67Medit. Mar. Sci., 8/1, 2007, 67-89

Introduction

The Mediterranean is one of the mostimportant enclosed seas in the world witha large shipping circulation but narrowcommunication with two oceans by theGibraltar strait (Atlantic Ocean) and bythe man-made Suez Canal (Indian Oceanthrough the Red Sea). It also has a freewater exchange with its annexed Black Seathrough the Dardanelles, the Sea of Mar-mara and the Bosporus, all of which, thatin spite of marked particularities are inreality annexed parts of the Mediter-

ranean. This whole frame of marine andbrackish waters conveys a large variety ofbiota, which enter the Mediterranean con-tinuously and include organisms frequent-ly described as ‘allochthonous’, ‘exotic’,‘alien’, ‘introduced’ by man or ‘vagrantvisitors’. According to dependable ecolog-ical parameters they have settled in theMediterranean for a few or a large num-ber of years, temporarily or permanently,with large or scarce populations, followingtheir ecological requirements and theshort or long term environmental oscilla-tions. Therefore, the Mediterranean is by

Mediterranean Marine ScienceVolume 8/1, 2007, 67-89

Allochthonous and vagrant ichthyofauna in Hellenic marine and estuarine waters

M. CORSINI-FOKA1 and P. S. ECONOMIDIS2

1 Hellenic Center for Marine Research,Hydrobiological Station of Rhodes, Cos Street, 85100 Rhodes, Hellas

2 Karakasi Street, 79, 54453 Thessaloniki, Hellas

e-mail: mcorsini-foka@hsr-ncmr.gr

Abstract

A review of the non-indigenous ichthyofauna occurring in the Hellenic marine and estuarine watersis presented, including Atlantic origin colonizers, aquaculture introduced and Lessepsian alien species. 34non-native species have been registered. The majority of allochthonous fish are Lessepsian immigrants,which represent approximately 80% of the ascertained non-native ichthyofauna in Hellenic waters; theirestablishment, spread, habitat, abundance and interaction with indigenous fish are discussed.

Keywords: Allochthonous fish species; Alien fish species; Greece; Ionian; Aegean; Mediterranean;Lessepsian migration.

Review Article

far the major recipient of exotic speciesand vagrant visitors like macrophytes,invertebrates and fish. Apart from therather free energetic movement throughthe above passages, ships’ ballast, fouling,clinging of ship hulls and aquaculture arealso important vectors of non-indigenousmarine species. Since the last glacial peri-od, it seems the Mediterranean has pre-sented a continuous exchange of faunawith the Atlantic and the Black Sea. Dur-ing the last 150 years since the opening ofthe Suez Canal, it has been presentingmany Indo-Pacific species, which migratethrough it. The rate of this so-calledLessepsian migration has increased in thelast decades, with ecological and economicimpacts, mainly in the Levantine basin.

The Mediterranean Ichthyofauna

In a rather simplified scheme, whichcould also be adapted and used for theHellenic seas, recent Mediterraneanichthyofauna is composed of four maingroups of species: (1) endemic(QUIGNARD & TOMASINI, 2000), (2)Atlantic, Boreal and Tropical (sensuTORTONESE, 1960, 1964, 1967, 1970;WHITEHEAD et al., 1984-86). TheseAtlanto-Mediterranean species occur ornot in the Mediterranean according toenvironmental factors, many of whichoscillate significantly depending on longor short term climatic changes. Altogetherthese species are called ‘autochthonous’.Several such species, with rare or periodicappearance, could be described as‘vagrant’ or ‘visitors’ but not ‘alien’ or‘exotic’, even if they are only present inlimited concentration and many of themare, in fact, detected with difficulty. Otherscould also be called ‘new colonizers’, par-ticularly when their spread is favoured by

special circumstances such as the tendencytowards a change in temperature and cur-rent regime in the NE Atlantic. (3) Intro-duced by man (sensu HUNTER &GIBBS, 2006) mainly for aquaculture oraquarium purposes, and which escape intothe wild or are carelessly released. This is arather obscure phenomenon and actuallythere is no or only very little elementaryknowledge of its impact on the naturalenvironment and genetic compatibilityafter mating with closely-related wildspecies. In this category should also beincluded the release with the ballast waterof various taxa, many of which maybe‘exotic’, and (4) the Lessepsian or Erythre-an, of Red Sea origin immigrants, whichare real ‘exotic’ or ‘alien’ species. In sev-eral cases, when a population explosionhas occurred, these last could be charac-terized as really ‘invasive’ (HUNTER &GIBBS, 2006), as in the case of the recentcolonizer Fistularia commersonii in manyareas of the Eastern Mediterranean.

Greek Allochthonous fish species

Among a number of about 500 nativemarine fish species in the Hellenic waters,34 allochthonous have been registered,while seven others are not yet confirmedfor various reasons (Table 1). Among thelast, six species belong to the category ofintroduced for aquaculture and their pres-ence in nature is based on unverifiedobservation of free swimming specimensor unidentified samples, strongly suggest-ing careless release or escape from fishfarms. From the remainder, the majority(27 species plus one uncertain, i.e. 79.4%)originate from the Indo-Pacific Ocean andthe Red Sea having penetrated into theMediterranean via the Suez Canal. Amongthe other seven species, four are usually

68 Medit. Mar. Sci., 8/1, 2007, 67-89

69Medit. Mar. Sci., 8/1, 2007, 67-89

Tab

le 1

Lis

t of

all

ocht

hono

us f

ishe

s in

Hel

leni

c w

ater

s.

O: O

rigi

n (I

P: I

ndo-

Pac

ific,

A: A

tlant

ic, A

q: a

quac

ultu

re).

ES:

Est

ablis

hmen

t suc

cess

(E

: Est

ablis

hed,

S: S

tock

ing,

C

: Cas

ual,

Q: Q

uest

iona

ble)

, FR

: Reg

ion

of fi

rst r

ecor

d.B

P: B

enth

opel

agic

, P: p

elag

ic, D

: dem

ersa

l, R

A: r

eef a

ssoc

iate

d, D

W: d

eep

wat

ers,

EP

: epi

pela

gic,

TE

: tem

pera

te, S

T: s

ubtr

opic

al,

T: t

ropi

cal,

F: f

ish,

BI:

ben

thic

inve

rteb

rate

s, I

: inv

erte

brat

es, C

: cru

stac

eans

, Z: z

oopl

ankt

on, M

: mol

lusc

s, G

A: g

reen

alg

ae,

Ph:

phy

topl

ankt

on, P

: pla

nkto

n, O

: om

nivo

rous

, H: h

erbi

voro

us.

