Aquatic Invasions (2007) Volume 2, Issue 4: 281-312
DOI 10.3391/ai.2007.2.4.2© 2007 The Author(s)Journal compilation © 2007 REABIC (http://www.reabic.net)
This is an Open Access article

Special issue “Alien species in European coastal waters”
Geoff Boxshall, Ferdinando Boero and Sergej Olenin (Guest Editors)

Research article

Seeing Red: Alien species along the Mediterranean coast of Israel
Bella S. Galil
National Institute of Oceanography, Israel Oceanographic & Limnological Research, POB 8030, Haifa 31080, Israel
E-mail: bella@ocean.org.il
Received 10 September 2007; accepted in revised form 20 November 2007

Abstract
An overview is presented of the metazoan alien biota recorded from the Mediterranean coast of Israel. Of the 296 alien species,
284 have been introduced from the Red Sea/Indo West-Pacific through the Suez Canal. A brief history of the research on
introduced species along the Israeli coast is presented along with the list of species, and their ecological and economic impacts
are discussed.
Key words: invasion, Israel, Mediterranean, Suez Canal, economic impact

A brief history of the research on marine alien
species off the Mediterranean coast of Israel
The first marine alien species from the coast of
Israel was found on the beach of Jaffa (Figure 1)
and described in 1891 as Cerithium levantinumby E. A. Smith (in Hart, 1891). That species waslater identified as C. scabridum Philippi, 1848
(Mienis, 1992b), a species known from the Red
Sea and the Indian Ocean, that had already been
described from the newly excavated Suez Canaland its Mediterranean terminal at Port Said
(Fuchs 1878, Keller, 1883). The first alien
species identified as such was the pearl oyster
Pinctada radiata (Leach, 1814) (as Meleagrinasavignyi Monterosato, 1878), mentioned from
Jaffa “ .. c’est là une espèce de la mer Rouge,
immigrée dans la Méditerranée depuis lepercement de l’Isthme de Suez” (Monterosato,
1899: 392). These early findings set a patternthat holds true to this day – 127 of the 297 alien

Figure 1. Map of the Levantine basin of the Mediterranean
and the Israeli coast

281

B.S. Galil

species recorded off the Israeli coast aremolluscs that had entered the Mediterranean
through the Canal.
Dr. med. et phil. Walter Steinitz from Breslau
was among the first to recognized the scientificsignificance of the movement of biota throughthe Canal. Dr. Steinitz, published a pamphlet at
his own expense, where he pointed out that theSuez Canal “…is the only place on earth where
two quite different zoological provinces havecome into direct touch with one another”, andthat the Canal “…connects two oceans having
each a very different fauna from the other, which
throws up a number of new problems: Does an
exchange of fauna take place between the twooceans by means of this canal? Do theimmigrants from the other side flourish on thisor die? Has such an immigration caused
important changes in the fauna of the easternbasin of the Mediterranean?” (W. Steinitz 1919:
8). Steinitz deplored the lack of faunistic
research in the Levantine Basin “… at thishighly important point of juncture there is noplace of observation!”, and forsook his
flourishing medical practice to study the marine
biota of Palestine in 1924 and 1925. Noting 15
species of Indo-Pacific origin, he urged thecreation of a Marine Station in the Levant
dedicated to the study of the transformation of
the Levantine biota (W. Steinitz 1929, 1936). In
1933 Dr. Steinitz immigrated to Israel andresumed his research in a space allotted him by
the Zevulun Seafaring Society, Tel Aviv, but theneed to provide for his family compelled him tojoin a communal village and breed poultry.
However, he passed his scientific interest to hisson, Heinz Steinitz, who studied with thepioneer ecologist Bodenheimer and received thefirst Ph.D. in zoology awarded by the Hebrew
University of Jerusalem.
Munro Fox, who would later head the ‘The
Cambridge Expedition to the Suez Canal’,
outlined in 1923 “the dates of the various stages
in the migration northward through the Canal of
the Red Sea crab (Neptunus pelagicus) [=Portunus pelagicus (Linnaeus, 1758)], and …
the apparent limits of its present distribution
along the Mediterranean coast” including thefish market of Haifa (Fox, 1924). Indeed,
Bodenheimer in his seminal “Animal life inPalestine” when discussing the “Indo-Pacificinflux through the Suez Canal” mentioned that
“Quite a number of [alien] fishes have not only
reached our shores, but some of them have evenincreased in such numbers, that they appear

282

regularly on the fish market” (Bodenheimer
1935: 457). He then presciently added “It isalmost certain that the Indo-Pacific influx is still
under way and it will be most interesting tostudy this process. The local increase will takeconsiderable time, even for those organisms
which thrive successfully in the new area”(Bodenheimer 1935: 460). The progress of
Erythrean biota in the Levant in the early 20th
century was noted in the inventories of molluscs(Monterosato 1899, Pallary 1912, 1938, Gruvel
and Moazzo 1931, Haas 1937, 1948, Steuer
1939); decapod crustaceans (Gruvel 1928, 1929,
1930a,b, 1931, Monod 1930, 1931, 1932, Steuer
1938) and fish (Gruvel 1929, 1931, Liebman1934), some of which report findings from theIsraeli coast.
The intensive investigations of the biota of
the continental shelf of Israel by the Sea
Fisheries Research Station (SFRS) in 1946-1956
resulted in a series of publications thathighlighted the extent the Erythrean taxa havebeen established along the coast (Carmin 1946,
Haas and H. Steinitz 1947, Haas 1948, 1951,
Gottlieb 1953, 1960, Holthuis and Gottlieb 1958,
Pérès 1958, Tebble 1959, Oren and H. Steinitz
1959, Lewinsohn and Holthuis, 1964). The
earlier community studies of the continental
shelf of Israel were mainly fishery research(Wirszubski 1953, Gilat-Gottlieb 1959), and it
was only in the 1960s that an extensive program
was undertaken by Gilat (1964) to describe themacrobenthic communities off the Israeli coast.
The SFRS fisheries biologist, Ben Tuvia, aformer student of H. Steinitz, published the firstlist of fishes from the Mediterranean coast of
Israel, focusing on the Erythrean alien species(Ben-Tuvia
1953a).
He
observed
thedevelopment of the Erythrean invasion in “a,appearance of new Erythrean species in theeastern part of the Mediterranean; b, thespreading of Erythrean species, alreadyestablished in the eastern Mediterranean close to
the Suez Canal, to new areas (a movement
following the coast of Israel, Syria, Turkey andRhodes); c, the increased abundance of theinvading species in areas where they wereformerly scarce.” (Ben-Tuvia 1953b: 464). Ben
Tuvia continued to update the list of Erythreanalien fish off the Israeli coast: 21 species (BenTuvia 1958), 30 species (Ben-Tuvia 1971), 41
species (Ben-Tuvia 1985). The steadily
increasing number of alien fish moved him to
write that “[v]ery little is known about how theinflux of Red Sea species has influenced theAlien species along the Mediterranean coast of Israel

ecology of the eastern Mediterranean. Certainly
these fish must play an important role in thefood chain and the general biological balance of
this area.” (Ben-Tuvia 1971: 3). Ben Tuvia
estimated, based on the catch statistics collatedby the Department of Fisheries, that between
1965 and 1969 Erythrean fish made up 8% of
the total landing (in weight) (Ben-Tuvia 1973),
and the amount doubled to 16% between 1980
and 1982 (Ben-Tuvia 1985).
In 1967, a joint program by the Smithsonian
Institution, the Hebrew University of Jerusalem,and the Sea Fisheries Research Station, Haifa,
was established to investigate the spread of theErythrean biota in the Levant (Israel, Cyprus,
Rhodes) and its impact on the native biota. The
scope of the program was expanded following
the ‘Six Day War’ to include the Sinai coasts
and the Suez Canal itself. H. Steinitz headed that
program with W. Aron, Director of the Oceanography and Limnology Program, Smithsonian
Institution. On the untimely death of Steinitz, his
deputy, Por, assumed the position. At the end of
the 3rd year of the program, with some 5300
samples collected and partially sorted andidentified, Steinitz published “A critical list of
immigrants via the Suez Canal” of 140 Erythrean
and Indo-Pacific species known to have crossed
the Suez Canal into the Mediterranean (H.
Steinitz 1970). By now, it was widely perceived
that the littoral and infralittoral biota of theLevantine basin has been undergoing a rapid andprofound change (Por 1978). Galil and
Lewinsohn (1981) noted that the shallow benthic
communities along the Mediterranean coast of
Israel had no known parallel outside the Levant
because of the great number of Erythrean aliens.
Over the past three decades Galil (a former
student of Lewinsohn), D. Golani (a former
student of Ben Tuvia), M. Goren and H. Mienis(among others) have continued to record the
spread, biology and impact of alien species
along the Israeli coast and the Levantine Basin(Galil 1986, 1993, 1997, 1999, 2000, 2004,
2005, 2006, 2007, Galil and Golani 1990, Galiland Lützen 1995. Galil et al. 1990; Golani 1987,
1992, 1993 a-c, 1996, 1998, 2000a,b, 2002,
2004, 2006, Golani and Ben Tuvia 1982, 1990,
Golani and Fine 2002, Golani and Sonin 1992,
2006; Goren and Aronov 2002, Goren and Galil
1989, 1998, 2001, 2005, 2006, Mienis 1976 a,b,
1977a,b, 1979, 1980a,b, 1984, 1985 a,b, 1987,
1990, 1992a,b, 1995, 1999, 2000a,b, 2002a-c,
2003 a-e, 2004a-d, 2006a-c, Mienis and Singer
1983, Mienis and Gat 1987, Mienis et al.

1993a,b ). Yet, no concerted effort had been
undertaken to survey the entire coast since theearly 1970s, and most of the records stem from
fortuitous finds. A national targeted effort tosurvey the presence and abundance of theErythrean species and study their biology andecology is wanting.
Aliens, vectors, trends
Research papers, conference abstracts, the local
Fishermen’s Bulletin, unpublished M.Sc. and
Ph.D. theses, and the author’s own databases
resulting from two decades of monitoring thebenthic communities off the Israeli coast, wereassembled and critically examined in order toconstruct an authoritative dataset tracing theorigin, date and mode of introduction, current
distribution, and impact of the 296 metazoan
alien species recorded along the Mediterranean
coast of Israel. Earlier records were reassessedand some were deleted, such as the seven sponge
species and three species of hydroids listed by
Por (1978) that, as tactfully suggested byZibrowius (1992), were in need of “criticalreevaluation”. Por (1978) also included in his
list seven species of ascidiaceans, that are either
widely distributed, or of “problematic identification. The list thus provides an inexact impression of lessepsian migration among ascidians”(Zibrowius 1992: 96).
The dataset includes the date of collection (ifunavailable, date of publication) of the firstrecorded specimens, the native range of thealien, its means of introduction, whether throughthe Suez Canal, vessels, or mariculture, and thecurrent population status. The date of collectionof the first specimen(s) is significant for the
study of the patterns and processes of invasion,
though we accept that the date of collection (or,when missing, the date of publication) may be
years behind the actual entry dates, and that
identification and publication may lag behindcollection: the gastropod Cerithium scabridum(Philippi, 1848) collected on Jaffa beach before
1891, was misidentified as C. levantinum (seeabove), and only when collected again in Jaffa in
1928-9 it was correctly identified as a “forme dela mer Rouge” (Gruvel and Moazzo 1931). The
pyramidellid gastropods Chrysallida maiae(Hornung and Mermod 1924) and Syrnola
fasciata (Jickeli 1882) were collected off theIsraeli coast in 1935 and 1949 and identified
only in 1992 (van der Linden and Eikenboom

