Marine alien species in Greek Seas: Additions and amendments by 2010

Medit. Mar. Sci., 12/1, 2011, 95-120 95

Marine alien species in Greek Seas: Additions and amendments by 2010

A. ZENETOS1, S. KATSANEVAKIS1, D. POURSANIDIS2, F. CROCETTA3,D. DAMALAS1, G. APOSTOLOPOULOS4, C. GRAVILI5, E. VARDALA-THEODOROU6

and M. MALAQUIAS7

1 Institute of Marine Biological Resources, Hellenic Centre for Marine Research, P.O. Box 712,19013 Anavissos, Hellas

2 Department of Marine Sciences, University of the Aegean, 81100, Mytilene, Hellas3 Laboratory of Cellular and Developmental Biology, Stazione Zoologica Anton Dohrn

Viale Antonio Dhorn, 80121 Napoli4 Law school, University of Athens, Ippokratus 33, 10680, Hellas

5 Dipartimento di Scienze e Tecnologie Biologiche ed Ambientali, Università del Salento, Lecce, Italy6 Department of Hydrobiology, Goulandris Natural History Museum – Gaia Center, 100 Othonos

Str., Kifissia, 14562, Hellas7 Phylogenetics, Systematics and Evolution Research Group, The Natural History Collections, Bergen

Museum, University of Bergen, 5020-Bergen, Norway

Corresponding author: [email protected]

Received: 09 December 2010; Accepted: 13 January 2011; Published on line: 11 March 2011

Abstract

An update of the inventory of alien marine species from the coastal and offshore waters of Greeceis presented. Records were compiled based on the existing scientific and grey literature, including theHCMR database of Greek alien species (ELNAIS), technical reports, scientific congresses, academicdissertations, websites, and unpublished/personal observations. 47 species were added to the inventory,including 34 invertebrates, one vertebrate (fish), three plants, eight protozoa, and one cyanobacterium.With the new records, the inventory of alien marine species of Greece now includes a total of 237species (33 macrophytes, 131 invertebrates, 42 vertebrates, two bacteria and 29 protozoans). Amongthese, the presence of the gastropod Hypselodoris infucata, the bivalves Dendrostrea frons and Septiferforskali and the chondrichthyan Rhizoprionodon acutus is reported here for the first time. Based onmolecular analysis, the occurrence of Bulla arabica in Greek waters is confirmed, and the suggestionthat previous records of Bulla ampulla in the Mediterranean should be considered as misidentificationof B. arabica is further supported. The acclimitization status of earlier records was revised in the lightof new data, and thus the fish Enchelycore anatina, Seriola fasciata and Tylerius spinosissimus, the redalgae Hypnea cornuta and Sarconema scinaioides, the scyphomedusa Cassiopea andromeda, thecephalopod Sepioteuthis lessoniana, the nudibranch Chromodoris annulata and the bivalves Gas-trochaena cymbium and Pseudochama corbieri were upgraded from casual records to established popu-lations. The increased rate of introductions of warm water species confirms previous findings, whichlink the rate of introduction in the eastern Mediterranean to climate change.

Keywords: Marine aliens; Greece; Hypselodoris infucata; Rhizoprionodon acutus; Dendrostrea frons;Septifer forskali; Bulla arabica.

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

Introduction

The eastern Mediterranean is especiallysusceptible to biological invasions becauseof its crossroads location between the Pon-to-Caspian and the Indian Sea/Red Sea re-gions, the maritime traffic through the Dar-danelles, Gibraltar and Suez, and the exis-tence of many fish and shellfish farms. Hence,Greek waters may function as a gateway tothe dispersal of marine alien species, eitherfrom the Levantine to the Adriatic and/orto the western Mediterranean or from theBlack Sea to the eastern Mediterranean.

The rate of introductions of alien ma-rine species in the Mediterranean Sea is in-creasing (ZENETOS, 2010) as is the spreadof alien species initially established in theLevantine, to other biogeographic areassuch as the Aegean. In addition, the sci-entific community has intensified its effortsto study alien invasions in coastal ecosys-tems. While new studies are being added tothe list, careful examination of historical da-ta on little studied taxa such as hydroids,polychaetes and foraminiferan further ex-pands the Mediterranean aliens’ list.

Following a series of scattered publi-cations dealing with single species or groups,a summary of marine alien biota in Greekwaters was prepared by PANCUCCI-PAPADOPOULOU et al. (2005a) and up-dated by PANCUCCI-PAPADOPOULOUet al. (2005b). ZENETOS et al. (2007), pro-vided some additions to the alien marinefauna of Greece. CORSINI-FOKA &ECONOMIDIS (2007) focused on the ma-rine and estuarine fish fauna, while TSIAMISet al. (2008, 2010) reviewed the alien ma-rine flora. The most up-to-date inventoryof the alien aquatic species of Greece, withcomments on the sources and patterns oftheir introduction is given by ZENETOS etal. (2009a). For Greek waters, the Hellenic

Centre for Marine Research maintains andcontinuously updates an online database ofalien marine species, with records of theirpresence in Greek localities and distribu-tion maps (ZENETOS et al., 2009b; ELNAIS,2010). Systematic study of Foraminifera inthe Greek Aegean Sea has provided addi-tional data on the distributional range ofalien Foraminifera in the Eastern Mediter-ranean (KOUKOUSIOURA et al., 2010).SIMBOURA et al. (2010) reported cases ofalien Polychaeta species that were previ-ously collected but had not been identifieduntil recently, or species not classified asaliens (ZENETOS et al., 2009a) althoughthey were present in Greek check lists.

Recently, MALAQUIAS & REID(2008) described a new species of the genusBulla (Mollusca: Opisthobranchia) from theRed Sea and Arabian Peninsula, that theynamed Bulla arabica; the authors hypothe-sized that the records of Bulla ampulla inthe Mediterranean Sea for the coasts of Is-rael, Greece, Cyprus and Turkey (BARASH& DANIN, 1982; ZENETOS et al., 2004;MIENIS, 2004), would probably belong toB. arabica. Both B. ampulla and B. arabicahave similar shells that can be misidenti-fied; however their anatomy is distinct andmolecular phylogenetics clearly separatesthese two species (MALAQUIAS & REID,2008). Furthermore, these species do notoccur sympatrically; whereas B. arabica isrestricted to the Red Sea and Arabian Penin-sula, the species B. ampulla is not presentin these areas but has a broader Indo-WestPacific distribution.