Fish

OES

FRRe

feren

ces*

Wor

ld ra

nge*

*Ec

ology

**Cl

imat

e**

Food

**Ac

ipens

er gü

ldens

tädtii

AqC

Evro

s estu

aries

[1]

Blac

k and

Cas

pian s

eas e

stuar

ies an

d rive

rsBP

, san

dy an

d mud

dyTE

F, B

I

Acipe

nser

stella

tus1

AqC

Evro

s estu

aries

[1]

Blac

k & C

aspia

n sea

s dra

inage

, nati

ve in

Evro

sBP

, san

dy an

d mud

dyTE

F, B

I

Acipe

nser

sturio

1Aq

QEv

ros e

stuar

ies[1]

Eu

rope

an es

tuar

ies, n

ative

in E

vros

BP, s

andy

and m

uddy

TEF,

BI

Alep

es dje

daba

2IP

CAe

gean

Sea

[2]

Indo

-Pac

ific,

Red S

ea, E

. Afri

caP,

insh

ore

STC,

F

Angu

illa ja

ponic

a ?Aq

QIo

nian S

eaPr

esen

t wor

kPa

cific

D, es

tuar

iesST

F, C

, inse

ct

Apog

on ph

arao

nisIP

ERh

odes

[3]

Indo

-Pac

ific.

Red S

ea, E

. Afri

ca co

asts

to A

ustra

liaD,

noctu

rnal,

rock

yT

Z

Athe

rinom

orus

lacu

nosu

sIP

CRh

odes

[4] [5

] W

ide In

do-P

acifi

c, Re

d Sea

RA, c

oasta

l, lag

oons

STZ

Callio

nymu

s fila

mento

sus

IPC

Rhod

es[6]

In

do-P

acifi

c, Re

d Sea

B, sa

ndy a

nd m

uddy

STBI

Ench

elyco

re an

atina

AC

Elafo

nisso

s[7]

Ea

stern

Atla

ntic

D, pe

lagic

eggs

STF,

C

Etru

meus

teres

IPE

Rhod

es, C

yclad

es[6]

[8]

Red S

ea to

E. A

frica

, Ind

ian O

cean

to A

ustra

liaP,

insh

ore

STZ

Fistu

laria

comm

erson

iiIP

ERh

odes

[9]

Wide

Indo

-Pac

ific,

Cent

ral a

nd So

uth A

meric

aRA

TF,

C, M

Gaidr

opsa

rus g

ranti

AQ

Rhod

es[10

] At

lantic

DWT

Palae

monid

praw

ns

Hemi

ramp

hus f

arIP

ERh

odes

[11] [

12] [

13]

Wide

Indo

-Pac

ific,

Red S

eaEP

, insh

ore

STZ,

GA,

Ph

Huso

huso

AqS

Evro

s estu

aries

[1]

Blac

k and

Cas

pian s

eas d

raina

geBP

TEF

Iniis

tius p

avo

IPC

Rhod

es[14

] W

ide In

do-P

acifi

c, Re

d Sea

inclu

ded,

and E

. Pac

ific

RAT

M, C

Lago

ceph

alus s

celer

atus

IPE

Rhod

es[14

]In

do-P

acifi

cB

(main

ly), o

n san

dy

TBI

Lago

ceph

alus s

padic

eus

IPE

Samo

s[15

] W

ide In

do-P

acifi

c, Re

d Sea

BPST

BI

Lago

ceph

alus s

uezen

sisIP

ERh

odes

[6]

Red S

ea en

demi

cB,

sand

y and

mud

dyTE

BI

Leiog

nathu

s klun

zinge

riIP

ERh

odes

[16]

Red S

eaD

TBI

Liza

carin

ata ?

AqQ

Amvra

kikos

Pres

ent w

ork

Wes

tern I

ndian

, Red

Sea

Coas

tal an

d bra

ckish

wate

rT

BI, P

Liza

haem

atoch

eila

AqE

Thra

cian S

ea[17

] Fa

r eas

tern A

sia fr

om R

ussia

to C

hina

Coas

tal, e

stuar

ies, f

resh

water

TEO

(con

tinu

ed)

70 Medit. Mar. Sci., 8/1, 2007, 67-89

Fish

OES

FRRe

feren

ces*

Wor

ld ra

nge*

*Ec

ology

**Cl

imat

e**

Food

**

Micr

opter

us sa

lmoid

esAq

QIo

nian S

eaPr

esen

t wor

kNo

rth A

meric

aFr

eshw

ater (

river

s and

Atla

ntic

drain

ages

)TE

F, B

I

Pagru

s majo

r ?Aq

Q?

Pres

ent w

ork

North

west

Pacif

icD,

ocea

nodr

omou

sST

BI

Parex

ocoe

tus m

ento

IPE

Rhod

es[16

] W

ide In

do-P

acifi

c fro

m Re

d Sea

to F

ijiEP

, insh

ore

TZ

Pemp

heris

vanic

olens

isIP

EKa

stello

rizon

[18]

Wide

Indo

-Pac

ific,

Red S

eaDa

ytime

in ca

ves,

night

insh

ore p

elagic

TPC

Petro

scirte

s anc

ylodo

nIP

CRh

odes

[6]

Red S

ea to

Ara

bian G

ulfD,

sand

y sho

reT

BI

Ptera

gogu

s pely

cus

IPE

Symi

[19]

Red S

ea to

E. A

frica

D, co

astal

with

sea g

rass

STBI

Sargo

centr

on ru

brum

IPE

Rhod

es[20

] In

do-P

acifi

c (E.

