Biological Invasions 3: 201–210, 2001.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Invasion note

The introduced green alga Caulerpa taxifolia continues to

spread in the Mediterranean

Alexandre Meinesz1,∗ , Thomas Belsher2 , Thierry Thibaut1 , Boris Antolic3 ,

Karim Ben Mustapha4 , Charles-François Boudouresque5 , Danièle Chiaverini1 ,
Francesco Cinelli6 , Jean-Michel Cottalorda1 , Aslam Djellouli7 , Amor El Abed3 ,
Carla Orestano8 , Antoni M. Grau9 , Ljiljana Ivesa10 , Andrej Jaklin10 , Habib Langar6 ,
Enric Massuti-Pascual9 , Andrea Peirano11 , Leonardo Tunesi12 , Jean de Vaugelas1 ,
Nevenka Zavodnik10 & Ante Zuljevic3
1
Laboratoire Environnement Marin Littoral, Université de Nice-Sophia Antipolis, 06108 Nice Cedex 2, France;
2
IFREMER, Centre de Brest, Département d’Ecologie côtière, BP 70, 29280 Plouzané Cedex, France;
3
Laboratory of Benthos, Institute of Oceanography and Fisheries, I. Mestrovica 63, 21000 Split,
P.P. 500, Croatia; 4 Institut National des Sciences et des Technologies de la mer (INSTM) 2025 Salammbô,
Tunisia; 5 Centre d’Océanologie de Marseille, UMR CNRS 6540, Campus de Luminy, 13288 Marseille Cedex 9,
France; 6 Dipartimento di Scienze dell’Uomo e dell’Ambiente, Università di Pisa, Via A. Volta, 6, 56121 Pisa,
Italy; 7 Faculté des Sciences de Tunis, Campus Universitaire, 1060 Tunis, Tunisia; 8 Laboratorio di
Ecologia Acquatica, Dipartimento di Scienze Botaniche, Università degli studi di Palermo, Via Archirafi,
38, 90123, Palermo, Italy; 9 Conselleria d’Agricultura i Pesca, C/. dels Foners, 10, 07006 Palma de
Mallorca, Spain; 10 Laboratory for Ecology and Systematics, Center for Marine Research, Obala Giordano
Paliaga 5, 52210 Rovinj, Croatia; 11 ENEA Ente per le Nuove Technologie, l’Energia e l’Ambiente,
Centro Ricerche Ambiente Marino, C.P. 224, 19100 La Spezia, Italy; 12 ICRAM Central Institute for
Applied Marine Research, via di Casalotti, 300, 00166 Rome, Italy;
∗
Author for correspondence (e-mail: meinesz@unice.fr; fax: +33-4-92076849)

Received 3 April 2001; accepted in revised form 27 September 2001

Key words: benthic survey, Caulerpa taxifolia, introduced species, invasive species, Mediterranean Sea

Abstract

The tropical green alga Caulerpa taxifolia in the Mediterranean has spread steadily since its introduction in 1984.
At the end of 2000, approximately 131 km2 of benthos had been colonized in 103 independent areas along 191 km
of coastline in six countries (Spain, France, Monaco, Italy, Croatia and Tunisia). Large regions neighboring the
invaded areas appear favorable to further colonization, and there is thus no reason to believe that spreading will
slow down in the years to come.

