Comparison between amphipod assemblages associated with Caulerpa racemosa var. cylindracea and those of other Mediterranean habitats on soft substrate

The spread of the invasive alga Caulerpa racemosa var. cylindracea in shallow-water habitats can present different faunal assemblage composition. We compared the amphipod assemblages associated with C. racemosa and natural habitats found on shallow-water Mediterranean soft substrata. Four vegetated habitats were compared: C. racemosa, Caulerpa prolifera, Cymodocea nodosa and Posidonia oceanica with unvegetated substrata. Samples were collected during two sampling periods (September 2004 and March 2005). A total of 63 amphipod species were recorded. The results showed that the vegetated habitats sampled, including C. racemosa stands, supported a higher abundance and species richness of amphipods. Furthermore, the assemblage structure differed between the different habitats, while the abundance of some species was significantly different, depending on habitat. For example, Microdeutopus obtusatus was favoured by C. racemosa habitat; Ampelisca diadema was associated with C. prolifera beds; and Hyale schmidti was more abundant in P. oceanica meadows. Habitat invasion by C. racemosa can exert an important influence on biotic assemblages, modifying habitat structure and associated fauna.     

Temporal and spatial variability in shallow- and deep-water populations of the invasive Caulerpa racemosa var. cylindracea in the Western Mediterranean

The invasive green alga Caulerpa racemosa var. cylindracea represents an important threat to the diversity of Mediterranean benthic coastal ecosystems by interfering with native species and modifying benthic assemblages. The present study deals, for the first time, with the temporal and spatial variability of the biomass and phenology of C. racemosa considering both deep- and shallow-water populations. Two sampling depths (30 and 10 m) were sampled at three different rocky bottom sites every 3 months in the Archipelago of Cabrera National Park (Western Mediterranean). All morphometric variables analysed showed a spatial variation and temporal patterns depending on depth. Between depths, C. racemosa biomass, stolon length, number of fronds and frond length were usually significantly higher at deep-water populations, suggesting that C. racemosa grows better in deep-waters. Deep- and shallow-water populations displayed a high temporal variation although no evidence of seasonal patterns was found, in contrast with what has been reported by other authors. The sources of this variability are still unknown but probably both physical factors and differential herbivory pressures display a key role.     

Juvenile bottlenecks and salinity shape grey mullet assemblages in Mediterranean estuaries

Previous research has suggested that competitive bottlenecks may exist for the Mediterranean grey mullets (Osteichthyes, Mugilidae) at the fry stage with the exotic Cyprinus carpio (Osteichthyes, Cyprinidae) playing a central role. As a consequence, the structure of grey mullet assemblages at later stages is thought to reflect previous competition as well as differences in osmoregulatory skills. This paper tests that hypothesis by examining four predictions about the relative abundance of five grey mullet species in 42 Western Mediterranean estuary sites from three areas (Aiguamolls de l'Empordà, Ebro Delta and Minorca) differing in the salinity level and occurrence of C. carpio. Field data confirmed the predictions as: (1) Liza aurata and Mugil cephalus were scarce everywhere and never dominated the assemblage; (2) Liza saliens dominated the assemblage where the salinity level was higher than 13; (3) Liza ramado always dominated the assemblage where the salinity level was lower than 13 and C. carpio was present; and (4) Chelon labrosus dominated the assemblage only where the salinity level was lower than 13 and C. carpio was absent. The catch per unit effort of C. labrosus of any size was smaller in the presence of C. carpio than where it had not been introduced, which is in agreement with the juvenile competitive bottleneck hypothesis. Discriminant analysis confirmed that the assemblage structure was linked to the salinity level and the occurrence of C. carpio for both early juveniles and late juveniles as well as adults. The data reported here reveal that the structure of grey mullet assemblages inhabiting Mediterranean estuaries is determined by salinity and competitive interactions at the fry stage.     