Afri

ca to

Samo

a and

Japa

n)No

cturn

al, da

ytime

in ca

ves

STF,

BI

Saur

ida un

dosq

uami

sIP

ENa

xos

[21]

Indo

-Pac

ific t

o Aus

tralia

and S

. Jap

anD,

sand

y or m

uddy

ST

F, C

, BI

Serio

la fas

ciata

AC

Rhod

es[14

] Ea

stern

a and

Wes

tern A

tlant

icYo

ung E

P, ad

ult B

PST

F, M

Sigan

us lu

ridus

IPE

Tilos

[22]

Red S

ea, E

. Afri

ca to

Ara

bian G

ulfRA

, roc

ky or

hard

with

vege

tation

STH

Sigan

us ri

vulat

usIP

ERh

odes

[23]

Red S

ea an

d Gulf

of A

den

RA, s

andy

, with

alga

e and

seag

rass

STH

Spho

eroide

s pac

hyga

ster

AE

Rhod

es[24

] At

lantic

, Ind

ian oc

eans

D, m

uddy

, san

dy an

d roc

ky bo

ttoms

STM

Sphy

raen

a chr

ysotae

nia (p

inguis

)IP

ERh

odes

[19]

Indo

-Pac

ific t

o Chin

a and

N. A

ustra

liaBP

, coa

stal

STF,

C

Sphy

raen

a flav

icaud

a (ob

tusata

)IP

CRh

odes

[6]

Wide

Indo

-Pac

ific,

Red S

ea, E

. Afri

caEP

, insh

ore

TF,

I

Steph

anole

pis di

aspr

osIP

ERh

odes

[11] [

12] [

13]

Red S

ea to

the A

rabia

n Gulf

D, ro

cky s

ubstr

ate w

ith ve

getat

ion.

TBI

Torqu

igene

r flav

imac

ulosu

sIP

CRh

odes

[25]

Red S

ea, A

rabia

n Gulf

, E. A

frica

, Sey

chell

esRA

, sha

llow

sand

y sho

res w

ith se

agra

ssT

BI

Tyler

ius sp

inosis

simus

IPC

Rhod

es[6]

In

do-W

est P

acifi

c, So

uthe

ast A

tlant

icDW

T

Tylos

yrus c

roco

dilus

IPC

Chalk

idiki

[26]

Wide

Indo

-Pac

ific

EP, c

oasta

lT

F

Upen

eus m

olucc

ensis

IPE

Rhod

es[27

] [28

] In

do-P

acifi

cB,

sand

y or m

uddy

ST

BI, F

Upen

eus p

ori

IPC

Rhod

es[6]

Re

d Sea

, Gulf

of O

man

B, sa

ndy a

nd m

uddy

STBI

(1):

As t

he sp

ecie

s is c

onsid

ered

as r

are

or e

xtin

ct fr

om it

s nat

ive

rang

e in

Evr

os d

rain

age,

the

new

reco

rds m

aybe

con

cern

spec

imen

s fro

m st

ocki

ng (s

ee K

OU

TRA

KIS

& E

CO

NO

MID

IS, 2

006)

.(2

): T

he o

ccur

renc

e of

Alep

es d

jedab

ain

Gre

ek w

ater

s is t

o be

asc

erta

ined

.*

[1] K

OU

TRA

KIS

& E

CO

NO

MID

IS, 2

006;

[2] B

INI,

1960

; [3]

CO

RSI

NI F

OK

A e

t al.,

200

4; [4

] QU

IGN

AR

D &

PR

AS,

198

6; [5

] CO

RSI

NI,

2004

, new

dat

a; [6

] CO

RSI

NI e

t al.,

2005

; [7]

GO

LAN

I et a

l.,20

02; [

8] K

ALL

IAN

IOTI

S &

LEK

KA

S, 2

005;

[9] C

OR

SIN

I et a

l.,20

02; [

10] Z

AC

HA

RIO

U-M

AM

ALI

NG

A, 1

999;

[11]

TO

RTO

NES

E, 1

946,

[12]

TO

RTO

NES

E, 1

947a

; [13

]TO

RTO

NES

E, 1

947

b; [1

4] C

OR

SIN

I et a

l.,20

06; [

15] A

NA

NIA

DIS

, 195

2; [1

6] K

OSS

WIG

, 195

0; [1

7] K

OU

TRA

KIS

& E

CO

NO

MID

IS, 2

000;

[18]

PA

PAC

ON

STA

NTI

NO

U &

CA

RA

GIT

SOU

,19

87; [

19] C

OR

SIN

I & E

CO

NO

MID

IS, 1

999;

[20]

LA

SKA

RID

IS, 1

948a

; [21

] ON

DR

IAS,

197

1; [2

2] K

AV

ALA

KIS

, 196

8; [2

3] B

RU

NEL

LI &

BIN

I, 19

34; [

24] Z

AC

HA

RIO

U-M

AM

ALI

NG

A &

CO

RSI

NI,

1994

; [25

] CO

RSI

NI F

OK

A e

t al,

2006

; [26

] SIN

IS, 2

005;

[27]

SER

BETI

S, 1

947;

[28]

LA

SKA

RID

IS, 1

948b

.**

Bas

ed o

n: G

OLA

NI e

t al.,

200

6 on

-line

; FR

OES

E &

PA

ULY

, 200

6

Tab

le 1

(co

ntin

ued)

considered to originate from the AtlanticOcean and three, two sturgeons and onemullet, come from aquaculture eitherfrom stocking or escape.

All these allochthonous species can beclassified in three main groups accordingto their origin: (a) Atlantic, (b) Aquacul-ture and (c) Indo-Pacific. The first groupof species could eventually be includedmore easily in the indigenous Mediter-ranean species, according to the aboveexpressed points. The list of species com-ing from aquaculture is expected to grad-ually increase, as the number of aquacul-ture installations is increasing in the Hel-lenic seas and in the neighbouring areas aswell. The Indo-Pacific group includes themajority of allochthonous species, whichare real alien species in the Mediter-ranean as they have penetrated an areaout of their original distribution. It isworth noting that the majority of thesespecies have quite a slow process ofspread and settlement, underlining theirthermophilous and stenohaline, tropicalor sub-tropical, character. In the Mediter-ranean such environmental conditions arecurrent in the eastern basin, mainly alongthe Asian coasts. In the Aegean Sea aclosely related situation occurs in the SEcorner around the Dodecanese continen-tal plateau. There is evidence that the con-tribution of the Indo-Pacific alien speciesto the Aegean Sea fish species composi-tion and ecosystem structure, is graduallyaccelerating, as many populations of thesespecies have already constructed commer-cial stocks, as did long ago the Upeneusand Siganus species in the Dodecaneseislands and more recently Fistularia com-mersonii (CORSINI et al., 2002;KALOGIROU et al., 2007) and Etrumeusteres in the Cyclades (KALLIANIOTIS &LEKKAS, 2005) and in the Dodecanese

as well (CORSINI-FOKA, unpubl. data).

