Behavioural response of Mullus surmuletus to habitat modification by the invasive macroalga Caulerpa taxifolia

Densities of red mullet Mullus surmuletus have declined since 1996 at Cap Martin, Alpes‐Maritimes, France, following colonization by Caulerpa taxifolia . Foraging M. surmuletus were rarely observed over C. taxifolia or the seagrass Posidonia oceanica , but were observed over bare sand and endemic macroalgae. Within colonized sites (where the mean cover of C. taxifolia was 30–100%), fish were concentrated on areas with low cover of C. taxifolia (<21%). A difference in the frequency of observations of foraging groups (17%) was observed between sites with and without C. taxifolia . The observed patterns were probably due to a physical barrier to foraging that is presented by meadows of C. taxifolia . Changes to habitat structure that occur when the substratum is colonized by C. taxifolia influence the accessibility to benthic food resources, and consequently the foraging activities of fish that feed on benthic invertebrates. The decline in density of M. surmuletus at Cap Martin could be the result of fish emigration from colonized sites (unfavourable to the species) to uncolonized sites.     

Enzymatic antioxidant response of a labrid fish (Coris julis) liver to environmental caulerpenyne

Exposure of marine animals to certain toxic compounds can enhance reactive oxygen species production with subsequent damage to macromolecules and alterations in oxidant defenses levels. Caulerpenyne is the major metabolite synthesized by Caulerpa species, used as chemical defense affecting several cellular and molecular targets. We assessed the changes produced by the presence of Caulerpa spp. in the activities of antioxidant enzymes as well as lipid peroxidation levels in liver of the teleost Coris julis. Fish were captured at two stations with Caulerpa species-Caulerpa taxifolia and Caulerpa prolifera-and at a region with the seagrass Posidonia oceanica as negative control. Caulerpenyne concentration was significantly higher in C. prolifera than in C. taxifolia (p<0.05). Glutathione S-transferase, glutathione peroxidase and glutathione reductase activities were significantly higher in both Caulerpa stations compared to the P. oceanica (p<0.05). No statistical difference (p>0.05) existed in catalase activity between groups. Glutathione reductase activity is significantly higher in C. prolifera station than in C. taxifolia (p<0.05). Despite the variations in the antioxidant enzyme activities, there was no significant difference in malondialdehyde concentration. In conclusion, the production of caulerpenyne by Caulerpa species could induce an antioxidant adaptation in the liver of C. julis in order to prevent oxidative damage.     

Variations in the concentration of phenolic compounds in the seagrass Posidonia oceanica under conditions of competition

The concentration of phenolic compound was measured in the seagrass Posidonia oceanica when interacting with two Bryopsidophyceae, Caulerpa taxifolia and Caulerpa racemosa, between May 1999 and May 2000. These measurements were performed on adult and intermediate leaves and in sheaths of the seagrass. Sampling was carried out at three stations subject to increasing levels of interaction with Caulerpa. The number of tannin cells was also analysed. Five phenolic compounds were identified in P. oceanica, with a predominance of caffeic acid in the adult and intermediate leaves. For a given level of interaction (and for both caulerpa sp.), a significant seasonal variation in phenolic compounds was shown in the adult leaves (higher in November and lower in September and March for example for the interaction with C. taxifolia). Only for two compounds (corresponding to a mixture containing ferulic acid and the ester methyl 12-acetoxyricinoleate) were significant differences observed as a function of the level of interaction with C. taxifblia, and only in the adult leaves: higher concentrations of phenols were observed with increasing level of interaction. Thus,adult leaves gave values of 55.5 +/- 14.1 microg g(-1) dm without interaction (OCt) and 94.9 +/- 23.4 microg g(-l) dm with high interaction (2Ct),corresponding to an increase of 70%. No significant difference was observed with intermediate leaves and sheaths, or for interaction with C. racemosa. The number of tannin cells (supposed to produce the phenolic compounds) largely increased in the adult and intermediate leaves when the degree of interaction with C. taxifolia increased: 90 mm above the base of the sheath (in adult leaves), 16.7 +/- 10.6 tannin cells cm(-2) were found without interaction (OCt), and 57.8 + 21.2 tannin cells cm(-2) with high interaction (2Ct). No significant difference was found for C. racemosa interaction. It thus appears that when the seagrass P. oceanica is in interaction with C. taxifolia, it accelerates its production of secondary metabolites so as to limit invasion of the beds.     

Are the Mediterranean ascoglossan molluscs Oxynoe olivacea and Lobiger serradifalci suitable agents for a biological control against the invading tropical alga Caulerpa taxifolia?