Assessment of substratum effect on the distribution of two invasive Caulerpa (Chlorophyta) species

Two-year monitoring of the invasive marine Chlorophyta Caulerpa taxifolia and Caulerpa racemosa var. cylindracea shows the great influence of substratum on their spatial distribution. The cover of C. taxifolia and C. racemosa was measured in shallow (<8 m) areas indicating that these species are more abundant in rocks with photophilic algae and in the dead matte of the seagrass Posidonia oceanica than in sand or inside the P. oceanica meadow. A short-term experiment comparing the persistence of C. taxifolia and C. racemosa planted either in a model of dead matte of P. oceanica or in sand shows that the persistence of these species was higher in the dead matte model than in sand. Correlative evidence suggests that C. taxifolia and C. racemosa tolerate near-bottom orbital velocities below 15 cm s^?1 and that C. taxifolia cover declines at velocities above that value. These results contribute to understand the process of invasion of these Caulerpa species predicting which substrata would be more susceptible to be invaded and to the adoption of appropriate management strategies.     

Expansion of the macroalga Caulerpa racemosa and changes in softbottom macrofaunal assemblages in Moni Bay,Cyprus

The recent expansion of the Red Sea macroalga Caulerpa racemosa and its impact on the diversity and abundance of macrobenthos were examined and compared in the summers of 1992 and 1997, in Moni Bay, Cyprus. The phytobenthic community of the bay in 1992 was dominated by the seagrass Posidonia oceanica while, in 1997, the Lessepsian migrant C. racemosa became the most dominant, forming extensive mats. Changes in the vegetation system in Moni Bay have caused significant compositional changes in macrofaunal assemblages. A total of 178 individuals of 62 species are recorded. The composition of the macrofauna in 1992 was dominated by gastropods (44 %), crustaceans (22%), bivalves (17 %), polychaetes (11 %) and echinoderms (6 %). In 1997, the gastropods and crustaceans had decreased to 13 % and 16 % respectively, while, polychaetes had increased to 38 % becoming the most dominant taxon. Bivalves and echinoderms also increased to 22 % and 11 %, respectively, in 1997. The proliferative growth of C. racemosa imposed successional changes on the macrofaunal assemblages in Moni Bay, Cyprus, between 1992 and 1997. It remains to be tested whether the expansion of C. racemosa is related to the increase of water temperature associated with global warming or nutrient inputs or with the differences in the life history characteristics of this migrant vs. native algal species.     

Functional changes due to invasive species: Food web shifts at shallow Posidonia oceanica seagrass beds colonized by the alien macroalga Caulerpa racemosa

Multiple stable isotope analyses were used to examine the trophic shifts at faunal assemblages within the invading macroalga Caulerpa racemosa in comparison to established communities of Posidonia oceanica seagrass meadows. Sampling of macrobenthic invertebrates and their potential food sources of algal mats and seagrass meadows in Mallorca (NW Mediterranean) showed differences in species composition of faunal and primary producers among seagrass and C. racemosa. Accordingly, changes in food web structure and trophic guilds were observed, not only at species level but also at community level. The carbon and nitrogen isotope signatures of herbivores, detritivores and deposit feeders confirmed that the seagrass provided a small contribution to the macrofaunal organisms. δ¹³C at the P. oceanica seagrass and at the C. racemosa assemblages differed, ranging from −6.19 to −21.20‰ and −2.67 to −31.41‰, respectively. δ¹⁵N at the Caulerpa mats was lower (ranging from 2.64 to 10.45‰) than that at the seagrass meadows (3.51–12. 94‰). Significant differences in isotopic signatures and trophic level among trophic guilds at P. oceanica and C. racemosa were found. N fractionation at trophic guild level considerable differed between seagrass and macroalgae mats, especially for detritivores, deposit feeders, and herbivores. Filter feeders slightly differed with a relatively lower N signal at the seagrass and CR values at community level and at trophic guild level were higher in the C. racemosa invaded habitats indicating an increase in diversity of basal resource pools. C. racemosa did seem to broaden the niche diversity of the P. oceanica meadows it colonised at the base of the food web, may be due to the establishment of a new basal resource. The extent of the effects of invasive species on ecosystem functioning is a fundamental issue in conservation ecology. The observed changes in invertebrate and macrophytic composition, stable isotope signatures of concomitant species and consequent trophic guild and niche breadth shifts at invaded Caulerpa beds increase our understanding of the seagrass systems.     