The aim of this work is to update theZENETOS et al. (2009a) list of marine alienspecies by a) adding all new records reportedin the literature or observed by the authorsup to December 2010; b) amending collec-tion dates based on literature; c) clarify-ing the nomenclature of Bulla ampulla; and

Medit. Mar. Sci., 12/1, 2011, 95-12096

d) revising the establishment success of ear-lier casual records in the light of new data.

Materials and Methods

Records of marine alien species fromcoastal and offshore waters of Greece arecompiled based on the existing scientificand grey literature, including ELNAIS (2010),technical reports, scientific congresses,academic dissertations, and websites/onlineforums (e.g. http://www.scubadive.gr,http://www.seaslugforum.net), as well as onunpublished/personal observations. Recordsfrom grey literature or personal observa-tions are included provided they are sup-ported by physical evidence such as speci-mens or photographs. Additional recordsof species found in the period July 2009-December 2010 or previous findings re-ported within 2010 are discussed. The up-dated list also includes some records re-ported before June 2009 but missed byZENETOS et al. (2009a).

Determination of establishment suc-cess follows terminology provided inZENETOS et al. (2009a) and KATSANE-VAKIS et al. (2009).

Established: Introduced or feral popu-lation of species settled in the wild with free-living, self maintaining and self-perpetu-ating populations unsupported by and in-dependent of humans. Species with at leasttwo records in Greece spread over time andspace (at least three records for fishes) arealso classified as established, in the senseof the CIESM atlas series.

Casual: Casual species are those, whichhave been recorded only once (no morethan twice for fish) in the scientific and greyliterature and are presumed to be non-es-tablished in Greece. In this paper ‘casual’is used in the same sense as ‘alien’ in theCIESM atlas series.

Questionable: Species with insufficientinformation – ‘suspects’. This also includesnew entries not verified by experts or specieswith taxonomic status unresolved.

Cryptogenic: Species with no definite ev-idence of their native or introduced statusaccording to CARLTON (1996) and specieswhose probable introduction occurred in‘early times’ and has not been witnessed,e.g. prior to 1800.

Invasive: Species defined as establishedaliens that have overcome biotic and abi-otic barriers and are able to disseminateaway from their area of initial introductionthrough 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.

For some species reported as casual inZENETOS et al. (2009a) there was evidenceof further expansion, which led us to changethe status of their establishment success.Nomenclature adopted in this paper followsthe World Register of Marine Species(WoRMS <www.marinespecies.org>) andcontributing databases (AlgaeBase,CLEMAM, FishBase).

To clarify the nomenclature of Bullaampulla, living specimens that provided thematerial for DNA analysis were collectedin May 2006, in Psili Ammos on Salamis Is-land of Saronikos Gulf and preserved inethanol. The taxonomic identification ofthis material was assessed by molecularmethods.

The results are structured in six units.A. The new species with a brief text on theirfindings; B. Species to be excluded; C. Changein establishment success with comments; D.Misidentifications and Nomenclaturalchanges; E. Change in introduction dates;and F. Spread of reported aliens and cur-

Medit. Mar. Sci., 12/1, 2011, 95-120 97

rent distribution of marine aliens in theGreek Seas.

Results and Discussion

A. New species

Table 1 presents the new species alongwith some information on their origin, spec-ulated mode of introduction, year of firstsighting, and literature. Among the newspecies 34 are invertebrates, 1 vertebrate(fish), 3 macroalgae, 8 protozoa, and 1cyanobacterium (Table 1). By adding these47 species, and excluding three species (seesection B), the inventory of alien marinespecies of Greece now includes a total of237 species of which 33 are macrophytes,131 invertebrates, 42 vertebrates, 2 bacte-ria, and 29 protozoans (Fig. 1). All the newrecords of invertebrates are zoobenthicspecies belonging to the taxa investigatedthe most intensively, i.e. Polychaeta, Mol-lusca and Crustacea. Fifteen of the recordsappear to have established viable popula-tions already. With the exception of twospecies [Anotrichium okamurae Baldock andChattonella verruculosa Y. Hara & M. Chi-hara], originating in the North Pacific, allthe new introductions are warm water species.Examination of the presumed modes of in-troduction, leads to the inference that someof the well established Lessepsian immi-grants in the Levantine Sea are spreadingnorthwards to the Aegean. However, therole of shipping bears equal responsibilityfor the transfer of warm water species tothe Aegean Sea, either via Suez or via Gi-braltar. It is also worth noticing that withthe addition of the 47 species reported here,the number of alien species has increasedby 24.4% since the last compilation (June2009). Overall, 84 new marine alien specieshave been collected/sighted since 2000, this

high rate of introduction being signifi-cantly correlated to climate change(RAITSOS et al., 2010). More details foreach species are provided in the text thatfollows.

CYANOPHYCEA

The cyanobacterium Trichodesmiumerythraeum is known from nutrient-poortropical and subtropical ocean waters (par-ticularly around Australia). It has beenreported from the Turkish Aegean, in Sici-ly (BARRONE, 2004), and in the Gulf ofGabes, Tunisia (DRIRA et al., 2009). InGreek waters it was found in Chalkida(Evvoikos Gulf) by METAXATOS et al.(2003), while blooms have been reportedfrom Rodos island (S Aegean) and Lesvosisland (NE Aegean) [Spatharis, pers. obs].

PROTOZOA

Marteilia refringens is a parasite whichaffects the digestive system of several bi-valve species, inducing physiological disor-ders and eventually results in death. Thefirst report in Greece dates back to 1997from oyster beds in Thermaikos Gulf(ANGELIDIS et al., 2001). According tothe Directorate General of Veterinary Ser-vices in Athens there have been five out-breaks of the disease in farmed musselsacross Greece until November 2010.