283

B.S. Galil

284

140

128

120
100
80

64

60

48

40
5

4

4

Ectoprocta

Chlorophyta

6Echinodermata

6

Cnidaria

8

Platyhelminthes

21

20

2

Phaeophyta

Rhodophyta

Annelida

Arthropoda

Chordata

Mollusca

0

Figure 2. Taxonomic classification of marine alien species
along the Mediterranean coast of Israel

160
140

142

120
100
72

80

51

60
40

5

4

1

2

Cosmopolitan

Atlantic W

Pacific W

Unknown

19

20

Pantropical

Red Sea

Indian Ocean0
Indo Pacific1992), and 1995 (Mienis 1995) respectively; thetube worm Hydroides brachyacantha Rijoa,
1941, was collected in Jaffa, Israel, in 1933, but
identified only nearly 60 years later (Ben-Eliahu
1991).
The 296 metazoan species identified in thepresent work as alien off the Israeli coast arelisted in the Annex. This particular list is limited
to multicellular organisms because the identityof many unicellular organisms is still in doubt,
as well as their native range and distribution. A
taxonomic classification of the alien species(Figure 2) shows that the alien phyla mostfrequently recorded are Mollusca (43%),
Chordata (22%), Arthropoda (16%), andAnnelida (7%). Phyla not represented in the list
include the little studied Porifera, Nemertea,
Priapula, Nematoda, Entoprocta, Pogonophora,
Sipuncula, Echiura, Brachiopoda and Phoronida.
The data are presumably most accurate for large
and conspicuous species, which are easily
distinguished from the native biota.
A geographic classification (Figure 3) showsthat the native range of the alien taxa recorded in
Israel is most commonly the Indo-Pacific Ocean(48%), the Indian Ocean (24%), the Red Sea(17%), and pantropical (6%). Caution should be
exercised when using these data: the true originof the Israeli populations of a species widelydistributed in the Indo-Pacific Ocean may beeither its populations in the Red Sea or further in
the Indian Ocean, or secondarily from otherestablished Levantine populations. The source
populations or means of introduction of alien
species in Israel have not yet been successfully
ascertained by molecular means (Terranova et al.
2006). But even taking into account thesecaveats, it is quite clear that most of the alien
species off the Israeli coast are thermophilic,originating in tropical seas.
A classification according to the means of
introduction shows that the great majority ofaliens off the Israeli coast entered through theSuez Canal, whereas vessels and mariculture
contributed only a small number of aliens. Heretoo, caution should be exercised when using
these data: only rarely are the means and route of
introduction known from direct evidence. Mostly
they are deduced from the biology and ecology(if known) of the species, the habitats andlocales it occupies in both the native andintroduced range, and its pattern of dispersal (ifknown), i.e., for a fouling species frequently
recorded from ports, shipping is assumed to be
the most probable vector. It is has been assumed

Figure 3. Native range of marine alien taxa recorded along
the Mediterranean coast of Israelthat Erythrean aliens progress through the Suez
Canal and along the coasts of the Levant as aresult of “natural” dispersal, by autochthonous
active or passive larval or adult movements,
unaided further either directly or indirectly byhuman activity. Indeed, a temporal succession of
directional (“stepping stones”) records from the
Red Sea, the Suez Canal, and along the coasts of
the Levant confirms a species status as anaturally dispersing Erythrean alien. However,vessel-transported dispersal is feasible as well:“It is, of course, well known that ships have inmore than one instance dispersed marineorganisms from one part of the world to another.
This must apply equally to transport through theSuez Canal. …. Possibly tugs and barges
permanently employed in the Canal may take alarger share than other vessels in this transport
from one end of the Canal to the other. There arecoal barges, for instance, which remain for somemonths at one end of the Canal and then are
towed through to stay for some months more at
the other end. The time spent at either end would
permit on the one hand of the settling of larvaeon the bottom of the barges, and on the otherAlien species along the Mediterranean coast of Israel

hand of the liberation of eggs or larvae frommature individuals” (Fox 1926:20). Even whererecords are consistent with long-shore dispersal,
there might be a degree of uncertainty wherefouling organisms (such as serpulid polychaetes
or mussels) are concerned, as they are more
susceptible to shipping-mediated transfer. In
some cases we suspect simultaneous mechanismsof transport.
Though the date of collection (or publication,
see above) may lag years behind the date of
introduction, the numbers of alien species
recorded off the Israeli coast each decade overthe past century are given in Figure 4. With the
exception of few scattered early records (seeabove), the records begin in the 1920s with theworks of W. Steinitz and the results of ‘The
Cambridge Expedition to the Suez Canal’,
whereas the 1930s saw the publication of several
regional lists (see above). The smaller number ofrecords in the 1940s probably reflects thedevastation of the Second World War and thestruggle for the independence of Israel. A surge
in the records in the 1950s through the 1970s
probably represents both an increase in introductions and an interest in their study, culminatingin the publications resulting from the joint
program by the Smithsonian Institution, theHebrew University of Jerusalem, and the Sea
Fisheries Research Station, Haifa. Though noconcerted effort was undertaken since, thenumber of records in the 1980s and 1990s were
35 and 36 respectively. More introductions wererecorded in the first years of the 21th century
than in the 1980s and 1990s.
Since the 1950s major political, economicaland societal changes have occurred in theLevant. Their impact on the rate and means of
introduction is apparent (Figure 5): though theSuez Canal remained the main access route foralien species, the rate of introductions by vessels
has been higher. Of the 56 alien species known
in 1950, 93% entered the Mediterranean throughthe Suez Canal, 7% were possibly canal/vesseltransported; of the 240 alien species recorded
since 1950, 80% were Erythrean aliens, 9% werecanal/vessel-transported, 10% and 1 % respectively were vessel-transported and mariculture
introductions. The increase in vessel-transported
aliens may be attributed to the increase inshipping volume throughout the region.
Since the probability of collecting a vagrant
incursion in the sea is diminishingly small, mostrecorded alien species are considered as‘established’ species that have self-maintaining

Number of alien species recorded in the Israel <1900-2007

50
40
30
20
10
0
<1900 1900- 1910- 1920- 1930- 1940- 1950- 1960- 1970- 1980- 1990- ≥2000

Figure 4. Number of marine alien species recorded per
decade along the Mediterranean coast of Israel <1900-2007.

Figure 5. Number of species introduced by each pathway
before and after 1950 along the Mediterranean coast of
Israelpopulations of some duration. It is recognizedthat some alien species may fail to maintainpopulations over time and thus a single record
dating back several decades may be consideredan ephemeral entry. We list 36 species known
from single records, most of them recorded in
the last decade. The distinction between the‘established’ and ‘ephemeral’ aliens is sometimes difficult to discern and circumscribed inlarge part by our ignorance of the marine
environment. Even once established in the Mediterranean, the alien species differ markedly intheir histories.
The parasitofauna of the alien species off theMediterranean coast of Israel
Absence of natural enemies, be it competitors,
predators, pathogens, or parasites, is one of the
explanations given for the success of alien biota(Wolfe 2002, Torchin et al. 2003). Diminishedparasitization of alien species may be attributed
to reduced probability of infestation due to thesmall numbers of the founding population, its285

B.S. Galilbeing composed of uninfected larval stages, or,in the case of heteroxenous parasites, theabsence of intermediate hosts in the new locale,and on the other hand, the host-specificity ofsome native parasites forestalling infection ofalien hosts.
The parasitofauna of the two Erythrean alien
siganids were examined in the 1970s and 1980s
and 11 parasite species were identified – allmonoxenous parasites with direct life cycles
requiring no intermediate host, including themonogenoids
Glyphidohaptor
plectocirra(Paperna, 1972), Tetrancistrum polymorphus(Paperna, 1972), T. suezicus (Paperna, 1972), T.
strophosolenum Kritsky, Galli and Yang, 2007
(Paperna 1972, Diamant 1989, Galli et al.,2007). A study of S. rivulatus collected at theport of Ashdod, next to fish farming facilities, inthe mid 1990s, revealed 7 Erythrean species ofmonoxenous parasites (Diamant et al. 1999).
However, Fischthal (1980) identified in Siganus
rivulatus (Forsskål, 1775) specimens from theMediterranean two species of heteroxenous
digenetic trematodes with complex life cycles
requiring more than one host. Paperna (1972:3)
suggested that endoparasitic helminthes spread
into a new area only if they are not highly hostspecific or if the suitable intermediate host has
arrived as well, though “in most cases the
migration of the fish host does not coincide withsimilarmigrationbytheintermediate
invertebrate hosts”. It is of interest that a recent
study of S. rivulatus has shown that theprevalence of its Erythrean monogenean ectoparasite Polylabris cf. mamaevi off the Mediterranean coast is three times as high as thatfound in the Red Sea populations (Pasternak et
al. 2007). These authors ascribe the heavier
infection to “Changes in the natural environment
and impact of anthropogenic factors encounteredby the rabbitfish in their new Mediterranean
habitats” (Pasternak et al. 724). However, thesiganids are immensely successful colonizers,
and it may be that the parasite high infestationlevels stem from their access to much denser
host populations.
Fischthal (1980) examined parasite loads of
500 fish of 59 species collected by gill-netting in
1977 off the Mediterranean coast of Israel. He
found that 29 of the 43 (67%) native Mediterranean species were infected with digenetic
trematodes, but only 5 of the 12 (42%) Erythreanalien species. Interestingly, many of the trematodes infecting the Erythrean hosts were ofAtlanto-Mediterranean origin and were probably286

acquired in the Mediterranean from native hosts.
These results support the ‘enemy release
hypothesis’ – alien species generally import only
a subset of their parasitofauna, thus releasing
themselves from the parasites of their native
habitat during introduction, and though subsequently accumulating parasites native to their
new environment, they harbor fewer parasites intheir new locale than in their native range(Torchin et al. 2001).
The Levantine populations of the Erythrean
swimming crab Charybdis longicollis Leene,
1938 have been parasitized recently by thesacculinid rhizocephalan, Heterosaccus dollfusi
Boschma, 1960 (Galil and Lützen 1995). The
swimming crab, can form as much as 70% of thebenthic biomass on sandy-silt bottoms off theIsraeli coast (Galil 1986). Release from parasites
may have contributed to its success - of thethousands of C. longicollis collected off theIsraeli coast in over three decades, none wasparasitized until 1992, when a few specimens
were discovered carrying the externae (reproductive part of the parasite) of the castratingsacculinid, itself an Erythrean alien. Along theIsraeli coast infection rates of up to 77% wererecorded, with up to 57.6% of the infected hosts
bearing more than one externa (Galil and
Innocenti 1999). The rapid spread and the highprevalence of H. dollfusi infestation may berelated to the dense population of the host andthe year-round reproduction of the parasite,causing recurrent infection. In its second decadein the Mediterranean, the population of H.
dollfusi seems stable: despite the high prevalence of the parasite and its injurious impact on
the host reproduction, there is no noticeable
reduction in the host population. It is suggested
that the high fecundity of the host females, the“size refuge” formed by parasite-free larger
males, and the “open” recruitment dynamics of
C. longicollis, keep its population density highenough for H. dollfusi, with its “closed” recruitment dynamics, to maintain its pandemic
infection rates (Innocenti and Galil 2007). The
Erythrean sacculinid had not been detected inany of the other portunid crabs, alien or native,
inhabiting the Levantine sublittoral, including
the congeneric C. helleri.
The ecological impact of alien species off theMediterranean coast of Israel
A single local extirpation of a native species isknown and several cases of sudden decline inAlien species along the Mediterranean coast of Israel

