Viral dynamics in two trophically different areas in the Central Adriatic Sea

To understand the activity of marine viruses, experiments on viral production, viral decay and the percentage of lytic and lysogenic bacterial cells among the total number of bacterial cells were carried out seasonally at two stations in the Adriatic Sea with different trophic conditions. Additionally, we are providing an insight on the enrichment with dissolved and particulate organic matter by viral lysis in the studied area. Viral production was higher at the coastal station than at the open-sea station. Viral decay rates were also higher at the coastal sea station than at the open-sea station, and accounted for approximately 40% of viral production at both investigated stations. The percentage of lysogenic infection was lower than that of lytical infection, which indicates the prevalence of the lytic cycle at both stations. Viruses had a significant influence on bacterial mortality through high daily removal of the bacterial standing stock at the coastal and open-sea station. The viruses contributed to the restoration of dissolved organic carbon, nitrogen and phosphorus in the microbial loop by lysing the bacterial cells at the studied stations. All the above suggest that viruses are important in the microbial food web and an important factor in the control of bacterial populations within the study area.     

Microplankton respiratory activity and CO2 production rates in the Otranto Strait (Mediterranean Sea)

Respiratory activity and metabolic CO₂production of the microplankton in the Otranto Strait (Mediterranean Sea) were determined by monitoring the Electron Transport System activity. Ten stations were repeatedly investigated during two oceanographic surveys in February–March and August 1994. Respiratory activity and CO₂ production, estimated from the surface to the bottom, were higher in the euphotic layers (0-200 m) during summer (mean values: Winter = 0.024 μg C h⁻¹ dm⁻³; Summer = 0.042 μg C h⁻¹ dm⁻³); in the aphotic zone (deeper than 200 m), the rates were similar throughout different seasons (0.013 and 0.014 μg C h⁻¹ dm⁻³, respectively). A comparison with data collected by other authors from the euphotic layers of the Mediterranean Sea was made. Respiratory activities decreased from Western to Eastern Mediterranean Basins. The values of CO₂ production, integrated between 200 and 1000 m in the Otranto Strait (mean value 237.7 mg C m⁻² d⁻¹), were compared with other data collected from the Mediterranean Sea as well as from the Pacific, Atlantic and Indian Oceans. The comparison showed the Otranto Strait to be a site of organic matter oxidation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Annual dynamics of bacterioplankton assemblages in the Gulf of Trieste (Northern Adriatic Sea)

Bacterioplankton community diversity was investigated monthly in coastal waters of the Gulf of Trieste (NE Adriatic Sea) throughout 2003. Superficial bacterial assemblages of two differently freshwater influenced stations were studied using PCR-DGGE fingerprinting techniques. Bacterial genetic diversity of the sampled area, as estimates of the number of DGGE bands was high (36-64) compared to that reported in other studies employing this fingerprint technique. The similarity index (Sorensen Index) between assemblages showed a defined operational taxonomic units (OTUs) succession pattern in the more typically marine station with stable winter communities and quickly changing summer ones. On the contrary in the station affected by riverine inputs no clear pattern was detected. In both sites, according to cluster analyses performed on the DGGE banding pattern, three seasonal assemblages were identified: winter-spring, summer and fall. Sequence analysis of fifty-six among the brightest gel bands led to the observation of bacteria affiliated to Gram positive, Cyanobacteria, Cytophaga-Flavobacteria-Bacteroides (CFB) lineages and the alpha-, gamma- and delta- subdivisions of the Proteobacteria. Gamma-Proteobacteria constituted the main fraction (60%) of sequences in the more typically marine station, whereas the river-influenced station was characterised by more heterogeneous assemblages (39% alpha-Proteobacteria, 32% Flavobacteria).     

Microplankton response to environmental conditions in the Alboran Sea (Western Mediterranean): One year sediment trap record