KOUKOUSIOURA et al. (2010) re-ported six cryptogenic foraminiferan speciesnamely: Amphistegina lobifera Larsen, Soritesorbiculus (Forssk l), Cymbaloporetta plana(Cushman), Triloculina fichteliana d’Or-bigny, Planogypsina acervalis (Brady) andCoscinospira hemprichii Ehrenberg fromGreek coastal areas. Their distribution canbe associated with several pathways, via theAtlantic during the interglacial warm

a

Medit. Mar. Sci., 12/1, 2011, 95-12098

Medit. Mar. Sci., 12/1, 2011, 95-120 99

Tab

le 1

Add

itio

nal a

lien

spec

ies

in G

reek

Sea

s to

tho

se r

epor

ted

in Z

EN

ET

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et a

l.(2

009a

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tebr

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tor

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gin

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eaun

know

nun

know

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opic

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200

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edes

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1997

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002

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Medit. Mar. Sci., 12/1, 2011, 95-120100

Tab

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Medit. Mar. Sci., 12/1, 2011, 95-120 101

Tab

le 1

(co

ntin

ued)

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gdom

Spec

ies

Tax

onE

stab

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ent

Vec

tor

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gin

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2000

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mal

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Sea

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read

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y, 1

900)

Cni

dari

a/es

tabl

ishe

dvi

a Su

ez /

Indo

-Pac

ific

MA

RIN

OPO

UL

OS,

19

77(i

nv)

Hyd

rozo

asp

read

ing

1979

Ani

mal

ia

Syna

lphe

us tu

mid

oman

us a

frica

nus

Cru

stac

ea/

cryp

toge

nic

Via

Sue

z?un

know

nK

OU

KO

UR

AS,

un

know

n(i

nv)

(Cro

snie

r &

For

est,

1965

)D

ecap

oda

1978

cont

inue

d

Medit. Mar. Sci., 12/1, 2011, 95-120102

Tab

le 1

(co

ntin

ued)

Kin

gdom

Spec

ies

Tax

onE

stab

lishm

ent

Vec

tor

Ori

gin

sour

ceF

irst

su

cces

sSi

ghti

ngA

nim

alia

A

terg

atis

rose

us (R

üppe

ll, 1

830)

Cru

stac

ea/

esta

blis

hed

via

Suez

/ In

do-P

acifi

cC

OR

SIN

I-FO

KA

&20

09(i

nv)

Dec

apod

asp

read

ing

PAN

CU

CC

I-PA

PA-

DO

POU

LO

U, 2

010

Ani

mal

ia

Gon

ioin

frade

ns p

auci

dent

atus

C

rust

acea

/ca

sual

unkn

own

Indo

-Pac

ific

CO

RSI

NI-

FOK

A20

10(i

nv)

(A. M

ilne

Edw

ards

, 186

1)D

ecap

oda

et a

l.,20

10a

Ani

mal

ia

Mac

roph

thal

mus

gra

effe

i C

rust

acea

/ca

sual

via

Suez

/ In

do-P

acifi

cPA

NC

UC

CI-

PAPA

-20

09(i

nv)

A. M

ilne

Ew

dard

s, 18

73D

ecap

oda

spre

adin

gD

OPO

UL

OU

et

al.,

2010

Ani

mal

ia

Car

dite

s aka

bana

(St

uran

y, 1

899)

Mol

lusc

a/ca

sual

via

Suez

/ R

ed S

eaM

AN

OU

SIS

2008

(inv

)B

ival

via

ship

ping

et a

l,20

10A

nim

alia

C

ham

a pa

cific

a B

rode

rip,

183

4M

ollu

sca/

ques

tiona

ble

via

Suez

/ In

do-P

acifi

cM

AN

OU

SIS

2008

(inv

)B

ival

via

spre

adin

get

al.,

201

0A

nim

alia

Se

ptife

r for

skal

i(D

unke

r, 18

55)

Mol

lusc

a/es

tabl

ishe

dvi

a Su

ez /

Indo

-Pac

ific

/ T

his

wor

k20

10(i

nv)

Biv

alvi

asp

read

ing

Red

Sea

Ani

mal

ia

Den

dros

trea

frons

Mol

lusc

a/es

tabl

ishe

dvi

a Su

ez /

Indo

-Pac

ific

/ T

his

wor

k20

10(i

nv)

(Lin

naeu

s, 17

58)

Biv

alvi

asp

read

ing

Red

Sea

Ani

mal

ia

Dip

lodo

nta

bogi

i va

n A

arts

en, 2

004

Mol

lusc

a/ca

sual

via

Suez

/ R

ed S

eaM

AN

OU

SIS

2008

(inv

)B

ival

via

spre

adin

get

al.,

2010

Ani

mal

ia

Dos

inia

ery

thra

ea R

oem

er, 1

860

Mol

lusc

a/ca

sual

via

Suez

/ R

ed S

eaM

AN

OU

SIS

2007

(inv

)B

ival

via

spre

adin

get

al.,

2010

Ani

mal

ia

Hyp

selo

doris

infu

cata

M

ollu

sca/

esta

blis

hed

via

Suez

/ In

do-P

acifi

c /

Thi

s w

ork

2007

(inv

)(R

üppe

ll &

Leu

ckar

t, 18

28)

Opi

sthob

ranc

hia

spre

adin

gR

ed S

ea

periods of the Pleistocene-Holocene or fair-ly recent invasions, now well established,most probably via the Suez Canal(KOUKOUSIOURA et al., 2010). Of these,Amphistegina lobifera and Sorites orbiculuswere included in the compiled list ofZENETOS et al. (2009a). Coscinospirahemprichii and Amphistegina madagascariensisare missing from the list although they havebeen known since the 1950s (BLANC-VERNET, 1969). Amphistegina madagas-cariensis was reported in the infralittoralzone of Peloponnesos, Kastellorizo, Kritiand Saronikos in Halophila meadows(BLANC-VERNET, 1969).

Coscinispira hemprichii is a symbiont-bearing foraminiferan exhibiting a globaltropical distribution. It is reported from theIndo-Pacific and the Red Sea. Widely dis-tributed in the Mediterranean, it is knownfrom the coasts of Turkey under the syn-onym Peneroplis arietinus (MERIC et al.,

2008). In Greece C. hemprichii was report-ed from Kriti (HOLLAUS & HOTTINGER,1997), and from the Aegean Sea under thename Spirolina arietina (BLANC-VERNET,1969). KOUKOUSIOURA et al. (2010)found C. hemprichii mainly in the centraland northern Aegean sites, yet in very lowabundances.

Cymbaloporetta plana thrives in the shal-low water environments of the Pacific Ocean.KOUKOUSIOURA et al. (2010) found C.plana mainly at the northern and centralAegean sites displaying a peak in relativeabundance at the site of Mavro Lithari (Sa-ronikos Gulf).

Planogypsina acervalis seems to be com-mon in the Atlantic, Indian and Red Seas.BLANC-VERNET (1969) described thisspecies from the coasts of Provence, France.KOUKOUSIOURA et al. (2010) reportedP. acervalis as rare at the central Aegeanand always in very low abundances.

