l, 1775)
Processa macrodactyla
Holthuis, 1952
Rhithropanopeus harrisii (Gould, 1841)
Rimapenaeus similis(Smith, 1885)
Scyllarus caparti
Holthuis, 1952
Scyllarus posteli
Forest, 1963
Sirpus monodi
Solenocera crassicornis
Sphaerozius nitidus
Sternodromia spinirostris**
Synalpheus tumidomanus
africanus**
Thalamita gloriensis
Thalamita indistincta
Thalamita poissonii
Trachysalambria
palaestinensis
Urocaridella pulchella
IsopodaAnilocra pilchardi p
Apanthura sandalensis
Cymothoa indicap
Mesanthura spp.
Paracerceis sculpta
Paradella dianae
400

Gordon, 1953
(H.Milne Edwards, 1837)
Stimpson, 1858
(Miers, 1881)
Crosnier & Forest, 1966

OriginWMED CMED ADRIA EMED
Indianest
est
Indo West PacificestIndian/Red Sea
est
NW Atlantic
est
est
cas
Indo West PacificcasIndo West PacificestIndo West PacificestIndo West Pacificcas
NE PacificcasIndo West Pacificest
W Atlantic
est
est
cas
est
Indo-PacificcasIndian/Red Sea
cas
est
est
Indo-PacificcasestIndo West Pacific
ques
est
cas
cas
Indiancasestest
Tropical East Atlantic est?
cas
Noth Atlantic
cas?
est
est
W Atlantic
estTropical East Atlanticcas
Temperate-tropicalcas
East Atlantic
Tropical East Atlanticcascas
Indo West PacificcasIndo West Pacificcas
Tropical East AtlanticcasTropical East Atlantic cry
cry

Crosnier, 1962
Apel & Spiridonov, 1998
(Audouin, 1826)(Steinitz, 1932)

Indo West PacificIndianIndo West Pacific
Red Sea

Yokes & Galil, 2006

Indo-Pacific

Bariche & Trilles, 2006
Stebbing, 1900
Schioedte & Meinert, 1884

Indo-Pacific
S Africa
Indo-PacificTropical/subtropical
Subtropical
NE Pacific(Holmes, 1904)(Menzies, 1962)

cas
cas
cry
est

cry
est

est

est
ques
est
est

ques
est
est?

est

cas
est
cas
ques
est
est

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

Table 2 (continued)
Species
AuthorSphaeroma venustissimum Monod, 1931
Sphaeroma walkeri
Stebbing, 1905
Stomatopoda
Clorida albolitura
Ahyong & Naiyanetr, 2000
Erugosquilla massavensis (Kossmann, 1880)
Erythrosquilla sp. (postlarvae)
Tanaidacea
Zeuxo (Parazeuxo) coralensis Sieg, 1980

OriginWMED CMED ADRIA EMED
Tropical/subtropicalest
E Atlantic
Indianest
est
est
Indo West PacificIndian/Red Sea
Indo-Pacific

est
est

est
cas

Circumtropicalest

MOLLUSCA
Bivalvia
Acar plicata
Afrocardium richardi
Alectryonella crenulifera
Anadara granosa
Anadara inflata
Anadara kagoshimensis**(Dillwyn, 1817)(Audouin, 1826)(Sowerby, 1871)(Linnaeus, 1758)(L. A. Reeve, 1844)(Tokunaga, 1906)

Anadara natalensis
Anadara transversa**
Angulus flacca
Antigona lamellaris
Atactodea glabrata
Brachidontes pharaonis
Callista florida
Cardites akabana
Chama asperella
Chama aspersa
Chama pacifica
Circe scripta
Circenita callipyga
Clementia papyracea
Crassostrea gigas
Dendrostrea frons
Diplodonta bogii**
Divalinga arabica
Dosinia erythraea
Electroma vexillum
Ensiculus cultellus
Fulvia australis
Fulvia fragilis(Krauss, 1848)(Say, 1822)(Roemer, 1871)
Schumacher, 1817
(Gmelin, 1791)(Fischer, 1870)(Lamarck, 1818)(Sturany, 1899)
Lamarck, 1819
Reeve, 1846
Broderip, 1834
(Linnaeus, 1758)(von Born, 1778)(Gray, 1825)(Thunberg, 1793)(Linnaeus, 1758)
Van Aartsen, 2004
Dekker & Gould, 1994
Römer, 1860
(Reeve, 1857)(Linnaeus, 1758)(Sowerby G.B., 1834)(Forssk a l, 1775)

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

Indo-Pacific/Red Sea
Red Sea
Pacific/Red Sea
Indo West Pacific
NW PacificTemperate North
Pacific
W Pacific/Red Sea
W Atlantic
Indo Pacific/Red Sea
Indo-Pacific/Red Sea
Indo Pacific/Red Sea
Indian/Red Sea
Indian/Red Sea
Red Sea
Indo-Pacific/Red Sea
Indo-PacificIndo-PacificIndo West Pacific
Red Sea
Indo-Pacific/Red Sea
NW PacificIndo Pacific/Red Sea
Red Sea
Persian Gulf/Red Sea
W Pacific/Red Sea
Indian/Red Sea
Indo West PacificIndo-Pacific/Red Sea
Indian

est

est

est

est

est

cas
est

est

est

est

est

est

est

est
cas

cas
est
ques
cas
cas
est
est
est
cas
cas
cas
est
cas
cas
cas
est
est
cas
cas
est
est
est
est
cas
est
est
cas
cas
est
401

Table 2 (continued)
Species
Gafrarium pectinatum
Glycymeris arabicus
Hiatula ruppelliana
Laternula anatina
Limopsis multistriata
Mactra lilacea
Mactra olorina
Malvufundus regula
Mercenaria mercenaria
Modiolus auriculatus
Musculista perfragilis
Musculista senhousia

OriginWMED CMED ADRIA EMED
Indo-Pacific/Red Sea
est
W Pacific/Red Sea
cas
Red Seacas
Indo-Pacific/Red Sea
est
Indo-Pacific/Red Sea
cas
Indo-Pacific/Red Sea
est
Red Sea
est
Indo-Pacific/Red Sea
estest
W Atlantic
est
estIndian/Red Sea
est
Indo-Pacific/Red Sea
est
Temperate North
est
est
est
est
Pacific
Mya arenaria
Linnaeus, 1758
N Atlantic
est
est
est
Mytilopsis sallei(Recluz, 1849)
W Atlantic
estNanostrea exigua
Harry, 1985
Indo-PacificestPaphia textile(Gmelin, 1791)
Indo-Pacific/Red Sea
est
Pedicirce sulcata(Gray, 1838)
Red Seaques
Petricola hemprichi(Issel, 1869)
Indo-Pacificest
Petricola pholadiformis
Lamarck, 1818
W Atlantic
est
Pinctada margaritifera(Linnaeus, 1758)
Indo-Pacific/Red Sea
ques
Pinctada radiata(Leach, 1814)
Indo-Pacific/Red Sea
est
est
cas
est
Psammotreta praerupta(Salisbury, 1934)
Indo-Pacific/Red Sea
cas
Pseudochama corbieri(Jonas, 1846)
Red Seaest
Ruditapes philippinarum (Adams & Reeve, 1850)
Temperate North
est
est
est
est
Pacific
Saccostrea commercialis (Iredale & Roughley, 1933) Australia
cas
Saccostrea cucullata(Born, 1778)
Indo-Pacificques
est
Septifer bilocularis(Linnaeus, 1758)
Indo-Pacificcas
Septifer forskali
Dunker, 1855
Red Seaest
Sphenia rueppelli
A. Adams, 1850
Indianest
Spondylus groschi
Lamprell & Kilburn, 1995 Indian/Red Sea
ques
Spondylus cf. multisetosus Reeve, 1856
Indo-Pacificques
Spondylus nicobaricus
Schreibers, 1793
Indo-PacificcasSpondylus spinosus
Schreibers, 1793
Indo-Pacific/Red Sea
est
Tellina valtonis
Hanley, 1844
Indian/Red Sea
est
Teredo navalis
Linnaeus, 1758
CircumtropicalcrycryTheora lubrica
Gould, 1861
Indo-Pacificest
estTimoclea marica(Linnaeus, 1758)
Red Seacas
Trapezium oblongum(Linnaeus, 1758)
Indo-Pacific/Red Sea
ques
Xenostrobus securis(Lamarck, 1819)
Tropical Atlantic
est
est
402

Author(Linnaeus, 1758)(Adams H., 1871)(Reeve, 1857)(Linnaeus, 1758)(Forssk a l, 1775)
Lamarck, 1818
Philippi, 1846
(Forssk a l, 1775)(Linnaeus, 1758)(Krauss, 1848)(Dunker, 1857)(Benson in Cantor, 1842)

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

Table 2 (continued)
Species
AuthorOriginWMED CMED ADRIA EMED
Zygochlamys patagonica** (King & Broderip, 1832)
W Atlanticcascas
Cephalopoda
Octopus aegina
Gray, 1849
Indo-Pacific/Red Sea
est
Octopus cyanea
Gray, 1849
Indo-Pacificcas
Sepia gibba
Ehrenberg, 1831
Red Seacas
Sepia pharaonis
Ehrenberg, 1831
Pacific/Red Sea
ques
Sepioeuthis lessoniana
Lesson, 1830
Indo-PacificestTremoctopus gracilis(Eydoux & Souleyet, 1852) Indo-Pacificcas
cas
Gastropoda
Acteocina crithodes
Melvill & Standen, 1907
Indo-Pacific/W Indiancas
Acteocina mucronata(Philippi, 1849)
Red Seaques
est
Alvania dorbignyi(Audouin, 1826)
Cosmopolitan
cry
cry
cry
Amathina tricarinata(Linnaeus, 1767)
Red Sea
est
Angiola punctostriata(Smith E.A., 1872)
Red Seaques
Anteaeolidiella foulisi**(Angas, 1864)
Circumtropicalcas
cas
Aplysia dactylomela
Rang, 1828
Circumtropical
est
est
est
Aplysia parvula
Guilding in Mörch, 1863
Circumtropical
cry
cry
cry
Atys angustatus
Smith, 1872
Red SeacryAtys cylindricus(Helbling, 1779)
Indo-Pacific/Red Sea
cas
Bostrycapulus odites**
Collin, 2005
Tropical Atlantic
estBulla arabica **Malaquias & Reid, 2008
Indo-PacificestBursatella leachii
De Blainville, 1817
Circumtropical
est
est
est
est
Caloria indica(Bergh, 1896)
Indo-PacificcasCanarium mutabile**(Swainson, 1821)
Indo-Pacific/Red Sea
ques
Cantharus tranquebaricus (Gmelin, 1791)
Indiancas
Cellana rota
(Gmelin, 1791)
Indian/Red Sea
cas
cas
est
Cerithidium diplax**(Watson, 1886)
Persian Gulf
est
Cerithidium perparvulum** (Watson, 1886)
Pacific
est
Cerithiopsis pulvis(Issel, 1869)
Red Sea
est
Cerithiopsis tenthrenois(Melvill, 1896)
Indianest
Cerithium columna
Sowerby 1834
Indo West Pacificques
Cerithium egenum
Gould, 1849
Indo-Pacific/Red Sea
est
Cerithium litteratum(Born, 1778)
W Atlantic
cas
Cerithium nesioticum
Pilsbry & Vannata, 1906
Indian/Red Sea
est
Cerithium nodulosum
Bruguière, 1792
Indo West Pacificques
Cerithium scabridum
Philippi, 1848
Indian/Red Sea
est
est
est
Chelidonura fulvipunctata Baba, 1938
Indo-Pacificest
est
Chromodoris annulata(Eliot, 1904)
Indianest
Chromodoris quadricolor (Rüppell & Leuckart, 1828) Red Sea
cas
cas
est
Chrysallida fischeri
Hornung & Mermod, 1925) Red Sea
est
est
Chrysallida maiae(Hornung & Mermod, 1924) Red Sea
estMedit. Mar. Sci., 11/2, 2010, 381-493

403

Table 2 (continued)
Species
AuthorChrysallida micronanaÖztürk & van Aartsen, 2006
Chrysallida pirintella(Melvill, 1910)
Cingulina isseli(Tryon, 1886)
Clypeomorus bifasciatus(Sowerby G.B. II, 1855)
Conomurex persicus**(Swainson, 1821)
Conus arenatus
Hwass, 1792
Conus fumigatus
Hwass, 1792
Conus inscriptus
Reeve, 1843
Conus rattus
Hwass, 1792
Coralliobia madreporarum (Sowerby, 1832)
Crepidula fornicata(Linnaeus, 1758)
Cuthona perca(Marcus, 1958)
Cycloscala hyalina(Sowerby, 1844)
Cylichnina girardi(Audouin, 1826)
Dendrodoris fumata(Rüppell & Leuckart, 1830)
Diala semistriata(Philippi, 1849)
Diodora funiculata(Reeve, 1850)
Diodora ruppellii(Sowerby, 1834)
Discodoris lilacina(Gould, 1852)
Doxander vittatus**(Linnaeus, 1758)
Echinolittorina punctata(Gmelin, 1791)
Elysia grandifolia
Kelaart, 1858
Elysia tomentosa
Jensen, 1997
Engina mendicaria(Linnaeus, 1758)
Ergalatax contracta(Reeve, 1846)
Ergalatax junionae**
Houart, 2008
Erosaria turdus(Lamarck, 1810)
Ethminolia hemprichi(Issel, 1869)
Favorinus ghanensis
Edmunds, 1968
Finella pupoides
A. Adams, 1860
Flabellina rubrolineata(O’Donoghue, 1929)
Fusinus verrucosus(Gmelin, 1791)
Gastrochaena cymbium(Spengler, 1783)
Gibborissoa virgata(Philippi, 1849)
Gibbula albida(Gmelin, 1791)
Godiva quadricolor(Barnard, 1927)
Halgerda willeyi
Elliot 1904
Haliotis pustulata cruenta Reeve, 1846
Haminoea cyanomarginata Heller & Thompson, 1983
Haminoea japonica**(Pilsbry, 1895)
Hinemoa cylindrica(de Folin, 1879)
404

OriginWMED CMED ADRIA EMEDRed Seacas
Red Sea
est
Subtropicalest
Indo-Pacific/Red Sea
casest
Persian Gulf
est
cas
est
Indo-Pacific/Red Sea
cas
Red Seacas
Indiancas
Indo-Pacificcas
Pantropicalcas
cas
NW Atlantic
est
cas
est
Tropical AtlanticcasIndo-Pacific/Red Sea
est
Indo-PacificestIndo Pacific/Red Sea
cas
Indo-Pacific/Red Sea
est
Indo-Pacificest
Indo-Pacific/Red Sea
cas
est
Indo-Pacific/Red Sea cry
cry
Indo-Pacificcas
Tropical Atlantic
Nativeest
NativeIndianest
Indo West PacificestIndo West Pacificcas
Red SeacasPersian Gulf/Red Sea
est
Indian/Red Sea
est
est
Red Seacas
Tropical Atlantic
est
Indo-Pacificest
Indo-Pacific/Red Sea
est
Indianest
Indo-Pacific/Red Sea
est
Indo-Pacificest
Adriatic Sea
est
NativeEastern PacificcasIndo West PacificcasIndian/Red Sea
cascas
Red Seaest
est
Pacific
est
Indo-Pacificcas
Medit. Mar. Sci., 11/2, 2010, 381-493

Table 2 (continued)
Species
AuthorOriginWMED CMED ADRIA EMED
Hypselodoris infucata(Rüppell & Leuckart, 1828) Indo-Pacific/Red Sea
est
Iolaea neofelixoides(Nomura, 1936)
Pacific
cas
Leucotina cfr eva
Thiele, 1935
Indo-Pacificcas
Leucotina natalensis**Smith, 1910
Indo-Pacific/Red Sea
est
Lienardia mighelsi
Iredale & Tomlin, 1917
Pacific
cas
Littorina saxatilis(Olivi, 1792)
Atlantic
cry
Melibe viridis**(Kelaart, 1858)
Indo-Pacific
est
est
est
est
Metaxia bacillum(Issel, 1869)
Red Seaest
Mitrella psilla(Duclos, 1846)
Tropical Atlantic
est
Monetaria annulus(Linnaeus, 1758)
Indo-Pacificques
Monotigma lauta**(A. Adams, 1853)
Indo-Pacific/Red Sea
est
Murchisonella columna(Hedley, 1907)
Indo-PacificcasMurex forskoehlii
Roeding, 1798
Persian Gulf/Red Sea
est
Nassa situla(Reeve, 1846)
Indo-PacificcasNassarius arcularia plicatus (Roeding, 1798)
Indian/Red Sea
cas
Nassarius concinnus(Powys, 1835)
unknowncasNassarius stolatus(Gmelin, 1791)
Indo-Pacificcas
Nerita sanguinolenta
Menke, 1829
Red Seacas
est
Notocochlis gualteriana** (Recluz, 1844)
Indo-Pacific/Red Sea
casest
Odostomia lorioli(Hornung & Mermod, 1924) Red Sea
cas
Oscilla jocosa
Melvill, 1904
Persian Gulfcas
Oxynoe viridis(Pease, 1861)
Indo West Pacific
est
Palmadusta lentiginosa(Gray, 1825)
Indiancas
lentiginosa
Parviturbo dibellai
Buzzurro & Celalupo, 2006 unknown
cry
Patelloida saccharina(Linnaeus, 1758)
Pacific
cas
Philinopsis cyanea(Martens, 1879)
Indian/Red Sea
cas
Planaxis griseus(Brocchi, 1821)
Red Seacas
Pleurobranchus forskalii
Rüppell & Leuckart, 1828 Red Sea
cas
Plocamopherus ocellatus Rüppell & Leuckart, 1830 Red Sea
est
Polycera hedgpethi
Marcus Er., 1964
NE Pacificcas
cas
Polycerella emertoni
Verrill, 1881
Pantropicalest
est
Pseudominolia nedyma
Melvill, 1897
Persian Gulf/Red Sea
est
Purpuradusta gracilis notata (Gill, 1858)
Indian/Red Sea
est
Pyrunculus fourierii(Audouin, 1826)
Indo-Pacific/Red Sea
est
Rapana rapiformis(Von Born, 1778)
Indo-Pacific/Red Sea
ques
Rapana venosa(Valenciennes, 1846)
Temperate North
est
est
est
Pacific
Retusa desgenettii(Audouin, 1826)
Red SeacasRhinoclavis kochi(Philippi, 1848)
Indo-Pacific/Red Sea
est
Rhinoclavis sinensis(Gmelin, 1791)
Indo West Pacificcas
Medit. Mar. Sci., 11/2, 2010, 381-493

405

Table 2 (continued)
SpeciesRissoina ambigua
Rissoina bertholleti
Rissoina spirata
Sabia conica
Siphonaria crenata

Author(Gould, 1849)
Issel, 1869
Sowerby, 1825
(Schumacher, 1817)
de Blainville, 1827

OriginWMED CMED ADRIA EMED
Indo-PacificcasIndian/Red Sea
est
Indo-Pacificcas
cas
Indo-Pacific/Red Sea
ques
ques
Persian Gulf/
estArabian Sea(Linnaeus, 1758)
Atlanto-Mediterranean Native
est(Montrouzier, 1863)
Indo-Pacific/Red Sea
est(A. Adams, 1861)
Indo-Pacificcas(Burrow, 1815)
Indo-Pacific/Red Sea
cas(Adams & Reeve, 1850)
Circumtropicalest
est
A. Adams, 1860
Indo-Pacific/Red Sea
cas(Jickeli, 1882)
Indo-Pacific
est(A. Adams, 1863)
Red Seaest(von Born, 1778)
Indiancas
est(Gmelin, 1791)
Indian/Red Sea
estAdam & Knudsen 1969
W Africa
ques
Schilder, 1932
Indo-PacificcasBrocchi, 1821
Persian Gulf
est
(Melvill, 1896)
Indo-Pacificest(Deshayes in Laborde, 1834) Indian/Red Sea
ques(Issel, 1869)
Red Seacas(Moazzo, 1939)
Red Sea
est(Melvill & Standen, 1901) Indian/Red Sea
est

Siphonaria pectinata
Smaragdia souverbiana
Sticteulima cf. lentiginosa
Stomatella impertusa
Syphonota geographica
Syrnola cinctella
Syrnola fasciata
Syrnola lendix**Thais lacera
Thais sacellum
Tornus jullieni
Trivirostra triticum
Trochus erithreus
Turbonilla edgarii
Vexillum depexum
Voorwindia tiberiana
Zafra savignyi
Zafra selasphora
Polyplacophora
Acanthopleura gemmata
Chiton hululensis(de Blainville, 1825)
Indo-Pacific/Red Sea(Smith E.E. in Gardiner, 1903) Indo-Pacific/Red Sea

cas
ques

MISCELLANEA INVERTEBRATA
Arthropoda/Pycnogonida
Ammothea hilgendorfi(Böhm, 1879)
Anoplodactylus californicus Hall, 1912
Anoplodactylus digitatus(Böhm, 1879)
Pigrogromitus timsanus
Calman, 1927
Bryozoa
Aeverrillia setigera(Hincks, 1887)
Arachnoidea protecta(Harmer, 1915)
Bowerbankia gracillima(Hincks, 1877)
Bugula fulva
Ryland, 1960
Bugula serrata(Lamarck, 1816)
Celleporaria aperta(Hincks, 1882)
Celleporaria brunnea(Hincks, 1884)
406

Circumboreal
Circum(sub)tropical
Circum(sub)tropical
Circum(sub)tropical
CircumtropicalIndo-Pacific
E Atlantic
NW Atlantic
Indo-PacificCircumtropical
NE Pacific

est
est

est
est
ques
cas

ques
cas
est
est

cas
cas
ques

cas

cas
est
est

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

Table 2 (continued)
SpeciesCelleporaria pilaefera
Celleporella carolinensis
Crepidacantha poissoni
Electra tenella
Hippaliosina acutirostris
Hippopodina feejeensis
Parasmittina egyptiaca
Parasmittina serruloides
Parasmittina spondylicola
Pherusella brevituba
Reteporella jermanensis
Rhynchozoon lareyi
Schizoretepora hassi
Scrupocellaria jolloisii
Smittina malleolus
Tricellaria inopinata
Chaetognatha
Aidanosagitta neglecta*
Ferosagitta galerita*
Chordata/Ascidiacea
Ascidia cannelata
Botrylloides violaceus
Cystodytes philippinensis
Distaplia bermudensis
Ecteinascidia styeloides
Ecteinascidia thurstoni
Herdmania momus
Microcosmus exasperatus
Microcosmus squamiger
Perophora multiclathtrata
Phallusia nigra
Polyandrocarpa zorritensis
Rhodosoma turcicum
Styela clava
Symplegma brakenhielmi
Trididemnum cf. savignii
Cnidaria/Anthozoa
Acabaria erythraea

Author(Canu & Bassler, 1929)
Ryland, 1979
(Audouin, 1826)(Hincks, 1880)
Canu & Bassler, 1929
(Busk, 1884)(Waters, 1909)
Harmelin, Bitar &Zibrowius, 2009
Harmelin, Bitar &Zibrowius, 2009
Soule, 1951
(Waters, 1909)(Audouin, 1826)
Harmelin, Bitar &Zibrowius, 2007
(Audouin, 1826)(Hincks, 1884)
d'Hondt & Occhipinti, 1985

OriginIndo-W Pacific
W Atlantic
Circumtropical
CircumtropicalIndo-PacificIndo-PacificIndianIndo-PacificAida, 1897
(Dallot, 1971)

Indo-PacificIndianIndo-Pacific
NE Pacific
Red Sea
Indo-Pacific
Red Sea?

