The spreading of the introduced seaweed Caulerpa taxifolia (Vahl) C. Agardh in the Mediterranean Sea: testing the boat transportation hypothesis

The Spreading of the Introduced Seaweed Caulerpa taxifolia (Vahl) C. Agardh in the Mediterranean Sea: Testing the Boat Transportation Hypothesis

Quaternary active tectonic structures in the offshore Bajo Segura basin (SE Iberian Peninsula Mediterranean Sea).

The Bajo Segura fault zone (BSFZ) is the northern terminal splay of the Eastern Betic shear zone (EBSZ), a large left-lateral strike-slip fault system of sigmoid geometry stretching more than 450 km from Alicante to Almería. The BSFZ extends from the onshore Bajo Segura basin further into the Mediterranean Sea and shows a moderate instrumental seismic activity characterized by small earthquakes. Nevertheless, the zone was affected by large historical earthquakes of which the largest was the 1829 Torrevieja earthquake (IEMS98 X). The onshore area of the BSFZ is marked by active transpressive structures (faults and folds), whereas the offshore area has been scarcely explored from the tectonic point of view. During the EVENT-SHELF cruise, a total of 10 high-resolution single-channel seismic sparker profiles were obtained along and across the offshore Bajo Segura basin. Analysis of these profiles resulted in (a) the identification of 6 Quaternary seismo-stratigraphic units bounded by five horizons corresponding to regional erosional surfaces related to global sea level lowstands; and (b) the mapping of the active sub-seafloor structures and their correlation with those described onshore. Moreover, the results suggest that the Bajo Segura blind thrust fault or the Torrevieja left-lateral strike-slip fault, with prolongation offshore, could be considered as the source of the 1829 Torrevieja earthquake. These data improve our understanding of present deformation along the BSFZ and provide new insights into the seismic hazard in the area.

Natural or naturalized? phylogeography suggests that the abundant sea urchin arbacia lixula Is a recent Colonizer of the Mediterranean

We present the global phylogeography of the black sea urchin Arbacia lixula, an amphi-Atlantic echinoid with potential to strongly impact shallow rocky ecosystems. Sequences of the mitochondrial cytochrome c oxidase gene of 604 specimens from 24 localities were obtained, covering most of the distribution area of the species, including the Mediterranean and both shores of the Atlantic. Genetic diversity measures, phylogeographic patterns, demographic parameters and population differentiation were analysed. We found high haplotype diversity but relatively low nucleotide diversity, with 176 haplotypes grouped within three haplogroups: one is shared between Eastern Atlantic (including Mediterranean) and Brazilian populations, the second is found in Eastern Atlantic and the Mediterranean and the third is exclusively from Brazil. Significant genetic differentiation was found between Brazilian, Eastern Atlantic and Mediterranean regions, but no differentiation was found among Mediterranean sub-basins or among Eastern Atlantic sub-regions. The star-shaped topology of the haplotype network and the unimodal mismatch distributions of Mediterranean and Eastern Atlantic samples suggest that these populations have suffered very recent demographic expansions. These expansions could be dated 94-205 kya in the Mediterranean, and 31-67 kya in the Eastern Atlantic. In contrast, Brazilian populations did not show any signature of population expansion. Our results indicate that all populations of A. lixula constitute a single species. The Brazilian populations probably diverged from an Eastern Atlantic stock. The present-day genetic structure of the species in Eastern Atlantic and the Mediterranean is shaped by very recent demographic processes. Our results support the view (backed by the lack of fossil record) that A. lixula is a recent thermophilous colonizer which spread throughout the Mediterranean during a warm period of the Pleistocene, probably during the last interglacial. Implications for the possible future impact of A. lixula on shallow Mediterranean ecosystems in the context of global warming trends must be considered.

Time and space: genetic structure of the common sea urchin paracentrotus lividus in the south-eastern iberian coast

Spatio-temporal variability in settlement and recruitment, high mortality during the first life-history stages, and selection may determine the genetic structure of cohorts of long-lived marine invertebrates at small scales. We conducted a spatial and temporal analysis of the common Mediterranean Sea urchin Paracentrotus lividus to determine the genetic structure of cohorts at different scales. In Tossa de Mar (NW Mediterranean), recruitment was followed over 5 consecutive springs (2006-2010). In spring 2008, recruits and two-year-old individuals were collected at 6 locations along East and South Iberian coasts separated from 200 to over 1,100 km. All cohorts presented a high genetic diversity based on a fragment of mtCOI. Our results showed a marked genetic homogeneity in the temporal monitoring and a low degree of spatial structure in 2006. In 2008, coupled with an abnormality in the usual circulation patterns in the area, the genetic structure of the southern populations studied changed markedly, with arrival of many private haplotypes. This fact highlights the importance of point events in renewing the genetic makeup of populations, which can only be detected through analysis of the cohort structure coupling temporal and spatial perspectives.

The reproductive cycle of the sea urchin Arbacia lixula in northwest mediterranean: potential influence of temperature and photoperiod

We studied the reproductive cycle of the sea urchin Arbacia lixula in a subtidal population from northeast Spain over four years using a gonadosomatic index (GSI) and gonad histology. Our results show that the GSI of A. lixula follows a seasonal cycle which peaks in May-July and attains its lowest values in October-November every year. The time course of the GSI matched closely the photoperiod cycle. We also found a remarkable inter-annual variability in the maximum value of GSI, which correlated with mean water temperature during the gonad growth period (winter and spring). Gonad histology was also in agreement with a single gametogenic cycle per year in this species. We explored the application of circular statistics to present and analyse gonadal development data, which allowed us to adequately handle the high intra-individual variability detected, with several developmental stages commonly found within the same gonad. The picture that emerged is one of a gametogenic timing driven by photoperiod, while the amount of reproductive output is determined by temperature. This is coherent with the tropical origin of the species and lends support to recent warnings about an increase in the abundance of this species in the Mediterranean as a result of global warming, with associated increased impact potential in sublittoral communities.