Medit. Mar. Sci., 12/1, 2011, 95-120 103

Fig. 1: Breakdown of marine alien species in Greek Seas per taxonomic group.

Triloculina fichteliana thrives in theshallow water environments of the AtlanticOcean, Pacific Ocean and Red Sea. In theMediterranean Sea, MERIC et al. (2008)reported it from the coasts of Turkey.KOUKOUSIOURA et al. (2010) report-ed it as rare at the southern and centralAegean and always found in very low abun-dances.

DINOFLAGELLATA

The species Chattonella verruculosa ap-peared in Amvrakikos in 1998, causing massfinfish mortality (NIKOLAIDIS et al., 2005).However, its presence has not been con-firmed by later findings and even its iden-tification is problematic (K. Aligizaki, pers.comm.). Blooms of Prorocentrum triestinumwere reported in Thermaikos Gulf during2000 and 2001 and were considered re-sponsible for water discoloration(NIKOLAIDIS et al., 2005 reported as Pro-rocentrum redfeldii).

MACROALGAE

TSIAMIS & BELLOU (2010) report-ed a first finding of the red macroalgaeApoglossum gregarium in Greece, which isalso the first record in the eastern Mediter-ranean basin. It was found on artificial sub-strata (in experimental frames, after oneyear of deployment) at 50 m depth nearSapientza Island, southeastern Ionian Sea,Greece (Lat 36o 44’33.6"N, Lon 21o42’32.5"E), on 2 April 2009. The Greek spec-imens were identical to the previous Mediter-ranean descriptions and were epiphyticon a bryozoan. Moreover, the Greek habi-tat is the deepest ever recorded for thespecies. The surrounding natural habitatwas sandy mud, and the invasive alga Cauler-pa racemosa var. cylindracea was the pre-

dominant species (TSIAMIS & BELLOU,2010).

Two taxa have probably been confusedunder the name of Anotrichium furcellatum:the native taxon described from Naples anda species (Anotrichium okamurae) originallydescribed from Japan, probably introducedfrom the Pacific at an unknown time andexhibiting invasive behaviour. In Greece,where A. furcellatum has frequently beenrecorded in the Ionian and Aegean Seas,three hypotheses have to be considered: ei-ther we have the native taxon, or the intro-duced species, or both taxa. Pending newinformation, the Greek records of A. fur-cellatum are provisionally attributed to A.okamurae as a debatable case (TSIAMIS etal., 2010).

In the Mediterranean Sea, Chondriapygmaea was found among the epiflora ofthe introduced Halophila stipulacea everytime that it was studied. Considering thewide distribution of the introduced seagrassin Greece, the occurrence of C. pygmaea ishighly probable. Pending confirmation, C.pygmaea was provisionally included in thequestionable category by TSIAMIS et al.(2010).

ASCIDIACEA

The ascidian Phallusia nigra is amongthe established alien species of the Levan-tine basin. Its presence in Greek waters wasfirst noted in 2009. Established populationswere observed by SCUBA diving offshorethe beach ‘Stegna' and in one of the har-bours of Rodos Island (KONDILATOSet al., 2010).

CRUSTACEA/DECAPODA

A single specimen of the crab Macroph-thalmus graeffei, was collected in 2009 by

Medit. Mar. Sci., 12/1, 2011, 95-120104

PANCUCCI et al. (2010) in the Gulf of Tri-anta (W. Rodos Island). The xanthid Ater-gatis roseus firstly recorded in 2009 at Plim-miris Bay (SE Rodos) at 6-8 m depth onsandy bottom with rocks covered by vege-tation (CORSINI-FOKA & PANCUCCI-PAPADOPOULOU, 2010) is now consid-ered established (CORSINI-FOKA et al.,2010). Both aforementioned species arewell established in the Levantine Basin.

CORSINI-FOKA et al. (2010a) docu-mented the first record for the Mediter-ranean Sea of the red swimming crab Go-nioinfradens paucidentatus, from specimenscollected at Rodos Island (southeasternAegean Sea), giving a detailed descriptionof samples and biotopes.

The Alboran populations of Synalpheustumidomanus africanus may represent a nat-ural expansion of the biogeographic dis-tribution range of the species from theAtlantic. However, the references of Sy-nalpheus tumidomanus africanus from Greeceand Turkey (KOUKOURAS &KATTOULAS 1974; KOCATA , 1981)could indicate a separate advance in theeastern Mediterranean littoral from the Is-rael populations. Recently it has been cap-tured in Sicily (BACCI et al., 2010). The un-resolved question is whether the easternMediterranean populations are ‘alien' ornot. Following ZENETOS et al. (2010) wehave classified it as cryptogenic.

MOLLUSCA

Dosinia erythraea is reported from twolive specimens with length of 35 mm and 44mm respectively, collected at a depth of 5-8 m, during 2007 at Aggelochori, ThermaikosGulf (MANOUSIS et al., 2010). The recordof Diplodonta bogii is based on one live spec-imen of 7 mm, found at 5m in Palioura,Epanomi, Thermaikos Gulf in March 2008

(MANOUSIS et al., 2010). Cardites aka-bana was collected from sandy-muddy bot-toms of the sublittoral zone of the southernThessaloniki Gulf and eastern ThermaikosGulf. Following the first finding at 0.5 mdepth in Cape, Epanomi in April 2008, sev-en live specimens and two shells (some bear-ing their periostracum) were found withtheir lengths ranging from 1.6 mm to 7.6mm (MANOUSIS et al., 2010).

Two established populations of Den-drostrea frons were discovered in June 2010in northern Kriti and in June 2010 in Astyp-alaia Island. The species, initially report-ed from southern Turkey ( EVIKER, 2001),appears to have spread eastwards and be-come established in Vai Bay, Astypalaia Is-land (36Æ35'13.75"N 26Æ24'10.52"E), wherethousands of live specimens were found ona big metallic cage brought by the sea to asandy-muddy bottom with Posidonia patch-es at 5-6 m of depth (cages in use were sit-uated 1.5 km away, just at the end of thebay) (R. Villa, pers. comm.) and in Kriti, asevidenced by the finding of ci 25 mature in-dividuals in Agia Pelagia ( D. Poursanidis,A. Zenetos, pers. obs).

About ten live Septifer forskali speci-mens were found byssed on Dendrostreafrons at Astypalaia Island, together withPinctada radiata and Brachidontes pharao-nis, constituting the first record of this speciesfor Greece. The species was sighted againin Agia Pelagia, Kriti in December 2010(Poursanidis, pers. obs.) and in Alimos, Sa-ronikos Gulf, attached on Acanthocardiaspinosa (P. Ovalis, pers. obs).