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

cas
cas

Indo-Pacific

cas
cas
est
ques

Indo-PacificCircumtropicalIndo-PacificOken, 1820
Indo-PacificOka, 1927
NW Pacific
Herdman, 1886
Circumtropical
Van Name, 1902
W Atlantic(Traustedt, 1882)
Circumtropical
Herdman, 1890
Indo-Pacific(Savigny 1816)
Indo-PacificHeller, 1878
Indo-PacificHartmeyer & Michaelsen, 1928 Circumtropical(Sluiter, 1904)
Circumtropical
Savigny, 1816
Circumtropical(Van Name, 1931)
E Pacific(Savigny, 1816)
Circumtropical
Herdman, 1881
NW Pacific(Michaelsen, 1904)
Indo-Pacific(Herdman, 1886)
Circum(sub)tropical(Ehrenberg, 1834)

WMED CMED ADRIA EMED
ques
est
ques
cas
ques
cas
est
est

est
est
est

est
est
est
est
est

cry
est
est

est
est

cry
est
est

est
est

est
est
est

est
est
cas
cas

est
cas

est

cas

est
est

est
407

Table 2 (continued)
Species
Diadumene cincta
Haliplanella lineata
Oculina patagonica
Cnidaria/Hydrozoa
Aequorea conica*
Amphogona pusilla*
Arctapodema australis*
Bougainvillia niobe

AuthorStephenson, 1925
(Verrill, 1870)
De Angelis, 1908

WMED CMED ADRIA EMED
est
est
est
est

Indo-PacificIndo-Pacific
Antarctic/Indo-PacificMayer, 1894(sub)tropicalW Atlantic
Campalecium medusiferum* Torrey, 1902
E Pacific/ Atlantic
Cirrholovenia tetranema* Kramp, 1959
CircumtropicalClytia hummelincki*(Leloup, 1935)
CircumtropicalClytia linearis**(Thornely, 1900)
Circum(sub)tropicalClytia mccradyi*(Brooks, 1888)
CircumtropicalCordylophora caspia(Pallas, 1771)
Circumglobal
Coryne eximia*
Allman, 1859
Circum(sub)tropical
Diphasia digitalis(Busk 1852)
Circumtropical
Dynamena quadridentata (Ellis & Solander, 1786)
Circumtropical
Eirene viridula*(Péron & Lesueur, 1810)
Circumtropical
Eucheilota paradoxica*
Mayer, 1900
Circumtropical
Eudendrium carneum
Clarke, 1882
Circumtropical
Eudendrium merulum
Watson, 1985
CircumtropicalEuphysora annulata*
Kramp, 1928
Indo-PacificEuphysora bigelowi*
Maas, 1905
Indo-Pacific
Filellum serratum*(Clarke, 1879)
Circum(sub)tropicalGarveia franciscana(Torrey, 1902)
Circum(sub)tropical
Gonionemus vertens*
A. Agassiz, 1862
Circumboreal
Kantiella enigmatica*
Bouillon, 1978
IndianLaodicea fijiana*
Agassiz & Mayer, 1899
Indo-PacificMacrorhynchia philippina (Kirchenpauer, 1872)
Circumtropical
Moerisia carina*
Bouillon, 1978
Indo-PacificMoerisia inkermanica*
Paltschikowa-Ostroumova, 1925 Circum(sub)tropicalNubiella mitra*
Bouillon, 1980
SW Pacific
Octotiara russelli*
Kramp, 1953
Indo-PacificOlindias singularis*
Browne, 1905
IndianParacytaeis octona*
Bouillon, 1978
W Indian
Russellia mirabilis
Kramp, 1957
Antarctic
Scolionema suvaensis*(Agassiz & Mayer, 1899)
Indo-Pacific
Sertularia marginata(Kirchenpauer, 1864)
Circum(sub)tropical
Sertularia thecocarpa(Jarvis, 1922)
Indo-PacificTetrorchis erythrogaster*
Bigelow, 1909
Circumtropical408

Browne, 1905
Hartlaub, 1909

OriginNE Atlantic
Circumboreal?SW Atlantic

cas
cas
cas

cas
est

est
est
est
est
est
cry
ques

est
cas
est
est

cas
cas
est

cas

cas
cas
est
est

est
est

cry

cry
ques

est
cas

est
cas
cas
cas
cas

est
est
cry
cas
est
est
est
est
est
cas
cas

est
est
cas
cas
est
est
cas
cas

cas
est
cas
cas
est
est

est
est
est

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

Table 2 (continued)
Species
Trichydra pudica*
Cnidaria/Scyphozoa
Cassiopea andromeda
Marivagia stellata
Phyllorhiza punctata*
Rhopilema nomadica*
Stomolophus meleagris*
Ctenophora
Beroe ovata*Mnemiopsis leidyi*
Sulculeolaria angusta*
Echinodermata
Acanthaster planci
Amphiodia (Amphispina)
obtecta
Amphioplus (Lymanella)laevisAquilonastra burtoni**
Asterias rubens
Diadema setosum
Eucidaris tribuloides
Ophiactis macrolepidota
Ophiactis savignyi
Protoreaster nodosus
Prionocidaris baculosa
Synaptula reciprocans
Porifera
Cinachyrella australiensis
Geodia micropunctata
Haliclona spinosella
Haliclona viridis
Hyrtios erecta
Lissodendoryx
schmidti
Mycale erythraeana
Paraleucilla magna
Siphonophora
Abyla trigona*

AuthorWright, 1858

OriginCircumtropical(Forssk a l, 1775)
Galil & Gershwin, 2010

Indo-Pacific?Indo-PacificCircumtropical
Circumtropical
Red Sea

von Lendenfeld, 1884
Galil, Spannier &Ferguson, 1990
(L. Agassiz, 1862)

WMED CMED ADRIA EMED
cas
cas

cas

est?

est
est

cas
cas

est
est

Atlantic/Pacificcas

Mayer, 1912
A. Agassiz, 1865
Totton, 1954

Circum(sub)tropical
NW Atlantic
Indianest(Linnaeus, 1758)
Mortensen, 1940

Indo-PacificIndo-Pacific(Lyman, 1874)

Indo-Pacificcas(Gray, 1840)
Linnaeus, 1755
(Leske, 1778)(Lamarck, 1816)
Marktanner-Turneretscher,
1887(Müller & Troschel, 1842)(Linnaeus, 1758)(Lamarck, 1816)(Forssk a l, 1775)

W Indian
NE Atlantic
Indo West Pacific(sub)tropical Atlantic:
Circumtropical

est
est
cas

Carter, 1886
Row, 1911(Thiele, 1905)(Keller, 1881)(Keller, 1889)(Ridley, 1844)

Indo-PacificIndo-PacificIndo-PacificCircum(sub)tropicalIndo-PacificIndo-Pacific

Burton, 1936
Klautau et al., 2004

Indo-Pacific
SW Atlantic

Quoy & Gaimard,1827

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

CircumtropicalIndo-PacificIndianIndo-Pacific

est

est
est

est
est
est

ques
est

est
est
cas
cas

est

est

cas
est
ques
ques
ques
ques
ques
ques
ques
cry

cry

cry
cas
409

Table 2 (continued)
Species
AuthorOriginSipuncula
Apionsoma (Apionsoma) (Ikeda, 1904)
Circumtropicalmisakianum
Apionsoma (Apionsoma) (Sluiter, 1902)
Circumtropicaltricocephalus
Aspidosiphon (Akrikos)(Murina, 1967)
Circumtropical
mexicanus
Aspidosiphon (Aspidosiphon) (Chamisso & Eysenhardt, Circumtropicalelegans
1821)
Phascolion (Isomya)(Sluiter, 1902)
Indo-Pacificconvestitum
Phascolosoma scolops(Selenka & de Man, 1883) Indo-Pacific
Platyhelminthes
Allolepidapedon fistulariae p Yamaguti, 1940
Indo-Pacific
Red SeaGlyphidohaptor plectocirrap (Paperna, 1972)(Pallas, 1774)
Atlantic
Hirudinella ventricosa p
Manter, 1969
Indo-PacificHysterolecitha sigani p
Lecithochirium
Fischthal & Kuntz, 1963
Red Seamagnicaudatum
Indo-PacificMonilicaecum ventricosum p Yamaguti, 1942
Neothoracocotyle(Meserve, 1938) Hargis, 1956 Atlantic/Pacificacanthocybii p
Nosema ceratomyxa p
Diamant & Paperna, 1985 Red Sea
Ogawa & Egusa, 1980
Indo West Pacific
Polylabris cf. mamaevi p
Red Sea
Tetrancistrum polymorphus p (Paperna, 1972)
Tetrancistrum
Kritsky, Galli & Yang, 2007 Red Sea
strophosolenum p
Tetrancistrum suezicus p(Paperna, 1972)
Red Sea

WMED CMED ADRIA EMED
ques

ques
cas
ques

cas

cas
est

est

cas
est

est

cas
est
cas
ques
cas
cas
cas

cas

cas

est
est
cas
est
est

FISH
Elasmobranchii
Carcharhinus altimus
Carcharhinus falciformis
Dasyatis marmorata
Galeocerdo cuvier
Himantura uarnak
Isurus paucus
Rhizoprionodon acutus
Sphyrna mokarran
Torpedo sinuspersici
Actinopterygii
Abudefduf vaigiensis
410(Springer, 1950)(Müller & Henle, 1839)(Smith, 1828)(Peron & Le Sueur, 1822)(Forssk a l, 1775)
Guitart Manday, 1966
(Rüppell, 1837)(Rüppell, 1837)
Olfers, 1831

Indo-PacificCircumtropicalTropical Atlantic
Tropical/subtropicalIndo-PacificCircumtropical
Circumtropical
CircumtropicalIndo-Pacific(Quoy & Gaimard, 1825)

Indo-Pacific

est
est
cas

est
ques
cas

ques
est

cas
cas
cas
ques
cas

cas

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

Table 2 (continued)
SpeciesAcanthurus monroviae
Alepes djedaba
Aluterus monocerus
Anarhichas lupus
Anguilla japonica
Apogon fasciatus
Apogon pharaonis
Apogon queketti
Apogon smithi
Arius parkii
Atherinomorus forskalii**
Beryx splendens
Bregmaceros atlanticus
Callionymus filamentosus
Centrolabrus exoletus
Cephalopholis taeniops
Champsodon nudivittis
Champsodon vorax
Chaunax suttkusi
Cheilodipterus
novemstriatus
Cheilopogon furcatus
Chilomycterus reticulatus
Chirocentrus dorab
Coryogalops ochetica
Crenidens crenidens
Cyclichthys spilostylus
Cyclopterus lumpus
Cynoglossus sinusarabici
Decapterus russelli
Diodon hystrix
Diplodus bellottii
Dussumieria elopsoides
Elates ransonetti
Enchelycore anatina
Epinephelus coioides
Epinephelus malabaricus
Equulites klunzingeri**Etrumeus teres
Favonigobius
melanobranchus**

AuthorSteindachner, 1876
(Forssk a l, 1775)(Linnaeus, 1758)
Linnaeus, 1758
Temminck & Schlegel, 1847
(White, 1790)
Bellotti, 1874
Gilchrist, 1903
(Kotthaus, 1970)
Gunther, 1864
(Rüppell, 1838)
Lowe, 1834
Goode & Bean, 1886
Valenciennes, 1837
(Linnaeus, 1758)(Valeciennes, 1828)(Ogilby, 1895)
Günther, 1867
Caruso, 1898
(Rüppell, 1838)

OriginWMED CMED ADRIA EMED
Tropical Atlantic
est
cas
cas
Indo-PacificestAtlanticcas
N AtlanticcasPacific
ques
Indo-Pacificest
Indo-PacificestIndianest
Indianest
Indo-Pacificcas
Tropical Atlanticest
est
Circumtropicalestest
Tropical Atlanticcas
est
Circumtropicalest
Tropical Atlanticcas
Subtropical Atlanticestcas
Tropical AtlanticcasIndo West PacificcasIndo-Pacificcas
cas
W Indiancas(Mitchill, 1815)(Linnaeus, 1758)
Forssk a l, 1775)(Norman, 1927)(Forssk a l, 1775)(Leis & Randall, 1982)
Linnaeus, 1758
(Chabanaud, 1913)(Rüppell, 1830)(Linnaeus, 1758)(Steindachner, 1882)
Bleeker, 1849
(Steindachner, 1876)(Lowe, 1839)(Hamilton, 1822)(Bloch & Schneider, 1804)(Steindachner, 1898)(DeKay, 1848)(Fowler, 1934)

Circumtropical
CircumtropicalIndo-Pacific
Red Sea
IndianIndo-PacificAtlantic
Red Sea
Indo-PacificCircumtropicalTropical Atlantic
Indo-PacificTropical PacificTropical Atlantic
Indo-PacificIndo-PacificIndianSubtropicalIndo West Pacific

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

cas
cas
cas
est
est
est

est
cas

est
est
ques

cas
est

est
cas

cas
cas

cas
est

cas

est
est
est
est
est
cas

411

Table 2 (continued)
Species
AuthorOriginFistularia commersonii(Rüppell, 1835)
Indo-PacificFistularia petimba
Lacepède, 1803
Tropical-Atlantic
Gephyroberyx darwini(Johnson, 1866)
Tropical Atlantic
Glaucostegus halavi(Forssk a l, 1775)
Indo-PacificGymnammodytes(Jourdain, 1879)
N Atlantic
semisquamatus
Halosaurus ovenii
Johnson, 1863
Tropical Atlantic
Hemiramphus far(Forssk a l, 1775)
Indo-Pacific
Heniochus intermedius
Steindachner, 1893
IndianHerklotsichthys punctatus (Rüppell, 1837)
Red Sea
Hippocampus fuscus
Rueppell, 1838
IndianHyporhamphus affinis(Gunther, 1866)
Indo-PacificIniistius pavo
Valenciennes, 1840
Indo-PacificKyphosus incisor(Cuvier, 1831)
Tropical Atlantic
Kyphosus sectator(Linnaeus, 1758)
Tropical Atlantic
Lagocephalus sceleratus(Gmelin, 1788)
Indo-Pacific
Lagocephalus spadiceus(Richardson, 1844)
Indo-Pacific
Lagocephalus suezensis
Clark & Gohar, 1953
Red Sea
Liza carinata(Valenciennes, 1836)
IndianLiza haematocheila(Temminck & Schlegel, 1845) Indo-Pacific
Lutjanus argentimaculatus (Forssk a l, 1775)
Indo-Pacific
Lutjanus jocu
(Bloch & Schneider, 1801) Atlantic
Makaira indica(Cuvier, 1832)
Indo-PacificMicrochirus boscanion
Chabanaud, 1926
Tropical Atlantic
Microchirus hexophthalmus (Bennet, 1831)
Tropical Atlantic
Monotaxis grandoculis(Forssk a l, 1775)
Indo-Pacific
Muraenesox cinereus(Forssk a l, 1775)
Indo-Pacific
Mycteroperca fusca(Lowe, 1838)
Tropical Atlantic
Nemipterus randalli
Russell, 1986
IndianOmobranchus punctatus (Valenciennes, 1836)
Indo-Pacific
Oplegnathus fasciatus(Temminck & Schlegel, 1944) Pacific
Oxyurichthys petersi(Klunzinger, 1871)
Red Sea
Pagellus bellottii
Steindachner, 1882
Tropical Atlantic
Pagrus major(Temminck & Schlegel, 1843) NW PacificPampus argenteus(Euphrasen, 1788)
Indo-Pacific
Papilloculiceps longiceps (Ehrenberg in Valenciennes, Indian
1829)
Parexocoetus mento(Valenciennes, 1846)
Indo-Pacific
Parupeneus forsskali(Fourmanoir & Guézé, 1976) Indian
Pelates quadrilineatus(Bloch, 1790)
Indo-PacificPempheris vanicolensis
Cuvier, 1831
Indo-Pacific412

WMED CMED ADRIA EMED
est
est
est
est
cas
cas
ques
cas
ques
est
est
est

cas

est
cas
est
est
cas
cas

cas
est
cas

cas
cas
est
est

est
est
est
est
est
cas

cas
cas
est
cas
cas
cas
est
cas
cas
est
cas

est
cas
cas

cas
est
ques
est

est

est
cas
est
est

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

Table 2 (continued)
Species
AuthorPetroscirtes ancylodon
Rüppell, 1838
Pinguipes brasilianus
Cuvier & Valenciennes, 1829
Pisodonophis semicinctus (Richardson, 1848)
Platax teira(Forssk a l, 1775)
Platycephalus indicus(Linnaeus, 1758)
Plotosus lineatus(Thunberg, 1787)
Pomacanthus imperator(Bloch, 1787)
Pomacanthus maculosus (Forssk a l, 1775)
Pomadasys stridens(Forssk a l, 1775)
Priacanthus hamrur(Forssk a l, 1775)
Priacanthus sagittarius
Starnes, 1988
Psenes pellucidus
Lutken, 1880
Pseudupeneus prayensis(Cuvier, 1829)
Pteragogus pelycus
Randall, 1981
Pterois miles(Bennet, 1803)
Rachycentron canadum(Linnaeus, 1766)
Rastrelliger kanagurta(Cuvier, 1816)
Rhabdosargus haffara(Forssk a l, 1775)
Rhynchoconger trewavasae Ben-Tuvia 1993
Sargocentron rubrum(Forssk a l, 1775)
Saurida undosquamis(Richardson, 1848)
Scarus ghobban
Forssk a l, 1775
Sciaenops ocellatus(Linnaeus, 1766)
Scomberomorus commerson Lacepède, 1800
Scorpaena stephanica
Cadenat, 1943
Selene dorsalis
Gill, 1862
Seriola carpenteri
Mather 1971
Seriola fasciata(Bloch, 1793)
Seriola rivoliana
Valenciennes, 1833
Siganus javus(Linnaeus, 1766)
Siganus luridus(Rüppell, 1829)
Siganus rivulatus
Forssk a l, 1775
Silhouetta aegyptia(Chabanaud, 1933)
Sillago sihama(Forssk a l, 1775)
Solea senegalensis
Kaup, 1858
Sorsogona prionota(Sauvage, 1873)
Sphoeroides marmoratus (Lowe, 1838)
Sphoeroides pachygaster(Müller & Troschel, 1848)
Sphyraena chrysotaenia
Klunzinger, 1884
Sphyraena flavicauda
Rüppell, 1838
Spratelloides delicatulus(Bennett, 1831)
Medit. Mar. Sci., 11/2, 2010, 381-493

OriginWMED CMED ADRIA EMED
Indianest
W Atlantic
cascas
Tropical Atlantic
est
est
cas
Indo-PacificcasIndo-Pacificcas
est
Indo-Pacificest
Indo-PacificcasIndo-PacificcasIndianest
est
Indo-Pacific
ques
ques
Indo-Pacificcas
Tropical Atlanticestcas
Tropical AtlanticcasIndianest
Indiancas
Circumtropicalcas
cas
Indo-PacificcasIndianest
Red Seacas
Indo-Pacificcas
est
Indo-Pacific
est
est
est
Indo-Pacificest
W AtlanticcasIndo-Pacific
?
est
est
Tropical Atlanticcas
W AtlanticcasTropical Atlantic
estTropical Atlantic
est
est
cas
Tropical AtlanticcasIndo-PacificcasIndiancas
est
est
Red Seaest
cas
est
Red Sea
est
Indo-Pacificest
Tropical Atlantic
estques
Indiancas
Atlanticcas
cas
Tropical Atlantic
est
est
est
est
Indo-Pacific
est
est
est
Indianest
est
Indo-Pacific/Red Sea
est
413

Table 2 (continued)
Species
AuthorStephanolepis diaspros
Fraser-Brunner, 1940
Synagrops japonicus(Doderlein, 1884)
Synaptura lusitanica
Capello, 1868)
Syngnathus rostellatus
Nilsson, 1855
Terapon jarbua(Forssk a l, 1775)
Terapon puta
(Cuvier, 1929)
Terapon theraps
Cuvier, 1829
Tetrosomus gibbosus(Linnaeus, 1758)
Torquigener flavimaculosus Hardy & Randall, 1983
Trachurus indicus
Necrasov, 1966
Trachyscorpia cristulata(Koehler, 1869)
echinata
Tridentiger trigonocephalus (Gill, 1859)
Trypauchen vagina(Bloch & Scheider, 1801)
Tylerius spinosissimus(Regan, 1908)
Tylosurus choram(Rüppell, 1837)
Tylosurus crocodilus(Peron & Le Sueur, 1821)
Upeneus moluccensis(Bleeker, 1855)
Upeneus pori
Ben-Tuvia & Golani, 1989
Vanderhorstia mertensi
Klausewitz, 1974
Zenopsis conchifera(Lowe, 1852)

1. THE WESTERN MEDITERRANEAN
The WMED occupies a key position
because it receives the influx of surface
waters from the Atlantic, through the Straitof Gibraltar. It is further compartmentalized into fairly isolated sub-basins with different climatic and hydrologic conditions.
These sub-basins have a different biogeographic character, which may affect invasion and settlement of aliens. The Alboran
Sea, situated immediately east of Gibraltar,
exhibits stronger Atlantic affinities, due tothe continued penetration of Atlantic floraand fauna with the incoming influx of water(HARMELIN & D'HONDT, 1993). In
return, most Mediterranean endemics arerare or missing. The incoming Atlantic
414

OriginWMED CMED ADRIA EMEDRed Seaest
cas
est
Indo-Pacificcas
Tropical Atlanticest
est
N Atlanticcasques
Indo-PacificcasIndo-Pacific/Red Sea
est
Indo-PacificcasIndo-PacificestIndianest
Indiancas
Tropical Atlanticestest
Tropical PacificIndo-PacificIndo-PacificIndo-PacificIndo West PacificIndo-PacificIndianIndo-PacificAtlantic/Pacific

est
est

cas
cas
est
cas
cas
est
est
est

cas

waters form a permanent clockwise gyre in
the Alboran Sea, which is separated fromthe remainder of the WMED by a wellmarked hydrographic front between Oranand Almer a. As a consequence of the
admixture of fauna and flora of different
origins, the westernmost part of the Alboran Sea, from the Ceuta region to Punta deCalaburras on the coast of Malaga, constitutes a hot spot of biodiversity. Its enrichedbenthic environment includes some species
linked with the tropical periods of the~A et al., 2009;
Mediterranean Sea (OCAN
GARC A RASO et al., 2010; URRA et al.,in press). The Tyrrhenian Sea is comparatively isolated from the rest of the WMED
and is surrounded by mountains reducing
the impact of the meteorological events
Medit. Mar. Sci., 11/2, 2010, 381-493

that strongly influence the internal conditions of the other Western Mediterraneanbasins (ASTRALDI et al., 1995). The Gulf
of Lions and the Ligurian Sea are the coldest parts of the WMED. They are characterized by a severe reduction of thermophilic species, while some cold temperate species, not found to the south, arepresent (BIANCHI & MORRI, 1993,
1994).
1.1. Alien protozoans in the Western
Mediterranean
Four pathogenic protozoans havebeen recorded in WMED. Three of themare pathogenic to shellfish: Marteilia refringens (RIERA et al., 1993), Bonamia ostreae(MONTES & LAMA, 1993), and Perkinsus olseni, formerly Perkinsus atlanticus(SAGRIST A et al., 1996). The fourth,Photobacterium damselae, formerly Pasteurella piscicida, causes pasteurellosis/photobacteriosis in fish: it was first isolated in mortalities occurring in natural
populations of white perch and stripedbass in 1963 in Chesapeake Bay, USA.
From 1990 it has caused economic losses
in different European countries. Cultured
gilthead seabream (Sparus aurata) andseabass (Dicentrarchus labrax) are the mostaffected species in the WMED (TORANZO et al., 1991).
Only four alien foraminiferal species
occur in the WMED. Planogypsina acervalis (BLANC-VERNET, 1969, as Planorbulina acervalis) and Cymbaloporetta plana
have been previously cited in the other sectors of the Mediterranean. Very rare specimens of Sorites orbiculus have beenreported for the Gulf of Naples(HOFKER, 1930) and from the Ligurian
Sea (BANCHETTI et al., 2009, as S. orbicularis). Schackoinella imperatoria, probably
cryptogenetic/casual, has been recorded
Medit. Mar. Sci., 11/2, 2010, 381-493

from the Gulf of Naples, South Tyrrhenian(SGARRELLA & MONCHARMONT,
1993), from the Ligurian Sea and northTyrrhenian (BANCHETTI et al., 2009)
and from the coast of Vulcano, Aeolian
Islands (CIMERMAN & LANGER, 1991,
as Conorbella imperatoria).
1.2. Alien macrophytes in the Western
Mediterranean
A total of 91 alien macrophytes havebeen hitherto reported in the WMED,
which represent 72% of the alien macrophytes reported from the Mediterranean
Sea. The major vectors of introduction are
the shellfish aquaculture industry and shiptraffic (fouling, ballast waters). As much as
75 alien macrophyte species are well established, seven are casual, and nine are cryptogenic/questionable. Casual taxa (i.e.
Cladophora hutchinsioides, Fucus spiralis,
Grateloupia patens, Lomentaria flaccida,
Rhodophysema georgei, Saccharina japonica and Sarconema filiforme) might attainestablished status in the future. The cryptogenic/questionable category includes
Acanthophora nayadiformis, Anotrichiumokamurae, Antithamnionella boergesenii, A.
elegans, the Atlantic strain of Asparagopsis
taxiformis, Ceramium bisporum, Ganonema farinosum, Osmundea oederi andPolysiphonia atlantica.
On the 21 invasive or potentially invasive macrophytes introduced in the
Mediterranean (Table 3), 16 are present in
the western basin mainly in coastal lagoons.
Codium fragile, invasive in the last century,
has become less abundant in the open sea.
The northern coasts are the mostimpacted region, with a maximum of alien
macrophytes in France (78 taxa out of 91).
On the other hand, the lowest number (23taxa) was recorded along the northern
Africa coast (Morocco and Algeria). Such415

a difference is probably due to insufficientinvestigations in the south and the hugeflood of introductions along with shellfish
transfers in the French coastal lagoons
harbouringshellfish
aquaculture.
VERLAQUE et al. (2007) listed up to 58
alien macrophytes in the Thau Lagoon(64% of alien macrophytes of the WMED
and 46% of the total of alien macrophytesof the Mediterranean), which is the leading site of shellfish aquaculture in the
Mediterranean Sea.
The majority of alien macrophytes ofthe WMED comes from temperate andcold regions (NE Atlantic and NW Pacific)
in relation with the major vector of introduction (shellfish transfer). Only 10 putative Lessepsian immigrants (i.e. Acanthophora nayadiformis, Chondria pygmaea,
Ganonema farinosum, Halophila stipulacea, Hypnea spinella, H. valentiae, Lophocladia lallemandii, Padina boergesenii, Sarconema filiforme and Ulva fasciata) havebeen hitherto recorded, mainly out of thecoldest zones (Gulf of Lions and Gulf ofGenoa). However, the only WMED reportof Sarconema filiforme (France) requires
confirmation. Acanthophora nayadiformis,
Ganonema farinosum, Hypnea spinella,
Padina boergesenii and Ulva fasciata could
alternatively originate from the Atlantic
Ocean. The strains of Ulva fasciata andHypnea valentiae introduced in the Thau
Lagoon most probably originate from thenorth-western Pacific (Japan or Korea).
The invasive behaviour of Lophocladia
lallemandii in the Balearic Islands contrasts with a non-invasive behaviour everywhere else in the Mediterranean, and adifferent origin for the western strain
deserves to be considered. Thus, there are
hitherto only two unquestionable Lessepsian macrophytes in the WMED: Chondria
pygmaea and Halophila stipulacea.
416