abundance concurrent with proliferation ofaliens had been recorded off the Israeli coast.
But even when populations of native Mediterranean species appear to have been wholly outcompeted or partially displaced from theirhabitat space by an alien, the causes cannot be
disentangled from potential confounding factors
such as the profound anthropogenic alteration of
the marine ecosystem through habitat destruction, pollution, and rising Mediterranean seawater temperature.
The Erythrean alien killifish, Aphanius dispar(Rüppell, 1829), is markedly euryhaline,
occurring in freshwater, and in salinities up to 4
times as high as seawater, whereas its periMediterranean endemic congener, A. fasciatus(Nardo, 1827), occurs mostly in brackishlagoons. The first Erythrean killifish wascollected off Tel Aviv, Israel, in the winter of
1943/44 (Mendelssohn 1947). The last specimens of the native killifish, A. fasciatus, along
the Israeli Mediterranean coast were collected on
August 13, 1976 (M. Goren, pers. comm.;
preserved in the National Collections, Tel Aviv
University, access number 6319). Naturally
occurring hybrids of the two killifish species
were described as “ ... common, and in somelocalities (Ashdod Harbour [southern coast of
Israel], parts of Bardawill Lagune etc.), these
populations comprise mostly hybrids” (Goren
and Rychwalski, 1978: 261). Within a generation
an endemic genotype appears to be locally lost
through hybridization, and A. dispar and its
hybrids replaced A. fasciatus along theMediterranean coast of Israel (Goren and Galil
2005).
The two species of siganid fish, Siganus
rivulatus and S. luridus (Rüppell, 1828), thatentered the Mediterranean from the Red Sea
through the Suez Canal, were first recorded off
the coast of Israel in 1924 (Steinitz 1927) and1955 (Ben Tuvia 1964), respectively. The
schooling, herbivorous fishes, able to settle in arange of habitats, comprise one third of the fishbiomass in rocky habitats along the Israeli coast(Goren and Galil 2001). The multitudinous
siganids have altered the community structure
and the native food web along the Levantine
rocky infralittoral. They replaced native herbivorous fish, such as Sarpa salpa (Linnaeus, 1758),
that had been abundant in trawl catches early inthe 20th century (Gruvel 1931, George andAthanassiou 1967, Bariche et al. 2004). Prior to
the invasion of the siganids, there were few
native herbivorous fish and invertebrates andtheir role in the food web off the Levantine
rocky habitats had been negligible. The siganids
increased the rate by which large amounts ofalgal material were recycled, these fish apparently accelerated the transfer of energy from theproducer to the consumer levels (in hours
through the fish gastrointestinal system rather
than weeks or months of decomposition), and byserving as major item of prey (up to 70 %) forlarger infralittoral predators such as groupers(Aronov 2002, Goren and Galil 2005). Theirgrazing pressure on the intertidal rocky algae
may have benefited the proliferation of an alien
Erytrean mussel by providing suitable substrate
for its settlement (see below). A comprehensive
study of the algal vegetation and the ecology ofthe fish community off a vermetid reef, south of
Haifa Bay, Israel, provided an opportunity tocompare the diets of S. rivulatus and S. luridus
collected at the site with the composition of thelocal algal community, and examine the importance of algal spatial variability and seasonality
in the dietary choices of both siganid fishes. An
analysis of the siganids’ gut contents, in conjunction with the spatial and seasonal compositionof the local algal community, showed that theirdiet has a significant impact on the structure of
the local algal community: it seems that byfeeding selectively they have nearly eradicatedsome of their preferred algae locally (Lundberg
et al. 2004).
A small Erythrean mytilid mussel, Brachidontes pharaonis (Fischer P., 1870), wasconsidered in the early 1970s “c. 250 times
rarer” than the native mytilid Mytilaster minimus(Poli, 1795), that formed dense ‘Mytilaster beds’
on intertidal rocky ledges along the Israeli
coastline, with up to 26 specimens per cm² (atPalmahim, Safriel et al. 1980, tab. 4). Studies
conducted in the late 1970s along the Israeli
coastline, while B. pharaonis was still relatively
rare, predicted it would neither establish dense
populations nor outcompete the native Mytilasterminimus: “B. variabilis [pharaonis] never forms‘beds’ in the eastern Mediterranean, and issingly or in small groups dispersed within M.
minimus”, and “… it did not displace M.
minimus” (Safriel et al. 1980: 39, 59). The
relative rarity of B. pharaonis was attributed tothe invader’s low intrinsic rate of increase incomparison to that of the native species, and tostrong density-independent mortality generated
by exposure to either high wave action or
sedimentation. By the end of the 1980s, following a series of experiments, it was determined287

B.S. Galilthat Brachidontes interferes with recruitment of
Mytilaster, and detrimentally affects its survivaland growth, yet the researchers maintained that
the Erythrean mytilid was not likely to outcompete the native species: “The two species
can coexist, both locally and regionally” (Safriel
and Sasson-Frostig 1988: 225). In the late 1990s
a survey conducted in some of the same sites
have shown a rapid shift in dominance, withsome dense populations of up to 300 specimens
per 100 cm² on rocky platforms “where mussel
beds were absent in the past” (Rilov et al. 2004:
347). More recently “the same rocks are ….
completely covered with the Erythrean B.
pharaonis, while M. minimus is only rarely
encountered.” (Mienis 2003d: 15). The
establishment of massive beds of Brachidontes
has had significant effects on the biota of the
rocky intertidal. As the presence of algae isnegatively correlated with the presence of theErythrean mytilid, and is considered to impede
the settlement of its postlarvae (Safriel andSasson-Frostig 1988), a shift in habitat conditions that reduces algal cover might havebenefited Brachidontes. As it happens, few
herbivores occurred in the rocky shores of thesoutheastern Levant prior to the arrival of theErythrean siganid fish (see above). During hightide schools of (mostly young) siganids feed on
the intertidal platforms (Lundberg et al. 2004). It
is suggested that the multitudinous siganids may
have triggered the population increase by
clearing the intertidal platform of algae. The
displacement of the native mussel by the larger,
thicker-shelled Erythrean mytilid may havechanged predation patterns so that the populationof the native whelk, Stramonita haemastoma(Linnaeus, 1758), that were found to preferentially prey on Brachidontes, increased greatly(Rilov et al. 2002).
An Erythrean limpet, Cellana rota (Gmelin,1791), first collected in the Mediterranean in1961 (Christiaens 1967), spread by 2000 along
the Israeli coast, occupying the less waveexposed intertidal sites, whereas the native
Mediterranean limpet, Patella caerulea Linnaeus, 1758, inhabited rocks exposed to the surf.
A recent survey along the Mediterranean coastof Israel found that the alien limpet dominates
the upper rocky littoral and has been replacing
the native limpet when “a few years ago Patellacaerulea was the only limpet inhabiting the samerocks at that locality [Ashdod]” (Mienis 2002b:
275). Along the southern coast of Israel it has
already completely replaced the native limpet,288

along the central coast it “has taken possessionof 40-50% of the available space.” (Mienis
2003d: 15). Similarly, the Erythrean jewel box
oyster Chama pacifica Broderip, 1834, outnumbered its native congener, C. gryphoides
Linnaeus, 1758: “The local S. gaederopus and C.
gryphoides are hardly even encountered as
epibionts on the new immigrant species.”(Mienis 2003d: 15). The native Mediterraneancerithiid gastropods, Cerithium vulgatum
Bruguière, 1792 and C. lividulum Risso, 1826,
respectively common and abundant in shallow
water along the coast of Israel until the 1970s,
were supplanted by the Erythrean cerithiids
Cerithium scabridum Philippi, 1848 andRhinoclavis kochi (Philippi, 1848) (Mienis
2003d: 15). The population of the Erythrean
narrow-barred mackerel, Scomberomorus comerson Lacepède, 1800, has greatly increased in the
1980s, coincidently with the decline of the onetime common native meager, Argyrosomus
regius (Asso, 1801), to the point where the latteris rarely encountered along the Israeli coast; theErythrean dragonet, Callionymus filamentosus
Valenciennes, 1837, replaced the native
callionymids C. pusillus Delaroche, 1809, and C.
risso LeSueur, 1814, along the Levantine upper
shelf (Golani 1998). The Erythrean snapping
shrimps Alpheus inopinatus Holthuis & Gottlieb,
1958, and A. audouini Coutière, 1905, are morecommon in the south-eastern Levantine rocky
littoral than the native A. dentipes GuérinMéneville, 1832, and on the muddy bottoms A.
rapacida de Man, 1908, is much more common
than the native A. glaber (Olivi, 1792)(Lewinsohn and Galil 1982, Galil 1986). The
native penaeid prawn, Melicertus kerathurus(Forskål, 1775) was commonly caught bytrawlers along the Israeli coastal shelf on sandyor sandy mud bottoms, and supported a commercial fishery throughout the 1950s (Holthuis andGottlieb 1958). It has since nearly disappeared,
and its habitat overrun by the Erythrean penaeid
prawns.
Competitive displacement may also modify
bathymetric ranges in populations of Erythrean
and native species: the native red mullet, Mullus
barbatus (Linnaeus, 1758) and the native hake,
Merluccius merluccius (Linnaeus, 1758) wereboth displaced into deeper, cooler waters by
their respective Erythrean competitors, Upeneus
moluccensis (Bleeker, 1855) and Saurida
undosquamis (Richardson, 1848). The goldband
goatfish, U. moluccensis, made up 10-15% of thetotal mullid catches off the Israeli coast by the

Alien species along the Mediterranean coast of Israel

late 1940s (Wirszubski 1953). Following theexceptionally warm winter of 1954-55, its
percentages increased to 83% of the catch,
replacing the native red mullet in the commercial
fisheries (Perlmutter, 1956). Both mullid species
have a similar diet, and occupy muddy bottoms
shallower than 75 m, but whereas the red mullet
spawns from April to June with a peak in May,
the goldband goatfish spawns from June to
September (Wirszubski 1953). The considerably
higher water temperatures may have resulted inpoor survival of the red mullet spawn whereas an
unusually large year class of the goldband
goatfish survived. In previous years, the young
red mullets would settle to the bottom during
July through September, where they have had adistinct size advantage over the later-spawned
goldband goatfish. The failure of the 1955 redmullet year class may have left their niche only
partly occupied, to the advantage of theErythrean species, and the unusually warm
waters enhanced the latter species’ survival rate
(Oren 1957a, b). The native mullet has eversince been displaced into deeper, cooler waters:
87% of the mullid catch off the Israeli coast
consisted of alien mullids at depth of 20 m, 50%
at 55 m, but only 20% in waters deeper than 70
m (Golani and Ben Tuvia 1995). The Erythrean
lizardfish, S. undosquamis, was first recorded
from the Mediterranean coast of Israel in 1952
(Ben Tuvia 1953a). The “abrupt rise in catch of
the lizard fish Saurida undosquamis, taken byotter trawlers with the usual gear on the regular
fishing grounds” (Oren 1957b: 1) was attributed
to the warm winter of 1954-55 (Ben Yami 1955,
Chervinsky 1959). In 1955-56 the lizardfish
became commercially important, constituting up
to one fifth of the total annual trawl catch along
the Mediterranean coast of Israel. This sudden
increase came at the expense of the native hake,
it too was displaced into deeper waters (BenYami and Glaser, 1974). Since the mid 1980s the
lizardfish’s share in catches has declined, andthat of the hake augmented. Similarly, The
Erythreanmantisshrimp,Erugosquillamassavensis, (Kossmann, 1880) is abundant off
the Israeli coast at depths of 20-60 m. Though on
occasion it is taken together with the native
Spottail mantis shrimp, Squilla mantis(Linnaeus, 1758), generally the latter occurs indeeper waters than Erythrean alien. The Spottail
mantis shrimp is fished commercially in shallow
waters elsewhere in the Mediterranean (Lewinsohn and Manning 1980), but off the Israeli
coast it is found mostly between 70 and 90 m.