The present work analyses the seasonal evolution of planktonic assemblages and particle fluxes through the water column in the Eastern Alboran Sea (Western Mediterranean) at 35º55.47'N/01º30.77'W. A Sediment trap was deployed below the influence of the Almeria-Oran Front (AOF), a semi-permanent geostrophic front, during July 1997 to June 1998. Overall, the temporal variability of coccolithophore, planktonic foraminifer, diatom, benthic and wind-carried biogenic particle fluxes is linked to the seasonal evolution of sea surface hydrological structures. Maximum planktonic fluxes were found during high-productivity periods and wind-induced upwelling, following a trimodal pattern, with maximum fluxes in July 1997, November–December 1997, and April–May 1998. These periods were characterized by vertical mixing and the full development of anticyclonic gyres in the Alboran Sea. The annual flux of coccolithophores was dominated by the “small Gephyrocapsa Group” and Emiliania huxleyi, whereas Turborotalita quinqueloba and Globigerina bulloides dominated the foraminiferal fluxes, and Chaetoceros Resting Spores (RS) were predominant in the diatom assemblage. Benthic specimens were also collected with the sediment trap, suggesting a variable influence of bottom water activity. Wind-driven particles (phytoliths and fresh-water diatoms) were collected along the year, but their fluxes followed the local wind regime.The high Sea Surface Temperature (SST) during fall due to weaker than usual westerly winds, and the pressure anomaly prevailing in the Alboran Sea during early winter, were reflected in the planktonic assemblages by the proliferation of warm, lower photic layer inhabitants and/or oligotrophic taxa of coccolithophores (Florisphaera profunda), planktonic foraminifers (Globigerinoides ruber and Globorotalia inflata) and diatoms (Leptocylindrus danicus). These unusual climatic conditions in the eastern Alboran Sea must have been caused by the 1997–1998 ENSO event.     

Daily variations of highly active bacteria in the Northern Adriatic Sea

Nowadays, it is recognized that only a fraction of aquatic bacteriaare actively growing, but there is little information aboutthe factors constraining their metabolism. Marine bacterioplanktoncan rapidly modify their metabolic activity level in responseto environmental changes. In this study, we focused on the dailychanges in abundance and activity of active bacterial fractionover a 20-day period preceded by intense rainfalls which slightlymodified water column conditions. Cells capable of reducingthe membrane-penetrable dye 5-cyano-2,3-ditolyl tetrazoliumchloride (CTC), estimated by epifluorescence microscopy, areconsidered very active (CTC+ bacteria). Total bacterial abundance(TBA) ranged from 0.8 to 2.4 x 10⁹ cells L^–1, whereasCTC+ bacteria were more variable (1.6–9.2 x 10⁷ cellsL^–1), accounting for 1.2–4.4% of TBA. Bacterialactivity (BA) quantified as the incorporation of [3H]-leucinevaried by more than one order of magnitude over the period (25.0–662.5pmol L^–1 h^–1). BA was strongly related to CTC+ bacteria,suggesting that they were mainly responsible for the bacterialcommunity metabolism. Nevertheless, cell-specific activity,scaled to only CTC+ cells, was very high, suggesting that afraction of cells not detectably CTC+ may be able to assimilate[3H]-leucine. The correlation between salinity and TBA, CTC+bacteria and BA supported the hypothesis of the active roleof freshwater input in enhancing cell activity. Our resultssuggest that freshwater inputs rather than phytoplanktonic bloomsare able to induce shifts in bacterial metabolism over a timescale of days in the area studied.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Annual survey of planktonic dinoflagellates and related cysts in the Gulf of Trieste (Northern Adriatic Sea)

Abstract

A survey of planktonic dinoflagellates and related cysts was carried out in the Gulf of Trieste throughout one year from April 1992 to March 1993. 113 taxa were recovered by the analysis of phytoplankton net samples. The most represented genera were Protoperidinium (34 species), Ceratium (24 species), Dinophysis (15 species), Gonyaulax (11 species) and Prorocentrum (8 species). A particular attention was given to potentially toxic species belonging to the genera Dinophysis, Alexandrium and Prorocentrum. The highest number of species (67 species) was recorded in July, and the lowest one (18 species) in February.

33 cyst morphotypes were recorded by the analysis of sediment samples. The most represented genera were Protoperidinium (8 morphotypes), Scrippsiella (3 morphotypes), Gonyaulax (3 morphotypes) and Alexandrium (2 morphotypes); the cysts most frequently found were those of Conyaulax polyedra and Alexandrium pseudogonyaulax.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Viral production in the Gulf of Trieste (Northern Adriatic Sea): Preliminary results using different methodological approaches

Although the temporal and spatial variability of virioplankton in the northernmost part of the Adriatic Sea has been repeatedly explored suggesting that viruses constitute an extremely dynamic component of the plankton community and hypothesizing their importance in marine food webs and mucilage events, there is still no information about viral replication rates. Hence, the contribution of viruses to bacterial mortality and the cycling of organic matter in this part of the Adriatic basin are still not fully comprehended. Assessment of the role of viral lysis requires a robust means of estimating viral production. Since, up to now, none of the available methods evolved to a state of a standard yet, in this preliminary study 3 different experimental approaches were simultaneously assayed (viral production estimated by radiotracer incorporation method [TdR], dilution technique for the estimate of viral production in already infected bacteria [DIL] and serial dilution method in manipulated phage-host assemblage [SER]). The present study provided the first evidence of viral production rates in this study area, that resulted in comprising between ∼ 3.5-15 × 10⁸ viruses L^− 1 h^− 1 and critically faced up the results obtained by different techniques with the consideration that they suffer from different biases. Based on TdR and DIL viral proliferation estimates, viral lysis was responsible for the loss of 54 to 95% of the bacterial standing stock, while the viral-induced mortality by SER (325% d^− 1) was likely consistently overestimated. These results indicate that viral lysis is a significant factor for prokaryotic mortality suggesting its implication as an important pathway for the cycling of dissolved organic matter in the Gulf of Trieste.     