Abbreviations: ENEA – Ente per le Nuove Technologie, l’Energia e l’Ambiente (National-Italian-Committee for
Research and Development of New Technologies and Environment); ICRAM – Istituto Centrale per la Ricerca
Scientifica e Technologica Applicata al Mare (Central Institute for Scientific and Technological Research on Sea);
IFREMER – Institut Français de Recherches pour l’Exploitation de la Mer (French Research Institute for Marine
Development); rDNA ITS – Internal Transcribed Spacer (ITS) of ribosomal DNA
202
Introduction
Marine algal invasions have played important roles
in community structure around the world (Ribera and
Boudouresque 1995). A major algal invasion was first
detected in the Mediterranean Sea in 1984 when the
tropical green alga Caulerpa taxifolia (Vahl) C. Agardh
was observed a few meters from the public aquarium
in Monaco, where it was used as a decorative alga and
from where it was accidentally released (Meinesz and
Hesse 1991).
Genetic studies based on sequence analysis of the
rDNA ITS region in a large number of Caulerpa sam-
ples collected both from the Mediterranean and public
aquaria (including samples originating from the algal
cultures cultivated in Monaco) have revealed the exten-
sive homogeneity of this invasive strain, which is now
available worldwide in the aquarist trade (Jousson et al.
1998; Olsen et al. 1998). These genetic studies, in addi-
tion to morphological, biogeographical and taxonomic
considerations (Meinesz et al. 1994; Meinesz and
Boudouresque 1996) confirm the identity of this alga as
C. taxifolia. It is not a cryptic species that has developed
from the warming of Mediterranean waters involving
the subsequent ‘metamorphosis’ of Caulerpa mexicana
(Sonder) J. Agardh (located in Israel and Lebanon,
eastern Mediterranean) as proposed by Chisholm et al.
(1995).
The worldwide availability of the invasive strain of
C. taxifolia through the aquarium trade forewarned of
additional aquarium-related introductions around the
world. In June 2000 an invasive strain of C. taxifolia
was discovered in San Diego and Los Angeles on the
southern California coast. C. taxifolia had not pre-
viously been reported from the Pacific coast of the
Americas (Dalton 2000; Kaiser 2000). Millar (2001)
observed a proliferation of C. taxifolia near Sydney,
Australia, approximately 600 km south of the known
range of autochthonous populations.
Sequence analysis (rDNA ITS region) and DNA fin-
gerprints of invasive, aquarium and natural worldwide
strains of C. taxifolia, and sequence analysis of 16S
rDNA of bacteria associated with C. taxifolia, revealed
that the California strains are identical to those found
in the Mediterranean and in many aquaria, and are very
similar to those collected from Australia (Jousson et al.
2000; Wiedenmann et al. 2001; Meusnier et al. 2001).
These results support the hypothesis that the invasive
strain originated in southern Australia and has been
disseminated via the aquarium trade to several parts
of the world. In order to reduce the risks of additional
invasions, it is now illegal to import or be in possession
of this alga in France, Australia, the USA and certain
regions of Spain.
In light of the discovery of invasive strains of
C. taxifolia in several non-Mediterranean localities,
and considering the global threat it may constitute for
temperate and tropical littoral communities, several
authors have recently attempted to describe the current
status of this alga in the Mediterranean Sea. The use of
qualitative adjectives, such as ‘carpeted’ or ‘infested’,
and older quantitative data, has resulted in confusion
regarding the history of the spread of C. taxifolia in the
Mediterranean Sea (Kaiser 2000; Dalton 2000; Gili
2000; Chisholm et al. 2000). In the present paper, we
clarify the current status and dynamics of the spread of
Caulerpa in the Mediterranean.
Centralized monitoring efforts, within the frame-
work of two European programs (Life DGXI), led to the
regular surveying of the C. taxifolia invasion until 1997
(Meinesz et al. 1997, 1998). Since then, the spread of
the alga has been monitored independently, by means
of standardized methods, by national organizations
and/or research institutes. The observations, emanating
from six Mediterranean countries concerned with the
C. taxifolia invasion, have been brought together here
in an effort to assess current invasion status.
Methods
Several international groups are involved in monitoring
the spread of C. taxifolia in the Mediterranean Sea.
Consequently, standardized methods were established
to measure the spread of this alga consistently on both
spatial and temporal scales (Vaugelas et al. 1999).
Regional or global status can thus be described and
compared from one year to the next. The standard-
ization of colonization characteristics has led to the
identification of three levels of colonization (Table 1
and Figure 1):
Level I is the first stages of colonization, in which
one to several colonies occurring less than 100 m
apart occupy a total surface area less than 1000 m2 .
Under these conditions it is relatively easy to esti-
mate the surface area covered by the alga (referred
to as the covered area) and to delimit the exact
perimeter of the area of dispersal by SCUBA
diving. The area included by the perimeter of the
established colonies is called the ‘affected area’.
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Table 1. Descriptive parameters for each level of colonization of C. taxifolia.