Hypselodoris infucata is very commonin the Levantine Sea where it can be foundall year round. This work reports the firstfinding of the species in Greek waters. Asingle specimen of Hypselodoris infucata(Ruppell & Leuckart, 1830) was pho-tographed on a rocky bottom at a depth

Medit. Mar. Sci., 12/1, 2011, 95-120 105

of 0.5 m in the west side of the port of Kastel-lorizo Island, on 28 August 2007. This specieswas sighted again at the same site in Septem-ber 2009.

The record of Chama pacifica is basedon one specimen (6.7 mm in length) foundin 2008 as epibiont of Pinna nobilis shell col-lected at Aggelochori, Thessaloniki Bay(MANOUSIS et al., 2010). However, thepicture shown in Fig. 2d of MANOUSIS etal. (2010) is not convincing. The specimenis too small to be accurately identified as C.pacifica and the record is based on a singleindividual. It is speculated here that thespecimen corresponds to Chama aspersa.

POLYCHAETA

The presence of Chaetozone corona inthe Mediterranean Sea was first reportedby INAR & ERGEN (2007) from IzmirBay who postulated that this species mightbe a cryptogenic species. According toSIMBOURA et al. (2010), earlier recordsdesignated as Chaetozone sp. B are currentlyidentified as C. corona. Thus, C. coronashows a wide distribution in North EvvoikosGulf, Thermaikos Gulf, Kyklades and Kri-ti in disturbed and undisturbed sites(SIMBOURA et al., 2010).

Marphysa disjuncta is a species firstrecorded in the Mediterranean Sea fromFethiye Bay (Levantine Sea, Turkey) byKURT AHIN & INAR (2009), pre-sumably introduced from the Pacific Oceanthrough ballast waters. In Greece Marphysadisjuncta was found in North Evvoikos Gulf(Aegean Sea) at 80 m depth on mud mixedwith metalliferous waste (slag), in the calderaof Santorini island (Kyklades, Aegean Sea)at 316 m depth on muddy gravel of volcanicmaterial, and in Saronikos Gulf at 60-70 mdepth in muddy sandy sediments(SIMBOURA et al., 2010).

Pseudopolydora paucibranchiata wasoriginally described from Japan and the Pa-cific Ocean and its establishment in theMediterranean was attributed to ballast wa-ters (DAGLI & INAR, 2008). In GreeceP. paucibranchiata was found in LarymnaBay (Evvoikos Gulf, Aegean Sea). It was al-so found casually among the endofaunaof a Zostera meadow in Geras Gulf (LesvosIsland) and in Elefsis Bay (Saronikos Gulf)(SIMBOURA et al., 2010).

Neopseudocapitella brasiliensis has beenknown since 1991 in the Evvoikos Gulf (fromChalkis to Larymna) and in Kriti(SIMBOURA, 1996). The species is alsoknown in the Levantine Sea (Cyprus, Turkey),Adriatic, Central and west Mediterranean(ZENETOS et al., 2010).

Lumbrineris perkinsi was first reportedas L. inflata by GIANGRANDE et al. (1981),who collected it from the Ischia Harbour(Tyrrhenian Sea, western Mediterranean).In Greece (as L. inflata), it was found inEvvoikos and Thermaikos Gulfs in the 1990s(ARVANITIDIS, 1994). Dense populationsof L. perkinsi on the southern coast of Turkey( INAR, 2009), Egypt (ABD-ELNABY,2009) and Greece (ARVANITIDIS, 1994)and the single record from the westernMediterranean might indicate that thisspecies could have been introduced to theMediterranean from the Suez Canal. How-ever, its occurrence in the Indo-Pacific areashould be checked.

The terebellid species Polycirrus twistiPotts, 1928 had been confused with Poly-cirrus plumosus Wollebaeck, 1912 becauseof incomplete or unclear descriptions andits presence in Greek Seas had been over-looked. SIMBOURA (2011) reports on itsoccurrence in Korinthiakos Gulf and dis-cusses previous findings in the Aegean Seasince the eighties.

A total of 14 questionable alien poly-

Medit. Mar. Sci., 12/1, 2011, 95-120106

chaetes (Capitellethus dispar, Dasybranchuscarneus, Dispio magnus, Dodecaceria capen-sis, Hyboscolex longiseta, Leocrates chinen-sis, Loimia medusa, Naineris quadraticeps,Polydora spongicola, Protodorvillea biartic-ulata, Scoletoma debilis, Sigambra con-stricta, Syllis schulzi and Timarete dasylophius)was reported by SIMBOURA & NICO-LAIDOU (2001) and ZENETOS et al. (2010)but they had not previously been classi-fied as aliens.

CNIDARIA

Clytia linearis was found on algae, Posi-donia oceanica, sponges, hydroids, bryozoansand on polychaete tubes, in shallow warmwaters (BOERO & FRESI, 1986). In theMediterranean, it has been recorded fromthe French coast, Ligurian Sea, TyrrhenianSea, Adriatic and Ionian Sea, Spanish wa-ters, Alboran Sea off the coast of Morocco,Greek waters, and Lebanese waters. InGreece, C. linearis was recorded from AspraSpitia (Korinthiakos Gulf) byMARINOPOULOS (1979) (as Clytia gravieri)(collection date: 1977) and from Kos Island(Aegean Sea) by MORRI & BIANCHI(1999) (collection date 1981).

FISH

The milk shark Rhizoprionodon acutushas a circumtropical distribution (TropicalAtlantic and Indian Ocean, Red Sea andJapan). To date it has been considered as avagrant fish in the Mediterranean, its recordbased on a single specimen captured in theGulf of Taranto, Italian Ionian Sea(PASTORE & TORTONESE, 1984). Thepresent work reports a second sighting ofthe species in the Greek SW Ionian Sea (Lat36Æ 52' N, Lon 20Æ 32' E). The individual,captured on 16 July 2004 by D. Damalas,

was measured and weighed. It was a malewith a total length 162cm, a fork length130cm, pectoral length 102cm and a roundweight of 21.5 Kg. A tissue sample was keptfor DNA analysis (code No 42), preservedin the Ichthyology Laboratory of the Univer-sity of Athens.