Lewinsohn and Manning (1980) questioned“whether temperature, bottom type, or pressure
from O. massavensis, or a combination of theseis responsible for its depth distribution”.
The economic impact of marine alien species
off the Mediterranean coast of Israel
Some Erythrean aliens have been exploitedcommercially almost as soon as they entered theLevant and their economic importance had been
acknowledged early : “... les passages définitifs
de ces espèces à travers la totalité du Canalprésentent un résultat économique [sic]également très important..... pour les marchés
palestiniens et syriens, un appoint non
négligeable et particulièrement intéressant, parconsequent, pour l’ensemble des populations deces deux Pays” (Gruvel 1936: 228, 229).
An early Erythrean invader, the swimming
crab Portunus pelagicus was on sale already by
the early 1900s in the fish market of Haifa (Fox
1924, Calman 1927), and during the British
Mandate the fishermen of Haifa and Acre sold
20 tons of the crab annually (Perlmutter 1956).
By mid-century the Erythrean fishes were an
important part of the Levantine fisheries. Insofar
as the Israeli fishing grounds were concerned,
the bulk of the trawler catch from 1950 to 1955,
was comprised of three species – the native redbream, Pagellus erythrinus (Linnaeus, 1758),
and hake, Merluccius merluccius, and theErythrean yellow striped mullet, Upeneus
moluccensis. The latter were fished commercially in the early 1940s only along the southerncoast of Israel, but by 1946-1947 were found allalong the coast (Gottlieb 1957), and by the late
1940s constituted an estimated 10-15% of thetotal mullid catch (Wirszubski 1953). In 1955
Israeli fishermen noticed greater numbers of theyellow striped mullet, and data assembled by the
Sea Fisheries Research Station, Haifa, indicatedthat their percentage in the mullid catch rose to20%, and to over 83% in early 1956 (Oren
1957a,b), and was considered “the most important commercial fish in the Israel trawl catches”(Gottlieb 1957:20). Since the total mullid catchhad remained constant, the yellow striped mullet
had in the early 1950s “almost completely
replaced the Mediterranean species, the redmullet, Mullus barbatus in the trawl catch”(Perlmutter 1956:4). In 1955, another Erythreanalien, the lizardfish, Saurida undosquamis
became an important part of the trawl catch289

B.S. Galil(Oren 1957b). In 1953 it was first recorded from
the Mediterranean coast of Israel (Ben Tuvia
1953a,b) as much rarer than the native Mediterranean lizardfish, Synodus saurus (Linnaeus,
1758). Within two years commercial catches
increased steadily, in November and December
1955, 22 and 27.5 tons respectively were taken,swelling to 40 and 46.8 tons in January andFebruary 1956 respectively, to a total of 266.5
tons for 1956 – 20% of the total annual trawl
catch (Oren 1957a, b). The sudden increase in
the populations of the lizardfish, the yellow
striped mullet, the red soldierfish, and Erythrean
penaeids was attributed to a rise of 1-1.5ºC in
sea temperature during the winter months of1955 (Ben Yami 1955, Chervinsky 1959).
Examination of the Israeli fisheries statistics
since the mid 1980s underscores the growing
prominence of the Erythrean aliens. The
Erythrean conch, Conomurex persicus Swainson,
1821, and on occasion the Erythrean spiny
oyster, are served in seafood restaurants in
Israel. Erythrean penaeid prawns make up mostof the shrimp catches along the SE Levantine
coasts. The Erythrean prawns, in particular
Marsupenaeus
japonicus(Bate,
1888),
Metapenaeus monoceros (Fabricius, 1798) andPenaeus semisulcatus de Haan, 1844, are highly
prized and beginning in the 1970s a shrimp
fishery developed off the Sinai coast, and since
the mid 1980s off the Israeli coast where a small
fleet of coastal “mini” trawlers has specializedin shrimping, bringing in a quarter of the total
trawl catch volume and a third of the trawl grossincome (Pisanty and Grofit 1991, Snovsky andShapiro 1999). Nearly half of the trawl catches
along the Israeli coast consist of Erythrean fish
(Golani and Ben Tuvia 1995). The dominantfishes in the inshore fisheries (trammel-netting
and hook-and-lining) are the siganids S.
rivulatus and S. luridus, the obtuse barracuda
Sphyraena chrysotaenia Klunzinger, 1884, andthe Erythrean jack, Alepes djedaba (Forsskål,1775). The above species, together with Sillagosihama (Forsskål, 1775) and S. commerson, two
species that underwent population explosion in
the early 1980s, are common in purse-seine
landings. The annual catch of the Erythrean
lizardfish which reached 400 tons in 1960 soonafter its arrival (see above), declined to 100 tons
in the mid 1960s, but has since increased, andcatch fluctuations are correlated with CPUE.
Catch statistics for mullids do not distinguishbetween the natives, M. barbatus and M.
surmuletus Linnaeus, 1758, and the Erythrean290

aliens Upeneus moluccensis and U. pori, but astudy of the frequency of the latter in trawl
catches conducted in the mid 1980s showed they
formed 87% of the mullid catch off the coast of
Israel at depths of 20 m, and 50% at 55 m,
whereas the native mullids are more abundant indeeper waters (Golani and Ben Tuvia 1995). The
percentage of the Erythrean mullids in the totalmullid catch has been increasing steadily, from
30% in 1980, 42% in 1984, to 47% in 1989
(Golani and Ben Tuvia 1995). Similarly, catch
statistics of sphyraenids do not separate the Red
Sea obtuse barracuda from the native Mediterranean species S. sphyraena (Linnaeus, 1758)
and S. viridensis Cuvier, 1829. However, examination of the landed catch showed that theErythrean barracuda had outnumbered the native
sphyraenids in inshore trawl and purse-seine
catches (Grofit 1987). In addition, two of thefour species of Erythrean clupeids that established populations in the Levant – Dussumieria
elopsoides Bleeker, 1849, and Herklotsichthys
punctatus (Rüppell, 1837) – are of importance in
the inshore-pelagic fishery. The increasing
exploitation of Erythrean aliens meant theshifting of the trawling grounds nearshore since
their densest populations occur at depths up to
50 m. Between 1980 and 1986 the Israeli
trawlers doubled their activity (measured as
fishing hours) in shallow waters (Pisanty and
Grofit 1991). The shoreward displacement of thefishing grounds coupled with the inexorable gainof Erythrean aliens raise the ratio of alien tonative taxa in the Levantine trawl landings.
But together with the commercially exploitable species, the Erythrean invasion swept
ashore the scyphozoan jellyfish, Rhopilema
nomadica Galil, 1990. Each summer since themid 1980s huge swarms of the Erythreanjellyfish have appeared along the Levantine
coast. These planktotrophic swarms, somestretching 100 km long, must play havoc withthe limited resources of this oligotrophic sea,
and when the shoals draw nearer shore, they
adversely affect tourism, fisheries and coastal
installations. Local municipalities report adecrease in holiday makers frequenting thebeaches because of the public’s concern over thepainful stings inflicted by the jellyfish. The local
newspapers and TV news report during thesummer months the presence of jellyfish along
the beaches. Coastal trawling and purse-seine
fishing are disrupted for the duration of theswarming due to net clogging and inability tosort yield “It is not uncommon that fishermen,

Alien species along the Mediterranean coast of Israel

especially purse seines, discard entire hauls dueto the overwhelming presence of poisonous
medusae in their nets” (Golani and Ben Tuvia
1995: 287). Jellyfish-blocked water intake pipes
pose a threat to cooling systems of port-boundvessels and coastal power plants: in the summerof 2001 Israel Electric removed tons of jellyfish
from its seawater intake pipes at its two largest
power plants, at estimated costs of 50,000 US$(M. Cohen, pers. comm.). Yet, the jellyfish
shelters among its nematocyst-laden tentacles
the juveniles of the Red Sea carangid fish,
Alepes djedaba (Galil et al. 1990), and may haveprecipitated the sudden population increase ofthis commercially important species (Grofit1987).
The recent spread of the silver stripe blaasop,
Lagocephalus sceleratus (Gmelin, 1789), andthe striped catfish, Plotosus lineatus (Thunberg,1787), pose severe health hazards. The blaasop’s
internal organs, and in particular the gonads
during the spawning season, contain a strong
paralytic neurotoxin. In the Suez City, on the
Red Sea, eight fatalities from tetrodoxin
poisoning associated with eating the fish havebeen described recently (Zaki 2004). Several
cases of poisoning were reported from Israel, but
none proved fatal. Injuries caused by the barbed
and venomous first dorsal spine and pectoral
spines of the striped catfish may produce pain
levels requiring hospitalization – injuries havebeen reported by local professional and amateur
fishermen.
The miners’ canary
Evidence is accumulating that changes inbiodiversity patterns in the Mediterranean arelinked to direct drivers such as climate change
and invasive species, in addition to the wellestablished drivers of habitat change, overexploitation and pollution. By the middle of the
century, climate change and invasive species
may be the dominant direct drivers of biodiversity loss and increased risk of extinction formany species, especially those already at risk
due to low population numbers, restricted or
patchy habitats, and limited climatic ranges.
The Erythrean aliens are thermophilic, originatingin tropical waters. It stands to reason that rising seawater temperature enhances the reproduction,
growth, and survival of the Erythrean aliens, andprovides them with a distinct advantage over thenative Mediterranean biota (Galil 2007). The

location of the Israeli coast “downstream” of theprevailing current from the opening of the Suez
Canal means that it is the first haven for theErythrean propagules. Most of the Erythrean aliens
known from the Mediterranean Sea have been firstrecorded from Israeli coast.
As both processes – sea surface temperature riseand influx of aliens – continue unabated, it isimperative to expand research into those issues. The
likely biotic change may impact the already teetering
fisheries, mariculture, and tourism through proliferation of alien parasitic, noxious and poisonous
species, displacement of commercially-importantnative species, or through alteration of the food web
and by causing phase shift in coastal ecosystems andchanging seascape patterns.
The profound changes in the biota off theIsraeli Mediterranean coast caused by the
intrusion of Erythrean aliens should serve as aminers’ canary - if unheeded and unchecked they
are likely to spread beyond the easternmost
Mediterranean Sea.
Acknowledgements
I am very grateful to my colleagues M. Gorenand H. Mienis for sharing with me unstintingly
their knowledge of alien marine species off theIsraeli Mediterranean coast. I thank Anna
Occhipinti-Ambrogi for greatly improving an
earlier draft of this manuscript. Thanks to E.
Apt, the IOLR librarian, who hunted down someof the “hard to find” literature, and to Limor
Shoval for her invaluable help with the
database.
Galil is a contributing partner in the EU-funded
Programme Delivering Alien Invasive Species
Inventories for Europe (DAISIE, SSPI-CT-2003511202, see also www.europe-aliens.org).
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Alien species along the Mediterranean coast of Israel

Annex
Aquatic alien species reported from the coastal waters of Israel
Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Galaxaura rugosa (Ellis &Solander) Lamouroux, 1816