Levels of Descriptive parameters

colonization
Covered Affected Concerned Number of
surface Surace Coastline Surface Coastline colonies

I
Threshold <1000 m2
Parameters + + + − − +
II
Threshold >1000 m2 <10 ha
Parameters − + + − − −
III
Threshold >1000 m2 >10 ha
Parameters − − − + + −

+: applicable, − : not applicable (after Vaugelas et al. 1999).

Figure 1. Theoretical configuration of the three levels of colonization of C. taxifolia. In level III the position of the dozen or hundred main
colonies, involving more than 100 ha and covering much more than 1000 m2 , is not presented (after Vaugelas et al. 1999).

Level II refers to the next step of spreading. It is area less than 10 ha. During this stage of the inva-
characterized by several colonies that occur within sion it is both time-consuming and futile to measure
250 m of each other, that cover a total area of more the covered area as the alga grows and spreads very
than 1000 m2 , and that are dispersed over a surface quickly. The best method to estimate the extent of
204
coverage is to delimit the perimeter that incorpo-
rates all of the colonies. We refer to this value again
as ‘affected area’.
Level III is attained when dozens or hundreds of
colonies of various sizes are dispersed over a sur-
face area greater than 10 ha, with a total cov-
ered area of more than 1000 m2 . At this stage,
it becomes impossible to map with precision the
location of each colony and to measure the cov-
ered areas. Rather, the suggested method is to
estimate the boundary by identifying the periph-
eral colonies. The surface area of this large zone
that incorporates all of the colonies is called the
‘concerned area’.
By summing the ‘affected’ (levels I and II) and
‘concerned’ areas (level III), we obtain an evaluation of
the overall status for a given region in terms of surface
area impacted by the colonization of C. taxifolia.
The linear extent of coastline adjacent to each
affected or concerned area constitutes an alternative
means to measure algal spread. Finally, the num-
ber of independent area of colonization (each area
has one of the three levels of colonization) also rep-
resents an important means of assessing population
status.
Using the methods outlined above, we have
compiled standardized yearly monitoring surveys
based on SCUBA diving and towed video transects
since C. taxifolia was first observed in each
of six affected countries in the Mediterranean.
Because public-awareness campaigns (distribution of
pamphlets and posters) resulted in some inaccurate
reports of C. taxifolia, only those reports that have been
verified by scientific divers are included in the results.
Finally, a brief description of the monitoring program
is given for each country.
Results
At the end of 2000, six countries were affected by
the invasion of C. taxifolia: Spain, France, Monaco,
Italy, Croatia and Tunisia (Table 2 and Figure 2). The
characteristic situation in each country, the changes
in surface area, the number of independent colonized
areas, the linear extent of coastline adjacent to the
colonies, the organization of the monitoring efforts and
the local eradication measures taken (if any) that may
have affected the spread are described here.
France and Monaco
The first discovery of C. taxifolia occurred in 1984 in
Monaco. Six years later, it was observed in France,
6 km east of Monaco (Meinesz and Hesse 1991). By
the end of 2000, 76 independent colonized areas were
inventoried off the coast of these two countries, repre-
senting a concerned area of 3184 ha. Two regions are
impacted with this invasion: the Languedoc-Roussillon
region (with one area (>5000 m2 ) in the harbor of
Saint-Cyprien located 30 km from the Spanish bor-
der), and the Provence-Alpes-Côte d’Azur region (with
the remaining areas dispersed along about 300 km of
coastline between the Italian border and Six-Fours-les-
Plages, 20 km west of Toulon).
Mapping efforts (by SCUBA diving) have been
made yearly since 1990 and annual mapping cam-
paigns using towed video cameras have been performed
by IFREMER in an attempt to assess the spread
of the alga in those areas deemed to be vulnera-
ble to high amplitude spread (Belsher et al. 1994).
During 1999–2000 a national Caulerpa survey pro-
gram was implemented by the French government,
which included public awareness campaigns and sur-
veys. Since the beginning of these surveys all of
the information generated has been compiled at the
University of Nice-Sophia Antipolis and has been
the subject of a number of publications and reports
describing the dynamics of the invasion (Meinesz et al.
1998).
All information gathered since 1990 for France has
been incorporated into a searchable database avail-
able on the Internet (http://www.caulerpa.org). Reg-
ular control of the algae only occurs in the waters of
the National Park of Port-Cros (under the aegis of the
Ministry of the Environment) where surveys and eradi-
cation efforts such as manual removal or the application
of a cloth soaked in copper salts, have been performed
annually since 1994 (Cottalorda et al. 1996; Robert
1996; Robert and Gravez 1998). In addition, 15 isolated
colonies measuring less than 5 m2 have been eradi-
cated successfully since the beginning of the invasion
through private initiatives.
Spain
C. taxifolia was first observed in Spain in 1992 at
Mallorca. At this site, 250 m2 were covered by the alga
at an anchoring site located in the vicinity of a port.
By the end of 2000, 62.8 ha were colonized in four
 205