Atlantic vagrant and new colonizer

As aforementioned, the vagrant or vis-itor and the new colonizer fish species ofAtlantic origin are not easy to enumeratebecause their flux has been continuing formore than 10,000 years. Various attempts,however, have been made at standardiza-tion of their origin and at the calculation ofthe number of Mediterranean fish species(TORTONESE, 1938, 1963, 1970, 1975,1982; QUIGNARD & TOMASINI,2000). Furthermore, as research in variouspart of the Mediterranean becomes moreintensive, many new species are registeredeach year, mainly in the western part of thesea, which is closest to the Atlantic. Thiscould also be attributed to other reasons,such as the warming trend of the Mediter-ranean, the expansion of maritime traffic,etc. (GOLANI et al., 2002) or combinationof all these reasons. There is widespreaddiscussion regarding the exact number ofAtlantic vagrant and new colonizer fishspecies (QUIGNARD & TOMASINI,2000). Recent estimations (up to 2006)have considered those species enteringthrough the Gibraltar straits and crossinginto the Mediterranean to be over 30(GOLANI et al., 2006 on-line; REINA-HERVA’ S et al., 2004), also reaching theAegean (ERYILMAZ et al., 2003;CORSINI et al., 2006) and the AdriaticSea as with the recently recordedCyclopterus lumpus (DUL

v

CIv

C &GOLANI, 2006). Some of them, particu-larly Sphoeroides pachygaster, Seriola fasci-ata and Seriola carpenteri have achievedsignificant biomass (ANDALORO, 2001;ANDALORO et al., 2005).

Three such species have been record-ed in the Aegean: Sphoeroides pachygaster,

71Medit. Mar. Sci., 8/1, 2007, 67-89

a species well established throughout theMediterranean, Enchelycore anatina andSeriola fasciata, while the consideration ofGaidropsarus granti as a non-indigenousAtlantic colonizer is questionable (Table 1).

The regular or rare appearance ofAtlantic species in the Mediterranean is amore complicated phenomenon, depend-ing mainly on hydro-climatic long-termchanges and short-term trophic relationsamong species. In a very preliminary andsimplified approach, when North Atlanticwaters become warmer, there is an accessof temperate, even sub-tropical, speciesinto the Mediterranean. On contrary,when Atlantic waters become colder,northern species enter and colonize theMediterranean. This play seems to havetaken place repeatedly during the glacierperiod dominating in the northern hemi-sphere of the planet during the last onemillion years. The inter-glacier periodfavours the first while the glacier the sec-ond movement (see also QUIGNARD &TOMASINI, 2000). Actually, there is arather clear range of these warm water or‘thermophilous’ and cold water or ‘psy-chrophilous’ fish fauna in the Mediter-ranean, as the first is distributed in thesouth mainly along the African and Asiancoasts, while the second is restricted to theEuropean coasts in areas where cold waterdominates (North Adriatic, NorthAegean, Black Sea, etc.), following theactual temperature and current regime.

Active movement according to trophicavailability concerns the errant largepelagic species such as sharks, variousscombrids, tetrapterids, swordfish, etc.Several such species are season regular orrare vagrant visitors. In some other cases,such as the sea lamprey (Petromyzon mari-nus), a parasitic mode of life forces it tomove far away from its normal distribu-

tion. Therefore, the recording of this lastspecies in the North Aegean Sea(ECONOMIDIS et al., 1999), where thereare no spawning grounds for permanentsettlement, could be qualified as acciden-tal and the species as ‘vagrant’ and not asexotic or as alien, because the area is with-in its large range. A similar example wouldbe any species of remoras, accompanyinglarge marine animals, such as the Remile-gia australis (= Remora australis), noticedlong ago in Hellenic waters(TORTONESE, 1946, 1947a), and which,according to GOLANI et al. (2002),although very rare could be considered aMediterranean resident.

In several other cases, such as theAtlantic muraenid Enchelycore anatina,spread from the Atlantic to the Mediter-ranean, where there are only two recordsone of which being from Elafonissos(Peloponnesus coasts) (GOLANI et al.,2002), could be explained by passive dis-semination of the leptocephali larvaeand/or via transport in ship ballast.

Aquaculture introduced

Uncontrolled and/or collapsed aqua-cultures are responsible for the releasingof specimens into nature from their rearedstocks. Some few cases have already beenreported in the Greek seas by picturedand/or preserved specimen, but informa-tion has not been yet published. One ofthese may concern some mullet specimens,apparently Liza carinata, captured in west-ern Greek estuaries (Table 1). Such aspecies was introduced long ago as fryfrom Egypt to a fish farm near Arta (riverArachthos estuary in the AmvrakikosGulf), which subsequently closed down(unpublished data). However, Liza carina-ta is listed among Indo-Pacific aliens

72 Medit. Mar. Sci., 8/1, 2007, 67-89

(GOLANI et al., 2002), since it has alsobeen recorded in the Levantine basin,including the Turkish Mediterranean andAegean coasts (TORCU & MATER,2000; ZAITSEV & ÖZTÜRK, 2001;BILECENOGLU et al., 2002). Conse-quently, its occurrence could be also dueto the expansion of this lessepsian immi-grant distribution from the EasternMediterranean to the Ionian Greek waterspreviously mentioned.

It is known that around Greek andother European estuaries, among cap-tured growing European eel (Anguillaanguilla) there also are some specimens ofthe elvers of the Japanese eel (Anguillajaponica), species largely used in fishfarms, which have escaped from eel farms(unpublished data). Another case con-cerns the largemouth bass (Micropterussalmoides) that was observed in 2003 andphotographed near the estuaries of thestream Lessini, flowing directly into theIonian Sea (unpublished data) (Table 1).Recent records concerning several speciesof sturgeons such as Acipenser güldenstän-di, Huso huso or various rearing hybrids, aswell as Acipenser sturio, captured in theThracian Sea, near the Evros estuaries orin the river itself, strongly suggest that theyoriginate from the release of fry orescapees from hatcheries lying in theEvros river catchment of Bulgaria and/orof Turkey, as there are no such installa-tions in the Hellenic part of the river(KOUTRAKIS & ECONOMIDIS, 2006).Therefore, some previous records ofAcipenser stellatus (ECONOMIDIS et al.,2000) maybe have had the same origin.Furthermore, there is unverified informa-tion that certain alien fish species (forinstance Pagrus major) or even fry of nativeMediterranean fish, such as Dicentrarchuslabrax and Sparus auratus, originating from