On the presence of Pseudo-nitzschia calliantha Lundholm, Moestrup et Hasle and Pseudo-nitzschia delicatissima (Cleve) Heiden in the Southern Adriatic Sea (Mediterranean Sea, Italy)

This investigation included a 2-year monitoring program aimedat assessing the abundance and distribution of harmful marinephytoplankton along the Southern Adriatic coast of Italy. Monthlysampling was conducted from April 1995 to March 1997 at foursampling stations along four transects, to determine the temporaland spatial presence/abundance of the potentially toxic Pseudo-nitzschiaspecies. The study focussed on the most abundant taxa Pseudo-nitzschiacalliantha and Pseudo-nitzschia delicatissima, the identitiesof which were confirmed by TEM on cleaned net material. Thedistribution patterns of these potential Amnesic Shellfish Poisoning(ASP) toxin producers were statistically analysed by GeneralisedLinear Model ANalysis Of VAriance, Principal Component Analysis(GLM ANOVA, PCA) and Spearman’s correlation analysis inorder to address relationships between environmental variableand population dynamics. Abundances displayed horizontal andvertical structure in the study area. Inter-annual variabilitywas also observed for both species that appeared to responddifferently to the environmental factors investigated. Distributionsof P. calliantha showed a stronger seasonality and was morecorrelated with winter water conditions than P. delicatissima,which in turn exhibited a broader temporal distribution andappeared independent from major environmental constraints. Thisis the first report of the occurrence and dynamics of P. callianthaand P. delicatissima populations in Southern Adriatic coastalwaters. First two authors equally contributed to this work.     

Age, growth and mortality of the Mediterranean amberjack Seriola dumerili (Risso 1810) from the south-eastern Adriatic Sea

The age, growth and mortality of the Mediterranean amberjack Seriola dumerili (Risso 1810) were determined in 298 specimens collected in the south-eastern Adriatic Sea (Donji Molunat Bay) from 17 May to 26 June 1997. The total length ranged from 32.0 to 160.0 cm and weight from 0.5 to 46.5 kg. Ten age classes, ranging from 1° to 10° years were defined by scale readings. Mean total length and weight-at-age data were used to estimate the growth parameters of the von Bertalanffy equation: L _∞=174.6 cm, K =0.190, t ₀=–0.314; W _∞=79.01 kg, K =0.139 and t ₀=–0.746. The length–weight relationship was estimated at: W =0.000123 ×  L ^2.847. The overall sex ratio was 1 : 1.06 in favour of males. Total ( Z ) and natural ( M ) mortality were found to be 0.41 years⁻¹ and 0.30 years^–1, respectively. The exploitation ratio ( E =0.27) indicates that the fishing pressure on the Mediterranean amberjack was low in the investigated region.     

Native and non-native ctenophores in the Gulf of Trieste, Northern Adriatic Sea

Observations of ctenophore species were made in the Gulf ofTrieste between 2003 and 2006. We examined native ctenophorespecies with special attention to representatives of the ordersLobata and Beroida, and we recorded among them two non-nativectenophores: Mnemiopsis leidyi A. Agassiz 1865 and Beroe ovatasensu Mayer, 1912. The validity of the Mediterranean speciesBeroe ovata is discussed. We determined that among the nativespecies, it is not Beroe ovata but rather Beroe cucumis sensuMayer, 1912 that occurs in the Mediterranean Sea.     