2003, Shikmona

alien

Pantropicalvessels

unknown

Hoffman et al.
2007

Hypnea cornuta (Kützing) J.
Agardh, 1851

unknown

alien

Atlantic
Ocean;Indian
Ocean

Suez Canal

unknown

Nemlich andDanin, 1964

Hypnea esperi Bory de SaintVincent 1828

unknown

alien

Indian Ocean

Suez Canal

unknown

Lipkin 1972

unknownalien

PantropicalSuez Canal

unknown

Gomez Garreta etal. 2001

1945, Tel Aviv

alien

Indo PacificSuez Canal

unknown

Rayss 1963

1957, Bat Yam

alien

Indo PacificSuez Canal

unknown

Rayss 1963

alien

PantropicalSuez Canal

unknownRamon andFriedman, 1965

alien

Atlantic
Ocean;Indian
Ocean

Suez Canal

unknown

Lundberg 1980 fide
Verlaque andBoudouresque
1991

Acetabularia calyculus
Lamouroux in Quoy & Gaimard, 1943, Haifa Bay
1824

alien

PantropicalSuez Canal

unknown

Rayss 1955

Caulerpa mexicana Sonder ex
Kützing, 1849

1939, Tel Aviv

alien

PantropicalSuez Canal

unknown

Rayss 1941

Caulerpa racemosa var
lamourouxii f. requienii(Montagne) Weber-van Bosse

1955, Haifa Bay

alien

Red SeaSuez Canal

unknown

Rayss and
Edelstein 1960

Caulerpa scalpelliformis(Brown ex Turner) C. Agardh,
1817

1929, Tel Aviv

alien

Cosmopolitan

Suez Canal

unknown

Carmin 1934

Cladophoropsis javanica(Kützing) P.C. Silva, 1996

unknown

alien

Indo PacificSuez Canal

unknown

Rayss 1955

Codium taylorii P.C. Silva,
1960

1958, Bat Yam

alien

Atlantic
Ocean;Indian
Ocean

Suez Canal

unknown

Silva 1960

1999, Hadera

alien

Red Sea

Vessels

unknown

Fine et al. 2005

Oculina patagonica de Angelis,
1993, Sdot Yam
1908

alien

Atlantic coastof SouthAmerica

Vessels

unknown

Fine and Loya
1995

Taxa
Macrophytes
Rhodophyceae

Lophocladia lallemandii(Montagne) F. Schmitz, 1893
Sarconema filiforme (Sonder)
Kylin, 1932
Solieria dura (Zanardini) F.
Schmitz, 1895
Phaeophyceae
Padina boergesenii Allender &Kraft, 1983
Stypopodium schimperi(Buchinger ex Kützing)
Verlaque & Boudouresque,1991

before 1965,
Mikhmoret
1973,
Mikhmoret

Chlorophyceae

Cnidaria
Hydrozoa
Acabaria erythraea Ehrenberg,
1834

299

B.S. Galil

Taxa
Anthozoa
Cassiopeia andromeda(Forsskål, 1775)
Phyllorhiza punctata vonLendendfeld, 1884
Rhopilema nomadica Galil,
1990

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

1988, Neve Yam

alien

Indo PacificSuez Canal

unknown

Galil et al. 1990

1965, Beit Yanai

alien

Indo PacificSuez Canal

unknown

Galil et al. 1990

Venomous,interferes withfisheries, coastal Galil et al. 1990
installations,seabathing

1976, Doralien

Indian Ocean

Suez Canal

Glyphidohaptor plectocirra(Paperna, 1972)

1971, unknown

alien

Red SeaSuez CanalparasitePaperna 1972

Hysterolecitha sigani Manter,
1969

1977, Haifa

alien

Indo PacificSuez Canalparasite

Fischthal 1980

Lecithochirium magnicaudatum
Fischthal & Kuntz, 1963

1977, Jaffa

alien

Red SeaSuez Canalparasite

Fischthal 1980

Monilicaecum ventricosum
Yamaguti, 1942

1977, Jaffa

alien

Indo PacificSuez Canalparasite

Fischthal 1980

Polylabris cf. mamaevi Ogawa& Egusa, 1980

1971, unknown

alien

Indo PacificSuez Canalparasite

Diamant 1989

Tetrancistrum polymorphus(Paperna, 1972)

1971, unknown

alien

Red SeaSuez CanalparasitePaperna 1972

Tetrancistrum strophosolenum
Kritsky, Galli & Yang, 2007

1971, unknown

alien

Red SeaSuez CanalparasitePaperna 1972

Tetrancistrum suezicus(Paperna, 1972)

1971, unknown

alien

Red SeaSuez CanalparasitePaperna 1972

Branchiomma boholense Grube,
1878

1927, unknown

alien

Indo PacificSuez Canal

unknown

Knight-Jones andErgen 1991

Ceratonereis mirabilis Kinberg,
1866

1974-5, Haifa
Bay

alien

PantropicalSuez Canal

unknown

Amoureux 1976

Eunice cf. indica Kinberg, 1865

1972, unknown

alien

Indo Pacific;
Red SeaSuez Canal

unknown

Ben Eliahu 1972b

Ficopomatus enigmaticus(Fauvel, 1923)

1954, swamp
near Akko,
Alexander river.

alien

southern
Hemisphere

Vesselsfouling

Fauvel 1955

Hydroides brachyacantha Rioja,
1941

1933, Jaffa

alien

Red SeaSuez Canal,
Vesselsfouling

Ben Eliahu 1991

Hydroides elegans (Haswell,1883)

1965-6, Haifa
port

alien

Pantropical

Vesselsfouling

Queiroz, 1968

Hydroides heterocerus (Grube,
1868)

1990, Akhziv

alien

Indian Ocean

Suez Canal,
Vesselsfouling

Ben Eliahu and ten
Hove 1992

Hydroides homoceros Pixell,1913

1955, Haifa

alien

Indian Ocean

Suez Canal,
Vesselsfouling

Ben Eliahu 1991

Platyhelminthes

Annelida
Polychaeta

300

Alien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Hydroides minax (Grube, 1878)

1990, Rosh
Hanikraalien

Indo PacificSuez Canal,
Vesselsfouling

Ben Eliahu 1991

Hydroides operculatus(Treadwell, 1929)

1959, Haifa

alien

Indian Ocean

Suez Canal,
Vesselsfouling

Ben Eliahu 1991

Leonnates decipiens Fauvel,
1929

1980-9, Haifa

alien

Indian Ocean

Suez Canal

unknown

Ben Eliahu 1989

Leonnates indicus Kinberg,1865

1991, off Haifa

alien

Indian Ocean

Suez Canal

unknown

Ilan et al. 1994

1937, Haifa

alien

Indian Ocean

Suez Canal

unknown

Ben Eliahu 1989

Lepidonotus tenuisetosus
Gravier, 1901

1969, Ashkelon

alien

Indo PacificSuez Canal

unknown

Barnich and Fiege
2003

Linopherus incarunculataPeters, 1858

before 1976,
unknownalien

unknown

Suez Canal

unknown

Ben Eliahu 1976

Metasychis gotoi Izuka, 1902

before 1972,
unknown

alien

Indo PacificSuez Canal

unknown

Ben Eliahu 1972b

Nereis persica Fauvel, 1911

1937, Haifa

alien

PantropicalSuez Canal

unknown

Ben Eliahu 1989

Pista unibranchia Day, 1963

1974-5, Haifa
Bay

alien

Arabian Gulf

Suez Canal

unknown

Amoureux 1977

Pomatoleios kraussii (Baird,1865)

1958, Hadera

alien

Indo PacificSuez Canal,
Vessels

unknown

Ben Eliahu 1992

Pseudonereis anomala Gravier,
1901

1937, Haifa

alien

Red Sea;
Arabian Gulf

Suez Canal

unknown

Ben Eliahu 1989

Terebella ehrenbergi Grube,
1870

1976, unknown

alien

Indo PacificSuez Canal

unknown

Ben Eliahu 1977

2003, Hadera
harbouralien

tropical andwarmtemperate seas

Vesselsfouling

F. Kerckhof pers.
com.

alien

Red SeaSuez Canalalien

Indo-PacificSuez Canal

unknown

Berdugo 1974

1965, Haifa

alien

Indian Ocean

Suez Canal

unknown

Berdugo 1968

before 1959,
unknownalien

CircumtropicalSuez Canal,
vessels

unknown

Ruffo 1959

before 1979,
Ashdod harbour

alien

Cosmopolitan

vessels

unknownkrapp-Schickel andMyers 1979

Photis lamellifera Schellenberg,
1951-2, Cesarea
1928

alien

Indian Ocean

Suez Canal

unknown

Ruffo 1959

Leonnates persica WesenbergLund, 1948

Crustacea
Cirripedia
Balanus reticulatus Utinomi,
1967

Heterosaccus dollfusi Boschma,1990, Palmahim
1960
Copepoda
Acartia centrura Giesbrecht,
1968, Netanya
1889
Calanopia media Gurney, 1927

parasite on Ch. Galil and Lützen
longicollis
1995

Amphipoda
Elasmopus pectenicrus (Bate,1862)
Gammaropsis togoensis
Schellenberg, 1925

Cumacea
Eocuma rosae Corbera & Galil,
2007

2005, Haifa Bay

singlerecordRed SeaSuez Canal

unknown

Corbera and Galil,
2007

Scherocumella gurneyi(Calman, 1927)

2005, Haifa Bay

alien

Red SeaSuez Canal

unknown

Corbera and Galil
2007

301

B.S. Galil

Date andlocation of firstrecordStatusNative range

Vector/Pathway

1951, Herzeliya

alien

Indo West
Pacific

Suez Canaldisplaced native Forest and Guinot
species
1956

Alpheus inopinatus Holthuis &Gottlieb, 1958

1951, Herzeliya

alien

Indian Ocean

Suez Canal

Forest and Guinot
displaced native
1958, Holthuis andspecies
Gottlieb 1958

Alpheus migrans Lewinsohn &Holthuis, 1978

1977, Palmahimalien

Red SeaSuez Canal

Alpheus rapacida de Man, 1908

1960, Tel Aviv

alien

Indo West
Pacific

Suez Canal

Ashtoret lunaris (Forsskål,1775)

1987, Haifa Bay

singlerecordIndo West
Pacific

Suez Canal

unknown

Galil and Golani
1990

Atergatis roseus (Rüppell,1830)

1961, Tel Aviv

alien

Indo PacificSuez Canal

unknown

Lewinsohn andHolthuis 1964

Callinectes sapidus Rathbun,
1896

before 1955,
unknownalien

Western
Atlantic

vessels

unknown

Holthuis andGottlieb 1955

Carupa tenuipes Dana, 1851

2002, Tel Aviv

alien

Indo PacificSuez Canal

unknown

Galil 2004

Suez Canal

unknown

Steinitz 1929

Taxa
Decapoda
Alpheus audouini Coutiere,
1905

Impactunknown

References

Lewinsohn andHolthuis 1978displaced native Lewinsohn andspeciesHolthuis 1964

Charybdis helleri (A. Milne
Edwards, 1867)

1925, Haifa Bay

alien

Indo West
Pacific

Charybdis longicollis Leene,
1938

1961, Tel Aviv

alien

IndianOcean;Persian
Gulf

Suez Canalnuisance tofisheries

Lewinsohn andHolthuis 1964

Coleusia signata (Paulson,
1875)

1953, Bat Yam

alien

Indo West
Pacific

Suez Canal

unknown

Holthuis 1956

Dorippe quadridens Fabricius,1793

2004, Ashdod

alien

Indo West
Pacific

Suez Canal

unknown

Galil 2005

Eucrate crenata de Haan, 1835

1994, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Galil 1997

Halimede tyche (Herbst, 1801)