The Mediterranean ascoglossan Oxynoe olivacea and Lobiger serradifalci, which are scarce in meadows of their usual food the alga Caulerpa prolifera, have become adapted to feeding on the invading tropical alga Caulerpa taxifolia. Grazing rates are low (O. olivacea destroys only a 5-cm C. taxifolia frond in 3-7 d). The grazing rates are a function of temperature. Despite a high spawning frequency (0.25 egg-mass.day-1) and a large number of eggs released per spawning event (O. olivacea: from 2,545 to 29,128 eggs.spawning-1; L. serradifalci: from 12,112 to 48,448 eggs.spawning-1), the recruitment on Caulerpa meadows is usually low due to the hazardous pelagic development of the larvae. The possible use of these molluscs as agents of biological control against C. taxifolia appears to be possible only through an artificial enhancement of their populations after cultivation of the veligers and release of juveniles during the winter season. This means of control is discussed and compared with the possible use of tropical ascoglossan species.     

Two novel aromatic valerenane-type sesquiterpenes from the Chinese green alga Caulerpa taxifolia

Caulerpal A (2) and B (3), two novel sesquiterpenes possessing an uncommon aromatic valerenane-type carbon skeleton, along with one known metabolite, caulerpin (4), have been isolated from the Chinese green alga Caulerpa taxifolia (Vahl) C. Agardh. Their structures and relative stereochemistry were elucidated on the basis of extensive spectroscopic analysis. Compounds 2-4 were evaluated for their inhibitory activity against hPTP1B and the result showed that only compound 4 had a strong PTP1B inhibitory activity with an IC50 value of 3.77 microM.     

Behavioural interactions between ecosystem engineers control community species richness

Behavioural interactions between ecosystem engineers may strongly influence community structure. We tested whether an invasive ecosystem engineer, the alga Caulerpa taxifolia , indirectly facilitated community diversity by modifying the behaviour of a native ecosystem engineer, the clam Anadara trapezia , in southeastern Australia . In this study, clams in Caulerpa -invaded sediments partially unburied themselves, extending >30% of their shell surface above the sediment, providing rare, hard substrata for colonization. Consequently, clams in Caulerpa had significantly higher diversity and abundance of epibiota compared with clams in unvegetated sediments. To isolate the role of clam burial depth from direct habitat influences or differential predation by habitat, we manipulated clam burial depth, predator exposure and habitat ( Caulerpa or unvegetated) in an orthogonal experiment. Burial depth overwhelmingly influenced epibiont species richness and abundance, resulting in a behaviourally mediated facilitation cascade. That Caulerpa controls epibiont communities by altering Anadara burial depths illustrates that even subtle behavioural responses of one ecosystem engineer to another can drive extensive community-wide facilitation.     

Influence of substrate and deployment time on fish assemblages on an artificial reef at Formentera Island (Balearic Islands,western Mediterranean)

The Migjorn artificial reef (MAR) was deployed in the summer of 1990 and is made by 48 concrete blocks set 30 m deep on a sea bed of sand with a small proportion of Posidonia oceanica (L.) Delile meadow. Visual censuses of blocks on both substrata were carried out during the months of February, May and September in 1992 and 1993. The MAR fish assemblages were influenced by the surronding substrate. Thus, a group of ubiquitous species, frequent on both substrata and typical of natural rocky habitats (mostly Sparidae) and a group associated with the samples recorded on the Posidonia meadow (mostly Labridae) could be distinguished. Season had only a secondary and slight effect on fish assemblages. The MAR was colonized by 74% of the species in the first two years after deployment. Positive and lineal relationships between species richness, diversity, abundance of necto-benthic species and time since deployment on sandy substrata were found. On Posidonia oceanica deployment time has only increased very sedentary and cryptic species. Although some ways in which the MAR would have a productive function are not rejected, the most obvious MAR working pattern is that of fish aggregation. This revised version was published online in September 2006 with corrections to the Cover Date.     

Adaptive colouration of Mediterranean labrid fishes to the new habitat provided by the introduced tropical alga Caulerpa taxifolia

Since the introduction of the invasive tropical alga Caulerpa taxifolia in the Mediterranean Sea. a new, uniformly green habitat has been available to local fish assemblages. Underwater visual censuses of three species, Symphodus ocellatus, Symphodus roissali and Coris julis , revealed a higher proportion of green morphs in the C. taxifolia meadows compared to those inhabiting their usual indigenous habitats. In a fourth species, Symphodus rostratus , the distribution of the different colour morphs did not differ between the two habitats investigated. Experiments in aquaria proved that S. ocellatus and S. roissali were homochromic species able to adapt their colouration to the environment. In contrast, no colour change was observed in S. rostratus during the experiments.     