Differential herbivory of invasive algae by native fish in the Mediterranean Sea

The potential role of generalist herbivores to serve as a source of biotic resistance against algal invasion in marine ecosystems has been poorly examined. The present study investigates the capacity of Mediterranean herbivorous fishes to consume three of the most invasive seaweeds of the Western Mediterranean (Caulerpa racemosa, Lophocladia lallemandii and Womersleyella setacea) and examines vertical and temporal variations of such consumption. Our results show that although fish feed throughout the depth gradient examined (5–35 m), they concentrate in shallow waters, and can consume high amounts of C. racemosa. Such high ingestion of C. racemosa does not appear to be random, since this alga is consistently chosen when offered in pairs with several native species. Conversely, L. lallemandii and W. setacea are barely eaten by fish even though they can be very abundant in the field throughout the year. Our results suggest that fish could be an important controlling agent that has been overlooked in temperate marine invasions, and they may be able to provide certain resistance to C. racemosa invasion. In contrast, they are unlikely to exert any important control effects on L. lallemandii or W. setacea.     

Differences in spatial and seasonal patterns of macrophyte assemblages between a coastal lagoon and the open sea

Although benthic macrophytes must be considered in monitoring programs to establish the ecological status of transitional and coastal waters in the European Union, the patterns of variability in species composition of macrophyte assemblages in Mediterranean coastal lagoons has scarcely been studied. In this work the spatial (both vertical and horizontal) and seasonal dynamics of macrophyte assemblages in a coastal lagoon (Mar Menor) are compared with those of open coastal assemblages in the SW Mediterranean to analyze any biological variability in lagoon assemblages and the factors that determine such variability. Different assemblages, characterized by well defined groups of species, can be described according to their isolation from the open sea and the type of substratum; at the same time, a vertical zonation pattern, similar to that found in all marine communities but more compressed, exists. This implies that when applying the EU Water Framework Directive or assessing environmental impact, a lagoon should not be considered spatially uniform and unique unit but as a mosaic of assemblages.     

Habitat preferences and life history of the red scorpion fish, Scorpaena notata, in the Mediterranean

Scorpaena notata is a small, sedentary scorpaenid species widely distributed in the Mediterranean and adjacent waters of the Atlantic. In the western Mediterranean it inhabits coastal continental shelf bottoms. In the Balearic Islands, these bottoms are characterised by the presence of the facies with red algae, including both Peyssonnelia and mäerl beds. These beds enhance the structural complexity, biodiversity and secondary production of the soft bottoms. Due to the oceanographic conditions of the Islands, the facies with red algae are especially rich in terms of biomass and algal coverage, and are widespread distributed between 40 and 90 m depth, where trawlers exploiting the continental shelf operate. The present work studies the biology of S. notata and its relationship with habitat characteristics. Special attention is focused on the aspects related to fish condition and growth as a tool to assess the importance of the facies with red algae for fish.     

Persistence of biological invasion effects: Recovery of macroalgal assemblages after removal of Caulerpa racemosa var. cylindracea

The present study evaluated the capability of recovery of Mediterranean macroalgal assemblages on rocky bottom invaded by Caulerpa racemosa. A manipulative experiment was performed to compare the structure of native assemblages that: (a) were invaded by the alga, (b) were invaded, but where the invasive alga was removed, and (c) had never been invaded. Macroalgal assemblages differed significantly between areas invaded by C. racemosa and control areas. Moreover, one year after removal of the invader, the recovery of community was minimal. Overall, recovery consisted of a significant increase in species richness and cover of macroalgae, but cover of each vegetation layer (encrusting, turf and erect) did not vary significantly between cleared and Invaded plots through time. Results showed that the effects of the colonization persist after the removal of the alga and the process of recovery of the assemblages appears to be quite slow. This finding might be an important contribution to evaluate invasion costs and thus potentially useful for an effective addressing of control strategies.     