B. Species to be excluded

The alga Neosiphonia sphaerocarpa(Borgesen) M.S. Kim & I.K. Lee was firstrecorded in the Mediterranean Sea fromTunisia and successively from Corsica, theBalearic Islands and Milos Island in Greece.CORMACI et al. (2004) considered N. sphae-rocarpa as an alien species. However, thespecies is distributed worldwide and mo-lecular data are needed to locate its placeof origin. Moreover, the identification ofthe species is not easy. Consequently theabsence of N. sphaerocarpa in the Mediter-ranean prior to 1970 could simply mean thatit was overlooked or mistaken for anotherPolysiphonia species. Pending further in-vestigations, TSIAMIS et al. (2010) provi-sionally consider this species as native in theMediterranean Sea.

The record of Circenita callipyga (Born1778) is based on one shell measuring 8.6mm in length, found in the Gulf of Thessa-loniki, Mikro Emvolo, at 5m in 2008(MANOUSIS et al., 2010). However, thepicture of the shell figured in MANOUSISet al. (2010) is not convincing in our opin-ion as it does not match the species de-scription (size, outline, sculpture). The ovaterather than trigonal outline and the radialsculpture are diagnostic features for thespecies. This is not clear from Figure 3aof MANOUSIS et al. (2010) where a wholebivalved specimen is depicted, presumablybelonging to a juvenile Gouldia minima(Montagu, 1803). After consideration of the

Medit. Mar. Sci., 12/1, 2011, 95-120 107

Mediterranean distribution of the specieswhich is restricted to Israel (ZENETOS etal., 2004), this record has been excludedfrom our list.

Two of the Atlantic invaders (Alop-ias superciliosus and Gaidropsarus granti)are excluded from the list of Greek alienspecies following ZENETOS et al. (2010),who consider them Atlanto-Mediterraneanspecies.

C. Change in establishment success

The species listed in Table 2 have beenreported from more than one site, and hencethey are now considered to be establishedin Greek waters. Details on their distribu-tion are provided below.

The establishment of the fangtoothmoray eel Enchelycore anatina in the southAegean is confirmed by the finding of a spec-imen in Rodos Island in 2010 (KALO-GIROU, 2010) and another one in KarpathosIsland (CORSINI-FOKA, 2010, pers. comm.)As an indication of its establishment, localfishermen state that they catch approxi-mately 4-5 individuals per vessel with gill-nets during the summer period along Ro-

dos Island’s rocky bottoms and even in Kastel-lorizo Island (KALOGIROU, 2010).

A second record of the small Indo-Pacific fish Tylerius spinosissimus (Te-traodontidae) (known as spiny blaasop) wasdocumented at the beginning of 2009 in thewaters of Rodos Island (south-eastern AegeanSea, Greece). The finding shows the puta-tive establishment of a population of thisalien fish in the Mediterranean around theDodekanisos islands (CORSINI-FOKAet al., 2010b).

Seriola fasciata (Bloch, 1793) is a speciesof amphi-Atlantic origin, which made itsfirst appearance in Rodos in 2004. Conse-quent findings in Rodos (2009) and Lesvos(2010) [ELNAIS, 2010] confirm its estab-lishment success in Greek waters.

The Lessepsian immigrant Hypnea cor-nuta, found only twice in the Aegean Sea[in 1894 in Rodos Island (as H. valentiae)and one century later, in 2007], was foundagain in the Saronikos Gulf (TSIAMIS etal., 2008) in August 2009; hence it appearsto have become established in the area(TSIAMIS et al., 2010).

The red alga Sarconema scinaioides,which was reported from Greece only in the

Medit. Mar. Sci., 12/1, 2011, 95-120108

Table 2Alien marine species in Greece classified as established.

Species Taxon SourceEnchelycore anatina (Lowe, 1839) Fish/Osteichthyes KALOGIROU, 2010Tylerius spinosissimus (Regan, 1908) Fish/Osteichthyes CORSINI-FOKA et al., 2010bSeriola fasciata (Bloch, 1793) Fish/Osteichthyes ELNAIS, 2010Hypnea cornuta (Kützing) J. Agardh 1851 Rhodophyta TSIAMIS et al., 2010Sarconema scinaioides B rgesen, 1934 Rhodophyta TSIAMIS et al., 2010Cassiopea andromeda (Forssk l, 1775) Cnidaria/Scyphozoa This workGastrochaena cymbium Spengler 1783 Mollusca/Bivalvia MANOUSIS et al., 2010Pseudochama corbieri (Jonas, 1846) Mollusca/Bivalvia MANOUSIS et al., 2010Chromodoris annulata (Eliot, 1904) Mollusca /Gastropoda This workSepioteuthis lessoniana Lesson, 1830 Mollusca/Cephalopoda This work

a

Saronikos Gulf in the early 1980’s, was foundagain at the same location after about 30years (see TSIAMIS et al., 2009); thus, re-vealing an established population in the areastill surviving after three decades (TSIAMISet al., 2010).

Cassiopea andromeda, which occursalong the Levantine coastline, was recent-ly reported from Malta (SCHEMBRI et al.,2010). SCHÄFER (1955) reported theoccurrence of very young specimens (2-30mm) on Neokameni, a small volcanic islandnear Santorini, Aegean Sea, where themedusae flourished in rocky pools with wa-ter temperatures reaching up to 36Æ Cdue to volcanic activity. The present workreports on the finding of >15 alive individ-uals of C. andromeda in Paros Island atdepths between 2–10 m (June 2010: S. Kat-sanevakis pers. obs.) and in S. Evvoikos,(Chalkoutsi, September 2010: A. Zenetospers. obs.).

From six stations of the sublittoral zoneof eastern Thermaikos Gulf, ThessalonikiGulf and Thessaloniki Bay, more than 30live specimens of Gastrochaena cymbium,5.4-5.7 mm in length, were dissected out oftheir calcareous capsules (MANOUSIS etal., 2010). Empty shells of G. cymbium wereextracted from Ostrea sp. shells collected inElefsis Bay, Saronikos Gulf (S. Katsanevakis,pers. obs.).

Two live specimens of Pseudochamacorbieri (Jonas 1846) measuring 13.1 mmand 19.1 mm, were found in April 2006 inEpanomi, Thermaikos Gulf, North Aegeanat 2 m depth (MANOUSIS et al., 2010).

The presence of Chromodoris annula-ta (Eliot, 1904) in Greece was documentedfrom a single sighting of the species in Sa-ronikos Gulf. The species was observed againin September 2010 in Madraki, Kastellori-zo at a depth of 0.5 m (G. Apostolopoulos,pers. obs.).