Polymerase chain reaction-single strand conformation polymorphism analyses of nuclear and chloroplast DNA provide evidence for recombination,multiple introductions and nascent speciation in the Caulerpa taxifolia complex

Independent lines of evidence support an Australian origin for the Mediterranean populations of the tropical alga Caulerpa taxifolia. To complement previous biogeographical studies based on nuclear rDNA internal transcribed spacer (ITS), a new chloroplast marker was developed--the cp 16S rDNA intron-2. Sequence variability for both nuclear and chloroplast markers were assessed in 110 individuals using single strand conformation polymorphism. Comparison of intrapopulation genetic diversity between invasive Mediterranean and 'native' Australian populations revealed the occurrence of two divergent and widespread clades. The first clade grouped nontropical invasive populations with inshore-mainland populations from Australia, while the second clustered all offshore-island populations studied so far. Despite our finding of nine distinct nuclear and five distinct chloroplast profiles, a single nucleocytoplasmic combination was characteristic of the invasive populations and sexual reproduction was found to be very rare. C. taxifolia is clearly a complex of genetically and ecologically differentiated sibling species or subspecies.     

Illegal trawling and induced invasive algal spread as collaborative factors in a Posidonia oceanica meadow degradation

Posidonia oceanica, a key seagrass species of the Mediterranean Sea, shows clear signs of regression throughout the Mediterranean and illegal trawling is recognized as one of the main causes. We examined the condition of a P. oceanica meadow in Alykes Bay (Zakynthos Island, western Greece), a typical Mediterranean littoral area where illegal trawling is common practice, in respect to the total area affected, and in terms of possible ecological substitution. A side scan sonar (SSS) survey of the seafloor provided an image of the condition of the meadow and biological sampling evaluated the ecological status in affected meadow areas. SSS images revealed that trawling has a serious effect on the meadow, with 11% of the vegetated area being abraded, and the affected areas were also found to be fully colonized by the invasive alga Caulerpa racemosa. Moreover, unusually high densities of the polychaete Sabella pavonina were detected in the affected areas among C. racemosa fronds. Recolonization by P. oceanica of the affected meadow areas that have been colonized by C. racemosa seems improbable considering the allelopathic interactions between the species, with the alga displaying phytotoxic properties through caulerpenyne production and deterioration of the sediment quality.     

Distribution and dynamics of Posidonia oceanica beds along the Albères coastline

The location and dynamics of Posidonia oceanica beds were studied in the marine natural reserve of Cerbère-Banyuls-sur-Mer (eastern Pyrenees, France), through the mapping of bottom assemblages (transects and image processing) and the monitoring of fixed markers installed at the Posidonia oceanica lower limit. The surface area occupied by continuous beds is relatively limited and localized, whereas the presence of important areas of dead 'matter' would seem to indicate that bottom cover by Posidonia oceanica was more extensive in the past. Although the limited extension of the seagrass beds seems to be linked to both a high level of turbidity and severe hydrodynamic conditions, it is hypothesized that the regressions observed are due to lack of sediment.     

The joint influence of environmental and anthropogenic factors on the invasion of two alien caulerpae in northwestern Mediterranean

Biological Invasions - Many factors are known to influence the invasion of alien species depending on the spatial scales considered. Caulerpa taxifolia and Caulerpa cylindracea are two green...     

Molecular characterization of variable tandem repeat sequence determined during RAPD analysis among Posidonia oceanica insular and coastline populations

The seagrass Posidonia oceanica plays a multifunctional role in the coastal area as an important and productive component of ecosystems in the Mediterranean Sea. We detected by RAPD analysis with two arbitrary primers genetic differences in P. oceanica collected from several sites in the Southern Mediterranean. By AMOVA analysis we observed a level of about 20% genetic difference among individuals within a population and 80% among populations. A common band of 200 bp was found in all the amplified samples. Cloning and sequencing analysis of this band revealed the presence of a simple tandem repeat sequence (minisatellite) that we called PoTR (Posidonia oceanica tandem repeat). Finally, the ability of PoTR to detect genetic variability in P. oceanica genome was demonstrated by the presence of amplification products of different lengths utilizing primers internal to this sequence.     

Role of Corollospora maritima in the degradation and bioconversion of leaf material from Posidonia oceanica

Posidonia oceanica supports mainly saprophytic marine flora, comprising predominantly lignicolous fungi. The frequency of occurrence of species recorded on this marine angiosperm, was high, indicating that they play a major role in the biological degradation of the sea grass Posidonia oceanica. In vitro experiments with Corollospora maritima (isolated from leaf material) were conducted in order to evaluate their role in the degradation of leaf material. Corollospora maritima actively degrade leaf material. Biophysical and biochemical changes (particle detritus formation, C and N variation), enzymatic activity involved and sterol production were studied during the transformation process of leaves to mycelial biomass.     