1.3. Alien polychaetes in the Western
Mediterranean
In the WMED, a total of only 49 alienpolychaete species have been reported todate, which is about 39% of the alien polychaete species reported from the Mediterranean Sea. Among them, 26 species are
established in WMED (see Table 2), 12
species are casual, 1 species (Paraprionospio
coora) is cryptogenic and 10 species arequestionable. Some species seem to haveinvasive character, forming dense populations in semi- or heavily polluted areas,
including harbours (Branchiomma luctuosum, Hydroides dianthus, H. elegans, Polydora cornuta, and others), brackish waters(Desdemona ornata, Ficopomatus enigmaticus, Polydora cornuta, etc.) or shallow-water
benthic habitats (Eunice antennata, Lumbrineris perkinsi, Notomastus mossambicus,
Prionospio (Aquilaspio) krusadensis, etc.).
The WMED has a relatively low number ofalien species, in comparison with the
EMED, especially the Levantine Sea, which
is densely colonized by alien polychaetes.
The majority of the established alien
species in the WMED are of Pacific, IndoPacific, Indian and/or Red Sea origin,whereas less originate in the tropicalAtlantic (six species) or West Atlantic (sixspecies).
A relatively low number of newcomers
originating in tropical seas and anthropogenically introduced via the Gibraltar
Strait are true alien species. Most of themappear to have arrived via shipping, especially in harbours, as also happens in the
EMED. Although the polychaetes of the
WMED are relatively well known, newreports are occurring continuously, making sometimes difficult to elucidate if they
are true invaders, or unrecorded or overlooked species. This is especially evident
with small, meiobenthic species; for examMedit. Mar. Sci., 11/2, 2010, 381-493

ple, in a recent work (DEL-PILAR-RUSO
et al., in press) two small syllids (Syllis mauretanica and Parapionosyllis macaronesiensis; not included in Table) are reported forthe first time in the Mediterranean; both
species were described from NE Africa
and Canary and Madeira islands, relatively close to the Strait of Gibraltar. There is
no way to know if these species were accidentally introduced, were overlooked inprevious papers, or simply never found.
1.4. Alien crustaceans in the Western
Mediterranean
In total, 52 alien species of crustaceans
have been found in the WMED, of which
20 have self-maintaining populations, 20
are casual records, eight are questionable
and four are classified as cryptogenic.
These species belong to amphipods (4),cirripedes (5), decapods (20), stomatopods(1), cumaceans (1), planktonic and parasitic forms of copepods (16), and isopods(5) (Table 2).
In the decapods, we have not considered alien the Atlantic species found in the
Alboran Sea and whose historical presence
is known in Morocco. For example,
Brachynotus atlanticus, a tropical African
species known in Morocco (FOREST &GANT S, 1960), was found in Europe forthe first time in the littoral of Malaga(GARC A RASO, 1984a), and later inthe Bay of Cadiz, in the Atlantic, where it
forms a stable population (GARC A
RASO, 1985). The Atlantic shrimps
Penaeopsis serrata and Hymenopenaeus
debilis, known in Moroccan waters (ZARIQUIEY, 1968; PÉREZ FARFANTE,
1979), were recently found in the western
Mediterranean Sea (perhaps due to their
deep-water habitat); apparently, they arebeing found in more areas of the Mediterranean; the former in the Alboran Sea
Medit. Mar. Sci., 11/2, 2010, 381-493(ABELL & TORRES, 1998) and Sardinia Channel (MURA et al., 2003), thelatter in Alboran Sea and Balearic Island(CARTES et al., 2000). A similar consideration can be made for Merhippolyte
ancistrota, a species occurring in the Alboran Sea but also known in the Gulf ofCadiz and Morocco (GARC A RASO,
1996; UDEKEM D'ACOZ & URI ,1996), and Plesionika ensis (A. Milne
Edwards, 1881), cited in the Alboran Sea(GARC A RASO, 1981) and Gulf ofCadiz (GARC A RASO, 1996). However,
new references from the island of Crete,
Aegean Sea (LABROPOULOU &KOSTIKAS, 1999) and its absence inintermediate Mediterranean areas raise
doubts about the status of the latter
species in the EMED. There is an old citation of the species Plagusia depressa(Fabricius, 1775) and Plagusia chabrus(Linnaeus, 1758), now Guinusia chabrus(Linnaeus, 1758), brought to the port ofMarseilles by an iron vessel in 1873
(STEBBING, 1893), but they have notbeen included in Table 2, because they
have never again been captured (there areno later references).
Half the species originate in the IndoPacific, Pacific, Indian Ocean or Red Sea,
while seven species are known to have acircumtropical distribution. The remainders have a tropical/subtropical Atlantic
origin, many of them being known in thewestern Atlantic.
Two species, Synalpheus tumidomanus
africanus and Necora puber, not includedin earlier Mediterranean alien lists, areadded in Table 2. Synalpheus tumidomanus
africanus (syn. S. hululensis africanus) was
first captured in Israel (LEWINSOHN &HOLTHUIS, 1964, as Synalpheus hululensis) and considered as alien; later it was
found in the Alboran Sea (GARC A
417

RASO, 1984b; L PEZ DE LA ROSA &GARC A RASO, 1992; GARC A
RASO, 1996), a characterization that subsequently changed following reclassification of the Mediterranean specimens within the subspecies S. tumidomanus africanus(or S. africanus?), known from Casablanca
and Rabat, Morocco (LAGARD RE,
1971). The Alboran populations may represent a natural expansion of the distribution range of the species, which enteredthrough the Strait of Gibraltar and colonized the southern coast of Spain (perhaps
with the help of the Atlantic surface water
inflow). The references from Greece andTurkey (KOUKOURAS & KATTOULAS 1974; KOCATA , 1981) could indicate a separate advance in the EMED littoral from the Israeli populations. Recently it has been captured in Sicily (BACCI etal., 2010). The unresolved question iswhether the EMED populations are alien
or not. At this time we have classified it ascryptogenic.
Necora puber is included as ‘alien’
species because (in agreement withZIBROWIUS, 1992) we believe that its
presence in several areas of the Mediterranean is due to introductions, most likely
related to its widespread food use rather
than an entry through ballast waters. Only
the specimens collected west of Malaga(GARC A RASO et al., 2010) could haveentered the Mediterranean through theStrait of Gibraltar by their natural meansof dispersion.
With regard to other groups of crustaceans we should note that the available
data are scarce and often difficult to assess.
This is partly due to the smaller number oftaxonomic specialists and studies developed on these groups in the different
Mediterranean sectors. These limitations
lead to a) a downwards-biased estimation
418

of aliens in the groups of amphipods, cirripedes,
cumaceans,
isopods
andtanaidaceans and b) a high number ofquestionable and cryptogenic species.
The amphipod Elasmopus pectenicrus
has a circumtropical distribution: Atlantic
Ocean, Red Sea, Indian Ocean, Pacific
Ocean (described from New Guinea); it
has gradually colonized the EMED, theAdriatic and the CMED (ZAKHAMASRAIEB & CHARFI-CHEIKHROUHA,
2010). The latest finding could be clearly
the result of increased scientific effort in
the area.
In the WMED the only species
markedly invasive is Percnon gibbesi, whichshows a rapid expansion of its geographical distribution range, e.g. in the east Spanish coast (DEUDERO et al., 2005): firstrecorded from the Balearic Archipelago in1999 (GARCIA & REVIRIEGO, 2000),
it established populations since 2002 inBarcelona, 2003 in Alicante and Murcia,
2006 in Almeria.
Among alien planktonic copepods
reported in the WMED Sea, three areclearly Lessepsian immigrants (Triconia
hawii, T. rufa and T. umerus) and were
found in the Gulf of Naples. Two otherLessepsian immigrants, Labidocera detruncata and Calanopia elliptica, were reported
from the Gulf of Naples (VIVES &SHMELEVA, 2007), but were never foundin the more than 20 years time series ofsamples collected bi-weekly to weekly inthe area (Mazzocchi, pers. comm.): therefore, their presence is considered as questionable. Pseudocalanus elongatus andParacalanus indicus seem to be cryptogenic
in the area; according to RAZOULS et al.
(2005-2010) the latter species could be confounded with Paracalanus parvus, a very
common native Mediterranean species.
Two copepods parasitic on oyster beds
Medit. Mar. Sci., 11/2, 2010, 381-493

(Myicola ostreae and Mytilicola orientalis)are well established in the WMED, after
their accidental introduction into the
French coast with the Pacific oyster Crassostrea gigas (CLANZIG, 1989).
1.5. Alien molluscs in the Western Mediterranean
The WMED has a low incidence ofalien Mollusca with 34 recorded species (22established, nine casual, two cryptogenic
and one questionable) in a total of around1500 native species. Two of these (Cerithium scabridum and Brachidontes pharaonis)
are found only very close to the Strait ofMessina and are merely outliers from theconfirmed area where these species haveestablished in the Ionian Sea. Others, like
Chromodoris quadricolor, Pinctada radiata,
Rissoina spirata and Thais lacera, had casual occurrences that are unlikely to persist,
although they may be firmly established in
the EMED. Leaving aside the outliers andthese casual species, the cryptogenic
species, and the questionable report of Saccostrea cucullata for Tunisia, we are left
with not more than 25 alien molluscs definitely relevant to the basin.
The north coast of Tunisia represents
a major step towards the colonization of
the WMED. Two of the hardiest Lessepsian immigrants are present there, having
made their way across the WMED to thecoast of Spain and are locally invasive.
These
are
Bursatellaleachii(WEITZMANN et al., 2009; RAMOSESPL et al., 2010) and Fulvia fragilis(TAMAYO-GOYA, 2008). B. leachii also
exists in the Atlantic, but the hypothesisthat WMED populations entered throughGibraltar does not hold since the species
does not exist in Morocco.
The largest pool of alien molluscs inthe WMED includes species that are
Medit. Mar. Sci., 11/2, 2010, 381-493

actively cultured (for example Crassostrea
gigas and Ruditapes philippinarum, to alesser extent Mercenaria mercenaria) andspecies that were probably introduced
accidentally along with aquaculture(Crepidula fornicata, Gibbula albida, Musculista senhousia, Xenostrobus securis),
thereby comprising one-fourth of thereported aliens. M. senhousia and X.
securis are locally invasive in the basin, but
C. fornicata, which is one of the most invasive aliens in the NE Atlantic, has difficulty in surviving where it has been introduced in the Mediterranean. Gibbula albida was originally an Adriatic endemic, but
is now introduced in the Ebro Delta(TRIGO, 1981) and the French Mediterranean lagoons (CLANZIG, 1989), as wellas on the Atlantic coast of France.
A unique and certainly recent featureis the occurrence in the vicinity of Tunis(Tunisia) of two alien species (Favorinus
ghanensis, Mitrella psilla) introduced
through shipping from tropical West
Africa (BEN SOUISSI et al., 2004; ANTIT
et al., 2010). More generally, harbours area focus for established populations ofaliens, with random origins. Other examples are the bivalve Theora lubrica in
Leghorn, Italy, and the gastropod
Bostrycapulus odites in Alicante, Spain,
which did not spread out to the neighbouring open sea.
On the whole, the incidence of alienmollusc species in the WMED is heavily
concentrated in marginal marine environments such as the lagoons of the French
Mediterranean coast, the Spanish Mar
Menor and Ebro Delta, and the Italianlagoons of Caprolace and Fusaro on the
Tyrrhenian coast. With the exception of
the recent report of the nudibranch Godiva quadricolor from the large harbour of
Algeciras (CERVERA et al., 2010), the419Alboran Sea is free of alien Mollusca, a situation which may be related to its exceptionally high species richness estimated as
about 1200 species. More generally, theopen sea in the WMED is hardly, if at all,impacted by alien species of molluscs.
Although a couple of alien molluscs in
the Mediterranean are of West African origin, there is no species yet that has gradually entered through the Strait of Gibraltar
and become newly established in the
Mediterranean. As already noted by
GOFAS & ZENETOS (2003), all theprevalently Atlantic species found in thewesternmost Mediterranean have a historical range since at least the 19th century andare, therefore, to be considered native
species. The WMED is nevertheless asource area for such species as Siphonaria
pectinata, native to Algeria and Spain but
introduced in Greece and recently expanding its range towards northern Tunisia.
Although not qualifying as ‘aliens’ and notincluded in the species counts, one must
keep in mind the fact that such species as
Eastonia rugosa (see ALBANO, 2006) are
expanding their range.
1.6. Miscellaneous invertebrates in theWestern Mediterranean
Bryozoa include five alien species in
the WMED, out of 23 known from thewhole basin. Arachnoidea protecta is an
Indo-Pacific species first recorded in the
WMED by CHIMENZ GUSSO et al.
(1998), further records being provided by
D'HONDT & CHIMENZ GUSSO(2006). The morphological divergenceobserved between the Indo-Pacific andMediterranean specimens led Harmelin(in ZENETOS et al., 2005) to believe thatthey represent two distinct species. However, OCCHIPINTI-AMBROGI et al.
(2010) maintain A. protecta as established.
420

Alien Ascidiacea are represented byeight species (out of 16), most of themestablished. Five species are circumtropical. Trididemnum cf. savignyi was firstrecorded by LAFARGUE (1972) at PortCros, France. LAFARGUE (1974) revised
the Trididemnum tenerum complex, assigning the record by PÉR S (1954) fromsouthern Tunisia to T. cf. savignyi. Subsequent records of this species include Catalonia and Columbretes Islands, Spain(RAMOS-ESPL , 1988). Perophora multiclathrata and Ecteinascia styeloides havebeen reported from Corsica (MONNIOT,
1983) and the northern Tyrrhenian Sea(MASTROTOTARO & TURSI, 2010),
respectively. Microcosmus squamiger, firstrecorded in Bizerte (as M. exasperatus) is
widespread in the WMED (synthesis in
TUR N et al., 2007). Cystodytes philippinensis, previously observed in southern
Tunisia (MÉLIANE, 2002), has beenrecorded recently in the Balearic Islands(D AZ-VALDÉS & RAMOS-ESPL ,2010). This cryptogenic species had probably been confused in the past with its congener C. dellechiajei, widely distributed in
the Mediterranean Sea (IZQUIERDO~OZ et al., 2009). The Western
MUN
Atlantic species Distaplia bermudensis andthe Eastern Pacific species Polyandrocarpa
zorritensis have been introduced by shellfish culture (TUR N & PEREA, 1988;
BRUNETTI & MASTROTOTARO,
2004; MASTROTOTARO & BRUNETTI, 2006). Styela clava, originatingfrom the north-western Pacific, has been
spreading along the European Atlantic
coast since 1954 and has only recently beenrecorded in the WMED; however, its
absence from harbours and marinas closed
to Gibraltar may suggest that this species
has also been accidentally transported into
the Mediterranean Sea by shellfish transMedit. Mar. Sci., 11/2, 2010, 381-493

fer (DAVIS & DAVIS, 2008).
Cnidaria are an exception to most taxa
in that the majority of aliens occur in the
WMED rather than in the EMED: 26
species (out of 46 listed for the wholebasin). Alien Anthozoa are represented by
two species, out of four reported for thewhole Mediterranean. Although firstrecordedinthe
Ligurian
Sea(ZIBROWIUS, 1974), Oculina patagonica
is abundant in the southern part of the
WMED, both in Spain (ZIBROWIUS &RAMOS, 1983; BALLESTEROS, 1998;
IZQUIERDO et al., 2007) and along theNorth African coast (SARTORETTO etal., 2008). Eighteen alien species (out of37) belong to Hydrozoa, a group well represented in the fouling assemblages(MORRI & BOERO, 1986) that colonize
the large ports of the WMED. Cordylophora caspia has been reported only forthe brackish lake of Fondi, Tyrrhenian Sea(MORRI, 1979). Two alien Scyphozoa
were found in the WMED: a single specimen of Phyllorhiza punctata was seen in
October 2009 off NE Sardinia (BOERO etal., 2009), while Stomolophus meleagris is
the only one recorded exclusively in the
WMED: this may suggest it entered fromthe Atlantic, although the species is also
known from the Pacific (MORAND &DALLOT, 1985).
Both species of alien Ctenophora
reported for the Mediterranean occur in
the WMED (BOERO et al., 2009;
FUENTES et al., 2009; MILLS, 2009),
while Sipuncula and Pycnogonida haveone alien species each (PANCUCCIPAPADOPOULOU et al., 1999; BARTOLINO & CHIMENZ GUSSO, 2010).
The only alien echinoderms recordedin the WMED are two seastars and one
ophiuroid of Indo-Pacific origin: Acanthaster planci, Protoreaster nodosus, andMedit. Mar. Sci., 11/2, 2010, 381-493

Ophiactis savignyi. The former has beenreported from Port Cros, France, in 2000
(ICES, 2006) with no further details(hence it is classified as questionable),
whereas two individuals of Protoreasternodosus, probably released accidentallyfrom a private aquarium, have been caught
by trawling off Majorca, Balearic Islands,
Spain, in 1981 (ALVARADO et al., 1986).
Casual is considered the presence of Ophiactis savignyi in Banyuls (GUILLE, 1969).
The only alien Porifera in the WMED
is Paraleucilla magna, which is proliferating
acrosstheMediterranean(GUARDIOLA et al., 2010). Platyhelminthes are represented by Allolepidapedon fistulariae, an endoparasite of Fistularia commersonii reported from Sardinia (PAIS et al., 2007).
1.7. Alien fish in the Western Mediterranean
MASSUTI et al. (2010) reported 38
new fish in the WMED since the middle
20th century. Here we report 45 species (20established, 25 casual) inventoried in the20th century. Most of them are of tropical(subtropical) Atlantic origin.
The entrance of alien fish in the
WMED has drawn the attention on the socalled effect of ‘tropicalization'. This phenomenon has expanded geographically,reaching the EMED. Such is the case for
the Atlantic origin species Carcharhinus
altimus, Acanthurus monroviae, Pisodonophis semicinctus, Sphoeroides pachygaster, Pagellus bellottii and Seriola fasciata,
which have recently entered the Mediterranean through the Strait of Gibraltar, andhave already reached the EMED.
Documentation on some species islimited so that those here reported as casuals could have already established populations. This could be the case of a) benthic421

species with limited swimming capacity
such as Microchirus boscanion, which has
been captured at different localities andtimes (MASSUTI et al., 2010); b) confusion with others, such as the soleids
Microchirus hexophthalmus and Synaptura
lusitanica known for the Iberian Peninsula
and the Gulf of Lions (MATALLANAS,
1984). Some species, such as Anarhichas
lupus, Aluterus monocerus, Scorpaena
stephanica, Fistularia petimba and Zenopsis
conchifera, are known only from single
observations and are still limited to the
WMED. The latest findings include Lutjanus jocu (VACCHI et al., 2010) andKyphosus incisor (ORSI-RELINI et al.,2010), both in the Ligurian Sea.
Within the alien Indo-Pacific species,
Pomadasys stridens was reported for the
first time in the WMED and then it was
known in the EMED, where it is common;
Abudefduf vaigiensis, a casual species in theLevantine basin, has been reported threetimes in the WMED could be similar. By
contrast, Siganus luridus, and Fistularia
commersonii, which were first reported in
the EMED, where they are now wellknown and even common, have expanded
westwards and are considered established.
Fistularia commersonii has expanded as far
north as the Ligurian Sea (OCCHIPINTIAMBROGI & GALIL, 2009) and as far
west as Algeria (HEMIDA & CAPAPÉ,2009) and Spain (S NCHEZ-TOCINO
et al., 2007).
In the WMED, only a few species
appear regularly in the catches of somefisheries. That could be the case of Solea
senegalensis, whose presence in the
WMED was recorded in 1920 off the Iberian coast and has presently extended toAlgeria, Tunisia and the Gulf of Lions, andGymnammodytes semisquamatus reported
only in the north-eastern Iberian coast,
422

where it is exploited in commercial fishery.
Diplodus bellottii and Pagellus bellottii,
reported at the Maghrebine and Iberian
coasts of the Alboran Sea, are occasionallycaptured by commercial fisheries(MASSUTI et al., 2010).
2. CENTRAL MEDITERRANEAN
The bulk of the CMED is represented
by the Ionian Sea, the least known of all
theMediterranean
sub-basins(ZENETOS et al., 1997). Local endemics,
mostly within molluscs, have been reported for its southern, non-European shores(SABELLI & TAVIANI, 1980) but also
for Malta (EVANS et al., 2010). The Ionian is connected to the WMED throughthe narrow Strait of Messina, a micro-sector that harbours a wealth of biogeographic peculiarities, including Pliocene Atlantic
remnants and local endemisms (FREDJ &GIACCONE, 1995), and the larger Straitof Sicily, the meeting point of native
Western and Eastern Mediterranean
species (BIANCHI, 2007), as well as ofaliens of either Atlantic or Indo-Pacificorigin (COLL et al., 2010). Reflecting thissituation at the crossroads, the composition of the alien faunas is much more balanced between different sources than in
the EMED. Its situation at the transition
between the eastern and western basins
make it a particularly sensitive place formonitoring the progression of the much
more numerous aliens already establishedin the EMED.
2.1. Alien protozoans in the Central
Mediterranean
Fish pasteurellosis from the protist
Photobacterium damselae was reported
from cultured seabass and seabream
installations in the Greek Ionian Sea andMedit. Mar. Sci., 11/2, 2010, 381-493

from Malta (BAKOPOULOS et al., 1995,
1997).
Only 4 alien foraminiferal species arereported for the CMED: Amphistegina lobifera, which probably has established apopulation around the island of Malta(YOKES et al., 2007), A. lessonii and A.
madagascariensis documented along thecoast of southern Tunisia (BLANCVERNET,
1969;
HOLLAUS&HOTTINGER, 1997). Planogypsina acervalis seems to have been recorded in thisarea only by BLANC-VERNET (1969).
2.2. Alien macrophytes in the Central
Mediterranean
In the CMED, 57 alien macrophytes
have been hitherto reported, which represent 46% of the alien macrophytes known
for the Mediterranean Sea as a whole. The
major vectors of introduction are the shiptraffic (fouling, ballast waters), the Suez
Canal, and, in the south Italian coastallagoons, the shellfish aquaculture. Amongthem, 35 alien macrophytes are well established, four are casual, 16 are cryptogenic/questionable, and two require confirmation. Casual taxa (i.e. Grateloupia
lanceolata, Hypnea anastomosans, Polysiphonia paniculata and Undaria pinnatifida)might turn into the established status in
the future. The occurrence of Batophora
sp. and Gracilaria arcuata in the CMED
requires confirmation.
Of the 21 invasive or potentially invasive macrophytes introduced in the
Mediterranean (Table 3), 13 have beenreported in the CMED. Undaria pinnatifida that was only reported from the Mar
Piccolo of Taranto (south Italy), seems tohave disappeared from the region(GRAVILI et al., 2010).
The Ionian coasts and Sicily are themost impacted regions, with 35 and 36
Medit. Mar. Sci., 11/2, 2010, 381-493

alien macrophytes respectively, followed
by Tunisia (25) and Libya (12). Such differences are probably due to a lower investigation effort, particularly in Libya.
The alien macrophytes of the CMED
come from temperate/cold regions (NEAtlantic and NW Pacific) and from tropical regions in almost equal proportions (51and 49%, respectively). A total of 17 putative Lessepsian immigrants have been
hitherto recorded in the region.
2.3. Alien polychaetes in the Central
Mediterranean
The CMED includes 36 alien speciesof polychaetes (17 established, nine casual,and ten questionable). Aliens alreadyestablished in the EMED have colonized
this area to subsequently spread to the
WMED: examples are provided by thethree Indo-Pacific species, and likely
Lessepsian immigrants, Eunice antennata,
Pseudonereis anomala (both established)
and Protodorvillea egena (questionable).
By contrast, Linopherus canariensis andOphryotrocha japonica, both established,
could be indicative of a passage from west
to east. The former is very abundant in the
Faro Lake, on the Messina Strait(COSENTINO et al., 2009), together withanother alien species: Syllis hyllebergi,
which is exclusively found in this area
(COSENTINO, in press). Other exclusive
species of the CMED are Eunice floridiana, Epidiopatra hupferiana hupferiana,
Epidiopatra hupferiana monroi and Isolda
pulchella, mostly collected in the Gulf ofNoto (Sicily), all considered casual, nonestablished
taxa(OCCHIPINTIAMBROGI et al., 2010). On the contrary,
Ophryotrocha japonica is very abundant in
the Mar Piccolo of Taranto, an area considered a hot spot of alien species diversity
within the Italian coasts (OCCHIPINTI423