1998, Palmahim

singlerecordIndo West
Pacific

Suez Canal

unknown

Galil 2000

Hyastenus hilgendorfi de Man,
1887

1960, Palmahim

alien

Indo West
Pacific

Suez Canal

unknown

Lewinsohn andHolthuis 1964,
Galil 2006

Ixa monodi Holthuis & Gottlieb,
1977, Palmahim
1956

alien

Red SeaSuez Canal

unknown

Galil and Golani
1990

Leptochela pugnax de Man,
1916

1947, Tel Aviv

alien

Indo West
Pacific

Suez Canal

unknown

Holthuis andGottlieb 1958

Macrophthalmus graeffei A.
Milne Edwards, 1873

2002, Tel Aviv,
Ashquelon

alien

Indo West
Pacific

Suez Canal

unknown

Ksiunin and Galil
2004

Marsupenaeus japonicus (Bate,
1888)

before 1951,
unknown

Melicertus hathor (Burkenroad,
1997, Haifa Bay
1959)
Metapenaeopsis aegyptia Galil
1987, Palmahim& Golani, 1990
Metapenaeopsis mogiensis
1996, Palmahim
consobrina (Nobili, 1904)
Metapenaeus monoceros(Fabricius, 1798)

302

before 1951,
unknowncommerciallyvaluable;
Gottlieb 1953displaced native
speciesalien

Indo PacificSuez Canal

alien

Indian Ocean

Suez Canal

unknown

Galil 1999

alien

Indo PacificSuez Canal

unknown

Galil and Golani
1990

alien

Indo West
Pacific

Suez Canal

unknown

Galil 1997

alien

Indo West
Pacific

Suez Canalcommerciallyvaluable;
Gottlieb 1953displaced native
speciesAlien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordStatusNative range

Vector/Pathway

Metapenaeus stebbingi (Nobili,
1904)

1958, Haifa Bay

alien

Indian Ocean

Suez Canal

Myra subgranulata Kossmann,
1877

1929, Jaffa

alien

Western IndianSuez Canal
Ocean

Notopus dorsipes (Linnaeus,
1758)

1962, Atlit

singlerecordIndo West
Pacific

Ogyrides mjoebergi (Balss,
1921)

1947, Cesarea

alien

Palaemonella rotumana(Borradaile, 1898)

1948, Haifa Bay

Panulirus ornatus (Fabricius,
1798)
Penaeus semisulcatus de Haan,
1844

ImpactReferences

commerciallyvaluable;
Lewinsohn anddisplaced native Holthuis 1964
speciesunknown

Monod 1930

Suez Canal

unknown

Lewinsohn andHolthuis 1964

Indo West
Pacific

Suez Canal

unknown

Holthuis andGottlieb 1958

alien

Indo West
Pacific

Suez Canal

unknown

Holthuis andGottlieb 1958

1988, Haifa Bay

singlerecordIndo West
Pacific

Suez Canal

unknown

Galil et al. 1989

before 1951,
unknown

alien

Indo West
Pacific

Suez Canal

1951, Haifa Bay

alien

Western IndianSuez Canal
Ocean

Portunus pelagicus (Linnaeus,
1758)

before 1924,
unknown

alien

Indo PacificSuez CanalcommerciallyvaluableThalamita poissonii (Audouin,
1826)

1952, Bat yam

alien

Indo West
Pacific

Suez Canal

unknown

Holthuis andGottlieb 1958

Trachysalambria palaestinensis(Steinitz, 1932)

1924-1925,
probably Haifa
Bay

alien

Red SeaSuez Canal

unknown

Steinitz 1929

Clorida albolitura Ahyong &Naiyanetr, 2000

2006, Ashdod

alien

Indo West
Pacific

Suez Canal

unknown

Ahyong and Galil
2006

Erugosquilla massavensis(Kossmann, 1880)

before 1938,
Khan Yunes

alien

Red Sea;
Persian Gulf

Suez Canal

Anoplodactylus californicus
Hall, 1912

1951, Cesarea

alien

Subcosmopolit
an

Vessels

unknown

Stock 1958

Anoplodactylus digitatus(Buhm, 1879)

1951, Cesarea

alien

Indo West
Pacific

Suez Canal

unknown

Stock 1958

Pigrogromitus timsanus
Calman, 1927

1960,
Mikhmoret

alien

Circumtropical

Vessels

unknown

Lipkin and Safriel
1971

Celleporaria aperta Hincks,
1882

1968, Acrealienpantropical

Vessels

unknown

Powell 1969

Reteroporella jermanensis(Waters, 1909)

1974, Haifa Bay

alien

Red SeaSuez Canal,
Vessels

unknownd’Hondt 1988

Scrupocellaria jolloisi(Audouin, 1826)

before 1988,
unknownalien

Red SeaSuez Canal

unknownd’Hondt 1988

Smittina malleolus (Hincks,
1884)

1985, Haifa

alien

circumtropical

Vessels

unknownd’Hondt 1988

Pilumnopeus vauquelini(Audouin, 1826)

commerciallyvaluable;
Gottlieb 1953displaced native
speciesunknown

Holthuis 1956
Fox 1924

Stomatopoda

displaced native
Steuer 1938
species

Pycnogonida

Ectoprocta303

B.S. Galil

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpact1934, unknown

alien

Indo West
Pacific

Suez Canal

unknown

Barash 1974

Acteocina mucronata (Philippi,
1849)

1986, Haifa Bay

alien

Red SeaSuez Canal

unknown

van Aartsen et al.
1990

Amathia tricarinata (Linnaeus,
1767)

2005, ShiqmonaalienIndo PacificSuez Canal

unknown

Mienis 2006b

Angiola punctostriata (Smith
E.A., 1872)

1950, DorsinglerecordRed SeaSuez Canal

unknown

Mienis 1980b

Atys angustatus Smith, 1872

1974, Haifa Bay

alien

Red SeaSuez Canal

unknown

van Aartsen and
Goud 2006a

Atys cylindricus Helbling, 1779

2002, Shikmona

singlerecordIndo PacificSuez Canal

unknown

Mienis 2004a

Bulla ampulla Linnaeus, 1758

1978, Ga’ash

alienIndo PacificSuez Canal

unknown

Barash and Danin
1982

Bursatella leachi De Blainville,1817

before 1939,
unknownalien

circumtropicalSuez Canal

unknown

O'Donoghue andWhite 1940

Caloria indica (Bergh, 1896)

1986, Ashqelon

alien

Indo PacificSuez Canal

unknown

Gat 1993

Canarium mutabilis Swainson,
1821

1991, HaBonim

alien

Indo West
Pacific

Suez Canal

unknown

Mienis 2001b

Cellana rota (Gmelin, 1791)

1961, Acre

alien

Indian Ocean

Suez Canal

Cerithidium diplax (Watson,
1886)

1961, Bat Yam

alien

Indo PacificSuez Canal

unknown

van Aartsen 2006

Cerithidium perparvulum(Watson, 1886)

2004, Rosh
HaNikraalien

Indo PacificSuez Canal

unknown

Bogi and Galil
2006

Cerithiopsis pulvis (Issel, 1869)

1978, Doralien

Red SeaSuez Canal

unknown

Mienis 1992

Cerithiopsis tenthrenois(Melvill, 1896)

1982, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

van Aartsen et al.
1989

Cerithium columna Soweby,1834

1966, Cesarea

singlerecordIndo PacificSuez Canal

unknown

Mienis 2003a

Cerithium egenum Gould,1849

1971, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Mienis 2001a

Cerithium nesioticum Pilsbry&Vanatta, 1906

1971, Shikmona

alien

Indo West
Pacific

Suez Canal

unknown

Mienis 1977

Cerithium scabridum (Philippi,
1848)

1891, Jaffa

alien

Red Sea,
Arabian Sea

Suez Canal

Chelidonura fulvipunctata
Baba, 1938

1986, Ashkelon

alien

Indo PacificSuez Canal

unknown

Mienis and Gat
1987

Chrysallida fischeri (Hornung& Mermod, 1925)

1974, Haifa Bay

alien

Red SeaSuez Canal

unknown

van Aartsen andCarroza 1979

Chrysallida maiae (Hornung &Mermod, 1924)

1935, Atlit

alien

Red SeaSuez Canal

unknown

van der Linden andEikenboom 1992

Taxa

References

Mollusca
Polyplacophora
Chiton hululensis (Smith E.A in
Gardiner, 1903)
Gastropoda

304

displaced native
Christiaens 1967
species

displaced native
Hart 1891
speciesAlien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Chrysallida pirintella (Melvill,
1910)

1984, Haifa Bay

alien

Red SeaSuez Canal

unknown

van Aartsen et al.
1989

Cingulina isseli (Tryon, 1886)

1980, Haifa Bay

alien

Atlantic
Ocean, Indian
Ocean

Suez Canal

unknown

van Aartsen andCarroza 1983

Clypeomorus bifasciatus(Sowerby G.B II, 1855)

1983, Akhziv

alien

Indo West
Pacific

Suez Canal

unknown

Mienis 1985a

Conomurex persicus Swainson,
1821

1983, Shikmona

alien

Persian Gulf,
Arabian Gulf

Suez Canal

unknown

Barash and Danin
1986

Dendrodoris fumata (Rüppell &Leuckart, 1830)

1980, Doralien

Red SeaSuez Canal

unknown

Barash and Danin
1986

Diala varia Adams A., 1860

1935, Jaffa

alien

Indo West
Pacific

Suez Canal

unknown

Mienis 1984

Diodora funiculata Reeve, 1850 1998, Palmahim

alien

Indo PacificSuez Canal

unknown

Mienis 2002b

Diodora ruppellii (Sowerby,1834)

before 1948,
Haifa Bayalien

Indo PacificSuez Canal

unknown

Haas 1948

Diplodonta bogii van Aartsen,
2004

1982, Hadera

alien

Red SeaSuez Canal

unknown

van Aartsen 2004

Discodoris lilacina Gould,1852

1974, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1977

Engina mendicaria Linnaeus,
1758

2001, Shikmona

alien

Indo PacificSuez Canal

unknown

Mienis 2004a

Ergalatax contracta Reeve,
1846

2001, Jaffa

singlerecordRed SeaSuez Canal

unknown

Mienis 2004a

2004, AkhzivRosh Hanikra

alien

Red SeaSuez Canal

unknown

Mienis 2006

Erosaria turdus (Lamarck,
1810)

1980, Doralien

Indian Ocean

Suez Canal

unknown

Barash and Danin
1986

Finella pupoides Adams A.,
1860

1958, unknown

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1977

Flabellina rubrolineata(O'Donoghue, 1929)

1988, Ashkelon

alien

Indo PacificSuez Canal

unknown

Gat 1993

Fusinus verrucosus (Gmelin,
1791)

1929, Haifa Bay

alien

Red SeaSuez Canal

unknown

Gruvel and Moazzo
1929

Gibborissoa virgata (Philippi,
1849)

1970, HaBonim

alien

Indo PacificSuez Canal

unknown

van Aartsen et al.
1989

Haliotis pustulata cruenta
Reeve, 1846

1961, Shikmona

alien

Red SeaSuez Canal

unknown

Talmadge 1971

Hypselodoris infucata (Rüppell& Leuckart, 1830)

1965, Caesarea

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1977

Leucotina natalensis (Adams
A., 1851)

1978, unknown

alien

Indo PacificSuez Canal

unknown

Mienis 1985b

Metaxia bacillum (Issel, 1869)

1978, Doralien

Red SeaSuez Canal

unknown

Mienis 1985b

Monotigma lauta (Adams A.,
1851)

1967, unknown

alien

Indo PacificSuez Canal

unknown

Lavaleye andBarash 1981

Murchisonella columna(Hedley, 1907)