The spread of Caulerpa racemosa var. cylindracea in the Mediterranean Sea: an example of how biological invasions can influence beta diversity

The present study aims to evaluate if invasion of Caulerpa racemosa var. cylindracea can affect both alpha and beta diversity in Mediterranean coastal system. The following hypotheses were tested: (i) measures of alpha diversity differ between invaded and non-invaded assemblages (ii) this pattern is consistent between different habitats, (iii) assemblages from habitats which are naturally very different tend to be more similar when invaded compared to non-invaded situations. Results showed that Mediterranean benthic assemblages invaded by C. racemosa var. cylindracea were characterized by lower values of alpha diversity than non-invaded assemblages; moreover, differences between deep (25 m) and shallow water (5m) assemblages appeared lower in invaded than in non-invaded areas, with a decrease of beta diversity. This homogenisation of habitats was related to a lack of several characteristic species and to a higher abundance of few opportunistic species in invaded areas, particularly turf-forming algae. Moreover, variability in taxa composition was lower in invaded than in non-invaded assemblages in both habitats.     

Detecting the intertidal morphologic change using satellite data

Previous research has suggested that competitive bottlenecks may exist for the Mediterranean grey mullets (Osteichthyes, Mugilidae) at the fry stage with the exotic Cyprinus carpio (Osteichthyes, Cyprinidae) playing a central role. As a consequence, the structure of grey mullet assemblages at later stages is thought to reflect previous competition as well as differences in osmoregulatory skills. This paper tests that hypothesis by examining four predictions about the relative abundance of five grey mullet species in 42 Western Mediterranean estuary sites from three areas (Aiguamolls de l'Empordà, Ebro Delta and Minorca) differing in the salinity level and occurrence of C. carpio. Field data confirmed the predictions as: (1) Liza aurata and Mugil cephalus were scarce everywhere and never dominated the assemblage; (2) Liza saliens dominated the assemblage where the salinity level was higher than 13; (3) Liza ramado always dominated the assemblage where the salinity level was lower than 13 and C. carpio was present; and (4) Chelon labrosus dominated the assemblage only where the salinity level was lower than 13 and C. carpio was absent. The catch per unit effort of C. labrosus of any size was smaller in the presence of C. carpio than where it had not been introduced, which is in agreement with the juvenile competitive bottleneck hypothesis. Discriminant analysis confirmed that the assemblage structure was linked to the salinity level and the occurrence of C. carpio for both early juveniles and late juveniles as well as adults. The data reported here reveal that the structure of grey mullet assemblages inhabiting Mediterranean estuaries is determined by salinity and competitive interactions at the fry stage.     

Deep-water stands of Cystoseira zosteroides C. Agardh (Fucales,Ochrophyta) in the Northwestern Mediterranean: Insights into assemblage structure and population dynamics

Populations dominated by Cystoseira zosteroides, an endemic and threatened Mediterranean seaweed, colonize deep-water rocky habitats down to more than 50 m depth. Assemblages dominated by this species display high algal and invertebrate species richness. Algal biomass averages 1134 g dw m⁻². Erect and turf algae account for only 25% of total algal dry weight, while encrusting corallines are responsible for the remaining 75%. Sponges, bryozoans and ascidians constitute the dominant sessile macrofauna. Cystoseira zosteroides is the dominant erect algae, with a mean biomass of 60.6 g dw m⁻², and densities ranging from 4 to 7 plants m⁻². The alien turf alga Womersleyella setacea has a biomass of 104.2 g dw m⁻² and covers most of the understory substrate. The size-frequency distribution of C. zosteroides populations shows differences over time. Mean annual growth of the main axis is around 0.5 cm and mean annual mortality rate is lower than 2%. Recruitment was almost nil during the studied period of time (10 years). Processes structuring these deep-water Cystoseira stands must be driven by episodic disturbances, after-disturbance recruitment pulses, and long periods of steady growth that last at least 10 years. However, it is also possible that recruitment is irreversibly inhibited by the alien alga W. setacea in which case these old-growth stands are faced with extinction. The highly diversified assemblages and the low growth and low mortality rates of C. zosteroides indicate high vulnerability to natural and anthropogenic disturbances, and call for effective measures to ensure their conservation.     