AMBROGI et al., 2010). In this locality
one of the most conspicuous populationsof Branchiomma luctuosum is present(LICCIANO et al., 2002), and it seems
here to compete with the native species
Sabella spallanzanii (Giangrande, pers.
observ.). Most of the other establishedspecies in this area are the same as thosethat are common in the whole basin suchas Ficopomatus enigmaticus, Hydroides
dianthus, H. diramphus, H. elegans,
Lysidice collaris, Metasychis gotoi, Neopseudocapitella brasiliensis, and Notomastus aberans.
2.4. Alien crustaceans in the Central
Mediterranean
A total of 46 alien crustacean arereported in the CMED (26 established, 13
casual, four questionable, three cryptogenic).
Three West Atlantic species are wellestablished at least in some regions of the
CMED: Percnon gibbesi, Libinia dubia,
and Rimapenaeus similis, which was discovered on the trawling grounds of the Gulf ofGabès (BEN HADJ HAMIDA-BEN
ABDALLAH et al., 2010). Percnon gibbesi
was first observed in 1999 at Linosa Island(RELINI et al., 2000). It rapidly spread inother localities of the CMED: Pantelleria(GALIL et al., 2002), Malta (BORG &ATTARD-MONTALTO, 2002), theshores of southern and north-western Sicily (MORI & VACCHI, 2002), the Strait ofMessina (BELLANTONI & CORAZZA,
2003) and the Gulf of Taranto (FACCIA& BIANCHI, 2007).
Several Lessepsian immigrants, such
as Trachysalambria palaestinensis, Eucrate
crenata, Metapenaeus monoceros andMetapenaeus stebbingi, established in the
EMED have spread westward as far as
Tunisian waters. Their apparent absence424

along the Libyan shores may well be consequence of a lower research effort, butthey are also absent from Malta and thesouthern Italian shores. Another IndoPacific species is Plagusia squamosa.
The following species were recorded
only once and with a single specimen:
Alpheus inopinatus, Dromia spinirostris, Sirpus monodi, Callinectes sapidus, Grapsus
granulosus. The presence of one adultspecimen of Paralithoides camtschaticus, astrictly boreal species, in this sector of theMediterranean (FACCIA et al., 2009) isreally puzzling.
The number of alien copepods is low(five) in the CMED and only two are
Lessepsian immigrants (Euchaeta concinna, Triconia umerus). The presence of the
Antarctic to Subantarctic species
Spinocalanus terranovae in the Malta area
is questionable (SCIBERRAS &SCHEMBRI, 2007).
The bentho-planktonic calanoid copepod Pseudocyclops xiphophorus was previously recorded only in coastal waters ofMozambique. The Mediterranean P.
xiphophorus specimens were collectedfrom fouling attached to submerged mooring posts and ropes in the brackish LakeFaro, eastern Sicily. It is classified as cryptogenic because according to ZAGAMI etal. ( 2005) it could represent a relict population of Tethyan origin. The genus Pseudocyclops has a worldwide distribution
from temperate to tropical shallow coastal
and brackish waters.
2.5. Alien molluscs in the Central Mediterranean
The CMED has reports for 38 alien
species of molluscs (19 established, 13 casual, two cryptogenic and four questionable)
in a total of 1000 to 1500 native species.
Among the species of Indo-Pacific origin,
Medit. Mar. Sci., 11/2, 2010, 381-493

there are 12 species that qualify as Lessepsian (e.g. Pinctada radiata, the earliest
reported Lessepsian mollusc) and six thatare definitely non-Lessepsian (e.g. Melibe
viridis which was first detected on the Ionian
coast of Greece), keeping the same proportion as in SE Turkey but far from the 90%
of Lessepsians along the Levantine coast.
There are scattered reports of species
from the temperate Atlantic (Crepidula
fornicata, Polycerella emertoni, Anadaratransversa, Zygochlamys patagonica) andNorth Pacific (Crassostrea gigas purposely
introduced for farming, Musculista senhousia, Ruditapes philippinarum) but most are
far less successful in this particular context,
than in the northern part of the Mediterranean (Ligurian Sea, Adriatic Sea) wheresome of them are invasive (see Table 3).
Considering the proximity of the EMED,
the total numbers are neverthelessremarkably low. Few species are invasive,
among them could be considered thebivalves Pinctada radiata and Fulvia fragilis
and the opisthobranch Bursatella leachii in
the Gulf of Gabès.
Tunisia holds altogether 24 alien
species (13 established, three casual, three
cryptogenic, five questionable) but a distinction must be made between the easternand southern coasts bordering the Gulf ofGabès, which are a continuation of the virtually unknown Libyan coast, and thenorth coast west of Cap Bon, which is considered as part of the WMED. A cowry,
Erosaria turdus, had a spectacular onset inrecent years and is so invasive that it willprobably displace the native Zonaria
pyrum. The limpet Cellana rota may be thenext spectacular invader in this part of theMediterranean.
Malta has possibly one of the beststudied molluscan faunas in the southernpart of the EU, and has reports for 17
Medit. Mar. Sci., 11/2, 2010, 381-493

species. The Ionian coast of Greece has
only nine species reported, of which two(Bursatella leachii and Pinctada radiata)
are among the most widespread Lessepsian immigrants and five are non-Lessepsian species of tropical Indo-Pacific origin.
The Ionian coast of Italy holds 14 species.
These three areas share very much of theiraliens including the most widespread
Lessepsian species (Bursatella leachii and
Pinctada radiata, Cerithium scabridum andFulvia fragilis in Italy and Malta) which
here are secondary introductions from the
EMED. Also shared are some of the tropical Indo-Pacific species which started
their spread in this area (Aplysia dactylomela, Haminoea cyanomarginata, Melibe
viridis). Some species, like Anadara transversa or Musculista senhousia, which areinvasive in the Adriatic, have spreadtowards the Italian Ionian coast.
2.6. Miscellaneous invertebrates in theCentral Mediterranean
Bryozoa are comprised of eightspecies (out of 23 known for the whole
Mediterranean), of various origins (threecircumtropical, three Pacific, twoAtlantic). The records of Celleporaria aperta and C. pilaefera at Malta have been considered questionable by SCIBERRAS &SCHEMBRI (2007).
Nearly half of the alien species ofAscidiacea inventoried in the Mediterranean (seven out of 16) are known for the
CMED. These aliens are of various origins(Western Atlantic, circumtropical, IndoPacific, Eastern Pacific) and both the alien
species of Microcosmus, the Indo-PacificM. exasperatus and the circumtropical M.
squamiger, occur in the CMED~OZ et al., 2009).
(IZQUIERDO-MUN
Most records of alien ascidians come from
Taranto harbour (BRUNETTI &425

MASTROTOTARO, 2004; MASTROTOTARO et al., 2004; MASTROTOTARO & BRUNETTI, 2006).
The CMED hosts nine out of 46 alien
species of Cnidaria known for the wholebasin. Hydrozoa include six species, all witha wide distribution in warm waters. The
cryptogenic species Cordylophora caspia
was found abundant in the low salinity
waters of the ‘Palude del Capitano’, Gulf of
Taranto (BIANCHI et al., 1994). Scyphozoa are represented by the three species
Cassiopea andromeda (SCHEMBRI et al.,2010a), Phyllorhiza punctata (ABEDNAVANDI & KIKINGER, 2007) andRhopilemanomadica(SIOKOUFRANGOU et al., 2006).
The only alien species of Ctenophora
is Mnemiopsis leidyi, recorded from Isola diCapo Rizzuto, Gulf of Taranto (BOERO
et al., 2009), whereas the only alien speciesof Porifera is Paraleucilla magna, originating from the SW Atlantic (ZAMMIT et al.,2009). In the case of Sipuncula, there arequestionable records of the circumtropical
Aspidoshiphon mexicanus from Malta andLampedusa(PANCUCCI-PAPADOPOULOU et al., 1999; SCIBERRAS &SCHEMBRI, 2007).
Alien Echinodermata are represented
by three species (out of 12 for the whole
Mediterranean). The record of Eucidaristribuloides from Malta was also the first of
this species for the Mediterranean(TANTI & SCHEMBRI, 2006).
2.7. Alien fish in the Central Mediterranean
In the CMED, there are records of 50
alien fish. Of them, 25 seem to be established, 19 are casual, whilst the presence ofsix species is questionable. The bulk of theserecords comes from the Sicily Strait (seeGUIDETTI et al., 2010 and references426

therein), especially from Pelagie Islands andMalta but also from Tunisia (BEN
SOUISSI et al., 2006a,b) and Libya(SHAKMAN & KINZELBACH, 2006;
BEN ABDALLAH et al., 2007). A few
records are also available from the northern
Ionian Sea (TORCHIO, 1963; MASTROTOTARO et al., 2007) and the Messina
Strait, as in the case of Galeocerdo cuvier(CELONA, 2000), Pinguipes brasilianus(ORSI-RELINI, 2002), Platycephalus indicus (CASTRIOTA et al., 2009) and Psenes
pellucidus (NAVARRA et al., 2008).
Evidence of established populations in
the CMED are mostly related to the occurrence of multiple records of the same
species, such as for Cephalopholis taeniops(GUIDETTI et al., 2010) and Pisodonophis
semicinctus (RAGONESE & GIUSTO,
2000) while a few species such as Siganus
luridus, Fistularia commersonii, Seriola carpenteri, Stephanolepis diaspros, Sphoeroides
pachygaster have been collected or observedin large numbers (RAGONESE et al., 1997;
PIZZICORI et al., 2000; AZZURRO &ANDALORO, 2004; BRADAI et al., 2004;
AZZURRO et al., 2007). Latest records
include Selene dorsalis (VELLA &DEIDUN, 2009) and Opleognathus fasciatus (SCHEMBRI et al., 2010b).
The geographical partitioning of
CMED alien species was: 19 of Atlantic
origin, 25 of Indo-Pacific or Pacific origin,and three circumtropical. If we look onlyat the established species, the number ofIndo-Pacific or Pacific fish (15) is almost
double than the number of Atlantic newcomers (eight) indicating unbalanced ratio
towards fish coming from the east. Indeed,
the progressive penetration westwards ofLessepsian immigrants is a continuous
phenomenon, which has accelerated during the last decades, as exemplified byUpeneus pori, Pempheris vanicolensis,
Medit. Mar. Sci., 11/2, 2010, 381-493

Sphyraena chrysotaenia and Siganus rivulatus (BRADAI et al., 2004; BEN SOUISSI
et al., 2006a,b).
3. THE ADRIATIC SEA
The Adriatic Sea is a rather uniqueand differentiated area within the
Mediterranean, with a strong contrast
between the predominantly linear sandy
shores along the western (Italian) side, andthe opposite complex coasts of the easternside (Slovenia, Croatia, Montenegro andAlbania) forming a maze of islands andinlets with rocky shores. The hydrographic
conditions are also peculiar, with very low
winter temperatures in the northern part,which is also quite shallow (40 m depth),and very hot summers in the southern part,which is much deeper. All these features
lead to differentiation between the northern and southern Adriatic areas.
From a biogeographic standpoint, theAdriatic Sea is divided in three sectors,
arranged more or less latitudinally. The
Northern Adriatic Sea is perhaps the mostpeculiar sector of the whole Mediterranean. Strong winter cooling, low salinity
due to significant river input, and comparatively great tidal range make it more similar to the Northern Atlantic than to therest of the Mediterranean. This is reflectedin the biota, which includes disjunctAtlantic-Adriatic species whose occurrence contributes to what has been called
the Northern-Adriatic ‘sub-Atlanticism’(SACCHI et al., 1985). Mediterranean
endemics are scarce and the overall diversity is the lowest of the Mediterraneanbasin, so that the phrase ‘North-Adriatic
gap’ has also been coined (SACCHI,
1983). In addition, peculiar to this sector
are some taxa with Black Sea affinity(SACCHI et al., 1985; BIANCHI et al.,Medit. Mar. Sci., 11/2, 2010, 381-493

2004). The Central Adriatic is characterized by the lack of both Northern-Adriatic
endemics and Atlantic-Adriatic disjunctions. Mediterranean endemics and thermophilic species are still scarce, so that this
sector is less diverse than the Gulf of Lionsand Ligurian Sea, to which it resembles.
However, it is significantly richer in species
than the Northern Adriatic. Finally, theSouthern Adriatic exhibits a transitional
character between the Adriatic and the
Ionian Sea, with which it communicatesthrough the Otranto Strait. Affinities with
the WMED are strongly reduced.
3.1. Alien protozoans in the Adriatic Sea
Three protozoans pathogenic to shellfish, namely Marteilia refringens, Bonamia
ostrea and Perkinsus olseni (formerly
Perkinsus atlanticus) have been reported
from shellfish cultures in the Adriatic Sea.
Serious outbreaks of pasteurellosis causedby Photobacterium damselae (formerly
Pasteurella piscicida) in the Adriatic arereported by CESCHIA et al. (1991).
Foraminiferal aliens from the Adriatic
are few (six species). Moreover, all of them
have been reported based on single specimens, in only one publication or site andcould be considered as cryptogenic/casual.
This is the case for warm water alien
species such as Archais angulatus(LANGER & HOTTINGER, 2000),
Clavulina angularis (BANCHETTI et al.,2009), Coscinospira hemprichii, Cymbaloporetta plana (as Trethomphalus bulloides)and Sorites orbiculus (CIMERMAN &LANGER, 1991). A. angulatus is only
reported from the Adriatic (LANGER &HOTTINGER, 2000). Schackoinella
imperatoria, described from the coast ofcentral Adriatic (CIMERMAN &LANGER, 1991), could be cautiously considered a cryptogenic/casual form.
427

3.2. Alien macrophytes in the Adriatic Sea
A total of 49 alien macrophytes has
been hitherto reported from the AdriaticSea, which represent 39% of the alien
macrophytes reported from the Mediterranean Sea. The major vectors of introduction are the shellfish aquacultureindustry in northern coastal lagoons andthe ship traffic (fouling, ballast waters).
Among them, 35 alien macrophytes arewell established, three are casual, and 11
are cryptogenic/questionable. Casual taxa
(i.e. Acrothamnion preissii, Hypnea valentiae and Polysiphonia paniculata) might turninto the established status in the future.
The cryptogenic/questionable category
includes 11 species (Table 2).
Of the 21 invasive or potentially invasive macrophytes introduced into theMediterranean (Table 3), 14 are present in
the Adriatic Sea. Among them, cold temperate taxa (i.e. Codium fragile, Gracilaria
vermiculophylla, Grateloupia turuturu,
Dasysiphonia sp., Sargassum muticum andUndaria pinnatifida) grow in north-western
coastal lagoons, especially in the Lagoonof Venice and the lagoons of the Po Delta,
while the warm temperate and tropicaltaxa (i.e. Asparagopsis spp., Caulerpa racemosa var. cylindracea, Caulerpa taxifolia,
Halophila stipulacea, Lophocladia lallemandii, and Womersleyella setacea) invade
the marine habitats of southern Italy andthe east coasts of the Adriatic (Croatia,
Albania), with the exception of Acrothamnion preissii. The invasive Gracilaria vermiculophylla is hitherto known only in the
lagoons of the Po Delta.
Northern Adriatic coasts are the mostimpacted, with 34 aliens out of a total of 49(69% of alien macrophytes of the Adriatic), while only 15 aliens were reported fromthe Central and Southern Adriatic.
SFRISO et al. (2009 and unpublished data)428

listed up to 33 aliens (67% of alien macrophytes of the Adriatic) in the Lagoon of
Venice. Such a difference is probably due
to a flood of introductions along with shellfish transfers (oysters, mussels, Manila
clams) on the north-Italian coasts whereaquaculture is extensively developed.
The majority of alien macrophytes of
the Adriatic basin (35 taxa) come fromtemperate and cold regions (NE Atlantic
and NW Pacific) in relation with the major
vector of introduction (shellfish transfer).
Only seven putative Lessepsian immigrants (i.e. Acanthophora nayadiformis,
Chondria pygmaea, Halophila stipulacea,
Hypnea spinella, Hypnea valentiae, Lophocladia lallemandii and Ulva fasciata) havebeen hitherto registered, mainly from thenorthern coasts. However, Acanthophora
nayadiformis and Hypnea spinella could
alternatively originate from the Atlantic
Ocean. The strains of Hypnea valentiae andUlva fasciata found in the Lagoon of
Venice may originate from the north-western Pacific as in the Thau Lagoon.
3.3. Alien polychaetes in the Adriatic Sea
The Adriatic hosts only 22 alien polychaete species (11 established, six casual andfive questionable), which represent 17% of
the Mediterranean polychaete alien species.
At least four species appeared exclusive of
the Adriatic: Fabriciola ghardaqa, Megalomma claparedei, Platynereis australis and Syllis
alose, the latter two being considered asquestionable taxa.
The established species are among
those well known and settled in the whole
Mediterranean area, and most of them arealso invasive, such as Branchiomma luctuosum, Ficopomatus enigmaticus, Hydroides
dianthus, H. elegans, but also Lysidice collaris, Metasychis gotoi, Neopseudocapitellabrasiliensis, and Notomastus aberans. The
Medit. Mar. Sci., 11/2, 2010, 381-493

most recently introduced species is B. luctuosum, present with very large populations
along the Mediterranean coasts. At present
this species seems to have reached also the~o Paulo, in the Bay ofBrazilian Coast of Sa
Santos, in the vicinity of the largest seaportin Latin America (DE MATOSNOGUEIRA et al., 2006), underlining its
ability to invade new areas, possibly transported by ships. An introduction via theSuez Canal and possible transfer via shipping, can be postulated not only for B. luctuosum, but also for the other two Red Sea
sabellid species, exclusive of this area andcollected on the hard bottoms of the Southern Adriatic coast: M. claparedei and F.
ghardaqa that were, however, found only
once with very few individuals and considered therefore casual (GIANGRANDE &MONTANARO, 1999; GIANGRANDE& LICCIANO, 2008). By contrast, theother sabellid Novafabricia infratorquata is
a Caribbean species that must have been
introduced from the Strait of Gibraltar,
appearing established in the WMED andcasual in the Southern Adriatic along theApuliancoast(LICCIANO&
GIANGRANDE, 2006).
3.4. Alien crustaceans in the Adriatic Sea
A total of 24 alien crustacean arereported from the Adriatic (11 established,
11 casual, two cryptogenic).
The first records of alien species introduced in the Northern Adriatic by maritime traffic date back to the years before
World War I, when STIASNY (1908)
recorded the capture of Plagusia squamosa(Herbst, 1790), Thenus orientalis (Lund,
1793) and Portunus sanguinolentus (Herbst, 1796), respectively in the harbours of
Trieste and Fiume (Rijeka), at that timethe main harbours of the Austro-Hungarian Empire (BABIC, 1913). The latter two
Medit. Mar. Sci., 11/2, 2010, 381-493

species are excluded from our list following GALIL et al. (2002), whereas P.
squamosa is kept as questionable because
thriving populations were recently reported from elsewhere in the Mediterranean:
Tunisia and Libya (ZAOUALI et al.,2007).
Only three crabs (Callinectes sapidus,
Dyspanopeus sayi and Rhithropanopeus
harrisii) are definitely established in thearea and all three originate from theAtlantic coast of USA. The spreading of
the two panopeid crabs in the Adriatic Sea
in recent years (FROGLIA & SPERANZA, 1993; MIZZAN & ZANELLA,
1996; ONOFRI et al., 2008) has been facilitated by the development of mussel aquaculture both in lagoons and open sea, withtransfer of mussels seed and half grownmussels among aquaculture plants.
The latest record is that of the IndoPacific crab Charybdis lucifera, caught six
miles off the Venetian coast (MIZZAN &VIANELLO, 2009).
The following species were recorded
only once and with single specimens: Marsupenaeus japonicus, Scyllarus caparti,
Callinectes danae, Charybdis lucifera, Eriocheir sinensis, Hemigrapsus sanguineus,
and Charybdis japonica (FROGLIA, inpress).
Only three planktonic copepod species
were recorded as aliens in the AdriaticSea, originating from the Atlantic andIndo-Pacific oceans.
3.5. Alien molluscs in the Adriatic Sea
The Adriatic Sea has possibly the oldest alien (treated as cryptogenic) marine
species in the Mediterranean Sea, viz. Littorina saxatilis, originally described from
Venice (OLIVI, 1792) before it could besuspected of being introduced from the
NE Atlantic (JANSON, 1985). The Adri429

atic holds a few endemic species, one ofwhich (Gibbula albida) has been introduced into the WMED and elsewhere.
The Adriatic holds only 27 alien
species (15 established, nine casual andthree cryptogenic) but the striking characteristic is the high proportion of themwhich have become invasive. Together
with the Levantine basin, the Adriatic may
be the part of the Mediterranean which
has been most transformed by the onset ofalien species. The most invasive species
include Anadara kagoshimensis (formerly
known as A. inaequivalvis), Musculista senhousia, Rapana venosa and Ruditapes
philippinarum, all originating from thetemperate North Pacific and therefore
comfortable despite seasonal lows in sea
water temperature. Anadara kagoshimensis, now coupled to the possibly Atlantic
Anadara transversa, has formed spectacular accumulations on the NE Italian coast(RINALDI, 1985) and has profoundly
impacted the sandy infralittoral bottoms
which are a habitat for commercial species
such as the venerid Chamelea gallina.
Farmed species include Crassostrea gigas
and Ruditapes philippinarum, and the activity related to farming may be responsible
for an important part of the introductions
to and from the Adriatic; the latter species
is invasive and likely to displace or to drive
locally extinct the native clam Ruditapes
decussatus. The Sidney rock oyster Saccostrea commercialis was introduced to the
Venice Lagoon in the 1980s but seems tohave dwindled and has not been reported
recently (MIZZAN, 1999).
Twenty-one of the alien species known
from the Adriatic are recorded in Italy,
and only the few tropical species Brachidontes pharaonis, Cellana rota, Melibe
viridis and Halgerda wileyi have records
exclusively along the eastern coast.
430