Table 2. Status of C. taxifolia invasion in the Mediterranean Sea.

Year Spain France and Monaco Italy Croatia Tunisia Total

A C L A C L A C L A C L A C L A C L
ha n km ha n km ha n km ha n km ha n km ha n km
1989 0 0 0 1 1 0.2 0 0 0 0 0 0 0 0 0 1 1 0.2
1991 0 0 0 30 9 3.5 0 0 0 0 0 0 0 0 0 30 9 3.5
1994 2 1 0.1 1383 13 21.7 153 9 8.9 0 0 0 0 0 0 1538 23 30.7
1997 30 2 2.6 1882 52 34 2704 17 43.6 13 8 2 0 0 0 4629 79 82.2
2000 63 4 9.4 3184 76 70.8 9415 14 103.7 41 8 5.4 350 1 2 13053 103 191.3
A – concerned areas (ha); C – number of colonies (n); L – concerned linear coastline (km) 0 – no data available.

Figure 2. Overall status of C. taxifolia in the Mediterranean Sea at the end of 2000.

different areas along the eastern coast of the island of
Mallorca, between Portocolom and Portopetro (these
two localities being separated by 15 km). These areas
are regularly surveyed, and all information concerning
the spread is kept in a centralized database (Conselle-
ria d’Agricultura i Pesca). Since the first discovery in
1992, regional authorities have led operations to slow
down the rate of spread through both the use of a benthic
vacuum and exposure to copper ions (Pou et al. 1993;
Riera et al. 1994; Grau et al. 1998).
Italy
In Italy the first discovery of C. taxifolia was made
in 1992 in a port of Imperia, located 40 km east of
Monaco (Relini and Torchia 1992). By the end of
206
2000 the alga had spread to five regions: Liguria
(eastern, 10 m2 ; western, 8623 ha), Toscania (south of
Livorno, 500 m2 ; Island of Elba, 51 ha), Sicily (Strait
of Messina, 480 ha), and Calabria (Strait of Messina,
260 ha), for a total of 9415 ha concerned area.
Since 1992, mapping data have been gathered and
verified by laboratories or institutions according to their
field of expertise (Liguria: Morucci et al. 1996; Torchia
et al. 2001, Belsher et al. 2001; Toscania: Cinelli and
Lavelli 1994, Piazzi et al. 2001; Sicilia and Calabria:
Fradà Orestano et al. 1994; Fradà Orestano and Calvo
1995; Orestano 1998; Orestano et al. 2001; Di Martino
2001). The frequency of these efforts is quite sporadic
and the degree of precision in the estimates made by
the divers is variable. A few compilations of these data
have been published (Morucci et al. 1994; Sandulli
et al. 1995). In 1999 an international mapping effort
was mounted using towed video cameras (Belsher et al.
2001). This allowed a more accurate appraisal of the
situation prevailing in western Liguria (IFREMER for
France, ICRAM and ENEA for Italy). With the excep-
tion of a few eradication attempts at Messina, Elba,
Chiavari and Livorno made at the onset of the invasion
(removal by hand of the alga by a few academics), no
control strategy has been set up by either the state or
the regions concerned.
Croatia
In Croatia C. taxifolia was observed for the first time
in 1994 close to a mooring dock in Stari Grad (Hvar
Island). In 1995 it was discovered again in a port
at Malinska (Krk Island) (Span et al. 1998; Zuljevic
et al. 1998; Zavodnik 1995; Zavodnik et al. 1998a,b,