Cytotoxicity screening of supercritical fluid extracted seaweeds and phenylpropanoids

Molecular Biology Reports - Detached leaves of Posidonia oceanica and Zostera marina creating nuisance at the shores were extracted by means of supercritical CO2 enriched with a co-solvent,...     

Unexpected response of the seagrass Posidonia oceanica to a warm-water episode in the north western Mediterranean Sea

The response of Posidonia oceanica (Linnaeus) Delile to the warm-water episode of summer 1999 was studied by means of the technique of lepidochronology. Study sites include three sites affected by the mass mortality event of benthic invertebrates and one not affected. The results showed a significant decline in some parameters (number of leaves and/or rhizome growth) for the three sites affected by the mass mortality event for the year following the warm-water episode (1999-2000). A similar decline was not observed for the unaffected site. The fact that high temperatures could have a negative impact on deep Posidonia oceanica near its cold limit of distribution is an unexpected result.     

175 Toward an Optical Biogeography of the Oceans

Invasions of marine communities by seaweeds are increasing globally. Vectors for introductions are primarily aquaculture and the aquarium trade. Attention to these vectors has been minimal, and few, if any, regulations exist to prevent seaweed introductions. Challenges in preventing, eradicating, and controlling invasive seaweeds will be discussed using the invasion of Caulerpa taxifolia in southern California as a model.     

THE PHYLOGENY OF CAULERPA BASED ON RDNA INTERNAL TRANSCRIBED SPACER SEQUENCES

Phylogenetic hypotheses for the pantropical marine green algal genus, Caulerpa , were inferred based on analyses of nuclear-encoded rDNA internal transcribed spacer (ITS) sequences. Results of these analyses were used to assess the correspondence between rDNA phylogeny and traditional sectional taxonomy, to identify synapomorphic morphological characters (including assimilator morphology and chloroplast ultrastructure), and to examine marine biogeographic hypotheses for the genus. Ribosomal DNA ITS sequences were aligned for thirty-three species and intraspecific taxa of Caulerpa. Results indicate limited correspondence between phylogeny and sectional taxonomy for the genus, (e.g., the sections Filicoideae and Sedoideae were not monophyletic). In contrast, chloroplast morphology could be mapped to the tree topology with limited homoplasy. Pantropical isolates of the filicoidean species, Caulerpa sertularioides and Caulerpa mexicana each formed monophyletic groups. Caulerpa reyesii was included as a derived taxon within the Caulerpa taxifolia clade, suggesting that these species were conspecific and affirmed the lack of correspondence between phylogeny and assimilator morphology. Isolates and various intraspecific taxa of Caulerpa racemosa did not form a monophyletic group. Instead, these taxa formed a heterogeneous assemblage with other sedoidean and filicoidean taxa. Within the C. sertularioides clade, Caribbean and Atlantic isolates formed a basal paraphyletic group, whereas eastern and western Pacific isolates formed a more derived monophyletic group. Therefore, these results are not consistent with an Indo-West Pacific origin of this species.     

Caulerpenyne from Caulerpa taxifolia has an antiproliferative activity on tumor cell line SK-N-SH and modifies the microtubule network.

Caulerpenyne, the major secondary metabolite synthesized by the green marine alga Caulerpa taxifolia, is cytotoxic against several cell lines. To identify possible targets of this toxin, we investigated the effect of caulerpenyne on the neuroblastoma SK-N-SH cell line. Caulerpenyne induced an inhibition of SK-N-SH cell proliferation with an IC50 of 10 +/- 2 microM after 2 hr of incubation.We observed no blockage in G2/M phase and an increase in cell death. On immunofluorescence microscopy, caulerpenyne affected the microtubule network in SK-N-SH cell line; we observed a loss of neurites and a compaction of the microtubule network at the cell periphery. In vitro, after 35 min of incubation, caulerpenyne inhibited the polymerization of pig brain purified tubulin or microtubule proteins, with an IC50 of 21 +/- 2 microM and 51 +/- 6 microM respectively. Analysis by electron microscopy indicated that caulerpenyne induced aggregation of tubulin, which may be responsible for inhibition of microtubule polymerization and bundling of residual microtubules.     

The Introduced Green Alga Caulerpa Taxifolia Continues to Spread in the Mediterranean

The tropical green alga Caulerpa taxifolia in the Mediterranean has spread steadily since its introduction in 1984. At the end of 2000, approximately 131km² of benthos had been colonized in 103 independent areas along 191km 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.