3.6. Miscellaneous invertebrates in the
Adriatic Sea
Three alien species of Bryozoa (out of
23 known for the whole Mediterranean)
have been recorded in the Adriatic. The
Indo-Pacific species Tricellaria inopinata
was originally described on material coming from the Lagoon of Venice by
D'HONDT & OCCHIPINTI-AMBROGI(1985). It has been subsequently found inmany European seas (DYRYNDA et al.,2000). The alien status of Bugula fulva is
supposed based on its patchy occurrence
nearlyrestrictedtoharbours(HAYWARD & MCKINNEY, 2002).
The only alien species of Ascidiacea isBotrylloides violaceus, introduced in the
Lagoon of Venice by the north-western
Pacific by shellfish culture (ZANIOLO etal., 1998).
Alien Anthozoa are represented onlyby the north-eastern Atlantic species Diadumene cincta (BIRKEMEYER, 1996),
which in a sense reinforces the NorthernAdriatic ‘sub-Atlanticism’.
Porifera are represented by Paraleucilla magna only (LONGO et al., 2007).
Fourteen alien species of Cnidaria(out of 46 for the whole Mediterranean)
are known for the Adriatic. Alien Hydrozoa are comprised of 13 species (out of
37). Most of them have circum(sub)tropical origin, but two estuarine species(Cordylophora caspia and Gonionemus
vertens) prefer temperate waters. The first
Mediterranean record of Garveia franciscana comes from the Venice Lagoon(MORRI, 1982). It is debatable whether
Coryne eximia can be considered as alien
species since the Mediterranean occurrence of this species seems to need re-confirmation (BOUILLON et al., 2004).
Two of the three alien species of
Ctenophora known for the Mediterranean
Medit. Mar. Sci., 11/2, 2010, 381-493

are established in the Adriatic: Beroe ovata
and Mnemiopsis leidyi (SHIGANOVA &MALEJ, 2009). These authors refer earlier citations of Beroe ovata in the Mediterranean to the native species B. cucumis,
and report the real B. ovata as a Western
Atlantic species first introduced to theBlack Sea and possibly from there to the
Adriatic.
Sipuncula number two alien species:
both are circum(sub)tropical (PANCUCCIPAPADOPOULOU et al., 1999). The
only alien pycnogonid is Ammothea hilgendorfi, first reported for the Mediterranean
from the Lagoon of Venice (KRAPP &SCONFIETTI, 1983).
3.7. Alien fish in the Adriatic Sea
AddingFistularia
commersonii(DUL IC’ et al., 2008), 11 Lessepsian fish
species had reached the Adriatic Sea by
2009 (DRAGI EVIC’ & DUL IC’,2010). The occurrence of Terapon theraps
in 2007 (LIPEJ et al., 2008) is of particularinterest since its record was the first for theMediterranean; furthermore, it was foundat great distance from its usual distribution
area. Similar are the cases of Pampus
argenteus and Epinephelus coioides.
The catch of Elates ransonnetii in
March 2010 in the eastern Adriatic(DUL IC’ et al., 2010) brought the number of Lessepsian fish immigrants thatwere recorded in the Adriatic Sea to 12.
Sphoeroides pachygaster, of tropicalAtlantic origin, has undoubtedly formed awell established population in the Adriatic (Albania, Slovenia, Montenegro, Croatia, Italy).
4. EASTERN MEDITERRANEAN
The EMED includes two major bodies
of water: the Levant Sea and the Aegean
Medit. Mar. Sci., 11/2, 2010, 381-493

Sea, together with the smaller Sea of Marmara, which connects it to the Black Sea.
The Levant Sea is warmer than the rest of
the Mediterranean and harbours a significant number of circumtropical species.
Atlantic-Mediterranean elements andMediterranean endemics are comparatively scarce (MORRI et al., 2009).
Since the construction of the Suez
Canal, the Levant Sea is experiencing animportant influx of Red Sea species. POR(1990) defined the geographical limits tothe expansion of Red Sea immigrants in
the Mediterranean as the ‘Anti-Psara line’to the north (Anti-Psara being an island in
the Aegean) and the Strait of Sicily to thewest: these boundaries match the 15 ÆC
surface isotherm for February (BIANCHI,
2007). The Aegean Sea has local
endemics, but the typical Mediterranean
biota is impoverished with respect to the
WMED (BIANCHI & MORRI, 1983);
this, however, may be partly due to insufficient inventory effort (MORRI et al., 1999;
KOUKOURAS et al., 2001). The Sea ofMarmara exhibits peculiar hydrologicalconditions, with low salinity waters comingfrom the Black Sea stratifying over saline
waters of Mediterranean origin on the bottom (ÜNLÜATA et al., 1990). This hydrological regime should facilitate the diffusion of Black Sea species into the Northern
Aegean rather than vice-versa, but ourknowledge on the exchanges between the
two areas is limited and their biotic affinity is low (KOUKOURAS et al., 2001). In
recent times, climatic change favoured anincrease of biotic penetration from the Seaof Marmara into the Black Sea, which
therefore has been undergoing a processof ‘mediterranization’ (TOKAREV &SHULMAN, 2007).
Some scientists (e.g., TORTONESE,
1973; OLIVERIO & TAVIANI, 2003;
431

POR, 2009) have argued that this basin,
particularly in its eastern stretches, suffered from an ‘ecological vacuum’ that is
now being filled by Red Sea immigrants.
This idea has been criticized by GALIL(2007).
4.1. Alien protozoans in the Eastern
Mediterranean
Fish pasteurellosis caused by Photobacterium damselae was reported from cultured seabass (Dicentrarchus labrax) in
Turkey (CANDAN et al., 1996).
Among the total of 50 alien
foraminiferal species presently known for
the Mediterranean, all except one arepresent in the EMED (LANGER &HOTTINGER, 2000). All of them havetropical affinity. Despite a significant number of species with circumtropical distribution (KOUKOUSIOURA et al., 2010b),most have Indo-Pacific origin and theirpenetration in the EMED is likely to havebeen favoured by the opening of the Suez
Canal (MERI et al., 2007a; LANGER,
2008). Off the west Turkish coast, alien
foraminifers showed high abundance at
hot submarine springs that allowed higher
bottom water temperatures in winter(MERI et al., 2010). Similar observations on alien macrophytes in the Aegean(DE BIASI & ALIANI, 2003) and in the
Tyrrhenian Sea (GAMBI et al., 2008) suggest that shallow water hydrothermal ventsin the Mediterranean Sea may act as stepping stones for the penetration of tropicalaliens (BIANCHI et al., in press).
Among the nine new taxa added tothose recorded by ZENETOS et al. (2008),
eight occur in the EMED (LANGER,
2008) and 4 of them have been reported
only from single or few localities: Brizalina
simpsoni and Euthymonacha polita fromthe coasts of Turkey (MERI et al., 2010),
432

Pegidia lacunata and Pseudolachlanella
slitella (as Quinqueloculina eburnea) from
north-eastern Africa (BLANC-VERNET
et al., 1979) and Cyprus (ABU TAIR &LANGER, 2010).
The new recovery of alien foraminifers
mainly comes from recent researches carried out on living coastal assemblages of
the EMED, which testify to the increasing
scientific interest in alien and Lessepsianimmigrants. These studies confirm thewide distribution of many large endosymbiont-bearing taxa, such as Amphisteginids, along the coasts of the Levantine
basin (LANGER, 2008), Greece(TRIANTAPHYLLOU et al., 2009;
KOUKOUSIOURA et al., 2010a), Turkey(MERI et al., 2010) and Cyprus (ABU
TAIR & LANGER, 2010). Amphistegina
lobifera appears to have established successful populations in Greek coastal
ecosystems (TRIANTAPHYLLOU et al.,2009; KOUKOUSIOURA et al., 2010a,b).
Taxa of Indo-Pacific origin, probably true
Lessepsian forms, are represented byBorelis sp., Coscinospira hemprichii, Cycloforina sp., Heterostegina depressa, Pegidia
lacunata, Pseudolachlanella slitella andSorites orbiculus.
Taxa undetermined at the specificlevel (Borelis sp., Cycloforina sp.) havebeen included as alien forms only whendescribed and figured in previous work on
Red Sea or Indo-Pacific areas(HOTTINGER et al., 1993; LANGER &HOTTINGER, 2000; LANGER, 2008
with references) or (Haddonia sp.) if pertain to endemic Indo-Pacific genera(LOEBLICH & TAPPAN, 1988).
At the present state of studies, it seems
prudent to exclude some very seldom
recorded species cited by ZENETOS et al.
(2008), yet very rare or poorly preserved,
but also known at various latitudes or as
Medit. Mar. Sci., 11/2, 2010, 381-493

fossils in the Mediterranean area. These
species are here represented by Acervulina
inhaerens, Iridia diaphana, Cymbaloporettasquammosa, Pyramidulina perversa andTriloculina affinis.
Schackoinella imperatoria, a small
species originally described from fossilmaterial of the Tertiary Basin of Vienna,
pertains to a genus now known only in the
Indo-Pacific Ocean (LOEBLICH &TAPPAN, 1988). Rare specimens havebeen recorded along the eastern coasts ofNorth Africa (BLANC-VERNET, 1969)
as Glabratella imperatoria. More data areneeded to verify its introduction by shipping through the Suez Canal. Its rareoccurrence suggests a cryptogenic/casualacclimatization status.
In addition to all these benthic species,
planktonic foraminifers are represented
only by Pulleniatina obliquiloculata, a thermocline dwelling species characteristic ofequatorial Atlantic and Indo-Pacific
Oceans (BE & TOLDERLUND, 1971;
KENNETT & SRINIVASAN, 1983). Its
occurrence has been reported only by
MERI et al. (2004) along the Turkish
coasts and its introduction was inferredfrom the Atlantic Ocean via Gibraltar(ZENETOS et al., 2008). Recently, P.
obliquiloculata has been reported from
Canary Islands, Eastern Atlantic (WILKE
et al., 2009). It is absent in the Red Sea, but
it has been frequently reported from the
Arabian Sea and Bay of Bengal (BE &TOLDERLUND, 1971; CHEN &FARRELL, 1991, inter alii). More data are
required to interpret its occurrence and to
exclude its transport within ballast waters.
4.2. Alien macrophytes in the Eastern
Mediterranean
A total of 63 alien macrophytes havebeen hitherto reported in the EMED
Medit. Mar. Sci., 11/2, 2010, 381-493

which represent 50% of the alien macrophytes reported from the Mediterranean
Sea. This high percentage is mainly due tothe proximity to the Suez Canal and denseinternational ship traffic in the area.
Among them, 35 alien macrophytes arewell established, nine are casual, 15 are
cryptogenic/questionable, and four requireconfirmation.
Casualtaxa(i.e.
Acrochaetium spp., Derbesia boergesenii,
Padina antillarum, Rhodophysema georgii,
Rhodymenia erythraea, Solieria dura andSpatoglossum variabile) might turn into theestablished status in the future. The cryptogenic/questionable category includes
both temperate taxa mainly registered in
the north of the basin, i.e. the Sea of Marmara and the Northern Aegean Sea(Anotrichiumokamurae,
Chondria
coerulescens, Cladosiphon zosterae, Ectocarpus siliculosus var. hiemalis, Halothrix
lumbricalis, Polysiphonia atlantica, P.
fucoides and Pylaiella littoralis), and tropical taxa distributed in the Aegean Sea andthe Levantine Basin (Acanthophora nayadiformis, Antithamnionella elegans, theAtlantic strain of Asparagopsis taxiformis,
Caulerpa racemosa var. turbinata, Ceramium bisporum, Ganonema farinosum andUlva fasciata). The presence of Antithamnionella spirographidis, Asparagopsis armata, Gracilaria arcuata and Hypnea flagelliformis in the EMED requires confirmation. The latest record is that of the seaweed Codium parvulum, recent blooms ofwhich were observed on the northernshores of Israel (ISRAEL et al., 2010).
On the 21 invasive or potentially invasive macrophytes introduced into theMediterranean (Table 3), 11 are reported
in marine benthic habitats of the EMED.
No data are available for coastal lagoons.
Cladophora cf. patentiramea and Codium
parvulum are hitherto known only for the
433

EMED, for Cyprus and Israel, respectively.
The majority of alien macrophytes of
the EMED (63%) come from tropicalregions, mainly the Red Sea and the IndoPacific Ocean. The major vector of introduction is the Suez Canal. Hitherto 35
putative Lessepsian immigrants (56% of
the total) have been registered, of which
Cladophora cf. patentiramea, Codium
parvulum, Halophila stipulacea, Lophocladia lallemandii and Stypopodium schimperi
are considered invasive. Acanthophora
nayadiformis, Ganonema farinosum andUlva fasciata were found in the Mediterranean before the opening of the Suez
Canal in 1869, but the occurrence, at leastin the EMED, of exotic strains comingfrom the Red Sea and the Indian Ocean
via the Canal is highly probable today. The
other alien macrophytes reported from the
EMED come from cold temperate regions(NE Atlantic and NW Pacific) and wererecorded along the north coasts of Greece
and Turkey (Sea of Marmara and Northern Aegean Sea). Some of these lastmacrophytes, i.e. Ectocarpus siliculosus
var. hiemalis, Polysiphonia fucoides andPylaiella littoralis, also occur in the BlackSea (TA KIN et al., 2008): so, a native status cannot be excluded in the northernpart of the EMED.
4.3. Alien polychaetes in the Eastern
Mediterranean
A total of 98 alien polychaete species
have been reported to date from the
EMED, almost 75% of the alien polychaete species reported from the Mediterranean Sea, mainly due to the proximity to
the Suez Canal and dense international
ship traffics in the area. Among them, 52
species have been well established in habitats of the EMED, 16 species are casual,
two species (Chaetozone corona and Para434

prionospio coora) are cryptogenic and 28
species are questionable. 22 species seemto have invasive character (see Table 3),
forming dense populations in semi- or
heavily polluted areas (including harbours) or shallow water benthic habitats.
There is a relatively high number of questionable species in the area. They include
the species only listed in ecological papers(i.e. Cossura coasta, Loimia medusa), those
identified as ‘cf.’ in the area (i.e. Oenone cf.
fulgida, Syllis cf. mayeri) or whose descriptions based on the Mediterranean specimens were insufficient (i.e. Lepidonotus
carinulatus), hindering us in deciding theirreal taxonomic positions. Future studies tobe performed in the region would enable
us to understand if these species reallyexist in the Mediterranean. A total of ten
casual species (i.e. Laonome elegans,
Nereis gilchristi) might turn into the established status if data regarding them areaccumulated. The native species, Chaetozone setosa (in part) and Paraprionospio
pinnata, previously widely reported from
the Mediterranean Sea in polluted anddeep waters, have been recently re-identified as C. corona and P. coora, respectively( INAR & ERGEN, 2007; YOKOYAMA et al., 2010). These species wereregarded as cryptogenic because the dataon their distributions are very limited; they
are relatively newly described species andcould have been in existence in the area formany years.
The Levantine Sea is densely colonized by alien polychaete species. In theregion, a total of 77 species (79% of thealien polychaetes from the EMED) havebeen hitherto reported. A total of 11
species are categorized as questionable.
The highest number of alien species (58species) are known from the Levantine
coast of Turkey, followed by the Israeli (29Medit. Mar. Sci., 11/2, 2010, 381-493

species) and Egyptian (28 species) coasts.
On the Syrian coast, only two alien species(Spirobranchus tetraceros and Spirorbis
marioni) were encountered. The majorityof species are Lessepsian immigrants (47species, 61% of total species), the others
were introduced to the area via ships,
except for Chaetozone corona which is a
cryptogenic species. A total of 66 species(86%) could have been originated in the
Red Sea and Indo-Pacific areas. The others (11 species) were introduced to thearea from the Atlantic Ocean (mainly
from the western part).
The Aegean Sea is on the route of theLessepsian species that have become wellacclimatized to the Mediterranean environment and have a large dispersal capacity. Up to date, 39 alien polychaetes werereported from the Aegean Sea, of which 16
species are Lessepsian immigrants. In thearea, 20 species have been well established, one species, Prionospio (Minuspio)
pulchra, is casual, 16 species are questionable and two species, Paraprionospio coora
and Chaetozone corona, are cryptogenic.
Ship-transported species dominate semior highly polluted areas, especially near
the large international harbours. The
opportunistic species Malacoceros fuliginosus and Capitella spp. seem to have been
replaced by the North Atlantic species
Polydora cornuta and Streblospio gynobranchiata and the Pacific species Pseudopolydora paucibranchiata ( INAR etal., 2005, 2006a). In the eastern part of the
Aegean Sea, 19 alien polychaete species
were found, of which two species are questionable. In the western part, 31 species
were reported, of which 14 species arequestionable.
Polychaetes from the Sea of Marmara
have been recently reviewed by INAR
(2010). He postulated that a total of 17
Medit. Mar. Sci., 11/2, 2010, 381-493

species of alien status were reported fromthe region. However, three species were
excluded from the list, eight species arequestionable and six species (Nereis persica, Paraprionospio coora, Polydora cornuta,
Streblospio gynobranchiata, Desdemonaornata and Ficopomatus enigmatus) havebeen established. The report of theLessepsian species N. persica by
RULLIER (1963) needs to be confirmed.
The other species could have been introduced to the area by ships.
4.4. Alien crustaceans in the Eastern
Mediterranean
A total of 119 alien crustaceans havebeen reported in the EMED among which
six are cryptogenic species (Cymadusa
filosa, Gammaropsis togoensis, Monocorophium sextonae, Pseudocalanus elongatus, Synalpheus tumidomanus africanus andThalamita poissonii). Introduced crustaceans belong to Amphipoda (10), Cirripedia (5), Copepoda (33), Cumacea (3),Decapoda (58), Isopoda (7), Tanaidacea(1) and Stomatopoda (2). These numbers
include both benthic and pelagic forms ofcopepods as well as parasites among cirripedes, copepods and isopods. The vast
majority of them occur in the Levantine
Sea (e.g. 68% of the benthic forms, allplanktonic taxa and all parasites). Twothirds of the species have colonized theLevantine and are spreading to the
Aegean, while almost one third is known
from single records. Very few species, suchas the decapods Calappa pelii, Metapenaeus affinis and Sirpus monodi, are limited to the Aegean Sea. Reasoning for the
relatively high number of questionablecopepod species (ca. 10%) is provided in
ZENETOS et al. (2005, 2008).
With the exception of a few species
originating in the Atlantic (Calappa pelii,435

Callinectes sapidus, Farfantepenaeus
aztecus, Necora puber, Percnon gibbesi, Processa macrodactyla, Sirpus monodi,
Synalpheus tumidomanus africanus andThalamita indistincta), the native range offour-fifths of the alien crustacean in the
Mediterranean is in the Indo-Pacific
Ocean, Indian Ocean and Red Sea. The
pathway of introduction of decapod
species is the Suez Canal, but penetration
has been either unintentional (Lessepsianmigration) or ship-mediated. Shipping
appears to be also responsible for theimmigration of species of Atlantic origin.
However, spreading of self maintaining
populations from the CMED appears tobe the mode of introduction of the west
Atlantic crab, Percnon gibbesi now present
in Greece, Turkey, Syria, Cyprus, Israel(KATSANEVAKIS et al., in press and references therein) and Egypt (AZZURRO
et al., in press). Thalamita poissonii could
be a Lessepsian species, but there arerecords of it before the Suez Canal opening (UDEKEM D’ACOZ, 1999), which
could support a pre-Messinian Mediterranean presence. If so, the alien status
should be downgraded to ‘questionable’.
Perhaps at this moment it is best to consider it as ‘crypogenic’.
The latest additions to the inventory of
the marine decapod species in the EMED
are the pilumnid crab Eurycarcinus integrifrons (ÖZCAN et al., 2010), the redswimming crab Gonioinfradens paucidentata (CORSINI-FOKA et al., 2010), bothof Indo-Pacific origin, and the western
Atlantic shrimp Farfantepenaeus aztecus(CENZIG et al., 2010).
The Indo-Pacific stomatopod Erugosquilla massavensis thrives along theLevantine coast and is expanding in the
Aegean (ÖZCAN et al., 2009).
Among the planktonic crustaceans,
436

information is available almost exclusively
for copepods, since it is the most studiedgroup (as in the entire Mediterranean
Sea). Most of the newcomers are Lessepsian immigrants, two originating in the
Atlantic (Calanopia biloba, Paracartia
grani), while the origin of Acartia tonsa,
Arietellus pavoninus, Centropages furcatus
and Parvocalanus crassirostris is questionable, since they are distributed in tropicalto subtropical waters both of the Atlantic
and Indian oceans and even of the Pacific.
The species Pontellina plumata, Corycaeus
speciosus and Subeucalanus crassus were
excluded from the list of alien species
given by ZENETOS et al. (2005) afterreconsideration of the available information on their distribution (RAZOULS etal., 2005-2010). The consideration of
Subeucalanus subcrassus as alien is questionable because it is present in most areasof the Mediterranean Sea as well as in thetemperate to tropical waters of theAtlantic and Pacific oceans. The presenceof Spinocalanus terranovae in the EMED
seems questionable, given its restricteddistribution in Antarctic and subantarctic
areas. Regarding the records of Triconia
rufa (originating from the Red Sea) in the
EMED (Lebanon) by MALT et al. (1989),
BÖTTGER-SCHNACK & SCHNACK(2009) assumed that the previous authors
may have confounded the specimens with
Triconia sp. 8. Finally, Pseudocalanus elongatus should be considered as cryptogenic
in the Mediterranean Sea: it has beeninterpreted as a boreal relict species(FURNESTIN, 1979) or an immigrant,
either from the Black Sea (SIOKOUFRANGOU, 1985) or the north-eastern
Atlantic (VIVES et al., 1981).
Two introduced cymothoid species of
Indo-Pacific origin, Anilocra pilchardi andCymothoa indica, typically parasitic ofMedit. Mar. Sci., 11/2, 2010, 381-493

teleost fish, have been reported from
Lebanon (BARICHE & TRILLES, 2006;
TRILLES & BARICHE, 2006) parasitizing mainly barracudas (Sphyraenidae).
Three parasitic copepods new to theMediterranean fauna were reported inwaters off the Egyptian coast near Alexandria by EL-RASHIDY & BOXSHALL(2009). Two of them, Mitrapus oblongus
and Clavellisa ilishae, are of Indo-Pacificorigin and are considered to have coinvaded the Mediterranean through theSuez Canal on Red Sea immigrant hosts.
The third parasite, Nothobomolochus
fradei, was previously known from the Gulf
of Guinea and the Arabian Gulf.
A rhizocephalan barnacle, Heterosaccus dollfusi, followed its portunid hostcrab, Charybdis longicollis, from the Red
Sea through the Suez Canal to theMediterranean Sea.
4.5. Alien molluscs in the Eastern Mediterranean
The EMED has the lion’s share ofalien molluscs, building up impressive
numbers and by far the highest proportion
worldwide. To date there are 190 alienmolluscs reported (105 established, 64
casual, 15 questionable and six cryptogenic), to be added to around one thousand native species. This represents morethan 90% of all alien mollusc species
reported in the Mediterranean as a whole,a percentage that remains remarkably stable if we consider established species only.
Six species are classified as cryptogenic(Alvania dorbignyi, Aplysia parvula, Atys
angustatus, Discodoris lilacina, Parviturbo
dibellai, Teredo navalis), as there are
doubts on whether they are native or introduced or perhaps represent very old introductions.
Predictably, the vast majority of these
Medit. Mar. Sci., 11/2, 2010, 381-493

species are of Indo-Pacific origin and theirpresence is related to the proximity of the
Red Sea and to the Suez Canal connection. Only six species (Siphonaria pectinata,
Mya arenaria, Cerithium litteratum,
Anadara transversa, Mytilopsis sallei, andPetricola pholadiformis) are Atlantic orAtlanto-Mediterranean in origin. Examples of Indo-Pacific species yet nonLessepsians, among the now well-established species, are the strombid Conomurex persicus, first recorded in Turkey
and only later arrived on the Levantine
coast, or the opisthobranch Melibe viridis,
first recorded in Greek waters. Some ofthese species, like Conomurex persicus, arenot even known from the Red Sea andtheir introduction must therefore have
involved maritime traffic at some stage.
POR (2009, 2010) expressed the viewthat the thermophilic species of IndoPacific origin should not be viewed asaliens but rather as a restoration of the
Tethyan faunal realm as it existed previously to the closure of the communication
with the Indian Ocean in the late Miocene.
Another quite distinct pool is made up
by species which originate from the temperate North Pacific, rather than from thetropical areas, and whose introduction islinked with transfers of living molluscs foraquaculture and/or maritime traffic.
The EMED has comparatively limited
aquaculture activity and this category
would hold, apart from the cultured
species proper Crassostrea gigas and Ruditapes philippinarum, very few species suchas Rapana venosa, Anadara kagoshimensis
or Musculista senhousia.
A particular aspect regarding molluscs
is the high incidence of unsupported
records, often based on single shells andcarried perpetuated in subsequent checklists. The rationale for their exclusion has
437been detailed in GOFAS & ZENETOS(2003). We have not considered in thisreport the records for 32 additional
species, including those found on an oil rig(MIENIS, 2004), among them oysters andother bivalves firmly attached to their substrate, considering that these have not yet
been found in the wild.
The Levantine Sea is the area whichholds the highest number of reports, with149 species (83 established, 45 casual, 5cryptogenic, 16 questionable); historicallythis is the area where pioneer reports weremade on alien species of Indo-Pacific origin (e.g. GRUVEL & MOAZZO, 1931;
HAAS, 1937).
There are 48 of the species recorded
from Israel, Lebanon and Syria, whichhave a record in the Suez Canal and may
be safely qualified as Lessepsian, against
only seven (Melibe viridis, Murchisonella
columna, Sticteulima cf. lentiginosa,
Smaragdia souverbiana, Conomurex persicus, Zafra savignyi, Zafra selasphora)
species of tropical Indo-Pacific origin
which are proven non-Lessepsians (whose
progression started elsewhere than on theLevantine coast and have no Suez Canal
record). The remainder cannot be positively assessed but the ratio of Lessepsianto non-Lessepsian among them should
probably remain high.
The coasts of southern Turkey and
Cyprus are the next largest centres wherealien species have been reported, but there
the exploration is much more recent andthe fauna had remained virtually unknown
until the 1980s. Here the totality of thespecies is of tropical Indo-Pacific origin(save the cryptogenic species) but theshare is more balanced between Lessepsian and non-Lessepsian. For the coast ofsouthern Turkey, we have 32 proven
Lessepsians previously recorded in the438