Feeding ecology of Mediterranean razorfish Xyrichthys novacula in the Tyrrhenian Sea (Central Mediterranean Sea)

The stomach contents of 196 razorfish ( Xyrichthys novacula ), collected in the Tyrrhenian Sea (Ponza Island, Italy) between July and December 1994, were examined in order to assess the diet and to analyse size-related dietary variation of the species. The diet of X. novacula showed a strict relationship with well-sorted fine sand (SFBC) benthic species, mostly Mollusca and Echinodermata, as pointed out by the analysis at the specific level of the stomach contents. Mollusca Pelecipoda, essentially Acanthocardium tuberculatum and Echinodermata, with Echinocardium cordatum , made up 90% of the volume of prey of X. novacula. Mysidacea and Gastropoda were frequent but volumetrically less important. The study showed that male and female X. novacula were not effectively segregated by trophic dimension in the study area. The niche overlap between males and females in prey composition was pronounced, as evidenced by the significantly high values of Schoener's index with larger males that showed a greater tendency towards predation of large prey. The increment in length (L.I.) in males, despite a decrease in the percentage body weight increment (B.W.I.), could represent the result of a male reproductive feature: the lack of sperm competition allows the males much more energy to devote to growth than do that of the females of this species.     

Seasonal variations in carbon and nitrogen content of Acartia clausi (Copepoda,Calanoida) in the Gulf of Trieste (Northern Adriatic Sea)

A yearly study was made on total and cephalothorax lengths, dry weight, carbon and nitrogen contents and C:N ratio of the species Acartia clausi (Copepoda, Calanoida) which is always abundant, particularly in spring, in the net zooplankton community of the Gulf of Trieste (Northern Adriatic Sea). The samples of net zooplankton were carried out every month at the permanent station located 200 m offshore and preserved for about one year in 4% buffered formalin to insure weight loss stabilization. A. clausi was separated from other zooplankton specimens; washed and dried. For each monthly sample the average total and cephalothorax lengths and the average dry weight of adult organisms of A. clausi were measured. The mean carbon and nitrogen contents, as a percentage of dry weight, were determined by using a CHN analyser. Linear regression models were computed on the log-transformed data in order to check the relationships between dry weight, total and cephalothorax lengths, carbon and nitrogen contents and C:N ratio of A. clausi. The best of the various established correlations, was between dry weight and carbon content. Our study pinpointed a high seasonal variability of the C:N ratio, mainly due to seasonal fluctuations in the nitrogen content.     

The family Xystonellidae (Ciliophora, Tintinnina) in the Adriatic Sea

This paper reports quantitative, horizontal and vertical distributiondata of the 11 species of the family Xystonellidae found inthe Adriatic Sea and in the Strait of Otranto. The samples werecollected in coastal and open waters, with a plankton net equippedwith a closing system, in the period from 1973 to 1986. At thedeeper stations of the South Adriatic Pit, hauls were takenin eight vertical layers, while at other stations sampling dependedon the respective depth of each station. Xystonellidae are oceanictintinnines, mainly present in the South Adriatic and the Straitof Otranto throughout the year. They are characteristic of SouthAdriatic and Eastern Mediterranean water masses. Horizontaldistribution of lower mesopelagic species is limited by the200 m isobath. Only sparse individuals of the upper mesopelagicand epipelagic species were found in the Jabuka Pit, probablydue to the late autumn and winter current. These species werenot registered to the north of this region. They were registeredbelow the 200 m depth as dominant tintinnines, while in summerthey represented an average of 81% of the total tintinnine population.     

Seasonal and spatial distribution of bacterial production and biomass along a salinity gradient (Northern Adriatic Sea)

The Adriatic Sea is a semi-enclosed ecosystem that receives in itsshallow part, the northern basin, significant freshwater inputs whichmarkedly increase its productivity with respect to the oligotrophic featuresof the Mediterranean sea. In this area, especially on the western coastwhere river plumes diffuse, high physical (density) and chemical (nutrients)gradients occur on a small scale, both horizontal and vertical. Results ofbacterial production as ³H-thymidine incorporation, bacterialabundance as DAPI direct count, autotrophic biomass as chlorophyll a andtotal biomass as ATP from three areas in the Northern Adriatic Sea arereported. The three sites, differently influenced by the river waterdiffusion, were sampled seasonally over two days, every 24 h, in foursurveys from April 1995 to January 1996. Bacterioplankton production,strongly correlated with primary production, was extremely high near thecoast in low-salinity, high-nutrient waters, mostly as an indirectconsequence of riverine inputs causing an increase in phytoplanktonproduction stimulated by physically driven nutrient inputs. In the warmmonths bacterial activity was higher than in cold months. While bacteriaabundance did not appear related to the salinity gradients, bacterialproduction (from 0.6 to 372 pM ³H-thymidine h^™1incorporated, corresponding to 0.01–8.2 μg C l^™1h^™1) and the relative generation times (from 0.2 to 35 days)showed a high range of values, representing a variety of situations, fromestuaries to the ocean. The resulting role of the bacterial community in thecarbon cycle is very consistent, processing amounts of carbon which havebeen estimated as high as the 80% and the 260% of thosesynthesized by autotrophs in summer and winter, respectively. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ecological advantages from light adaptation and heterotrophic-like behavior in Synechococcus harvested from the Gulf of Trieste (Northern Adriatic Sea)