distant areas (i.e. Spain), are used widelyin marine Greek fish farms. Consequently,very often these species of different genet-ic stocks are released or escape as fry oradults into the wild. But there is a con-firmed record of the Indo-Pacific nativemullet, Liza haematocheilus (Mugil soiuy),which was reared in Black Sea cage fishfarms. Many of these farms collapsed inthe nineties and specimens were carelesslyreleased in the Azov and Black Sea areas(STARUSHENKO & KAZANSKY,1996). Soon after, the species was observedin the Turkish Aegean Sea (Gulf of Smyr-na: KAYA et al., 1998) and in the ThracianSea (KOUTRAKIS & ECONOMIDIS,2000) (see also HARRISON, 2004). Actu-ally, it forms fished population in variousThracian and Macedonia lagoons and it iscaptured and appears frequently in the fishmarkets of North Greece. (After MINOSet al., 2007 and in press).

Lessepsian Aliens

The movement of various marineorganisms from the Red Sea to theMediterranean via the Suez Canal wasnamed Lessepsian migration by POR(1969), a term widely used in literaturethereafter; it is very often also called Ery-threan migration. According to GALIL &ZENETOS (2002), the Eastern Mediter-ranean, open to Atlantic, Pontic and Ery-threan biota, is particularly prone to inva-sion. Concerning fish, the Lessepsianspecies in the Mediterranean are real alienspecies because they belong to anotherecological status (thermophilous) and to adifferent bio-geographical zone (Indo-Pacific), and the Mediterranean is out oftheir native distribution.

In recent years intensive research hasbrought to light more alien species which

73Medit. Mar. Sci., 8/1, 2007, 67-89

might have been present in low densitiesand thus escaped our attention(CORSINI & ECONOMIDIS, 1999).The immigration of such alien species isfar from ceasing after approaching a stan-dard level, since in practice the flux ofinvaders is continuous (STREFTARIS etal., 2005). The dynamics of invasions andextension of the distribution of alienspecies obviously need a continuousupdate and a monitoring procedure(GOLANI et al., 2004; ZENETOS et al.,2005; PANCUCCI-PAPADOPOULOUet al., 2005a, b).

Establishment. According to the morerecent inventories the Lessepsian fishmigrants in the Mediterranean are oneElasmobranchii species and more than 60Teleostei species (GOLANI et al., 2006on-line; GOKOGLU et al., 2003;CORSINI et al., 2005; AKYOL et al., 2005;CORSINI et al., 2006; BILECENOGLU& KAYA, 2006; CINAR et al., 2006;GOLANI, 2006; ERYLMAZ &DALYAN, 2006; GOLANI & SONIN,2006). Once arrived in the MediterraneanSea through the Suez Canal, these speciesmove either westwards to the African

74

Fig. 1: The Dodecanese islands and the Hellenic Seas.

Medit. Mar. Sci., 8/1, 2007, 67-89

coasts or usually eastwards first and thennorthwards, following mainly the Asiaticcontinental shelf. They normally establish,fast or slow, dense or scarce populations,in the Levantine Sea where they influenceor even modify essentially the compositionand the structure of the local marineecosystems. The common way for furthercolonization of the Mediterranean is main-ly by using the South coast of Turkey andits narrow continental shelf. Besides, thewater masses around the DodecaneseIslands form the most important gates forentering and establishing in the AegeanSea and the main pathway of spread with-in the Mediterranean (PANCUCCI-PAPADOPOULOU et al., 2005a, b) (Fig.1). A number of 28 Lessepsian immigrantfish species has entered Greek waters(Table 1). Three additional ones havebeen reported from the SE Aegean Sea,along the Turkish coasts, Scomberomoruscommerson (BUHAN et al., 1997) andOxyurichthys petersi (AKYOL et al., 2006)in Gokova Bay and Sillago sihama in Datçapeninsula (BILECENOGLU, 2004).Older records of Lessepsian fish inHellenic waters amounted tothirteen species until 1990(PAPACONSTANTINOU, 1987, 1990;also References in Table 1). An evidentincrease in records has been observed inthe last decade, when 15 other such specieswere added. A similar picture is presentedby the Erythrean crustaceans, which sud-denly occurred in the SE Aegean in thepast decade and, according to GALIL &KEVREKIDIS (2002), this may be due tothe more extensive inflow of the AsiaMinor Current in the area.

CORSINI & ECONOMIDIS (1999)mention that several Lessepsian specieswere reported in the Dodecanese almostsimultaneously as along the coasts of

Israel. Such are the cases of Siganus rivula-tus, Upeneus moluccensis, Sargocentronrubrum, Pteragogus pelycus and Fistulariacommersonii. On the contrary, as for exam-ple the cases of Sphyraena chrysotaenia,Apogon pharaonis, Etrumeus teres andsome others, the advance along the Anato-lian coasts was gradual and the immigrantsreached this area after a relatively longtime, depending on biotic and abiotic fac-tors (GOLANI, 1998a; CORSINI &ECONOMIDIS, 1999; CORSINI et al.,2002, 2005; CORSINI FOKA et al., 2004)(Table 1). There is a direct relationbetween the environmental conditions andthe speed of spread and adaptation of thenew settler. In fact, species have to face acombination of unfavourable ecologicalconditions (water temperature, substra-tum, currents, trophic conditions etc.)when following the narrow continentalshelf of the South Anatolian coast as themain way for westward spread, as themajority of species are coastal. Conse-quently, once they reach the continentalshelf of the Dodecanese islands, they donot usually advance any further; some ofthem are also commercially exploited.

On the other hand, a large number ofLessepsian immigrants, being quite com-mon in the Levantine, have not beenrecorded in the SE Aegean Sea. This dif-ferent trend could be owing to many bioticand/or abiotic factors as, for example, thetemperature regime and the thermal toler-ance of the colonizing species, food avail-ability, competition with indigenousspecies, local pathogens and the suitablesubstrate (GOLANI, 1998a). The exten-sion of the spawning season, the aggressivebehaviour of the new species, their foodpreferences, could also be added. Coloniz-ers may also occupy free niches alreadyexisting or created by over-fishing of local

75Medit. Mar. Sci., 8/1, 2007, 67-89

species. All these, like many other factors,need a more accurate survey to obtain acomplete view of the real situation.