Sepioteuthis lessoniana Lesson, 1830 isone of the most commercially importantsquid species for inshore fisheries through-out its distributional range. It is commonin the Levantine basin and first appearedin Greek waters at Dodekanisos in 2009(LEFKADITOU et al., 2009). Ever since,occasional reports e.g. in Kastellorizo (G.Apostolopoulos pers. obs.) and elsewhere(ELNAIS, 2010) testify to its establishmentsuccess and eastward spread.

The presence of Alepes djedaba(Forssk l, 1775) in Greek water has to bedowngraded to questionable. The specieswas not reported in PANAGIOTOPOULOS(1916), as mentioned in ZENETOS et al.(2009a). However, under its synonym Caranxgallus it was reported by MALDURA (1938)in Rodos, a citation that was later correct-ed by LASKARIDIS (1948) as the nativeAlectis alexandrinus. It appears thatPAPACONSTANTINOU (1988) cites A.djedaba as questionable based onTORTONESE (1952) who has repeatedMALDURA (1938). Although it is estab-lished in the Levantine Sea and easternAegean Sea, its occurrence in Greek wa-ters, based only on BINI (1960), remains tobe confirmed.

D. Misidentifications and Nomenclaturalchanges

Attention was paid to recent nomen-clatural updates (Table 3). These are theresult of the latest taxonomic and/or mo-lecular studies. Such is the case for the com-mercially important swimming crab Por-tunus pelagicus (Linnaeus, 1758) and thegastropod Bulla ampulla (Linnaeus, 1958).Based on morphological and DNA charac-ters as well as biogeographical considera-tions P. pelagicus is in fact Portunus segnis,a species confined to the western Indian

a

Medit. Mar. Sci., 12/1, 2011, 95-120 109

Ocean from Pakistan to South Africa (LAIet al., 2010).

With regard to Bulla ampulla, DNA wasextracted and amplified for the mitochon-drial genes cytochrome c oxidase subunitI (COI) and 16S rRNA. COI amplificationwas not successful but the 16S yielded goodquality sequences. Those were blasted inGenBank and results retrieve maximumsimilarity with the two sequences of Bullaarabica available in this database, includingwith the holotype of the species (BMNH20060103/1, United Arab Emirates; GenBankaccession number: DQ986575.1). The re-sults of molecular analysis confirm the oc-currence of Bulla arabica in the Mediter-ranean Sea and support the suggestion by

MALAQUIAS & REID (2008) that previ-ous records of Bulla ampulla in this basinshould be considered misidentifications ofB. arabica.

E. Change in introduction dates

Recent literature and/or re-examina-tion of other sources make it clear thatthe collection date of species listed in Table4 needs to be corrected.

F. Distribution of reported aliens

Besides those species considered as ca-sual and reported in other locations, whichreveals their establishment success, many

Medit. Mar. Sci., 12/1, 2011, 95-120110

Table 3Misidentifications and nomenclature changes.

New name Old name SourceEquulites klunzingeri (Steindachner, 1898) Leiognathus klunzingeri FISHBASE

(Steindachner, 1898) Portunus segnis (Forssk l, 1775) Portunus pelagicus LAI et al., 2010

(Linnaeus, 1758)Conomurex persicus (Swainson, 1821) Strombus persicus WoRMS

(Swainson, 1821)Bulla arabica Malaquias & Reid, 2008 Bulla ampulla This work

(Linnaeus, 1758)

a

Table 4Correction of collection dates reported in ZENETOS et al. (2009a).

Species Correct collection Sourcedate

Alepes djedaba (Forssk l, 1775) 1960 not 1916 BINI, 1960Siganus rivulatus Forssk l, 1775 1925 not 1932 ZACHARIOU MAMALINGA, 1990Apogon pharaonis Bellotti, 1874 1982 not 2002 ZACHARIOU-MAMALINGA, 1990Pempheris vanicolensis Cuvier, 1831 1983 not 1985 ZACHARIOU-MAMALINGA, 1990Amphistegina lessonii d’Orbigny, 1826 1974 not 1997 Hottinger pers. comm.Conomurex persicus (Swainson, 1821) 1983 not 1986 VERHECKEN, 1984

aa

of the established species have expandedtheir geographic distribution range north-wards or have spread rapidly within the samearea, revealing an invasive character. Ex-amples are presented below:

The recent records of Lagocephalus scel-eratus (Gmelin, 1788) verify that the speciesis now very common along the central, south-ern and southeastern coasts of the AegeanSea. Recent captures confirm the spread-ing of the species in the NW Aegean Sea,increasing the number of its northern recordsin the whole Mediterranean (MINOS et al.,2010) (Fig. 2a).

Findings of the recently recorded Pet-roscirtes ancylodon Rüppell, 1838, Calliony-mus filamentosus Valenciennes, 1837, Torquigen-er flavimaculosus Hardy & Randall, 1983 andScomberomorus commerson Lacepède, 1800from Rodos in summer 2008, confirm theirquick establishment (CORSINI-FOKA, 2010).Torquigener flavimaculosus and P. ancylodonwere also spotted in Kastellorizo (G. Apos-tolopoulos, pers. obs.).

Stephanolepis diaspros Fraser-Brun-ner, 1940, well established in the southAegean Sea (CORSINI-FOKA &ECONOMIDIS, 2007; CORSINI-FOKA,2010), was captured in boat seine haulscarried out in the period 2008-2009 in theAegean Sea, occurring in up to 60% ofthe hauls (LEFKADITOU et al., 2010).Its current distribution is illustrated inFig. 2b.

KOUKOUSIOURA et al. (2010) pos-tulated that Amphistegina lobifera, is thedominant foraminifer species at the south-ern and central Aegean usually comprisingmore than 30% of total specimens of theforaminiferal assemblages. In the northernsite (N. Kallikratia) A. lobifera was presentat very low abundances.

Sorites orbiculus was regularly found atthe central Aegean and southern Aegean

sites with somewhat higher percentages atFalasarna (7.5% of the foraminiferal fau-na) (KOUKOUSIOURA et al., 2010).

Two bivalvia species of the Chamidaefamily were collected from rocky bottomsof Thermaikos Gulf: Chama asperella La-marck 1819 (four live specimens with lengthranging from 8.2 mm to 17.6 mm) at 5mParalia, Epanomi, in October 2008, on Pin-na nobilis, and at Palioura, Epanomi, in De-cember 2008, and Chama aspersa Reeve1846 (two live specimens 15 mm and 26 mmin length, respectively), in 2008, from thesea shore at Paralia, Epanomi, ThermaikosGulf (MANOUSIS et al., 2010). Chama as-persa was also collected in September 2010from Elefsis Bay, Saronikos Gulf (S. Kat-sanevakis, pers. obs.).