Suez Canal and along the Levantine coast,
all established except the casual Psammotreta praerupta versus 20 species of thetropical Indo-Pacific which are definitely
not Lessepsian and started their Mediterranean settlement in the area.
The Aegean Sea is the natural route ofprogression for the most successful IndoPacific invaders, both Lessepsian andthose introduced first in the southernTurkey/Cyprus area. Nevertheless it has
also its own pools of species introduced
through other pathways. Contrary to thesouth coast of Turkey, there are several
species that originate from the Atlantic(Anadara transversa, Crepidula fornicata,
Cerithium litteratum, Petricola pholadiformis, Mya arenaria) or the temperateNorth Pacific (Ruditapes philippinarum,
Rapana venosa) and are involved neither
in the Lessepsian migration nor in the SE
Turkey focus of introductions. Siphonaria
pectinata, introduced in the Gulf ofSaronikos, is a clear example of intraMediterranean introduction.
As may be derived from this, theimportance of Lessepsian species is moderate, and even the proven Lessepsianspecies found there may be secondary
introductions from populations settled
elsewhere in the EMED.
Among the 18 species which could
hold as Lessepsian because they wererecorded in the Suez Canal, species like
Cellana rota, Fulvia fragilis, Murex
forskoehlii, first showed up in theSaronikos Gulf heavily impacted by maritime traffic: Cerithium scabridum
appeared in Greece (ZENETOS et al.,2009) much later than in Sicily, where it
was obviously a secondary introduction.
Only a few Lessepsian species such as
Pinctada radiata or Trochus erithreus may
have arrived in the Aegean as a result of
Medit. Mar. Sci., 11/2, 2010, 381-493

regular progression inside the Mediterranean.
4.6. Miscellaneous invertebrates in the
Eastern Mediterranean
Fourteen alien species of Bryozoa, outof the 23 recorded in the whole Mediterranean, are known for the EMED. Most ofthem are of Indo-Pacific origin, and mostare established. Three recently described
species from Lebanon have presumably
penetrated from the Red Sea: this seems
sufficiently documented for Parasmittina
serruloides
andP.
spondylicola(HARMELIN et al., 2009) but the status
as a Levantine endemic cannot be excluded for Schizoretepora hassi (HARMELIN
et al., 2007).
Seven of the 16 alien species of Ascidiacea known for the Mediterranean are
established in the EMED. They are in
majority Indo-Pacific, or alternatively circumtropical.
Rhodosoma
turcicum,Ecteinascidia thurstoni, Phallusia nigra,
Ascidia cannellata, Symplegma brakenhielmi, Herdmania momus, and probably
Microcosmus exasperatus are regarded as
Red Sea immigrants ( INAR et al.,2006b; BITAR et al., 2007; IZQUIERDO~OZ et al., 2009; KATSANEVAKIS
MUN
et al., 2009; SHENKAR & LOYA, 2009;
KONDILATOS et al., 2010).
Cnidaria are represented by 32 alien
species out of 46 known for the whole
Mediterranean. Anthozoans are represented by two species: the Indo-Pacificgorgonian Acabaria erythraea has beenfound in the harbour of Hadera, Israel(FINE et al., 2004), whereas the scleractinian coral Oculina patagonica, possibly originating from the SW Atlantic, has beenrecorded from many sites through thewhole basin (BITAR & ZIBROWIUS,
1997; FINE et al., 2001; INAR et al.,Medit. Mar. Sci., 11/2, 2010, 381-493

2006b; SALOMIDI et al., 2006). Paradoxically no Red Sea coral species has yet penetrated in the EMED (BIANCHI et al.,2010). In the case of the Hydrozoa, the
EMED harbours two thirds of the alien
species hitherto known for the Mediterranean (26 out of 37) in accordance withwhat is known for virtually all the othergroups of organisms, which typically showa preponderance of aliens in that basin.
Among the 26 alien planktonic hydrozoa,
17 were recorded in the EMED. Clytia linearis and Eirene viridula are the only established alien species in the entire Mediterranean Sea. Most of the alien Hydrozoa in
the EMED have a circumtropical distribution, the remainders have Indo-Pacific origin. Macrorhynchia philippina is the mostwidespread and abundant (MORRI,
2008). Four Scyphozoa aliens known for
the Mediterranean supposedly came inthrough the Suez Canal and are now established along the coast of Israel (GALIL etal., 1990, 2010). Cassiopea andromeda has
extended its penetration range to Turkey(ÖZGÜR & ÖZTÜRK, 2008) andGreece (ZENETOS et al., in press). Abyla
trigona is the single alien siphonophore in
the Mediterranean Sea, found in Egypt(ZAKARIA, 2006).
After having successfully invaded theBlack Sea, with severe impact on both thenative ecosystem-functioning and the fishery yield, the Western Atlantic combjelly
Mnemiopsis leidyi started to spread in theMarmara and North Aegean Sea andreached Israel in 2009 (GALIL et al.,2009). Its predator Beroe ovata followed
the same path and arrived in the Aegean
Sea (SHIGANOVA et al., 2007). Amongthe three alien ctenophore species in theentire Mediterranean Sea, Sulculeolaria
angusta was found only in the Egyptian
coasts (ZAKARIA, 2006).
439

Seven species of alien Porifera with
Indo-Pacific origin have been traditionallyconsidered as Red Sea immigrants(TSURNAMAL, 1969). More recently,
this has been questioned by VACELET etal. (2007), who evidenced taxonomic problems with all of them except perhaps Geodia micropunctata. According to
VACELET et al. (2007), sponges withtropical affinities hitherto known only for
the EMED are more likely to be remnantsof the thermophilic biota that thrived in
the Mediterranean during warmer periodsof the late Pliocene or Pleistocene and thatdisappeared from the rest of the basin during the Würmian cooling (BIANCHI et al.,2010).
All six alien species of Sipuncula
occurring in the Mediterranean have beenrecorded for the EMED. Phascolion(Isomya) convestitum and Phascolosoma
scolop are of Indo-Pacific origin, theremaining four are circumtropical(PANCUCCI-PAPADOPOULOU et al.,1999). All are known from a single or few
localities, where they can however beabundant (A IK, 2007; 2008).
KRAPP et al. (2008) reviewed the Pycnogonida of the EMED: three species arealien, although the status of Pigrogromitus
timsanus is questionable. Anoplodactylus
californicus and A. digitatus are established.
The echinoderm fauna of the EMED
includes eight alien species (out of twelve
for the whole Mediterranean). Most ofthem are likely to have penetrated throughthe Suez Canal, but Asterias rubens originated from the North Atlantic and was
first introduced in the Black Sea(KARHAN et al., 2008). Two conspicuous
Indo-Pacific species, Diadema setosum(YOKES & GALIL, 2006) and Synaptula
reciprocans(ANTONIADOU&
440

VAFIDIS, 2009) might exert a considerable impact on Mediterranean ecosystems.
The EMED hosts the vast majority of
the alien platyhelminthes, all reported as
fish parasites, all with an Indo-Pacificand/or Red Sea origin. Of the nine monogeneids introduced into this basin, five areconsidered established. Both host and parasite are Lessepsian immigrants that haveco-invaded the Mediterranean Sea via theSuez Canal. The first documented case ofa monogeneid invading a new biogeographical region by ‘natural’ extension of
its host range is that of the gill ectoparasite
Polylabris cf. mamaevi infecting the rabbitfish Siganus rivulatus. The greater abundance of P. cf. mamaevi in the invading(Mediterranean) populations is probably
due to the changed, new environment,
possibly impacting on host resistance tothe parasite and encouraging heavier
infections (PASTERNAK et al., 2007). A
possible explanation for the unexpectedcolonization success of immigrant fish parasite species belonging to the class Myxosporea, despite their putative complex
life cycles, is discussed in DIAMANT(2010).
4.7. Alien fish in the Eastern Mediterranean
A total of 106 alien fish (nearly 70% of
the alien fish known in the Mediterranean)
has been reported from the EMED, themajority of which originate from the IndoPacific and Red Sea regions. Details on
their distribution per country, origin andmode of introduction are provided in
GOLANI & APPELBAUM-GOLANI(2010). New data extending the distribution of already reported alien species inthe Levantine basin include: Pisodonophis
semicinctus and Pomadasys stridens(BILECENOGLU et al., 2009); Apogon
Medit. Mar. Sci., 11/2, 2010, 381-493

smithi and A. queketti (GOREN et al.,2008); Enchelycore anatina, Lagocephalus
spadiceus and Lagocephalus suezensis(KATSANEVAKIS et al., 2009).
The complicated flood of confamilial
species is prominent and deserves special
interest, as it is unlikely to be coincidental.
Alien cardinal fishes represent a good
example of this, since only a single species
was known for over 60 years and four
species have been added to the fauna within the last five years (GOREN et al., 2009a;
GOREN et al., 2010; TURAN et al., 2010).
Two species of the family Champsodontidae were observed from Turkey (Champsodon nudivittis) and Lebanon (Champsodon vorax), interestingly both without
records from the Red Sea ( I EK &BILECENOGLU, 2009; BARICHE,
2010a). Among recent alien gobies, Vanderhorstia mertensi was found off southern
Turkish coasts and became quite common
shortly after its first observation(BILECENOGLU et al., 2008; YOKES etal., 2009), while the far-east chameleon
goby Tridentiger trigonocephalus is reported from Israel (GOREN et al., 2009b).
Sudden occurrence of two alien angelfishes, Pomacanthus imperator and Pomacanthus maculosus, from the Israeli and
Lebanese coasts, respectively, is also noteworthy (BARICHE, 2010b; GOLANI etal., 2010). Among the latest records is thatof the Indo-Pacific goby Trypauchen vagina
in Israel (SALAMEH et al., 2010).
The appearance of Mycteroperca fusca,a large Atlantic fish, in Israel in 2010, andwithout having been spotted on the way,raises the question of the route of its
arrival. A reasonable possibility is that M.
fusca entered the Mediterranean throughthe Strait of Gibraltar, as many Atlantic
species do, then expanded its distribution
along the North African coast but was
Medit. Mar. Sci., 11/2, 2010, 381-493

overlooked or confused with Mycteroperca
rubra. Alternatively, it was introduced in
the ballast waters of a ship (HEEMSTRA
et al., 2010).
5. PHYTOPLANKTON
The introduction of marine microalgae in the Mediterranean Sea is hard to
document. Theoretically, these microscopic organisms can easily be transported bycurrents, but also by the feet of migratory
birds, or introduced through human commerce of marine animals, ship’s ballast
waters or fouling organisms. The consequences of a microalgal invasion are only
evident if there are conspicuous changes inan area, e.g. water discoloration or
mucilage, or where the species is implied
in toxicity events or other harmful impacts(ZINGONE & WYATT, 2005), whileinvasions go totally unobserved in case ofrare species, which are a conspicuous partof the phytoplankton biodiversity in allseas. In addition, to prove that a species isan alien requires very sound backgroundknowledge of the species of a given area.
Unfortunately, the diversity of marine
microalgae is scarcely known in wide areasof the Mediterranean Sea, e.g. the southern shores, where only a few sites havebeen investigated, or the offshore waters,
where studies are limited to occasional
sampling during cruises. Even in the northern Mediterranean waters the knowledgeof the distribution of these unicellularorganisms in a given area is far from being
exhaustive, for several reasons. First, phytoplankton is difficult to sample properly,
as most species have an ephemeral occurrence and a discontinuous spatial distribution. The resolution of sampling programs
rarely matches the space and time scale ofoccurrence of microalgae, and new species
441

show up when more detailed or integrative
sampling approaches are applied. For
example, many cysts of the toxic dinoflagellate Alexandrium andersonii were foundin sediment traps deployed in the Gulf of
Naples (MONTRESOR et al., 1998). The
species was not known from the Mediterranean Sea before, and has never been
recovered in any other plankton study inthe area since. Probably Alexandrium
andersonii appears in the plankton for very
short periods, or it is restricted to specificdepths of the water column that are notcovered during investigations. Anothermajor problem concerns all organisms thatare difficult to preserve and either breakor loose their peculiar characteristics with
common fixatives such as formalin andlugol. Micromonas pusilla, a 2-3 Ìm naked
prasinophyte, is easily identifiable when it
grows in cultures due to a very typicalswimming pattern. It can reach concentrations of up to 106 cells l-3 in coastal
Mediterraneanwatersinwinter(THRONDSEN & ZINGONE 1994;
ZINGONE et al., 1999). However, its cells
generally burst upon preservation. This isalso the case for much larger flagellates,e.g. the raphidophyte Chattonella subsalsa(30-40 Ìm) or naked dinoflagellates, which
explode or lose their shape when fixed.
Finally, many species require particularskills or methods to be identified. This is
the case for cryptomonads, generally very
abundant in inshore waters, which requirecultivation, electron microscopy and attimes molecular techniques to be identified at the species level. While their abundance as a group can be traced with pigment analyses (HPLC) that allow alloxanthin to be detected, or by flow-citometry,individual species are only identified by
combining a number of different techniques (CERINO & ZINGONE, 2006).
442

For the reasons stated above, it canrarely be excluded that a suspected
microalgal invader was already present as
part of the rare, hidden and unsampled
phytoplankton. These factors are probablyat the origin of the extremely low numberof proven invaders among phytoplankton
species (WYATT & CARLTON, 2002).
Nonetheless, a number of cases of possibly introduced microalgal species in the
Mediterranean Sea have been brought to
the general attention over the last few
years. One of the first cases was provided
by the toxic planktonic dinoflagellate
Alexandrium catenella, appearing in highconcentrations in a French coastal lagoon,
the Thau Lagoon, in 1998 (LILLY et al.,2002). Based on the molecular similarity ofthe species with strains from Japan, it was
proposed that the species had been transported recently to the Mediterranean Sea.
The species had been seen before in the
Balearic Sea in 1983 (MARGALEF &ESTRADA, 1987), along the Spanish coast
since 1996 (VILA et al., 2001) and along
the Sardinian coast in 1999 (LUGLI etal., 2003). Apparently its range is expanding across the Mediterranean Sea, as it has
been found recently on the Tunisian coast(TURKI & BALTI, 2007). On the other
hand, more recent studies using alternative
molecular tools have questioned the close
relationships between Japanese andMediterranean populations, and hence theidentification of Japan as the source area of
A. catenella (MASSERET et al., 2009).
While the knowledge of planktonic
species distribution poses many problems
in terms of sampling and detection, benthic and epiphytic microalgae should be easier to collect. Yet these organisms havebeen studied very rarely in the Mediterranean Sea until the end of the last century, when they became popular mainly due
Medit. Mar. Sci., 11/2, 2010, 381-493

to health and environmental problems
caused by a toxic species Ostreopsis cf.
ovata, which forms conspicuous blooms inmany coastal areas (e.g., TOTTI et al.,2010; MANGIALAJO et al., in press). Alsoin this case, the sudden emergence of aconspicuous problem was initially attributed to a new invasion, probably in relation
to climate change, as Ostreopsis species
were previously known mainly from tropical areas. However, an Ostreospis species
did form a conspicuous bloom in Villefranche Bay in 1972, where macroalgae
appeared to be covered by a brown mucous
layer (Taylor, pers. comm.), but it was justrecorded due to the curiosity of an expert
taxonomist swimming in the area. The
species was reported as O. siamensis, theonly species known at the time (TAYLOR,
1979), but it was likely to be the same as the
one reported in more recent years in thesame area, i.e. O. cf. ovata. From themolecular point of view, the Mediterranean populations of this species were
known to be similar only to Brazilian populations (PENNA et al., 2010), but comparable ribotypes have very recently beenfound also in Japanese waters (SATO et al.,2010). Molecular analyses on other markers are required to establish where, among
the three sites, the molecular diversity ofOstreopsis cf. ovata is the highest, which
would indicate the source site for these different populations around the world.
Species of the benthic dinoflagellate
genus Gambierdiscus, which produce toxins that are responsible for the neurological syndrome ciguatera, had never beenrecorded in the Mediterranean Sea until
2003, when specimens were found in
Greek waters off Crete (ALIGIZAKI &NIKOLAIDIS, 2008). No other record isconfirmed from anywhere else in the
Mediterranean Sea, despite the intensive
Medit. Mar. Sci., 11/2, 2010, 381-493

microphytobenthos sampling carried out
to study O. ovata at several sites over thelast decade. In this case, an introduction
from the Indian Ocean could be hypothesised, but the source population would be
missing for the time being, as the taxonomic identity of the species is still uncertain. On the other hand, data on benthic
microalgae are now more complete than
10 years ago, which would allow one to test
a predictable range expansion westward.
Indeed, marginal range expansion havebeen observed for other species, such as
Alexandrium catenella (LUGLI et al.,2003), G. catenatum (G MEZ, 2003), anda number of diatom species (G MEZ,
2008), which could be related to the climatic changes observed in the Mediterranean Sea or to other unexplained longterm fluctuations in species abundance.
A sounder background knowledge on
phytoplankton distribution is available forplaces where time series exist and accurate
identification is performed over a longterm period. At the LTER-MC station in
the Gulf of Naples, two species neverrecorded despite their relatively easy identification, have suddenly appeared inrecent years. These were Skeletonema tropicum (ZINGONE et al., 2003; KOOISTRA
et al., 2008), which is easily identifiable as
it is the only species in the genus having
many chloroplasts, and Pseudo-nitzschia
multistriata (ZINGONE et al., 2003),
which has a typical sigmoid shape. The two
species had never been recorded before
1995 and 2002, respectively, and haveshown increasing concentrations since
their first finding. In both cases, an introduction could be hypothesised but it cannot be excluded that these are other examples of marginal range expansion from thesouthern Mediterranean Sea shore northward, or that these species were extremely443

rare for many years, suddenly finding
appropriate environmental conditions toincrease their abundance. The latterhypothesis takes into consideration longterm fluctuations in species abundance
and is partially supported by the absenceof discoveries since 1992 of another
Pseudonitzschia species, P. subpacifica.
This species was relatively common in the
1984-1987 period in the Gulf of Naples,
then it became rarer and eventually disappeared. Although P. subpacifica could still
be in the Gulf of Naples, this case indicatesthat there is always the possibility of unexplained local extinctions, and more generally points to important changes in species
abundance over the long term.
Another relevant case is that of species
showing discontinuous geographicalranges. The dinoflagellate Alexandrium
balechii is only found in the Gulf of Mexico
and in Tyrrhenian waters, suggesting thatthe species originated in one of the twoplaces and was transported to the other,although there are no data to establish
where the species should be considered analien. Several other dinoflagellates are only
found in the Indo-Pacific area and in the
Mediterranean Sea so far (G MEZ,
2006), but the suspicion that they wereintroduced through the Suez Canal should
be substantiated when their absence corresponds to their actual absence in the
Atlantic Ocean and not to our limited
knowledge of protist biogeography. For
similar reasons, the high number of flagellate species that have been described in the
Mediterranean Sea and never recorded
elsewhere cannot be considered as proof of
their endemism in the Mediterranean Sea(THRONDSEN & ZINGONE, 1994).
In synthesis, in the Mediterranean Sea
a number of planktonic and benthic
microalgae has appeared over the last444

decades and continue to do so, forming
conspicuous blooms in places where they
were not recorded at all before. However,whether these species are really alien
should be tested effectively, considering atleast two alternative hypotheses: 1) thatthey belong to the rare component of theplankton and that they were missed in previous studies; 2) that their range expandedfrom unexplored southern or eastern
Mediterranean areas.
6. INVASIVE SPECIESThe ongoing incoming of immigrants
and their increasing success in the
Mediterranean basin imply a profoundand continuous alteration in the species
distribution patterns that seems to haveaccelerated in the last decade. Up torecent times the influence of these immigrants was geographically limited to theareas close to the source of introduction,
but today the idea of biogeographicalboundaries to Lessepsian invasions (POR,
1981; QUIGNARD & TOMASSINI,
2000) seems to be overcome.
The most invasive species per MSFD
are listed in Table 3; their share per taxonomic group is depicted in Figure 2.
A total of 134 species are classified as
invasive or potentially invasive in the
Mediterranean. The present list is based ondata on worst invasive marine species collated from various sources for the Mediterranean (STREFTARIS & ZENETOS,
2006; GALIL, 2007; STREFTARIS et al.,2008), European Seas (EEA, 2007b;
DAISIE: VIL A et al., 2008), and globally(Global Invasive Species Database), managed by the Invasive Specialist Species
Group (ISSG: www.issg.org). The list wasenriched with species such as the bigfin reefsquid Sepioteuthis lessoniana, an invader
Medit. Mar. Sci., 11/2, 2010, 381-493

Table 3
Distribution of invasive (++) or potentially invasive alien species (+) in the Mediterranean MSFD
areas. (-) Species absent from the area, *marketable species.
SPECIESWMED CMED ADRIA EMED

Foraminifera
Amphistegina lobifera

+

+

+

++

++
++
++
++
++
++
++
+
+
+
+
++
++
+
++
++

+
+
++
++
++
++
+
+
++
+
+
+
++

+
+
+
++
++
++
+
++
+
+
+
++
++
+

+
++
++
++
++
++
++
++
+
++
+

+
+
+
++
++
++
+

++
+
++
++
++
-

+
++
++
++
++
-

++
++
++
++
++
++
++
++
++
++
++
++
++
++

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

Polychaeta
Branchiomma bairdi
Branchiomma luctuosum
Ceratonereis mirabilis
Desdemona ornata
Dorvillea similis
Eunice antennata
Ficopomatus enigmaticus
Hydroides dianthus
Hydroides elegans
Hydroides operculatus
Leonnates indicus
Leonnates persicus
Laonome triangularis
Notomastus mossambicus
Medit. Mar. Sci., 11/2, 2010, 381-493

445

Table 3 (continued)
SPECIES
Polydora cornuta
Pomatoleios kraussii
Prionospio (Aquilaspio) krusadensis
Prionospio (Prionospio) saccifera
Pseudonereis anomala
Pseudopolydora paucibranchiata
Spirorbis (Dexiospira) marioni
Streblospio gynobranchiata