In comparison, along the Turkish terri-torial waters of the Aegean and the EastMediterranean, 38 species of teleosts ofRed Sea origin have been recorded(GOLANI et al., 2006 on-line; BILECE-NOGLU et al., 2002; CINAR et al., 2005,2006; BILECENOGLU & KAYA, 2006).

A number of 25 species is common inthe Hellenic and the Turkish waters.

It is obvious that several fish speciesalready detected and established along theLevantine Turkish coasts can be expectedalso in Hellenic waters, especially Dussum-ieria elopsoides, Herklotsichthys punctatus,Pelates quadrilineatus, Cynoglossussinusarabici, Liza carinata.

Spreading. Lessepsian migrant speciesonce having reached the Dodecanese con-tinental shelf seem to have difficulties infurther spread to the rest of the AegeanSea, mainly to the north, or in continuingtheir expansion to all the Mediterranean.Among the above 28 Lessepsian fishspecies (Table 1), only nine have beenrecorded further north than the Dode-canese plateau, five before the last decadeand the other four during the last sevenyears (Table 2). The recording of the Tylo-surus crocodilus in the Chalkidiki peninsu-la, based on a dead fish found on Gerakinibeach (SINIS, 2005), needs a special men-tion as it is a recording which merits fur-ther investigation. This is a first recordingin the Mediterranean and its derivation asa rejected specimen from undelivered for-eign stock should also be considered.

Spreading to the south, central or west-ern Greek waters of the Aegean and theIonian seas seems to have the same low suc-cess, as only fourteen and six species respec-tively followed these routes (Table 2).

This statement shows that some abiot-ic (water temperature, salinity, currentregime, size of continental shelf, etc.)and/or biotic factors (food quality andquantity, spawning facilities, etc.) areimpeding the spread of Erythrean fish.

In general, the species which havespread out of the Dodecanese ‘refuge’(Table 2) have a wide native distributionarea (Table 1). Although tropical or sub-tropical fish, they are able to colonize tem-perate regions quite easily, the water tem-perature not prohibiting their spread.

Although the Cretan continental shelflies in the south Aegean Sea in a ratherwarm water zone, colonization by Lessep-sian immigrants presents some interestingparticularities. Since these species areregarded as thermophilous, it would beexpected that they would settle more easi-ly in the coastal waters of Crete. However,unfavourable factors obviously seem to bean interdiction for their large scale settle-ment there as only eleven fish species wererecorded, two before 2002, Stephanolepisdiaspros and Saurida undosquamis, andanother nine in the last four years: Siganusluridus, Fistularia commersonii, Siganusrivulatus, Pempheris vanicolensis, Sargocen-tron rubrum and recently, Lagocephalussceleratus, Etrumeus teres, Upeneus moluc-censis and Pteragogus pelycus (Table 2).These new records are the result of recentmore intensive local investigations, but stillthe number of the species remains low.This phenomenon could be attributed tothe fact that although the continental shelfof Crete is accessible to Lessepsian immi-grants through the Kasos-Karpathos corri-dor, its colonization shows a rather slowprogress mainly because it is very narrowand delimited by exceptionally deepwaters, thus isolating the coastal zone ofCrete.

76 Medit. Mar. Sci., 8/1, 2007, 67-89

Several of the Erythrean fish are stillnot only distributed in the Levant Basin,since the populations of at least nine ofthem already established in the easternMediterranean, both along the Asiatic andAfrican coasts as well as the Greek seas,have expanded westward. According toGOLANI et al. (2002, 2004), in some casesthey have reached the Central Mediter-ranean, such as Siganus luridus, Siganusrivulatus, Stephanolepis diaspros. Siganusluridus has enlarged its distribution to thesouthern coasts of Sicily (AZZURRO &ANDALORO, 2004) and the northern

coasts of Sicily in the Tyrrhenian sea(CASTRIOTA & ANDALORO, 2005),as have also Stephanolepis diaspros, whichhas reached the Palermo coasts(CATALANO & ZAVA, 1993), Sphyrae-na chrysotaenia which has been reportedfrom Malta (LANFRANCO, 1993),Parexocoetus mento and Fistularia com-mersonii from Tunisia (BEN SOUSSI etal., 2004), the last species also from south-ern Sicily (AZZURRO et al., 2004), theSW shore of Sicily (MILAZZO et al.,2006) and Central Tyrrhenian waters(MICARELLI et al., 2006; PAIS et al.,

77Medit. Mar. Sci., 8/1, 2007, 67-89

Species North South Ionian References*Aegean Central Sea

AegeanEtrumeus teres + [1] [2] Fistularia commersonii + + [3] [4] [5] Lagocepalus sceleratus + + [6] [2] [17]Lagocephalus spadiceus + [7] Leiognathus klunzingeri + + [3] Parexocoetus mento + + [3] Pempheris vanicolensis + [5] Saurida undosquamis + + + [8][9] [3] [10] Siganus luridus + + + [9] [3] [5] Siganus rivulatus + + + [11] [3] [5] Stephanolepis diaspros + + [3] [12]Tylosurus crocodilus + [13] Upeneus moluccensis + + + [14] [15] [3] [16]Pteragogus pelycus + [18]Sargocentron rubrum + [19]

Table 2Spreading of Lessepsian fishes in the Aegean and Ionian seas,

out of the Dodecanese continental shelf.

* [1] Cyclades, Crete, KALLIANIOTIS & LEKKAS, 2005; [2] Crete, KASAPIDIS et al., 2007; [3] GOLANI et al., 2006 on-line; [4]Chalkidiki, KARACHLE et al., 2004; [5] Crete, TINGILIS et al., 2003; [6] Izmir, BILECENOGLU et al., 2006; [7] Samos,ANANIADIS, 1952; [8] ZAITSEV & ÖZTÜRK, 2001; [9] ONDRIAS, 1971; [10] Crete, TSIMENIDIS et al., 1991; [11] Izmir,GELDIAY, 1969; [12] Crete, ECONOMIDIS & BAUCHOT, 1976; [13] Thermaikos, SINIS, 2005; [14] Kusadasi, KAYA et al., 1999;[15] Patraikos, KASPIRIS, 1976; [16] various areas, PERISTERAKI et al., 2006; [17] Lesvos, KOUTSOUMBA, D., 2007 (pers.comm.); [18] Crete, STERIOTI, A., 2005 (pers. comm.); [19] Crete, TINGILIS, G., 2003 (pers. comm.).