Brachidontes pharaonis (Fischer P.,1870), previously known only from Rodosand Saronikos Gulf, was also observed inSE Kriti (P. Ovalis, ELNAIS, 2010) andAstypalaia Island in June 2010. Pinctadaradiat· (Leach, 1814), known as north asthe Evvoikos Gulf, was collected at Palioura,Epanomi, in March 2005, at Cape, Epano-mi in August 2010 (S. Mitsoudi, pers.comm.) and at Astypalaia Island in June2010 (F. Crocetta, pers. obs.).

Trochus erithraeus Brocchi, 1821 previ-ously known from Kriti, (COSENZA &FASULO, 1997) has been spotted in 2010in central Aegean Sea. One shell was col-lected on the coast of Syros Island and de-posited in the collection of the Goulan-dris Natural History Museum.

An established population of Synaptu-la reciprocans (Forssk l, 1775) measuring20-100 cm in length was found at Loutro,SW Kriti (35Æ12' 1.68"N 24Æ 4' 53.70"E)at depths more than 5 m (D. Poursanidis,ELNAIS, 2010) (Fig. 2c).

Further to its spread in the North Aegeanand Ionian Sea (KATSANEVAKIS &

a

Medit. Mar. Sci., 12/1, 2011, 95-120 111

TSIAMIS, 2009; KATSANEVAKIS et al.,2011), and its population explosion along thecoasts of Rodos in 2010 (M. Corsini-Foka,pers. obs.), Percnon gibbesi (H. Milne Ed-wards, 1853) was found at 10 sites in SW, W,

and NW Kriti during summer and autumn2010 (D. Poursanidis, pers. obs.) and maythus be considered as established along theentire Cretan coastline; it was also found inPesada, S. Kefallonia Island, Ionian Sea in

Medit. Mar. Sci., 12/1, 2011, 95-120112

Fig. 2 (·-d): Distribution of some of the most invasive species. Asterisks indicate findings after ZENETOSet al. (2009).

·. b.

c. d.

July 2010 (A. Panou, pers. comm,). One ofus (G. Apostolopoulos) has observed andphotographed P. gibbesi along the north Kastel-lorizo coast repeatedly since 2004 (Fig. 2d).

Figure 3 shows aliens’ zoogeographi-cal patterns in Greek waters. The highestnumber (93 species) was reported in theDodekanisos area, SE Aegean, which tes-tifies to the importance of the area as theentrance point of Lessepsian immigrantsspreading towards the Aegean Sea. Par-ticular notice should be given to the 86species reported in the wider area of theSaronikos Gulf, a hotspot area for bioin-vasions in Greek waters, closely relatedto the route of ships towards Peiraias, thebiggest Greek port. Relatively high (40species) is also the number of alien speciesin the Thermaikos Gulf (related to the portof Thessaloniki). Many of the species en-

countered in Saronikos and/or ThermaikosGulfs present a limited distribution in theaforementioned areas.

Conclusions

> Following the review by ZENETOS etal. (2009a) on marine alien species inGreek Seas, 47 additional species are re-ported herewith, bringing the total to 237,which is a 24.4% increase. Twenty one ofthe listed species were reported for thefirst time in 2009-2010, whereas 21 species(mostly Polychaeta) although they ex-isted in the literature, had never been clas-sified as aliens previously.> The species Hypselodoris infucata, Den-

drostrea frons, Septifer forskali and Rhizo-prionodon acutus are reported in this workfor the first time in Greek waters.

Medit. Mar. Sci., 12/1, 2011, 95-120 113

Fig. 3: Pattern of alien species distribution in Greek waters.

> Four species reported in previous lists arenow excluded. These are: the rhodophyteNeosiphonia sphaerocarpa, the bivalveCircenita callipyga, and the fish Alopiassuperciliosus and Gaidropsarus granti.> Based on molecular analysis, we confirm

the occurrence of Bulla arabica in theMediterranean Sea and support the sug-gestion by MALAQUIAS & REID (2008)that previous records of Bulla ampulla inthe Mediterranean should be consideredas a misidentification of B. arabica.> In 2010, ten species previously known as

casual and/or questionable immigrants,have established viable populations andspread to many localities in the Greekcoasts. To-date, of the 237 alien speciesreported in Greek waters, 127 are estab-lished; 59 are casual records; 20 are cryp-togenic and 31 are questionable records.> The majority of alien species belong to

Mollusca (47 species: 19.8%), followedby fish (41 species: 17.3%), Polychaeta(38 species: 16.0%), macroalgae (33 species:13.9%), and Crustacea (31 species: 13.1%).>Most of the newly introduced and/or re-

ported species originate in tropical sub-tropical areas of the Pacific, Indian or At-lantic Oceans.> The increased rate of introductions of

warm water species confirms the previ-ous findings linking the rate of introduc-tion in the eastern Mediterranean toclimate change.> Although many of the newly reported

species are Lessepsian immigrants spread-ing in the Aegean Sea, the role of ship-ping is gaining significance as a vector ofalien species’ transfer in the Aegean Sea.

Ackowledgements

The authors would like to express theirthanks to all ELNAIS contributors. Special

thanks are due to Sofia Galinou-Mitsoudi(Alexander Technological Educational In-stitute of Thessaloniki); Nomiki Simboura(HCMR, Anavissos); Antonella Pancucci-Papadopoulou (HCMR, Anavissos); Eu-genia Lefkaditou (HCMR, Agios Kosmas);Yiannis Issaris (HCMR, Agios Kosmas);Mariolina Corsini-Foka (HCMR, Rodos)and Panayotis Ovalis (Athens) for provid-ing collection details for some of the newspecies. We also thank Kimon Moschan-dreou, Katerina Aligizaki (Aristotle Univer-sity of Thessaloniki), Sofia Spatharis (Univer-sity of the Aegean, Mytilene, Greece), Rai-mondo Villa (Italy) and Francis Kerckhof(Royal Belgian Institute of Natural Sciences)for profitable correspondence and exchangeof information. Sincere thanks are due toTassos Eleftheriou (Kriti) for helpful anduseful comments that improved significantlythe manuscript.

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Marine alien species in Greek Seas: Additions and amendments by 2010

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