A preliminary study was carried out on a picocyanobacterial mixed culture harvested from the Gulf of Trieste (Northern Adriatic) and identified as Synechococcus spp. both by transmission electron microscopy observations, biliprotein composition and molecular analyses. Absorption and fluorescence spectra revealed phycourobilin and phycoerythrobilin chromophores, suggesting the presence of both CU- and C-phycoerythrin, besides phycocyanobilin chromophores typical for phycocyanins and allophycocyanins. Both biliprotein analyses and molecular identification indicated the presence of at least two Synechococcus subgroups presumably differing either in phycoerythrin type or in physiological traits. Among the exoenzymatic activities acting on different substrates, only aminopeptidase showed high hydrolysis rates and the uptake of organic molecules was positive for leucine but not for thymidine. The protein carbon mobilized was high compared with the leucine incorporation rates, resulting in low percentages of newly mobilized carbon utilized by cultures. The organic carbon incorporated as leucine was compared with the photosynthetically produced one, and the balance between the phototrophic- and heterotrophic-like processes was c. 3 : 1. Our findings suggest that the Synechococcus heterotrophy plays an important role in cell's metabolism, and that the photoheterotrophic behavior, together with their chromatic adaptation capability, might represent the key for the absolute dominance of this genus in the Adriatic Sea.     

Microphytobenthic Primary Production and Sedimentary Carbohydrates Along Salinity Gradients in the Lagoons of Grado and Marano (Northern Adriatic Sea)

Primary production of the microphytobenthic community and carbohydrates concentrations were studied in the lagoonal system of Grado and Marano, located in the Northern Adriatic coast. Sediment samples were collected along a salinity gradient. Abundance and species composition of the microphytobenthic communities were analysed and the benthic microalgal biomass was estimated as Chlorophyll a (Chl a). Primary production of benthic diatoms was estimated using ¹⁴C-tracer. Extracellular carbohydrates were extracted from the sediment and separated in two operationally defined fractions (colloidal and EDTA-extractable). Salinity was higher in the Grado lagoon, where the benthic microalgal community was mainly composed of marine diatoms. In the Marano lagoon, which has a lower salinity, freshwater species were also found. In both lagoons, photosynthetic efficiency showed an inverse relationship with salinity and a direct relationship with the main biological variables. Photosynthetic activity was directly related to Chl a and abundance of benthic microalgae, suggesting that in the benthic system microalgal community is responsible for primary production. Overall, salinity was also influent on the microphytobenthic primary production, which was greater in the more saline Grado lagoon.     

The Colonisation of Human-made Structures by the Invasive Alga Codium fragile ssp. tomentosoides in the North Adriatic Sea (NE Mediterranean)

Human-made structures, such as groynes, breakwaters, seawalls, pier pilings and floating pontoons, are becoming common features of the landscape in urbanised coastal and estuarine areas. Despite this tendency few studies have focused on their ecology or on their potential impacts on natural assemblages of organisms. When artificial structures are introduced in areas with little or no hard substrata, they not only provide novel habitats, which enables the colonisation of sandy areas by hard-bottom dwelling species, but they can also provide suitable habitats for exotic species. Along the north-east coast of Italy, sandy shores are protected from erosion by a line of breakwaters, which runs almost uninterrupted for about 300 km. These structures provide habitat for a variety of macroalgae and invertebrates and also for the invasive green alga Codium fragile ssp. tomentosoides. The aim of this study was, therefore, to investigate patterns of distribution of this alga on breakwaters in Cesenatico. In particular, we compared the density of thalli, biomass, length and degree of branching of C. fragile ssp. tomentosoides between the landward and the seaward sides of breakwaters, to test the hypothesis that sheltered habitats (landward) represent more suitable habitats than exposed habitats (seaward). In general, the landward side of breakwaters supported greater numbers of thalli of C. fragile ssp. tomentosoides than seaward sides. Thalli grew longer and more branched in sheltered habitats, leading to an overall larger biomass of the alga on the landward side of breakwaters. The presence of sheltered human-made hard substrata in the vicinity of major trading ports and sources of eutrophication could enhance the dispersal of invasive species across regional and geographic scales. Thus, the effects of artificial structures and introduced species on coastal assemblages cannot be evaluated separately, but their synergistic nature should be considered in planning strategies for conservation of biodiversity in coastal habitats.