2007). Some Lessepsian fish arefurthrmore colonizing the coasts of theAdriatic Sea, like Siganus rivulatus,Stephanolepis diaspros, Sphyraena chryso-taenia, Saurida undosquamis, Parexocoetusmento, Leiognathus klunzingeri and Hemi-ramphus far (DUL

v

CIv

C et al., 2003).In conclusion the westward spread of

Lessepsian species will proceed slowly, butmay accelerate if the increase in watertemperature continues. A rearrangementof the ecosystem structure is alreadyappearing in the eastern basin and thiscould be an important factor for fisherymanagement.

Habitat. At present most ErythreanIndo-Pacific alien invertebrates occupythe Mediterranean littoral and infralit-toral zones to a depth of approximately50m, and are hardly ever found in deeperwaters (GALIL & ZENETOS, 2002).This is also true for the Lessepsian fishspecies, which are in the majority coastalspecies dwelling in rather shallow, sandyor muddy habitats (GOLANI, 1993) orrocky shores, often covered by sea-grass.

In Rhodes and other Dodecanesecoastal areas the majority of Erythreanfish are collected by trawlers at depths ofup to 50m, more frequently on sandy-muddy bottoms covered by well-devel-oped Chlorophyceae beds (Caulerpa pro-lifera and Caulerpa racemosa) andPhanerogames prairies, mainly Posidoniaoceanica and Halophila stipulacea(CORSINI & ECONOMIDIS, 1999;CORSINI et al., 2005, 2006). Otherspecies are encountered on sandy-rockybottoms, such as Pempheris vanicolensis,Sargocentron rubrum and schools of juve-nile Shyraena chrysotaenia or in very shal-low sandy areas, as for Iniistius pavo. Onthe other hand, as assessed by CORSINI

et al. (2005), Tylerius spinosissimus wascollected at a lower depth than those it isfound in in its original tropical waters, butdeeper than that usual for Lessepsianmigrant fish in the SE Aegean Sea. Thisprobably shows that there may be severalunexplored niches suitable for the newcolonizers. The first finding of this youngpufferfish in Rhodes, far from the SuezCanal, indicates that a certain populationof the species is already established in theeastern Mediterranean, especially closerto the Suez Canal areas; its spread followsquite unusual paths, as observed in thecase of Tetrosomus gibbosus (SPANIER& GOREN, 1988). However, the possibil-ity that other vectors (ship ballast, aqua-culture or aquaria purposes transport),different from the natural pathway, maybe responsible for the introduction ofsuch Indo-Pacific species as the spinyblaasop Tylerius spinosissimus should notbe undervalued, in common with othertaxa also introduced in this way into Hel-lenic waters (PANCUCCI-PAPADOPOULOU et al., 2006).

Abundance and interactions. The abun-dance of some Lessepsian immigrant fishhas assumed economic importance in thesouth-eastern Levantine and Anatolianfisheries (GUCU et al., 1994; TORCU &MATER, 2000; GOLANI et al., 2002;CICEK & ASVAR, 2003; HARMELIN-VIVIEN et al., 2005), while other Erythtre-an invaders are considered as economicburden by GOREN & GALIL (2005).

Populations of Lessepsian fishthroughout the coastal waters of theDodecanese continental shelf generallycorrespond to species that have been pre-viously established there and they havebeen large enough to be detectable(CORSINI et al., 2005). This is confirmed

78 Medit. Mar. Sci., 8/1, 2007, 67-89

by the fact that such various species havenormally been present in fishery activitiesfrom their first recording in the regionuntil today (CORSINI FOKA et al., 2004).Among the established species listed inTable 1, the majority are caught regularly,but these species cannot be consideredabundant. They are: Hemiramphus far,Stephanolepis diaspros, Upeneus moluccen-sis, Sargocentron rubrum, Sauridaundosquamis, Atherinomorus lacunosus,Pempheris vanicolensis, Pteragogus pelycusand Apogon pharaonis. On the contrary,some other successfully established speciesare common and have acquired a commer-cial importance, i.e. Siganus luridus, S. rivu-latus and Sphyraena chrysotaenia; the latteris normally confused with the two indige-nous species S. sphyhraena and S. viridensiscoexisting in the same coastal area(CORSINI & ECONOMIDIS, 1999), andalso with the new Erythrean colonizer S.flavicauda. The quickly increasing popula-tion of the recent colonizer Etrumeus teresis already contributing to fishery in theDodecanese (CORSINI-FOKA, unpubl.data) and in the Cyclades Islands(KALLIANIOTIS & LEKKAS, 2005), asalso has been observed in Israel, Cyprusand Turkey (GOLANI et al., 2002, CICEK& AVSAR, 2003).

Local indigenous fish, especiallysmall-sized and fry, are subjected to aremarkable predation pressure. This factpresents the other side of the coin, per-haps suggesting that several invaders andnative species are in competition. Conse-quently, the impact on the local exploitedpopulations by Lessepsian immigrantsseems to be serious and accelerated insome cases. Thus a more detailedapproach on this matter is needed.

The permanent establishment of aLessepsian species in the area, and conse-

quently how harmful this would be for thenative fish, is related to its reproductivesuccess. In many cases such a successfulbreeding, in combination with food avail-ability and other favourable environmen-tal factors, leads to a population explo-sion. Such a case of population explosionwas observed in the area during the1940's for Upeneus moluccensis (LASKA-RIDIS, 1948a), followed by a dramaticcrash shortly after, so that nowadays thisspecies is rarely found in the area(CORSINI & ECONOMIDIS, 1999).