Temporal and cross-shelf distribution of ichthyoplankton in the central Cantabrian Sea

Environmental variables have been measured and sampling for ichthyoplankton has been conducted monthly, since April 2001, at three stations, located at the inner (1), middle (2) and outer (3) shelf of the central Cantabrian Sea. This paper presents the results of the study of the ichthyoplankton collected from July 2001 to June 2004. Fish larvae from 99 species, belonging to 37 families, were identified. Families with higher number of species were Gadidae, Sparidae and Labridae. The larval fish assemblage was dominated by pelagic fish species, with Sardina pilchardus, as the most abundant. There was a pronounced spring peak in larval abundance, dominated by S. pilchardus. A smaller peak, dominated by S. pilchardus and Micromesistius poutassou, was recorded in late winter at Stns 2 and 3. This pattern was evident for the three-year study. Results also indicate that this study was limited to the coastal larval fish assemblage inhabiting the central Cantabrian Sea shelf. This assemblage was temporally structured into other three assemblages: winter, late winter–spring and summer–autumn. Temperature was apparently a key factor in larval fish assemblage succession. In a scenario of global warming, this study constitutes a basis to evaluating the ongoing changes in the pelagic coastal ecosystem of the central Cantabrian Sea.     

Effect of a simulated oil spill on natural assemblages of marine phytoplankton enclosed in microcosms

Two microcosm experiments were carried out to simulate the effect of sporadic oil spills derived from tanker accidents on oceanic and coastal marine phytoplankton assemblages. Treatments were designed to reproduce the spill from the Prestige, which took place in Galician coastal waters (NW Iberia) in November 2002. Two different concentrations of the water soluble fraction of oil were used: low (8.6 ± 0.7 μg l⁻¹ of chrysene equivalents) and high (23 ± 5 μg l⁻¹ of chrysene equivalents l⁻¹). Photosynthetic activity and chlorophyll a concentration decreased in both assemblages after 24–72 h of exposure to the two oil concentrations, even though the effect was more severe on the oceanic assemblage. These variables progressively recovered up to values close or higher than those in the controls, but the short-term negative effect of oil, which was generally stronger at the high concentration, also induced changes in the structure of the plankton community. While the biomass of nanoflagellates increased in both assemblages, oceanic picophytoplankton was drastically reduced by the addition of oil. Effects on diatoms were also observed, particularly in the coastal assemblage. The response of coastal diatoms to oil addition showed a clear dependence on size. Small diatoms (<20 μm) were apparently stimulated by oil, whereas diatoms >20 μm were only negatively affected by the high oil concentration. These differences, which could be partially due to indirect trophic interactions, might also be related to different sensitivity of species to PAHs. These results, in agreement with previous observations, additionally show that the negative effect of the water soluble fraction of oil on oceanic phytoplankton was stronger than on coastal phytoplankton.     

Spatial variability of epifaunal communities from artificial habitat: Shipwrecks in the Southern Bight of the North Sea

We investigated the cover, community structure and abiotic environment of nine shipwrecks lying at increasing distance from the Belgian coast. Results indicated that all shipwrecks were strongly dominated by cnidarians in terms of biomass and by amphipods in terms of abundances. Based on their epifaunal composition, three groups of shipwrecks could be determined. Metridium senile dominated a species poor community of the coastal sites. On the same sites, a Tubularia larynx community with a more species-rich assemblage was also developing. The T. larynx community had a lower biomass value (102 g AFDW m⁻²) and significantly lower species richness compared to the other sites. The coastal sites were characterized by periodic salinity decreases, large seasonal temperature fluctuation, high total suspended matter load and reduced current velocity. Channel water masses influence the offshore sites causing a more stable temperature and salinity environment, less turbid waters and high current speed. Tubularia indivisa dominated this community, with an average biomass of 229 g AFDW m⁻². Intermediate sites were also dominated by T. indivisa, but a higher biomass (424 g AFDW m⁻²) was observed. They showed intermediate results for the abiotic parameters and fast current velocities. Hypotheses for the observed variation in community structures are discussed in the light of the abiotic characterization of the shipwrecks.     