2001). Control measures were implemented annually
by covering isolated colonies with black plastic sheets
in Stari Grad Bay and by using a benthic vacuum in
Malinska (Zuljevic and Antolic 2001b; Zavodnik et al.
1998a,b, 2001). Despite these efforts it continues to
spread, particularly in the Hvar region.
The last record of an isolated colony of C. taxifolia
in Croatia was in 1996 at Barbat Channel near Dolin
Island, an area subject to frequent moorings (Zuljevic
and Antolic 2001a). C. taxifolia was eradicated from
this area in September 1996 by manual removal.
However, it was recorded here again in June 2001.
Since the initial discovery of C. taxifolia in 1994,
its spread has been regularly mapped by scientists
from the Oceanographic station of Rovinj (for the
colonized areas of Malinska) and from the Institute
of Oceanography and Fisheries of Split (for the areas
colonized on Hvar island). At the end of 2000 the total
concerned area was 40 ha for Hvar Island and 1.3 ha
for Krk Island.
Tunisia
In Tunisia the alga has recently been detected (January
2000) in front of the port of Sousse, where a large num-
ber of boats anchor before entering the port (Langar
et al. 2000). Divers from the Tunisian ‘Institut National
des Sciences et des Techniques’, estimated the perime-
ter of the concerned area as 350 ha, in depths of
15–18 m. No control strategies have been implemented
to date.
Discussion and conclusion
The status of C. taxifolia in the Mediterranean at the
end of 2000 can be described as follows. C. taxifolia
is present in 6 countries with 103 independent areas
of colonization, involving 131 km2 of concerned area
along 191 km of coastline. In the Mediterranean
C. taxifolia continues to spread and at the present
time it is still in the expansion phase (Ribera and
Boudouresque 1995).
It is remarkable that 90% of the concerned areas and
80% of the areas colonized are found along 500 km of
coastline between Toulon (France) and Genoa (Italy).
This distribution, in addition to the earlier historical
spreading in this region, further supports the hypothesis
that the origin of the introduction is in the center of this
region (Monaco).
Hill et al. (1998) simulated the alga invasion on
several levels. The simulation utilized data collected
by divers (natural dispersion around main colonies,
growth rates of the stolons, behavior of C. taxifolia on
different substrates, depth or hydrodynamic situation)
and attempted to estimate accurate rates of spreading
given certain known physical considerations (different
substrates, and direction and velocity of the main cur-
rents). Results indicate that each area has its own rate of
spread, which can then be enhanced by anthropogenic
activities (mooring or fishing), diminished by control
efforts (eradication of peripheral colonies) and always
limited by the depth boundaries (C. taxifolia colonizes
only to depths of 50 m).
Some local examples indicate that from the first
stages of colonization to 8 years later, 50–500 ha are