WMED CMED ADRIA EMED
+
++
++
++
++
+
++
+
++
++
++
Crustacea

Callinectes sapidus*
Charybdis helleri
Charybdis longicollis
Dyspanopeus sayi
Erugosquilla massavensis
Heterosaccus dollfusi
Marsupenaeus japonicus*
Melicertus hathor
Metapenaeus monoceros*
Metapenaeus stebbingi*
Penaeus semisulcatus*
Percnon gibbesi
Portunus segnis*
Rhithropanopeus harrisii

+
+
++
+
+

+
+
+
+
++
+
+

++
++
+
+
++

++
++
++
++
++
++
++
++
++
++
++
++
-

+
+
+
++
++
+
+
++

+
+
++
+
++
+
-

++
++
+
++
+
++
++

+
++
++
++
++
++
+
++
+
++
++
-

-

-

-

++

++
++

+
++

++
++

++

Mollusca/Bivalvia
Anadara kagoshimensis
Anadara transversa
Brachidontes pharaonis
Chama pacifica
Dendrostrea frons
Fulvia fragilis
Musculista senhousia
Pinctada radiata*
Ruditapes philippinarum*
Septifer forskali
Spondylus spinosus
Xenostrobus securis

Mollusca/Cephalopoda
Sepioeuthis lessoniana*

Mollusca/Gastropoda
Aplysia dactylomela
Bursatella leachii
446

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

Table 3 (continued)
SPECIES
Cellana rota
Cerithium scabridum
Conomurex persicus*
Crepidula fornicata
Ergalatax junionae
Erosaria turdus
Melibe viridis
Pseudominolia nedyma
Rapana venosa
Rhinoclavis kochi

WMED CMED ADRIA EMED
+
+
++
+
+
++
+
+
++
+
+
+
++
++
+
+
++
++
++
++
+?
+
+
++

Porifera
Paraleucilla magna

++

+

+

-

Cnidaria/Anthozoa
Oculina patagonica+++

Cnidaria/Hydrozoa
Clytia hummelincki
Clytia linearis
Garveia franciscana
Macrorhynchia philippina

++
++
+
-

++
++
-

++
++
++
-

++
++

-

+
+
+

-

+
+
++

+

+

+

++

+
-

+
+

+
++

+
-

-

-

-

++
+
++

+
++
+
+
+

+
+
+
+
+
-

+
-

+
++
+
-

Cnidaria/Scyphozoa
Cassiopea andromeda
Phyllorhiza punctata
Rhopilema nomadica

Ctenophora
Mnemiopsis leidyi

Bryozoa
Bugula fulva
Tricellaria inopinata

Echinodermata
Aquilonastra burtoni
Asterias rubens
Synaptula reciprocans

Ascidiacea
Botrylloides violaceus
Distaplia bermudensis
Microcosmus exasperatus
Microcosmus squamiger
Phallusia nigra
Polyandrocarpa zorritensis
Rhodosoma turcicum
Styela clava
Medit. Mar. Sci., 11/2, 2010, 381-493-

447

Table 3 (continued)
SPECIES
Symplegma brakenhielmi

WMED CMED ADRIA EMED+Fish
Apogon pharaonis
Atherinomorus forskalii*
Diplodus bellottii*
Etrumeus teres*Fistularia commersonii
Gymnammodytes semisquamatus
Lagocephalus sceleratus
Pagellus bellottii*
Parexocoetus mento
Pempheris vanicolensis
Plotosus lineatus
Pteragogus pelycus
Sargocentron rubrum*
Saurida undosquamis*
Scomberomorus commerson*
Seriola carpenteri*
Seriola fasciata
Siganus luridus*
Siganus rivulatus*
Sillago sihama*
Solea senegalensis*
Sphoeroides pachygaster
Sphyraena chrysotaenia*
Sphyraena flavicauda*
Stephanolepis diaspros
Upeneus moluccensis*
Upeneus pori*of the last decade that presented a population explosion within 2010 (ZENETOS etal., in press).
A total of 108 species are locally invasive or merely present in the EMED, 75 in
the CMED, 53 in the Adriatic and 64 in
the WMED (Fig. 2). The WMED hosts
most macrophytes, whereas the EMED
has the lion’s share in polychaetes, crustaceans, molluscs and fish.
448

++
+
++
++
+
+
++
++
-

++
++
++
+
+
+
+
+
++
++
+
++
++
+
++
++
+
++
+
+

+
+
+
+
+
+
+
-

++
++
++
++
++
+
++
++
++
++
++
++
++
+
++
++
++
++
++
++
++
++
++

The share of invasive species does notsupport POR’s (2009, 2010) statement thatspecies from the tropical Indo-Pacificshould behave more like one more elementof the native fauna and be less aggressive intheir onset than real aliens, e.g. those originating from the temperate Atlantic or
Pacific. Rather, it would conform to therule of thumb that one out of every ten
established alien species becomes invasive.
Medit. Mar. Sci., 11/2, 2010, 381-493

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

In addition to documented impacts on
biodiversity, some of the invasive species,
among decapods, molluscs and fish, haveturned to be beneficial to man and be commercially exploited in several countries(Table 3). Though no extinction of a native
species is known, sudden decline in abundance, and even local extirpations, concurrent with proliferation of aliens, has beenrecorded (GALIL, 2007).
Yet, there is no way to predict invasibility of ecosystems. However, community‘invasibility' appears to be the result of therelationship between native species richness and alien species ability to colonize
new habitats (BULLERI et al., 2008). This
concept implies that habitats with high levels of diversity are difficult to invade. In
contrast, species-poor communities, orstressed ecosystems, are arguably more
prone to invasion, primarily due to lack ofbioticresistance(OCCHIPINTIAMBROGI & SAVINI, 2003). At
Mediterranean basin scale, this appears to
be true (see chapter 7).
Medit. Mar. Sci., 11/2, 2010, 381-493

Little has been done to evaluate therisk posed by the introduction and spreadof invasive species, or to minimize the riskof introducing additional alien species.
Effective legislation on alien species exists
only in few countries and for selective vectors e.g. in EU countries for aquaculture
introductions; Egypt, France, Spain and
Syria for introductions in ballast waters(IMO/ BWC, Ballast Water Convention).
6.1. Macrophytes
Most spectacular has been the arrivalof several invasive exotic varieties of green
and red algae. These algae have flourishedat the expense of benthic Mediterranean
organisms. Their dense and creeping
meadows - which feature up to 27,000
erected blades per square meter in Caulerpa racemosa var. cylindracea - overwhelm
the benthic assemblages. Their success is
usually explained by a fast growth, permanent meadows and synthesis of secondary
metabolites active against natural enemies(herbivorous, epiphytes, pathogens, etc.).
449

Once they have spread, species like thisare impossible to eliminate.
Among the 125 alien macrophytes, 21
are invasive or potentially invasive (17% of
total taxa) (Table 3). The total number ofinvasive or potentially invasive alien
macrophytes decreases eastwards: WMED
(16 taxa), ADRIA (14 taxa), CMED (13taxa), and EMED (11 taxa). Only six invasive alien macrophytes have colonized thefour basins: the Indo-Pacific strain of
Asparagopsis taxiformis, Caulerpa racemosavar. cylindracea, Codium fragile, Halophila
stipulacea, Lophocladia lallemandii and
Womersleyella setacea.
6.2. Polychaetes
A total of 22 species (17% of the total
alien polychaetes) seem to have invasive
character, dominating shallow-water benthic habitats, estuarine areas, polluted
areas and harbour environments. All ofthem are invasive in the EMED, five arepresent in the CMED, six in the Adriaticand nine in the WMED.
Polluted soft bottoms especially near
harbours greatly facilitate the establishment of ship-mediated species such as
Polydora cornuta, Streblospio gynobranchiata and Pseudopolydora paucibranchiata( INAR et al., 2005, 2006a; DAGLI &INAR, 2008). Polluted environments,
where competition among species is low
and food is ample, provided these species
with an excellent opportunity to build up
dense populations. The maximum densityof P. cornuta was calculated as 3170 ind.m-2,
that of S. gynobranchiata as 60480 ind.m-2
and that of P. paucibranchiata as 6180
ind.m-2 in Izmir Bay ( INAR et al., 2005;DAGLI & INAR, 2008).
In a study by INAR (2006), alien
species comprised more than 95% of the
serpulids found on hard substrata (rocks,
450

molluscs, dock’s pilings, ropes and tires)
along the Levantine coast of Turkey.
Pomatoleios kraussii formed a densely populated belt in shallow-water areas in
Mersin Bay ( INAR, 2006). The invasive
nereid species Pseudonereis anomala is adominant component of the algal communities in the Levant Sea, attaining a population density of 2475 ind.m-2 on the alga
Jania rubens in Iskenderun Bay ( INAR
& ALTUN, 2007). BEN-ELIAHU (1989)
reported that a native nereidid species ofthe Mediterranean, Perinereis cultrifera(Grube), was excluded from the habitats of
the Levant Sea by P. anomala. This species
expanded its distributional range to Izmir
Bay, North Aegean Sea ( INAR &ERGEN, 2005) and the coasts of Greece(KAMBOUROGLOU & NICOLAIDOU, 2006). The eunicid worm Euniceantennata was found to be a common
species on the southern coast of Turkey(KURT SAHIN & INAR, 2009) and
was also previously reported from thecoasts of Italy (CANTONE, 1993).
6.3. Crustacea
The Indo-Pacific prawns, in particular
Marsupenaeus japonicus, Metapenaeus
monoceros, and Penaeus semisulcatus, arehighly prized and are considered a boon to
the Levantine fisheries (DURUER et al.,2008). Portunus segnis, formerly known as
P. pelagicus, is commercially exploited in
Egypt (ABDEL-RAZEK, 1987). It is alsocommercially important but caught insmall quantities in Turkey (DURUER etal., 2008). The latest record of P. segnis isfrom the northern Tyrrhenian Sea(CROCETTA, 2006).
The invasive crab Percnon gibbesi is
one of the most recent and successful
invaders in the Mediterranean Sea. Its distribution has been recently updated byMedit. Mar. Sci., 11/2, 2010, 381-493

KATSANEVAKIS et al. (in press) who
report that the relative abundance of thespecies in the Ka s -Kekova Marine Protected Area (SE Turkey) reached 112 individuals in 2010, from two individuals in
2006.
The blue crab Callinectes sapidus wastransported into the Mediterranean in ballast tanks from the north-east coast of theUSA. After the first Mediterranean record(Venice, Italy, 1949), this species has been
widely recorded in different Mediterranean regions, especially in the EMED
(Turkey, Greece, Syria, Lebanon, Israel
and Egypt) where it became invasive and iscommercially exploited (STREFTARIS &ZENETOS, 2006; DURUER et al., 2008).
A decade after its introduction,
Heterosaccus dollfusi, a parasite on Charybdis longicollis, became invasive in the south
Levantine coast. Despite the high prevalence of the parasite and its injurious
impact on the host reproduction, the invasive host-parasite pair reached a modus
vivendi with no noticeable reduction in thehost population (INNOCENTI & GALIL,
2007).
6.4. Mollusca
The gastropods Cerithium scabridum,
Rhinoclavis kochi, Conomurex persicus andBursatella leachi and the bivalves Pinctada
radiata, Brachidontes pharaonis, Fulvia
fragilis, Chama pacifica and Spondylus
spinosus have been reported as locallyinvasive in the EMED by ZENETOS et al.
(2005, 2008). Conomurex persicus has beenreported to achieve densities of tens ofspecimens per m2 at some sites(FISHELSON, 2000) on the Israeli coast.
To these may be added the limpet Cellanarota, reported as thriving to the point of
displacing the native limpet Patellacaerulea (MIENIS, 2002, 2003).
Medit. Mar. Sci., 11/2, 2010, 381-493

Rapana venosa, one of the most invasive species worldwide, has been rapidly
increasing in the Adriatic Sea, thus
enhancing the risk of invasion and further
spreading. Maximum estimated population densities of > 500 ind..km-2 werereached near the shores of Cesenatico(SAVINI et al., 2007).
The spread of the alien species acrossthe Mediterranean is not easy. Only six ofthe ca. 50 Lessepsian species have crossed
the Sicily Strait westwards and only two(Bursatella leachii and Fulvia fragilis) havereached the Spanish coast so far.
None of the other species of IndoPacific origin, so successful in the EMED,
have a real invasion success in the westernbasin. Some of the tropical Indo-Pacificspecies which started in the CMED
(Aplysia dactylomela, Haminoea cyanomarginata, Melibe viridis) are now spreading to
Malta (BORG et al., 2009; SCHEMBRI,
2009) or have even reached the western
Mediterranean. Several individuals of
Melibe viridis were recently observed in
Gulf Aranci, NE Sardinia (DONEDDU &TRAINITO, 2008).
The case history of the forerunners
indicates that the spread across theMediterranean involves a complex historyof natural spread with larvae and transport
via shipping from the original point of
introduction. In any case the distributionof these aliens in the western and central
parts of the Mediterranean is patchy anddoes not suggest a continuous progression.
The limiting factors may be on the one
hand, the natural species richness of thewestern basin; on the other, the severe difference in sea surface temperatures (POR,
2009, 2010) which may make it inhospitable to thermophilic species such as
Pinctada radiata.
The impact of invasive species is com451

parable within these two main poles ofalien molluscs in the Mediterranean, but
the pool of invasive species is not the same.
To the south, an aggressive onset is seenfor some tropical Indo-Pacific species suchas Conomurex persicus, Cellana rota or (atthe beginning of the invasion) Pinctada
radiata, which fail to become massively
established in other parts of the basin. Oneof the latest introductions is Sepioteuthis
lessoniana, a commercially important
squid species for inshore fisheries throughout its distributional range, which wasobserved in Iskenderun Bay (SALMAN,
2002) and has rapidly spread in the Levantand Aegean seas (KATSANEVAKIS etal., 2009; LEFKADITOU et al., 2009;ZENETOS et al., in press). In the northernparts (Northern Aegean, Adriatic, andFrench Mediterranean lagoons) invasive
species like Anadara kagoshimensis, Ruditapes philippinarum or Rapana venosa originate from temperate areas, and none ofthese are reported as invasive in the southern parts of the Mediterranean. Therefore
the climatic gradient in the Mediterranean
is likely to promote natural boundaries forthe alien species, which should reach a stable range at some time.
6.5. Miscellaneous invertebrates
The ascidian Microcosmus squamiger,
first recorded at 1963 in Tunisia, is widely
spread in the Western Mediterranean(France, Spain, Italy, North Africa: see
synthesis in TUR N et al., 2007), andlocally in the CMED (Malta:~
OZ et al., 2009).
IZQUIERDO-MUN
This species forms dense populations
reaching up to 2300 ind..m-2 and its presence has strong implications in the structure and functioning of the native communities (RIUS et al., 2009). Microcosmus squamiger is considered a global
452

marine invader with a high invasive
potential.
A recent immigrant, Styela clava,
among the worst invasives in Europe, hasbeen accidentally transported into theWestern Mediterranean by shellfish transfer (DAVIS & DAVIS, 2008). Styela clava,
a solitary, hermaphroditic ascidian, is considered to be an aggressive invader
throughout the world; this species is amajor pest to the mussel farming industryof Canada (ARSENAULT et al., 2009). Its
recent discovery in the Mediterranean Sea
caused a considerable alarm to mussel
farms.
Phallusia nigra is an old invader in the
Mediterranean Sea. The quite sudden
appearance of the species in the Aegean
Sea and the extended distribution that itdemonstrated within a short time along
the coasts of Rodos and Kriti(KONDILATOS et al., 2010; ZENETOS
et al., in press) support the invasive character of the species, as already pointed out by~
OZ et al. (2009).
IZQUIERDO-MUN
Other species such as Botrylloides violaceus, Symplegma brakenhemi and possibly Polyandrocarpa zorritensis and Distaplia
bermudensis are locally invasive in alteredhabitats (harbours, aquaculture rafts, Po
and Ebro deltas).
The scleractinian coral Oculina patagonica, a species of uncertain, but probably
Atlantic origin, has taken advantage of thewarming sea and has explosively expandedaround the southern Mediterranean during the last years, building coral pavements(see latest updates in SARTORETTO etal., 2008).
Of the 37 alien hydrozoan species
recorded for the Mediterranean, four canbe considered invasive. The occurrence ofClytia hummelincki in the Mediterranean
Sea was only discovered in 1996 in the
Medit. Mar. Sci., 11/2, 2010, 381-493

northern Ionian (BOERO et al., 1997),
and within the subsequent decade further
records came from the Southern Adriaticand the Tyrrhenian seas (BOERO et al.,2005; GRAVILI et al., 2008). At present,
C. hummelincki is extremely abundant in
the summer in shallow rocks and is rapidly expanding its range thanks to the dispersal capacity with currents of themedusa stage. Clytia linearis was recordedin the Suez Canal as early as in the 1930s(BILLARD, 1938) and rapidly spreadthrough the whole Mediterranean to get
its north-western reaches by the 1950s(PICARD, 1951): thus, it is probably oneof the first and most successful Lessepsianimmigrants. The medusae of the two
species of Clytia are relatively large andare produced in great number: their voracious predatory behaviour on coastal zooplankton may imply a significant impact
on native Mediterranean ecosystems bythese two invasive species (GRAVILI etal., 2008).
The first Mediterranean record ofGarveia franciscana came from the Lagoonof Venice in 1978, where it showed among
the most abundant hydroids (MORRI,
1981). In subsequent years it was also
found in other estuarine habitats of theNorth Adriatic and Catalonia, NE Spain(MORRI, 1982; GILI, 1986). Its spread inmany brackish environments world-wide
and its occurrence in ship-hull fouling(MORRI & BOERO, 1986), suggest highinvasive potential for this species: lack ofrecent studies on Mediterranean brackishwater hydroids, however, prevents any
solid evaluation of its present degree ofsuccess.
Macrorhynchia
philippina
has
occurred in the EMED since at least theearly 1990s, when has been found in polluted waters near Beirut (BITAR &Medit. Mar. Sci., 11/2, 2010, 381-493

BITAR-KOULI, 1995). It is now widespread and frequent in the whole Levant
Sea ( INAR et al., 2006b). Its diffusion
on coastal rocks from just below the surface to more than 40 m and the frequent
reproductive status (MORRI et al., 2009)
suggest a high invasive potential. Its
colonies, up to 30 cm tall and stinging to
the touch, are easily recognizable, so thatits further expansion in the Mediterranean should be easily tracked (MORRI,
2008).
One of the top 100 worldwide invasive
species, the comb jelly, Mnemiopsis leidyi,
was first seen in the Saronikos Gulf
(EMED) in 1990 (SHIGANOVA et al.,2001). Sporadic sightings ever since fromthe Turkish and Greek coasts raised noconcern about its notorious impacts on
the ecosystem and the fisheries, until 2009
when swarms if it spread across theMediterranean, from Israel to Spain(FUENTES et al., 2010). Each summer
since the mid-1980s swarms of the invading jellyfish, Rhopilema nomadica, haveappeared along the Levantine coast.
These swarms of voracious planktotrophs
can stretch up to 100 km long. As theydraw nearer to shore, they adversely affecttourism, fisheries, and coastal installations.
6.6. FishClassically, Lessepsian species were
expected to be limited to the EMED, due
to hydrological conditions, but the spreadto the western basin is now a reality. Sofar, only a few Lessepsian fish species havebeen observed in the Western Mediterranean: Abudefduf vaigiensis, Siganus
luridus, and Fistularia commersonii. Thislatter is probably the fastest-spreading
species in the Mediterranean: since thefirst record in Mediterranean ten years
453

ago, F. commersonii is now reported moreor less in the whole Mediterranean (Table2). The very recent records of F. commersonii and S. luridus in the north-western
Mediterranean (S NCHEZ-TOCINO etal., 2007; DANIEL et al., 2009) could thenbe the hint of future extensive spreadingof Lessepsian species. The opposite direction was followed by Atlantic invaders
such as Seriola fasciata (SONIN et al.,2009), Pagellus bellottii (SAAD &SBAIHI, 1995) and Sphoeroides pachygaster (GOLANI, 1996) that have reached
the Levantine basin.
Today, some of these species constitute an important part of local fisherycatches; 16 such species are listed (Table3) among the most invasive species in the
Mediterranean, Siganus luridus andSiganus rivulatus though marketable, they
are classified as venomous species in
FishBase. Lessepsian immigrants constituted 19% of the total abundance of fish
captured at the Lebanese rocky coast. In
decreasing order of importance, were
Atherinomorus lacunosus (11.5%), Siganus
luridus (2.3%), S. rivulatus (2.3%), Pempheris vanicolensis (1.6%), Sargocentron
rubrum (1.0%), Stephanolepis diaspros(0.2%), and Apogon nigripinnis (presently
Apogon pharaonis) (0.02%) (HARMELIN-VIVIEN et al., 2005). Siganus luridus
and S. rivulatus contributed 23% to thespecies composition captured in purse
seine operating during the daytime in
Abu-Qir and El-Mex Bays, Alexandria(Egypt) (AKEL, 2005).
Invasive species such as: Apogon
pharaonis, Fistularia commersonii, Gymnammodytes semisquamatus, Parexocoetus
mento, Pempheris vanicolensis, Pteragogus
pelycus, Sphoeroides pachygaster, Stephanolepis diaspros, are just a nuisance to fisheries. Others as the highly venomous454

striped catfish, Plotosus lineatus, and thepufferfish Lagocephalus sceleratus are athreat to human health.
7. SYNTHESIS: STATUS
GEOGRAPHICAL ORIGINAND

A total of 955 alien species occur inthe Mediterranean, the vast majority ofthem having being introduced in the
EMED (718), fewer in the WMED (328)and CMED (267), and least in the Adriatic (171). Of these, 535 species (56%) havebeen established in at least one area. We
calculated that the established species
accounted for 57.6% of the total numberof alien species in the WMED, 51.5%, in
the CMED, 52.6% in the Adriatic Sea and56.3% in the EMED. The casual records
constitute 27-29% of the alien species at
each basin. On the contrary, the number ofquestionable species in the EMED is threetimes as many as those of the WMED.
Macrophytes, polychaetes and crustaceans
have the highest number of questionablespecies. The questionable status for
macrophytes is generally due to the uncertainty regarding their origin. For polychaetes and crustaceans, the reason for thehigh percentage is the lack of precise taxonomic works in the area and the corollarythat these species were largely reported
from ecological studies and seldom documented by museum specimens or illustrations.
The number of species per establishment success category along with the contribution of alien taxa distributed at each
MSFD area is illustrated in Figure 3. Further details per taxonomic group are provided below.
Figure 4 shows the origin of alien
species for each sub-basin. Species originally distributed either in the Indo-Pacific
Medit. Mar. Sci., 11/2, 2010, 381-493

Fig. 3: a: Establishment success per basin (all taxa); b, c, d: Partitioning of alien taxa per establishment
success.
Medit. Mar. Sci., 11/2, 2010, 381-493

455

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

or Indian or Red Sea are clustered together. Similarly, species originating in either
the east or west of the tropical Atlantic aregrouped together, and so are those originating from the North Atlantic or the
North Pacific.
It is estimated that almost 60% of thealien species in the Mediterranean Sea
come from the Suez Canal. By adding
those species introduced via shipping
and/or aquaculture and trade, it is quite
clear that most of the alien species in the
Mediterranean are warm water species,
originating in (sub)tropical seas. In particular, thermophilic species, (Indo-Pacific,
Indian Ocean, Red Sea, tropical Atlantic,
tropical Pacific, circum(sub)tropical),account for 88.4% of the introduced
species in the EMED, 72.8% in the
CMED, 59.3% in the WMED and 56.1%
in the Adriatic. The respective figures for
Indo-Pacific species only are 47.5% for
EMED, 24.9% for CMED, 22.8% for the
Adriatic and 20% for WMED, whereas
tropical Atlantic species show a sharp
decrease from WMED (10.1%) to EMED
456

(2.7%) (Fig. 4). Cold water species, circumboreal, N Atlantic, N Pacific, make upa small percentage ranging between 4.2%
and 21.6%, being more pronounced in the
Adriatic and less so in the EMED.
While maritime traffic and otherhuman activities such as aquaculture areimportant vectors for the introduction ofalien species worldwide (RUIZ et al.,2000), in the Mediterranean they are not
the main responsibles for the large differences observed among the four basins. In
the EMED the human intervention
responsible for most of the aliens is the reestablishment of the connection with the
Indo-Pacific through the Suez Canal(1869), rather than the actual transfer ofthe invaders. In addition, with the present
climate change (BELKIN, 2009), the tropical features and temperature of the waters
are increasing more quickly in the EMED,
implying dramatic modifications of thebiota (POR 2009, 2010). As a consequence,Indo-Pacific species (regardless of themode introduction) have found optimal
environment for settlement in the EMED.
Medit. Mar. Sci., 11/2, 2010, 381-493