1997, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Bogi and Galil
1999

1932, Haifa

alien

Red SeaSuez Canal

unknown

Settepassi 1967

Ergalatax obscura Houart, 1996

Murex forskoehlii Roding, 1798

305

B.S. Galil

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Nassarius obvelatus (Deshayes,
1968, Haifa Bay
1834)

singlerecordIndian Ocean

Suez Canal

unknown

Barash and Danin
1977

Notocochlis gualteriana(Recluz, 1844)

1966, Caesarea

alien

Indo West
Pacific

Suez Canal

unknown

Mienis 2000b

Oscilla jocosa Melvill, 1904

1984, Haifa Bay

alien

Indo PacificSuez Canal

unknown

van Aartsen et al.
1989

Palmadusta lentiginosa (Gray,
1825)

1990, Tel Aviv

alien

Indian Ocean

Suez Canal

unknown

Mienis 1990

Planaxis griseus (Brocchi,
1821)

1997, unknown

alien

Red SeaSuez Canal

unknown

Gianuzzi-Savelli etal. 1997

Pleurobranchus forskalii
Rüppell & Leuckart, 1828

1975, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1977

Plocamopherus ocellatus
Rüppell & Leuckart, 1828

1977, Nizzanim

alien

Red SeaSuez Canal

unknown

Barash and Danin
1982

Pseudominolia nedyma (Melvill,
1897)

1966, Atlit

alien

Red Sea,
Persian Gulf

Suez Canal

unknown

Barash and Danin
1973

Purpuradusta gracilis notata(Gill, 1858)

1981, Hadera

alien

Indian Ocean

Suez Canal

unknown

Mienis and Singer
1983

Pyrunculus fourierii (Audouin,
1826)

before 1989,
Haifa Bayalien

Indo West
Pacific

Suez Canal

unknown

van Aartsen et al.
1989

Retusa desgenettii (Audouin,
1826)

1997, Haifa Bay;
Ashkelon; Acre

alien

Red SeaSuez Canal

unknown

Bogi and Galil
2002

Rhinoclavis kochi Philippi, 1848 1963, Haifa Bay

alien

Indo West
Pacific

Suez Canal

Rhinoclavis sinensis Gmelin,
1791

2003, Palmahim

singlerecordIndo PacificSuez Canal

unknown

Mienis 2004a

Rissoina bertholleti Issel, 1869

1958, Haifa

alien

Indian Ocean

Suez Canal

unknown

van Aartsen 1963

Rissoina spirata Sowerby, 1825

1997, Haifa

singlerecordIndo PacificSuez Canal

unknown

Gianuzzi-Savelli
1997

Sabia conica (Schumacher,1817) [doubtful]

1980, Doralien

Indo PacificSuez Canal

unknown

Barash and Danin
1986

1965, Shikmona

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1973

Sticteulima cf. Lentiginosa(Adams A., 1861)

2004, Rosh
Hanikraalien

Indo PacificSuez Canal

unknown

Bogi and Galil
2006

Stomega lorioli (Hornung &Mermod, 1924)

1974, Haifa Bay

alien

Red SeaSuez Canal

unknown

van Aartsen 1987

Syrnola fasciata (Jickeli, 1882)

1949, Tantura &Haifa Bayalien

Indo PacificSuez Canal

unknown

Mienis 1995

Syrnola lendix A. Adams, 1863

1994, Palmahim

alien

Indo PacificSuez Canal

unknown

Bogi and Galil
1997

1928, Jaffa

alien

Indian Ocean

Suez Canal

unknown

Mienis 1977

Thais sacellum (Gmelin, 1791)

2004, Segavyon
I.

alien

Indian Ocean

Suez Canal

unknown

Mienis 2006c

Trochus erithraeus Brocchi,
1821

before 1973,
Shikmona

alien

Red Sea,
Persian gulf

Suez Canal

unknown

Barash and Danin
1973

Turbonilla edgarii (Melvill,
1896)

1984, Haifa

alien

Indo PacificSuez Canal

unknown

van Aartsen et al.
1989

Taxa

Siphonaria crenata Blainville,
1827

Thais lacera (Born, 1778)

306

displaced native Barash and Danin
species
1973

Alien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordVexillum depexum (Deshayes in
1964, Akhziv
Laborde, 1834)
Voorwindia tiberiana (Issel,
1997, Haifa Bay
1869)

StatusNative range

Vector/PathwayImpactalien

Red SeaSuez Canal

unknown

Mienis 2004aalien

Red SeaSuez Canal

unknown

Bogi and Galil
1999

References

Zafra savignyi (Moazzo, 1939)

1954, Shavei
Zion

alien

Red SeaSuez Canal

unknown

Mienis 1976b

Zafra selasphora (Melvill &Standen, 1901)

1980, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

van Aartsen 1997

1978, Netanya

singlerecordIndo PacificSuez Canal

unknown

Barash and Danin
1986

Afrocardium richardi (Audouin, 1997, Haifa Bay
1826)

alien

Red Sea,
Arabian Sea

Suez Canal

unknown

Bogi and Galil1999

Alectryonella crenulifera(Sowerby, 1871)

2005, HaderaPortsinglerecordIndian Ocean

Suez Canal

unknown

Sharon et al. 2005

Anadara natalensis (Krauss,1848)

1935, unknown

alien

Indian Ocean

Suez Canal

unknown

Haas 1937

Angulus flacca Romer, 1871

1997, Haifa Bay

singlerecordIndo PacificSuez Canal

unknown

Mienis 2004a

Atactodea glabrata (Gmelin,1791)

1973, Netanya

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1977

Barbatia trapezina Lamarck,1819

2003, Ashkelon

singlerecordIndo Pacificvessels

unknown

Mienis 2004a

Brachidontes pharaonis(Fischer P., 1870)

1935, unknown

alien

Indian Ocean

Suez Canal,
vessels

Chama asperella Lamarck, 1819 2003, Ashkelon

singlerecordIndo Pacificvessels

unknown

Mienis 2004a

Chama aspersa Reeve, 1846

2002, Tel Aviv

alien

Indo PacificSuez Canal

unknown

Mienis 2004c

Chama brassica Reeve, 1846

2003, Haifa Bay

singlerecordIndo Pacificvessels

unknown

Mienis et al. 1993b

Chama pacifica Broderip, 1834

1988, Haifa Bay

alien

Indo PacificSuez Canal

Circenita callipyga (von Born,
1778)

1972, Netanya

alien

Red SeaSuez Canal

unknown

Mienis 2000a

Clementia papyracea (Gray,
1825)

1937, Gaza

alien

Indo PacificSuez Canal

unknown

Haas 1948

Divalinga arabica Dekker &Goud, 1994

1976, Netanya

alien

Red Sea,
Persian Gulf

Suez Canal

unknown

Mienis 1979

Fulvia australis (Sowerby G.B.,
1834)

before 1948,
Haifa Bayalien

Indo PacificSuez Canal

unknown

Haas 1948

Fulvia fragilis (Forsskål in
Niehbur, 1775)

1955, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

Barash and Danin
1973

Gafrarium pectinatum(Linnaeus, 1758)

1984, Palmahim

alien

Indo PacificSuez Canal

unknown

Mienis 1999

Bivalvia
Acar plicata (Dillwyn, 1817)

displaced native
Haas 1937
speciesEngineering
species,
Mienis et al. 1993b
displaced native
species307

B.S. Galil

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

1954, Palmahim

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1973

Glycymeris arabicus (Adams
H., 1871)

1966, DorsinglerecordIndian Ocean

Suez Canal

unknown

Barash and Danin
1977

Hyotissa hyotis Linnaeus, 1758

2003, Ashkelon

singlerecordIndo Pacificvessels

unknown

Mienis 2004a

Isognomon ephippium Linnaeus, 2003, Ashkelon
1758

singlerecordIndian Ocean

vessels

unknown

Mienis 2004a

Laternula anatina (Linnaeus,
1758)

1958, Shikmona

alien

Indo PacificSuez Canal

unknown

Barash and Danin
1973

Leiosolenus hanleyanus Reeve,
1857

2003, Ashkelon

singlerecordIndo Pacificvessels

unknown

Mienis 2004a

Limopsis multistriata (Forsskål,1775)

1965, DorsinglerecordIndo PacificSuez Canal

unknown

Barash and Danin
1977

Mactra lilacea Lamarck, 1818

2001, Acre

alien

Indo PacificSuez Canal

unknown

Mienis 2002c

Mactra olorina Philippi, 1846

before 1973, Tel
Aviv

alien

Red SeaSuez Canal

unknown

Barash and Danin
1973

Malvufundus regulus (Forsskål,1775)

before 1937,
unknown

alien

Indo PacificSuez Canal

unknown

Haas 1937

Modiolus auriculatus (Krauss,1848)

1935, Acre

alien

Indian Ocean

Suez Canal

unknown

Haas 1937

Musculista perfragilis (Dunker, 1960, Bat Yam
1857)

alien

Indian Ocean

Suez Canal

unknown

Barash and Danin
1973

Musculista senhousia (Benson
in Cantor, 1842)

1960, Tel Aviv

alien

NW PacificAquaculture,
vessels

unknown

Barash and Danin
1971

Nanostrea exigua Harry, 1985

2004, Haifa Bay

singlerecordIndo PacificSuez Canal,
vesselunknown

Lubinevsky andMienis 2005

Paphia textile (Gmelin, 1791)

1946, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Haas 1948

Parahyotissa imbricata
Lamarck, 1819

2003, Ashkelon

singlerecordIndian Ocean

vessels

unknown

Mienis 2004a

Pinctada margaritifera(Linnaeus, 1758)

2003, Ashkelon

alien

Indo Pacificvessels

unknown

Mienis 2004a

Pinctada radiata (Leach, 1814)

before 1899,
unknown

alien

Indo PacificSuez Canal

unknown

Monterosato 1899

Planostrea pestigris Hanley,1846

2003, Ashkelon

singlerecordIndian Ocean

vessels

unknown

Mienis 2004a

Plicatula chinensis Morch, 1853 2003, Ashkelon

singlerecordIndian Ocean

vessels

unknown

Mienis 2004a

Pseudochama corbieri (Jonas,
1846)

1963, Jaffa

alien

Red SeaSuez Canal

unknown

Barash and Danin
1973

Redicrice sulcata Gray, 18381970, Ashdod

singlerecordIndo Pacificvessels

unknown

Mienis 2004a

Taxa

Gastrochaena cymbium(Spengler, 1783)

308

Alien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactSeptifer bilocularis (Linnaeus,
1758)

2003, Ashkelon

singlerecordRed Seavessels

unknown

Mienis 2004a

Sphenia ruppellii Adams A.,
1850

1978, Netanya

alien

Red Sea,
Arabian Sea

Suez Canal,
vessels

unknown

Barash and Danin
1986

Spondylus groschi Lamprell &Kilburn, 1995

before 1998,
unknown?