colonized (concerned). Examples include the Island of
Elba (600 m2 in 1993 and 51 ha in 2000), the Straits
of Messina, where the dissemination is amplified by
strong currents (10 m2 in 1993 and 480 ha in 2000),
Island of Hvar, where most of the peripheral colonies
were eradicated (7 ha in 1995 and 40 ha in 2000),
and the Island of Mallorca, where regular eradica-
tion efforts are made (250 m2 in 1992 and 65 ha in
2000). In the region of Monaco (20 km west and east
of this principality), the alga spread from 1 m2 in 1984
to 30 ha at the end of 1991 and 427 ha by the end
of 1992.
With the exception of new invasions via aquariums
and in the absence of any sexual reproduction (Meinesz
et al. 1993; Carvalho et al. 1998; Zuljevic and Antolic
2000), the invading strain of C. taxifolia spreads either
as the result of small pieces of the plant (cuttings) that
are transported over short distances along the bottoms
by hydrodynamics, or through the transport of cuttings
by boat anchors or fishing nets, the latter over poten-
tially great distances (Meinesz 1992; Sant et al. 1996;
Relini et al. 2000).
Sixteen years after the introduction of the alga in
Monaco, it is remarkable to note that all the first set-
tlements of the colonized areas at Mallorca, Saint-
Cyprien, Livorno, Elba, strait of Messina, Hvar, Krk,
Barbat and Sousse, other than the initial introduction
region (between Toulon and Genoa), are either ports
or frequently used anchoring sites. The geographic
locations of these areas of colonization do not reveal a
dissemination gradient between them and between the
initial colonized area. Modeling of the spreading (Hill
et al. 1988) shows also that there is a gradient of natu-
ral fragment dispersal only for short distances (several
hundred meters). If long distance spread was the result
of cuttings floating over short or great distances, as
has been suggested by Smith and Walters (1999) and
Chisholm et al. (2000), strong gradients of distance
between the invaded areas such as the advancing-wave
model (Hengeveld 1989) should occur. These consider-
ations would tend to confirm the hypothesis of long dis-
tance anthropogenic spread by random dissemination
in harbors or mooring bays (anchorages).
The characteristics of a colonized area (number of
colonies; covered, affected or concerned areas), when
discovered, and data allowing the modeling of the natu-
ral development and spread of the alga (Hill et al. 1998)
make it possible to back-date the arrival of C. taxifolia.
In the colonized areas other than the initial introduc-
tion region (between Toulon and Genoa), the arrival of
207
the alga is estimated to be in the period between 1991
to 1993. Three hypotheses may explain the long-range
spread of the alga during this period, and the possible
decrease in the number of new populations appearing
at long distances away from established populations:
(i) the growing awareness of pleasure-boaters and fish-
ermen to the problem, as the result of public aware-
ness campaigns started in 1991, brought with it a
decrease in long-range spread;
(ii) some remote colonized areas, with a more recent
invasion date, and therefore a small surface area,
remain undiscovered, especially if they are located
in regions rarely visited by SCUBA divers or where
public awareness is low. The discovery in 2000 of
an invaded area involving 350 ha in Tunisia fol-
lowing a local public awareness campaign is an
example of this;
(iii) long-range spread can be attributed to a few boats
sailing between 1991 and 1993 that contained
clumps of C. taxifolia in their anchor locker (as
observed on occasion by some of us) originating
from highly invaded areas at that time (Monaco,
French and Italian Riviera). Accidental release
from a yacht possessing an onboard aquarium with
the alga may also be possible.
A continued monitoring of the spread of C. taxifolia
in the Mediterranean should allow these different
hypotheses to be assessed.
Large regions neighboring the areas now invaded by
C. taxifolia appear favorable to further colonization.
There is thus no reason to believe that spreading will
slow down in the years to come.
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