On the contrary, the Adriatic Sea,
which is topographically a dead end, is thearea with the lowest number of aliens,
receiving them among those alreadyestablished in the EMED and CMEDthat spread northwards, or among those
introduced via shipping or aquaculture in
hot spot areas such as the Venice Lagoon(OCCHIPINTI-AMBROGI et al., 2010).
A significant number of Indo-Pacificspecies reaches the WMED, which isenriched by ship-transferred species ofPacific origin mostly among macrophytes.
The Strait of Gibraltar is essentially different from the Suez Canal as a potential
pathway for alien species. First of all, it isan ancient waterway, believed to haveoriginated 5.33 million years ago(HILGEN & LANGEREIS, 1993), compared to the 142 years of the Suez Canal.
Therefore, the current distribution ofAtlantic species, tropical or not, with partof their range in the Mediterranean is theresult of a natural process over a longtime; these species do not in any casequalify as aliens, even if their discovery in
the Alboran Sea comes later than theirfirst description in the Atlantic. Secondly,
the Atlantic coast of Morocco is swept by
a prevalently southward oceanic circulation that prevents many potential newcomers to approach the Strait of Gibraltar.
7.1. Alien foraminiferans in the Mediterranean
A total of 50 foraminiferan species
are reported in this work, all of them
being warm water species. The high diversity of Indo-Pacific species in the EMED
probably implies that many of them are
Lessepsian immigrants. Data on shallow
water foraminifers are very few and scattered. Most works concern deeper assemMedit. Mar. Sci., 11/2, 2010, 381-493

blages, almost totally made up by taxa ofAtlantic affinity, more useful for the biostratigraphical and paleobathymetricalapplications that make foraminifers soimportant for micropaleontologists. As aconsequence, it is very difficult to recognize new Atlantic immigrants entering
the Mediterranean. On the basis of the(scarce) geological data the assumptionthat some more tolerant large foraminifers, such as amphisteginids, could
have survived during the Pleistocene in
the southern Mediterranean cannot be
ruled out.
7.2. Alien macrophytes in the Mediterranean
Compared to previous lists of alien
macrophytes in the Mediterranean Sea(RIBERA & BOUDOURESQUE, 1995;
BOUDOURESQUE & VERLAQUE,
2002; RIBERA SIGU N, 2002;
CORMACI et al., 2004; ZENETOS et al.,2008), the present revised checklist has
taken into account all the possible introductions of exotic genotypes. Several
species already represented in the
Mediterranean Sea by putative native
populations, such as Cladosiphon zosterae,
Desmarestia viridis, Ectocarpus siliculosus
var. hiemalis, Pylaiella littoralis, have been
only considered here if introduction
events are locally evident (e.g. Thau
Lagoon, Lagoon of Venice). A total of
125 taxa (79 Rhodophyta, 24 Fucophyceae, 20 Chlorophyta, 1 Pelagophyceae and 1 Magnoliophyta) has been
listed on the basis of recent works(FURNARI et al., 2010; SFRISO et al.,2010; TSIAMIS et al., 2010; CECERE etal., in press; VERLAQUE et al., in press)
and on-going researches. Among them,
97 taxa (78% of alien macrophytes) areestablished in at least one of the four
457

Mediterranean basins, the remainders(22%) belong to casual and cryptogenic/questionable categories.
As far as the major vectors of introduction are concerned, the transfer ofshellfish stands the first (46% of alien
macrophytes), followed by the shipping
and the Suez Canal (29% and 27%,
respectively each). Some species havebeen probably introduced several times bydifferent vectors. The predominantly
donor regions are temperate and cold
regions (54% of alien macrophytes) vstropical regions (46%). Such a result may
be an artefact due to the unequal effort ofinvestigation between the WMED and the
EMED. Introductions along with shellfish
transfers from north-eastern Atlantic andnorth-western Pacific have been thoroughly studied in the north-western basin
and the Adriatic, while in the EMED
smallest alien macrophytes remain probably insufficiently studied (most of reported Lessepsian migrants are large-sized
species).
7.3. Alien polychaetes in the Mediterranean
Compiling data on alien polychaete
species reported from the Mediterranean Sea revealed a total of 129
species belonging to 26 families, ofwhich 59 species have become established in the region, 31 species are casual, 37 species are questionable and 2
species (Chaetozone corona and Paraprionospio coora) are cryptogenic. The
majority of alien polychaete species havebeen found in the EMED (97 species),
followed by the WMED (49 species), theCMED (35 species) and the Adriatic Sea(22 species). In the Sea of Marmara, 14
alien polychaete species are known, of
which 6 species have become establishedin the area and 8 species are question458

able ( INAR, 2010).
The Mediterranean Sea is largely
under the siege of polychaete species,
mainly introduced from the Red
Sea/Indo-Pacific areas (99 species, 77% of
total species) and Atlantic Ocean (24species). The Suez Canal is one of themain vectors for the introduction of thespecies. The EMED habitats are largely
dominated by the Red Sea immigrants.
Polychaete species that, most probably,
entered the Mediterranean Sea via theSuez Canal and expanded their distribution to the WMED are Nereis jacksoni,
Lysidice collaris, Lumbrineris perkinsi,
Notomastus aberans, N. mossambicus,
Metasychis gotoi, Pista unibranchiata, Streblosoma comatus and Branchiomma luctuosum.
The knowledge about alien polychaete species in the Mediterranean Sea
has greatly increased recently. A total of
59 species were reported after 2005. This
number also includes some species (i.e.
Sigambra constricta, S. parva, Dasybranchus carneus) that were not classified
as alien species by ZENETOS et al. (2005,2008). Due to the lack of precise taxonomic works on polychaetes, there is ahigh number of questionable species.
These species were largely reported from
ecological studies. The future taxonomical
studies to be held in the whole basinwould enable us to realize the real numberof alien polychaete species in the Mediterranean Sea.
7.4. Alien crustaceans in the Mediterranean
A total of 153 alien crustacean species
have entered the Mediterranean. The
majority are decapods (78 species), followed by copepods (42), amphipod (11),isopods (8), cirripedes (7), cumaceans (3),stomatopods (3) and tanaids (1).
Medit. Mar. Sci., 11/2, 2010, 381-493

Approximately 77% of all alien
Mediterranean decapod species have an
Indo-Pacific/Indian/Red Sea origin, while
only 23% are from the Atlantic. The dominance of these ‘eastern alien species’
over the total is obvious; it mainly reflects
the situation (environment characteristics
mainly) in the EMED and CMED where
82% of all Mediterranean alien species
could be found (only the following alien
species are not present: Actumnus globulus, Charybdis feriata, Charybdis japonica,
Charybdis lucifera, Cryptosoma cristatum,
Dyspanopeus sayi, Eriocheir sinensis, Hemigrapsus sanguineus, Herbstia nitida,
Menaethius monoceros, Scyllarus caparti,
Scyllarus posteli and Thalamita gloriensis).
In the WMED, the situation is more balanced, where only 52.6% of the alien
decapods are Indo-Pacific and theremainders are Atlantic. The Adriatic is apeculiar sector with 13 alien species, six ofwhich (Callinectes danae, Charybdis
japonica, Charybdis lucifera, Dyspanopeus
sayi, Herbstia nitida and Scyllarus caparti)
are exclusive to this sector. The EMED
and WMED share only 10 species(12.8%), three of which inhabit estuarine
environments or are related to aquaculture or food (Rhithropanopeus harrisii,
Marsupenaeus japonicus and Necora
puber) and other two are invasive(Callinectes sapidus and especially Percnon gibbesi).
The contribution of Indo-Pacificspecies reaches 86% in copepods vs 11.6%
of Atlantic origin and one debatable case:
the Antarctic visitor (Spinocalanus terranovae). Recent taxonomic studies haveshed some light on alien microcopepods
in the Mediterranean (BÖTTGERSCHNACK & SCHNACK, 2009). The
contribution of warm water species is alsohigh among alien amphipods, excepting
Medit. Mar. Sci., 11/2, 2010, 381-493

Parhyale explorator, a NE Atlantic species
found in 2005 on the coast of Iskenderun
Bay (BAKIR et al., 2008). Yet, the lack ofstudies to determine the actual original
distribution of species such as Cymadusa
filosa, Gammaropsis togoensis and Monocorophium sextonae has led to the assignment of the cryptogenic status to almost
30% of the species.
7.5. Alien molluscs in the Mediterranean
Molluscs are one of the major groups
in the marine fauna worldwide and continue as the first contributors to the alienfauna in the Mediterranean Sea. The total
number of reported alien species amounts
to 212, a small but steady increase ofabout 8% since the recent review of
ZENETOS et al. (2005). More than halfof the species (115 in total) are establishedin at least one of the four basins, 74 arecasual and the remaining 24 are questionable or cryptogenic.
There is a clear decrease in the importance of aliens from the Levantine coast
where extra-Mediterranean species may
represent 10 to 20% of the local fauna,towards the Alboran Sea where only one
species (Godiva quadricolor) is reported,
i.e. less than 1% of the total. Neverthelessthe numbers drop dramatically if we disregard as aliens the Lessepsian immigrants,
following POR’s (2009, 2010) view thatthese species have made their way by theirown means following the restoration of anancient waterway, and therefore should
not be regarded as aliens. In this perspective, there are hardly some thirty aliens in
the Mediterranean which originate fromdonor regions other than the tropicalIndo-Pacific. These have arrived mainly as
a consequence of shipping and transfers ofliving spat for aquaculture, and the mostheavily impacted areas are not the Levan459

tine basin, but the marginal marine environments of the northern part of thebasins, including the Adriatic and thelagoons of the WMED.
It is noteworthy that the Adriatic Sea
shares more established species of molluscs (12 out of 16) with the WMED thanwith any of the other parts of the Mediterranean, even the neighbouring Ionian Sea(8 shared established species and 2 casual). This reflects not only the climatic similarity, with colder sea water temperaturein the northern parts, but also the fact thatmost of these aliens are estuarine species
having some connections with aquaculture, therefore likely to be transported
from the Venice area to the French
Mediterranean lagoons or vice versa.
7.6. Alien miscellaneous invertebrates in
the Mediterranean
The prevailing taxa among alien
invertebrates other than polychaetes,
crustaceans and molluscs, are thecnidarians (46 species: 37 hydrozoans, 4anthozoans, 5 scyphozoans) followed bybryozoans (23), ascidians (16), echinoderms (12), flatworms (12), sponges (8),sipunculans (6), pycnogonids (4),ctenophores (3), chaetognaths (2) andsiphonophores (1). With the exceptionof ascidians, the remaining taxa werepoorly treated by COLL et al. (2010), if
addressed at all.
The current information is always
proportional to the degree of expertise in
the area. Along this line, cnidarians andbryozoans appear to be the best studied
taxa represented by many alien species,
while sponges, although well studied in
the region, have relatively few representatives, and the reported ones are either
questionable or cryptogenic forms.
Alien Anthozoa are represented by460

four species, the most widespread ofwhich is Oculina patagonica. It is a speciesof presumably temperate SW Atlantic
origin, probably introduced into theMediterranean by shipping. Invasive innatural and artificial habitats, especiallyalong the southern WMED and EMED
coasts (SARTORETTO et al., 2008). Five
species coming from the subtropical eastern Atlantic which have recently expanded their range into the Mediterranean
and are presently restricted to the Albo~
A et al., 2007, 2009) areran Sea (OCAN
excluded from our list; yet we keep somereservations on their mode of introduction. Dendrophyllia laboreli is a coral in
expansion at the Canaries and also thecontinental Atlantic coast of Africa. Paramuricea grayi, Eunicella gazella and Eunicella labiata are mainly known from theAtlantic coast of Africa including a wide
tropical area (GRASSHOFF, 1992), buthave not been recognized as part of theMediterranean fauna by SHUCHERT etal. (2003). According to recent observations, boats have borne larvae of Eunicella spp. from the Atlantic coast of Africa intheir ballast waters. Antipathella wollastoni is a typical Macaronesian faunal element with clear tropical affinities~
A, 2004). Its recent(BRITO & OCAN
settlement in the Alboran Sea may havebeen favoured by the present climate
warming.
The 16 alien ascidian species hitherto
recorded in the Mediterranean basin represent about 7% of the known species in
the whole basin. In many cases (13 out of
16 species) the introduced ascidian species
form established populations in antagonism with the endemic species of theMediterranean benthos (ascidian or not).
The species Phallusia nigra, Herdmania
momus, Symplegma brakenhielmi, RhoMedit. Mar. Sci., 11/2, 2010, 381-493

dosoma turcicum, Ascidia cannellata andprobably Microcosmus exasperatus are considered immigrants via the Suez Canal~OZ et al., 2009),(IZQUIERDO-MUN
while species such as Botrylloides violaceus,
Polyandrocarpa zorritensis, Distaplia
bermudensis and Styela clava have been
recently introduced by shellfish culture(BRUNETTI & MASTROTOTARO,
2004; MASTROTOTARO & BRUNETTI,
2006; DAVIS & DAVIS, 2008;~
OZ et al., 2009). InIZQUIERDO-MUN
many cases the first record has beenreported when the species was alreadypresent with dense populations often inantagonism with native species (e.g.
BRUNETTI & MASTROTOTARO,
2004; MASTROTOTARO & BRUNETTI, 2006; DAVIS & DAVIS, 2008).
The majority of echinoderms originate in the Indo-Pacific or are circumtropical species and their distribution islimited to the EMED; only Ophiactis savignyi reaches the WMED. Asterias rubens,
originating in the N Atlantic, is limited to
the Marmara Sea where it was introduced
from an established population in theBlack Sea (KARHAN et al., 2008).
Chaetognaths are represented by two
species, both of Indo-Pacific origin, limited to the innermost part of the EMED.
The presence of the parasitic nematode Anisakis simplex s.str., in some individual hosts (Gaidropsarus granti) fromthe WMED, which seemed to be theresult of immigration of these fish fromthe Atlantic into the WMED
(MATTIUCCI & NASCETTI, 2006), hasbeen excluded, because, according to
MANFREDI et al. (2000), it is a common
parasite in many Mediterranean fishes.

form a significant portion of the Mediterranean marine alien species list. The rateof introduction is significantly high, where
25 species were added since April 2008
(ZENETOS et al., 2008), bringing thetotal number of alien fish species to 149.
This figure reflects the relevance ofMediterranean invasion that has no equivalents on other seas. The richness of
Mediterranean alien fish species isimpressive if compared with native ones.
Even if there is no general consensus on
the total number of Mediterranean fish for example FREDJ & MAURIN (1987)
listed 612 species while QUIGNARD &TOMASINI (2000) numbered 664 species– this review can give a more precise ideaof how the original identity of Mediterranean fish fauna has changed following
the arrival of these immigrants.
So far, no species extirpation has been
documented. Therefore the ingression of
new species generates an increase of
species richness, the main routes contributing to Mediterranean fish diversity
being the Suez Canal and the Strait ofGibraltar. Almost half of the alien fish
species that have been recorded in the
Mediterranean have given origin to permanent populations. The magnitude of
the phenomenon of fish invasion in the
Mediterranean Sea raises concern on theecological and economic consequences
and solicits the urgent need of ongoing
monitoring, especially over the southern
rim of the basin were research effort is lessintense.

7.7. Alien fish in the Mediterranean
In terms of number of species, fish

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

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

8. THE STATE OF THE ART IN
MEDITERRANEAN MARINE ALIEN
SPECIES

461

duced and colonize the Mediterranean ata fast rate, while cold water species aresettling at a lower rate. The net outcome isa jump in species richness.
Compared to the latest lists of alien
biota (COLL et al., 2010; COSTELLO etal., 2010) the present work includes a larger number (954) of species (Table 4). The
divergence in numbers is more obvious infish (149 this work vs. 116 in COLL et al.,2010), polychaetes (129 this work vs. 75 in
COLL et al., 2010), bryozoans (23 thiswork vs. one in COLL et al., 2010), andcnidarians (46 this work vs three in COLL
et al., 2010) (Table 4). Here, we also
include aliens among Porifera, Platyhelminthes and Foraminifera, taxa nottreated by COLL et al. (2010). In contrast
to ZENETOS et al. (2008), we do notaddress microalgal species (either pelagic
or benthic) for the reasons described inchapter 5. This implies that despite thecollective effort of experts attempted inthis work, the number of introduced
species remains underestimated. It is clearthat the introduced species have increased
the biodiversity of the Mediterranean Sea,
as an entity, by 5.9% excluding phytoplankton and microzooplankton (Table4). The figure is even higher reaching
27.9% in fish and 9% on average for thestudied taxa. While species richness isincreasing at whole basin scale, leading toa higher Á diversity, cases of local replace-

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

Taxon
Protozoa (excluding Foraminifera)
Foraminifera
Rhodophyta
Phaeophyta & Pelagophyceae
Chlorophyta
Magnoliophyta
Polychaeta
Crustacea
Mollusca
Cnidaria
Bryozoa
Ascidiacea
Echinodermata
Porifera
Platyhelminthes
Other Invertebrates
Fish
Total
Average %COLL et al., 2010
aliensnative
0
0
>600
0
600
657
73
584
277
23
254
190
17
173
7
1
6
1172
75
1097
2239
106
2133
2113
200
1913
757
3
754
388
1
387
229
15
214
154
5
149
681
0
681
1000
0
1000
2168
2
2166
650
116
534
637*
3.3%
all

This work
aliens% aliens
4
50
8.3
79
13.5
24+1
9.8
20
11.6
1
16.7
129
11.8
153
7.2
212
11.1
46
6.1
23
5.9
16
7.5
12
8.1
8
1.2
12
1.2
16
0.7
149
27.9
954*
5.9%

*excluding monocellular algae

462

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

ment have been reported (GALIL, 2007),
which may imply an alteration of · diversity. In addition, the spatial overlap
between alien and native species causes
biotic homogenization (BEN RAISLASRAM & MOUILLOT, 2009), andhence a depression in ‚ diversity. Thus,
the relationship between the entering ofalien species and threats to biodiversity is
not straightforward (BIANCHI et al.,2010).
While biodiversity is currently a key
indicator of an ecosystem's health, statistics tell too simple a story in the Mediterranean Sea. In a recent study (2008)
assessing fish assemblages associated with
Posidonia oceanica meadows in theRodos Island (EMED), the silververstripe blaasop, Lagocephalus sceleratus,
ranked among the 10 most abundantspecies, totalling 2% of total biomass,
while bluespotted cornetfish, Fistularia
commersonii, and reticulated leatherjacket, Stephanolepis diaspros contributed
0.6%
and
0.3%,
respectively(KALOGIROU et al., 2010). Of the nearly 30000 specimens collected by commercial benthic trawler off the central Israeli
coast at depths between 15 and 30 m, in
October
2008(OCCHIPINTIAMBROGI & GALIL, 2010), only 9%were native Mediterranean species, therest consisted of Erythrean aliens, such as
the highly venomous striped catfish, Plotosus lineatus (11437 specimens), theblotchfin dragonet, Callionymus filamentosus (5745), the silver sillago, Sillagosihama (1423), the kuruma prawn, Marsupenaeus japonicus (1154) and the velvet
shrimp Metapenaeopsis consobrina(1138).
The faunal survey of large harbours
has been largely neglected and is likely toyield many more localized, but estabMedit. Mar. Sci., 11/2, 2010, 381-493

lished, aliens if undertaken seriously.
The application of molecular methods recently developed will be of great
help in the definition of the relationships between populations from different areas, and in the assessment of possible routes of introduction and immigration. In addition, metagenomic techniques will presumably allow a much
more complete if not exhaustive knowledge of the microbial species in given
areas through their molecular signatures. In the meantime, classical taxonomic studies should be intensified toinvestigate the still wide proportion of
unreported alien species of small-sizedtaxa in the Mediterranean Sea including
microalgae.
Acknowledgements
The work was initiated under the auspices of the SEBI2010 - Streamlining
European 2010 Biodiversity Indicators Expert Group 5: Numbers and costs ofinvasive alien species.
Further research on the distributionof alien species in the Mediterranean Sea
has been carried out in the frame of theIntegrated Project ‘SESAME’ (Southern
European Seas: Assessing and Modelling
Ecosystem changes: Project no: 036949)
funded by the EU [A. Zenetos, I. Siokou
and N. Streftaris] and in the frame of theresearch project ‘The impacts of biological invasions and climate change on thebiodiversity of the Mediterranean Sea’
(C.N. Bianchi and C. Morri), an ItalyIsrael co-operation funded by the Italian
Ministry for the Environment. Special
thanks are due to C. Salas Casanova and
A. Logan whose constructive criticism
and suggestions have improved the manuscript.
463

ANNEX: Name changes
Current nameProtozoa
Perkinsus olseni Lester & Davis
Macrophytes
Caulacanthus okamurae Yamada
Codium fragile subsp. fragile (Suringar) Hariot
Hypnea anastomosans Papenfuss, Lipkin & Silva
Saccharina japonica (Areschoug) C.E. Lane, C. Mayes,
Druehl & G.W. Saunders
Spongoclonium caribaeum (B rgesen) M.J. Wynne
Polychaeta
Apoprionospio pygmaea (Hartman, 1955)
Linopherus canariensis Langerhans, 1881
Lumbrineris perkinsi Carrera-Parra, 2001
Prosphaerosyllis longipapillata (Hartmann-Schroeder, 1979)
Streblosoma comatus (Grube, 1856)
Crustacea
Austrominius modestus Darwin, 1854
Amphibalanus eburneus (Gould, 1841)
Glabropilumnus laevis (Dana, 1852)
Linguimaera caesaris Krapp-Schickel, 2003
Parvocalanus crassirostris Dahl, 1894
Pilumnus minutus De Haan, 1835
Portunus segnis (Forssk a l, 1775)
Subeucalanus subcrassus Giesbrecht, 1888
Sternodromia spinirostris (Miers, 1881)
Synalpheus tumidomanus africanus Crosnier & Forest, 1966
Mollusca
Anadara kagoshimensis (Tokunaga, 1906)
Anadara transversa (Say, 1822)
Anteaeolidiella foulisi (Angas, 1864)
Bostrycapulus odites Collin, 2005
Bulla arabica Malaquias & Reid, 2008
Diplodonta bogii Van Aartsen, 2004
Canarium mutabile (Swainson, 1821)
Cerithidium diplax (Watson, 1886)
Cerithidium perparvulum (Watson, 1886)
Conomurex persicus (Swainson, 1821)

464

Old /others name(s) used
Perkinsus atlanticus
Feldmannophycus okamurae
Codium fragile subsp. tomentosoides
Hypnea esperi
Laminaria japonica
Pleonosporium caribaeum
Prionospio pygmaea
Pseudeurythoe acarunculata
Lumbrineris inflata
Sphaerosyllis longipapillata
Streblosoma hesslei
Elminius modestus
Balanus eburneus
Heteropanope laevis
Maera hamigera
Paracalanus crassirostris
Pilumnus hirsutus
Portunus pelagicus
Eucalanus subcrassus
Dromia spinirostris
Synalpheus hululensis
Anadara inaequivalvis / Arca/Scapharca
Anadara demiri
Aeolidiella indica
Crepidula calyptraeiformis
Bostrycapulus aculeatus
Bulla ampulla
Diplodonta cf. subrotunda
Strombus mutabilis
Clathrofenella fusca
Clathrofenella ferruginea
Cerithiopsis tenthrenois
Strombus persicus
Strombus decorus raybaudii

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

ANNEX (continued)
Current name
Doxander vittatus (Linnaeus, 1758)
Ergalatax junionae Houart, 2008
Haminoea japonica (Pilsbry, 1895)
Leucotina natalensis Smith, 1910
Melibe viridis (Kelaart, 1858)
Monotigma lauta (A. Adams, 1853)
Notocochlis gualteriana (Recluz, 1844)
Syrnola lendix (A. Adams, 1863)
Zygochlamys patagonica (King & Broderip, 1832)
Miscellaneous invertebrates
Clytia linearis (Thornely, 1899)
Aquilonastra burtoni (Gray, 1840)
Fish
Atherinomorus forskalii (Ruppell, 1838)
Equulites klunzingeri (Steindachner, 1898)
Favonigobius melanobranchus (Fowler, 1934)

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