Several Erythrean invaders may havefound vacant ecological niches and obvi-ously they do not compete with localnative species. Thus, Siganus luridus andS. rivulatus were well established in theentire Levantine basin initially and theninvaded the Central Mediterranean, dueto the presence of few herbivorous com-petitors and/or abundant available food.Furthermore, Upeneus moluccensis ispresent but it does not dominate Mullusbarbatus. Apogon pharaonis is regularlycaught in trawl nets, as well as its indige-nous counterpart Apogon imberbis, whilePteragogus pelycus occurs with otherlabrid species of similar size (for exampleSymphodus sp., Coris julis, Thalassomapavo). Schools of Sphyraenidae are alsonoted, both belonging to the indigenousspecies Sphyraena sphyraena and S. viri-densis but also to the Lessepsian S. chryso-taenia and S. flavicauda (see above). Sau-rida undosquamis is very rare comparedto Synodus saurus. On the other hand, nolocal fish species disappeared in theRhodes marine area as assessed for therest of the Eastern Mediterranean(GOLANI, 1998b).

Fistularia invasion. In terms ofimpact, the recent invader, blue cornet-

79Medit. Mar. Sci., 8/1, 2007, 67-89

fish, Fistularia commersonii, caught nor-mally by trawl-nets up to 50-60m, hasdeveloped an important population and isconsidered one of the worst invasive fishspecies in the Mediterranean Sea(PANCUCCI-PAPADOPOULOU et al.,2005b; STREFTARIS & ZENETOU,2006). This species presents a very fastexpansion along the coasts of the Levan-tine basin (GOLANI, 2000; CORSINI etal., 2002) and recently up to the northAegean Sea (Chalkidiki peninsula), to thenorthern coasts of Crete and westward tothe Central Mediterranean and Tyrrhen-ian Sea (GOLANI et al., 2006 on-line;Table 2; ICES, 2006; PAIS et al., 2007).The phenomenon is alarming becausethis fish reproduces and grows very rapid-ly, reaching a remarkably large size. Sinceits first appearance in the Rhodes marinearea in 2001, the blue cornetfish occurs ata number of 5 to 20 specimens occasion-ally in a catch of any trawl net operation,mainly at a depth of 20-25 m, but also inthe very shallow waters of beaches andharbours (KALOGIROU et al., 2007).

Additionally, the species presents anextreme feeding activity and a clearaggressive behaviour when in schools. Asa free-swimming carnivore, it is merelydwelling over reefs and sea-grass beds. Asit has no or little commercial value, it issubjected to a very low fishing pressure,being more a ‘by catch’ fish than a target.Consequently, it is free to form large pop-ulations, which seriously affect economi-cally and ecologically important nativespecies not only in the limited area of theSE Aegean Sea, but in all occupied newhabitats, as in the South Aegean Sea(unpublished data). It mainly feeds onfish fry, gobiids and several exploitedpopulations of economically valuablenative fish, namely Spicara smaris, Mullus

spp. and Boops boops (CORSINI et al.,2002; KALOGIROU et al., 2007).Anatomical features of mouth and feed-ing behaviour observed in nature andfood items found in stomach contentreveal that Fistularia commersonii shouldnot be ranked in the highest trophic lev-els. Various sized individuals of thespecies could be seen rather as an easyprey-fish for any energetic predator, if itaccepts and/or appreciates them.

Inventorying Lessepsian migrants.Lessepsian alien fish species occurring inGreek waters belong to 20 families. Thenon-indigenous species have added sevenfamilies to the Hellenic ichthyofauna:Siganidae, Hemiramphidae, Fistularidae,Holocentridae, Leiognathidae, Pem-pheridae, Monacanthidae, while otheraliens increased the species number offive already well-represented families(ECONOMIDIS, 1973; WHITEHEADet al., 1984-1986; BAUCHOT, 1987;PAPAKONSTANTINOU, 1988) (Fig. 2).The remaining aliens increased thespecies number in families represented byone or two species, with the exception ofthe Tetraodontidae, which needs particu-lar attention. Until 1994, only theautochthonous Lagocephalus lago-cephalus, the Lessepsian Lagocephalusspadiceus and Sphoeroides pachygasterfrom the Atlantic were known in the area.A rapid increase of records in the last twoyears added four more Indo-Pacifictetraodontid species: Tylerius spinosis-simus, Lagocephalus suezensis, Lago-cephalus sceleratus and Torquigener flavi-maculosus (Table 1). Among these lastalien tetraodontid colonizers, the popula-tion of the large sized L. sceleratus, whichis not marketable since it may cause foodpoisioning, is increasing rapidly (AKYOL

80 Medit. Mar. Sci., 8/1, 2007, 67-89

et al., 2005; GOLANI & LEVY, 2005;CORSINI et al., 2006; BILECENOGLUet al., 2006; KASAPIDIS et al., 2007) as L.suezensis as well. This fact is also sus-tained by very recent unpublished datawhich show the presence of a large num-ber of juveniles of these two species infishery activities along the coasts ofRhodes Island.

Conclusions

Almost the entirety of alien fishspecies introduced for aquaculture andtheir presence in nature is based onunverified observations of free-swimmingspecimens which strongly suggest carelessrelease or escape from fish farms. The listof species is expected to graduallyincrease, as the number of aquacultureinstallations in the Hellenic seas and inthe neighbouring areas has been increas-

ing. Apparently, no serious precautionsare undertaken in order to avoid any acci-dental escape from farm stocks.

Concerning Lessepsian aliens, thepresence of other species may be ascer-tained only through a comprehensive andregular study along the coasts, alsoincluding habitats fished infrequently,where a species at a low density popula-tion although it is a regular inhabitantmay be undetected, (GOLANI et al.,2002), or habitats are not exploitedbecause their resources are considered ashaving minimal or zero commercial valueor they are unreachable by usual fishingmethods, however, they may be importantas nursery grounds and biodiversity reser-voirs.

Taking into account the increasedinterest of the scientific community inLessepsian colonization, a higher rate ofinvasion of the Levantine coasts by tropi-

81Medit. Mar. Sci., 8/1, 2007, 67-89

Fig. 2: Number of Lessepsian fish species and families added to the Hellenic ichthyofauna.

cal fish species and their successfulacclimatization is evident, confirming that‘the littoral and infralittoral biota of theLevantine Sea is undergoing a profoundchange due to the influx of Erythreaninvaders’ (GALIL & ZENETOS, 2002),and furthermore giving support to theindication of a tropicalization of the area(BIANCHI & MORRI, 2003). In thelong run, this phenomenon will little bylittle increase the complexity of the entireeastern Mediterranean ecosystem andwill most probably lead to a new equilib-rium, which should be characterized by ahigher productivity and stability.

Ackowledgements

The authors are grateful to two anony-mous referees for their constructive com-ments and advice on the manuscript.

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Accepted in 2007