Short-term variation in copepod community and physical environment in the waters adjacent to the Kuroshio Current

A continuous survey examined short-term variations in the zooplankton community and physical ocean environment from the northeastern Izu Islands to Boso Peninsula in Japan. High copepod abundance and small upwellings in the surface layer and salinity minimum layer in the subsurface were observed on the north side of coastal fronts in the westernmost transect, moving southward as the Kuroshio Current left the Boso Peninsula. Thus, the salinity minimum layer might be a key factor forming upwelling and the fronts, leading to large abundance of coastal copepods off the northeastern Izu Islands. A community structure analysis of calanoid copepods revealed an intermediate belt assemblage between coastal and offshore (Kuroshio) assemblages. Copepod abundance was remarkably low and Ctenocalanus vanus dominated (nearly 37%) in the intermediate belt zone, indicating that C. vanus has a relatively high tolerance to adverse environments for calanoid copepods. As the Kuroshio Current left the Boso Peninsula, the coastal assemblage expanded in the same direction, and the intermediate belt assemblage off the northeastern Izu Islands disappeared. The largest population of Calanus sinicus was found along the two western transects off the northeastern Izu Islands (>1000 m depth), which was assumed to be transported from Sagami Bay and advanced southwestward while growing from copepodite stages CIII to CV. Larvae of C. sinicus would be an important food for fish larvae in addition to Paracalanus parvus s.l., the numerically dominant species in the coastal assemblage, and C. vanus under the adverse conditions for coastal copepods.     

Physiological response of the sea urchin Paracentrotus lividus fed with the seagrass Posidonia oceanica and the alien algae Caulerpa racemosa and Lophocladia lallemandii

The aim was to determine the effects of alien algae feeding on biomarkers of oxidative stress in the sea urchin Paracentrotus lividus. Sea urchins were fed during three months with the native seagrass Posidonia oceanica, and the alien macroalgae Caulerpa racemosa and Lophocladia lallemandii and biochemical analysis were performed in the gonads. A control group was immediately processed after sampling from the sea. Antioxidant enzyme and glutathione S-transferase activities and GSH concentration were significantly higher in sea urchins fed with alien algae when compared with the control group and the one fed with P. oceanica group. This response was more intense in the group fed with L. lallemandii respect to the C. racemosa group. The concentration of MDA, protein carbonyl derivates and 8-OHdG reported no significant differences between treatments. In conclusion, the invasive algae C. racemosa and L. lallemandii induced an antioxidant response in P. lividus without evident oxidative damage.     

Ecological classification of European transitional waters in the North-East Atlantic eco-region

A new methodology to classify European North-East Atlantic transitional waters into ecological types has been developed based on the most important hydrological and morphological features that are likely to determine the ecology of aquatic systems in transitional waters. Hydrological indicators help identifying if a transitional water area is dominated by fresh or sea water and/or by intertidal or subtidal areas, while morphological indicators allow an estimation of the complexity of the transitional water and the diversity of the habitats involved. Twelve transitional waters of the southern Bay of Biscay were classified using this methodology and the five hydro-morphological types obtained were validated with benthic macro-invertebrate data. Transitional waters with a complex morphology showed the highest values of species diversity, while those with a smaller tidal influence showed lower species diversity. The ‘Scrobicularia’ and ‘Abra’ assemblages, previously identified in the study area, were found to be related to different types of transitional waters. The ‘Abra’ assemblage only appeared in estuaries with a complex morphology and dominated by tidal influences, while the ‘Scrobicularia’ assemblage was detected in all the transitional waters except for a single coastal lagoon. This classification of transitional waters may therefore be useful to establish the biological reference conditions needed for European Directives.