Indian Ocean

Suez Canal,
vessels

unknownLamparell 1998

Spondylus nicobaricus
Schreibers, 1793

2002, Akhziv

alien

Indo Pacificvessels

unknown

Mienis 2004a

Spondylus spinosus Schreibers,1793

1988, Haifa Bay

alien

Indo PacificSuez Canal,
vessels

Theora lubrica Gould, 1861

2006, Haifa Bay

alien

Indo Pacificvessels

Timoclea marica (Issel, 1869)

1997, Palmahimalien

Red SeaSuez Canal

Octopus aegina Gray, 1849

1934, Atlit

alien

Indo PacificSuez Canal

unknown

Mienis 2004a

Octopus cyanea Gray, 1849

1997,
Mikhmoret, Acre

alien

Indo PacificSuez Canal

unknown

Mienis 2003b

Sepia gibba Ehrenberg, 1831

2001, Ashdod

singlerecordRed SeaSuez Canal

unknown

Mienis,unpublished

Sepia pharaonis Ehrenberg,
1831

2003, Rishon
leziyyon to
Shiqmona

alien

Indian Ocean

Suez Canal

unknown

Mienis 2003c

Sepioteuthis lessoniana Lesson,
1830

2004, Akhziv

alien

Indo PacificSuez Canal

unknown

Mienis 2004a

Asterina burtoni Gray, 1840

1953, Haifa

alien

subcosmopolitanSuez Canal

Ophiactis parva Mortensen,
1926

1968, Haifa Bay

alien

Red SeaSuez Canal

unknown

Tom and Galil
1991

Ophiactis savignyi Müller &Troschel, 1842

1948, Atlit

alien

circumtropicalSuez Canal

unknown

Tortonese 1953

Synaptula reciprocans Forskål,1775

1971, Rosh
HaNikraalien

Red SeaSuez Canal

unknown

Cherbonnier 1986

1954, unknown

alien

Indo PacificSuez Canal

unknown

Ben Tuvia 1955

1997, Rosh
Hanikraalien

Indo PacificSuez Canal

unknown

Goren and Galil
1998

Alepes djedaba (Forsskål, 1775) 1924, Haifa Bay

alien

Indo PacificSuez Canal

Aphanius dispar (Rüppell,
1829)

alien

Indian Ocean

Suez Canal

References

Engineering
species,
Mienis et al. 1993adisplaced native
speciesBogi and Galil
unknown
2007
Bogi and Galil
unknown
1999

Cephalopoda

Echinodermata
displaced native
Tortonese 1966
species

Chordata
Chondrichthyes
Himantura uarnak (Forsskål,1775)
Osteichthyes
Abudefduf vaigiensis (Quoy &Gaimard, 1825)

1944, Tel Aviv

Commercially
Steinitz 1927
valuable
Hybridizes with
the native A. Mendelssohn 1947
fasciatus

309

B.S. Galil

Taxa

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Apogon pharaonis Cuvier, 1828

before 1947,
unknown

alien

Indo PacificSuez Canal

unknown

Haas and Steinitz
1947

Atherinomorus lacunosus(Forster in Bloch & Schneider,1801)

1924, Haifa

alien

Indo PacificSuez Canal

Bregmaceros atlanticus Goode& Bean, 1886

2004, PalmahimaliencircumtropicalSuez Canal,
vessels

Callionymus filamentosus
Valenciennes, 1837

before 1953,
unknown

alien

Indo PacificSuez Canal

Chilomycterus spilostylus Leis
& Randall, 1982

1992, Ashdod

alien

Indo PacificSuez Canal

unknown

Golani 1993

Crenidens crenidens (Forsskål,1775)

before 1992,
unknown

alien

Indian Ocean

Suez Canal

unknown

Golani 1992

Cynoglossus sinusarabici(Chabunaud, 1913)

before 1953,
unknownalien

Red SeaSuez Canal

unknown

Ben Tuvia 1953

2005, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Golani 2006

Dussumieria elopsoides
Bleeker, 1849

before 1949,
unknown

alien

Indian Ocean

Suez Canalcommerciallyvaluable

Lissner 1949

Epinephelus coioides(Hamilton, 1822)

1966, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Ben Tuvia andLourie 1969

Epinephelus malabaricus(Bloch & Schneider, 1804)

1966, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Ben Tuvia andLourie 1969

Etrumeus teres (DeKay, 1842)

1961, Haifa, Tel
Aviv

alien

Cosmopolitan

Suez Canal

Fistularia commersonii Rüppell,1835

2000, Jaffa

alien

Indo PacificSuez Canal

Hemiramphus far (Forsskål,1775)

1924, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Steinitz 1927

Herklotsichthys punctatus(Rüppell, 1837)

before 1943,
unknownalien

Red SeaSuez Canallimitedcommercial
interest

Bertin 1943

Hippocampus fuscus Rüppell,1838

2001, Hadera
Port

alien

Indian Ocean

Suez Canal

unknown

Golani and Fine
2002

Lagocephalus sceleratus(Gmelin, 1789)

2004, Jaffa

alien

Indo West
Pacific

Suez Canalpoisonous

Golani and Levy
2005

Lagocephalus spadiceus(Richardson, 1844)

1951, unknown

alien

Indo PacificSuez Canalpoisonous

Ben Tuvia 1953

Lagocephalus suezensis Clark &1987, Haifa Bay
Gohar, 1953

alien

Red SeaSuez Canalpoisonous

Golani 1993b

Decapterus russelli (Rüppell,
1830)

Commerciallyvaluable in some Norman 1927
countries
unknown

Goren and Galil
2006

displaced native
Ben Tuvia 1953
speciescommerciallyvaluable
limitedcommercial
interest

Whitehead 1963
Golani 2000b

Leiognathus klunzingeri(Steindachner, 1898)

before 1934,
unknownalien

Red SeaSuez Canal

unknown

Liebman 1934

Liza carinata (Valenciennes,
1830)

Before 1970,
Haifa Bayalien

Indian Ocean

Suez Canalcommerciallyvaluable

Ben Tuvia 1971

310

Alien species along the Mediterranean coast of Israel

Taxa

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactReferences

Muraenesox cinereus (Forsskål,1775)

1979, Tel Aviv

singlerecordIndian Ocean

Suez Canal

unknown

Golani and Ben
Tuvia 1982

Nemipterus japonicus (Bloch,
1791)

2005, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Golani and Sonin
2006

Omobranchus punctatus
Valenciennes, 1836

2003, Ashdod

singlerecordIndo PacificSuez Canal,
vessels

unknown

Golani 2004

Oxyurichthys petersi(Klunzinger, 1871)

1982, Tel Aviv

alien

Red SeaSuez Canal

unknown

Ben Tuvia 1983

Papilloculiceps longiceps(Ehrenberg in Valenciennes,
1829)

1986, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

Golani and Ben
Tuvia 1990

Parexocoetus mento(Valenciennes, 1846)

before 1935,
unknown

alien

Indo PacificSuez Canallimitedcommercial
interest

Bruun 1935

Pelates quadrilineatus (Bloch,1790)

1969, unknown

alien

Indo PacificSuez Canal

unknown

Ben Tuvia andLourie 1969

Pempheris vanicolensis Cuvier,1821

1979, Jaffa

alien

Indo PacificSuez Canal

unknown

Ben Tuvia 1985

Petroscirtes ancylodon Rüppell,
1988, Nizzanim
1835

alien

Red Sea,
Persian Gulf

Suez Canal

unknown

Goren and Galil
1989

Platycephalus indicus(Linnaeus, 1758)

before 1953,
unknown

alien

Indo PacificSuez Canal

unknown

Ben Tuvia 1953

Plotosus lineatus (Thunberg,
1787)

2001, AshdodAshkelon

alien

Indo PacificSuez Canalvenomous

Golani 2002

Pomadasys stridens (Forsskål,1775)

1971, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

Ben Tuvia 1977

Pteragogus pelycus Randall,
1981

1991, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

Golani and Sonin
1992

Pterois miles (Bennett, 1803)

1991, Herzeliya

alien

Indian Ocean

Suez Canalvenomous

Golani and Sonin
1992

Rachycentron canadum(Linnaeus, 1766)

1978, Haifa Bay

alien

PantropicalSuez Canal

unknown

Ben Tuvia 1985

Rastrelliger kanagurta (Cuvier,
1967, Haifa Bay
1816)

alien

Indo PacificSuez Canal

unknown

Collete 1970

Rhabdosargus haffara(Forsskål, 1775)

1991, Tel Aviv

alien

Red Sea,
Persian Gulf

Suez Canal

unknown

Golani 1992

Rhynchoconger trewavasae
Ben-Tuvia, 1993

before 1993,
Haifa

singlerecordRed SeaSuez Canal

unknown

Ben Tuvia 1993

Sargocentron rubrum(Forsskål, 1775)

1947, Haifa Bay

alien

Indo PacificSuez Canal

unknown

Haas and Steinitz
1947

Saurida undosquamis(Richardson, 1848)

1952, Haifa Bay

alien

Indo PacificSuez Canal

Scarus ghobban Forsskål, 1775

2001, Shikmona

alien

Indo PacificSuez Canal

Sciaenops ocellatus (Linnaeus,
1766)

1999, Hadera
portsinglerecordW Atlantic
Ocean

Aquaculture

Commerciallyvaluable,
Ben Tuvia 1953displaced native
species
Goren and Aronov
unknown
2002
unknown

Golani and Mires
2000

311

B.S. Galil

Date andlocation of firstrecordStatusNative range

Vector/PathwayImpactScomberomorus commerson
Lacepede, 1800

before 1935,
unknown

alien

Indo PacificSuez Canal

Commerciallyvaluable

Siganus luridus Rüppell, 1828

before 1964,
unknown

alien

Indian Ocean

Suez Canal

Siganus rivulatus Forsskål,1775

1924, Haifa Bay

alien

Red SeaSuez Canal

Silhouetta aegyptia (Chabanaud,
1933)

before 1998,
Ashdodalien

Red SeaSuez Canal

unknown

Sillago sihama (Forsskål, 1775)

1977, unknown

alien

Indo PacificSuez Canalcommerciallyvaluable

Ben Tuvia 1985

Sorsogona prionota (Sauvage,
1873)

1946, unknown

alien

Indian Ocean

Suez Canal

unknown

Haas and Steinitz
1947

Sphyraena chrysotaenia
Klunzinger, 1884

before 1931,
unknown

alien

Indo PacificSuez Canalcommerciallyvaluable

Spicer 1931

Sphyraena flavicauda Rüppell,1838

1991, Jaffa

alien

Indo PacificSuez Canal

unknown

Golani 1992

Spratelloides delicatulus(Bennett, 1831)

1973, Atlit

alien

Indo PacificSuez Canal

unknown

Ben Tuvia 1978

1924, Haifa Bay

alien

Red Sea,
Persian Gulf

Suez Canal

unknown

Steinitz 1927

Tetrosomus gibbosus (Linnaeus,
1987, Shikmona
1758)

alien

Indo PacificSuez Canal

unknown

Spanier and Goren
1988

Torquigener flavimaculosus
Hardy and Randall, 1983

1987, Haifa Bay

alien

Indian Ocean

Suez Canal

unknown

Golani 1987

Tylosurus choram (Rüppell,1837)

2001, Hadera

alien

Indian Ocean

Suez Canal

unknown

Golani and Levy
2005

Upeneus moluccensis (Bleeker,
1855)

before 1947,
unknown

alien

Indo PacificSuez Canal

Upeneus pori Ben-Tuvia &Golani, 1989

before 1953,
unknownalien

Red Sea, Gulfof OmanSuez Canal

unknown

Ben Tuvia 1953

Herdmania momus Savignyi,
1816

1955, Haifa Bay

alien

Indo Pacificvessels

unknown

Peres 1958

Phallusia nigra Savigny, 1816

1952, Caeserea

alien

W Atlantic
Oceanvessels

unknown

Peres 1958

Taxa

Stephanolepis diaspros FraserBrunner, 1940

References

Hornell 1935

LimitedBen Tuvia 1964
commercial
value, venomousCommercially
Steinitz 1927
valuable,
venomous
Golani 1998

commerciallyvaluable,
Haas and Steinitz
displaced native 1947
species

Chordata
Tunicata, Ascidiacea

312