Bioavailability of sinking organic matter in the Blanes canyon and the adjacent open slope (NW Mediterranean Sea)

Submarine canyons are sites of intense energy and material exchange between the shelf and the deep adjacent basins. To test the hypothesis that active submarine canyons represent preferential conduits of available food for the deep-sea benthos, two mooring lines were deployed at 1200 m depth from November 2008 to November 2009 inside the Blanes canyon and on the adjacent open slope (Catalan Margin, NW Mediterranean Sea). We investigated the fluxes, biochemical composition and food quality of sinking organic carbon (OC). OC fluxes in the canyon and the open slope varied among sampling periods, though not onsistently in the two sites. In particular, while in the open slope the highest OC fluxes were observed in August 2009, in the canyon the highest OC fluxes occurred in April-May 2009. For almost the entire study period, the OC fluxes in the canyon were significantly higher than those in the open slope, whereas OC contents of sinking particles collected in the open slope were consistently higher than those in the canyon. This result confirms that submarine canyons are effective conveyors of OC to the deep sea. Particles transferred to the deep sea floor through the canyons are predominantly of inorganic origin, significantly higher than that reaching the open slope at a similar water depth. Using multivariate statistical tests, two major clusters of sampling periods were identified: one in the canyon that grouped trap samples collected in December 2008, oncurrently with the occurrence of a major storm at the sea surface, and associated with increased fluxes of nutritionally available particles from the upper shelf. Another cluster grouped samples from both the canyon and the open slope collected in March 2009, concurrently with the occurrence of the seasonal phytoplankton bloom at the sea surface, and associated with increased fluxes of total phytopigments. Our results confirm the key ecological role of submarine canyons for the functioning of deep-sea ecosystems, and highlight the importance of canyons in linking episodic storms and primary production occurring at the sea surface to the deep sea floor.

Metabolic effects of insulin and IGFs on gilthead sea bream (Sparus aurata) muscle cells.

Primary cultures of gilthead sea bream myocytes were performed in order to examine the relative metabolic function of insulin compared with IGF-I and IGF-II (insulin-like growth factors, IGFs) at different stages in the cell culture. In these cells, the in vitro effects of insulin and IGFs on 2-deoxyglucose (2-DG) and L-alanine uptake were studied in both myocytes (day 4) and small myotubes (day 9). 2-DG uptake in gilthead sea bream muscle cells was increased in the presence of insulin and IGFs in a time dependent manner and along with muscle cell differentiation. On the contrary, L-alanine uptake was also stimulated by insulin and IGFs but showed an inverse pattern, being the uptake higher in small myocytes than in large myotubes. The results of preincubation with inhibitors (PD-98059, wortmannin, and cytochalasin B) on 2-DG uptake indicated that insulin and IGFs stimulate glucose uptake through the same mechanisms, and evidenced that mitogenesis activator protein kinase (MAPK) and PI3K-Akt transduction pathways mediate the metabolic function of these peptides. In the same way, we observed that GLUT4 protein synthesis was stimulated in the presence of insulin and IGFs in gilthead sea bream muscle cells in a different manner at days 4 or 9 of the culture. In summary we describe here, for the first time, the effects of insulin and IGFs on 2-DG and L-alanine uptake in primary culture of gilthead sea bream muscle cells. We show that both MAPK and PI3K-Akt transduction pathways are needed in order to control insulin and IGFs actions in these cells. Moreover, changes in glucose uptake can be explained by the action of the GLUT4 transporter, which is stimulated in the presence of insulin and IGFs throughout the cell culture.

Multiple processes regulate long-term population dynamics of sea urchins on Mediterranean rocky reefs

We annually monitored the abundance and size structure of herbivorous sea urchin populations (Paracentrotus lividus and Arbacia lixula) inside and outside a marine reserve in the Northwestern Mediterranean on two distinct habitats (boulders and vertical walls) over a period of 20 years, with the aim of analyzing changes at different temporal scales in relation to biotic and abiotic drivers. P. lividus exhibited significant variability in density over time on boulder bottoms but not on vertical walls, and temporal trends were not significantly different between the protection levels. Differences in densities were caused primarily by variance in recruitment, which was less pronounced inside the MPA and was correlated with adult density, indicating density-dependent recruitment under high predation pressure, as well as some positive feedback mechanisms that may facilitate higher urchin abundances despite higher predator abundance. Populations within the reserve were less variable in abundance and did not exhibit the hyper-abundances observed outside the reserve, suggesting that predation effects maybe more subtle than simply lowering the numbers of urchins in reserves. A. lixula densities were an order of magnitude lower than P. lividus densities and varied within sites and over time on boulder bottoms but did not differ between protection levels. In December 2008, an exceptionally violent storm reduced sea urchin densities drastically (by 50% to 80%) on boulder substrates, resulting in the lowest values observed over the entire study period, which remained at that level for at least two years (up to the present). Our results also showed great variability in the biological and physical processes acting at different temporal scales. This study highlights the need for appropriate temporal scales for studies to fully understand ecosystem functioning, the concepts of which are fundamental to successful conservation and management.

The El Masnou Infralittoral sedimentary environment (Barcelona province, NW Mediterranean Sea): morphology and Holocene seismic stratigraphy

A detailed analysis of the morphology and the Holocene seismic and sequence stratigraphy and architecture of the infralittoral sedimentary environment of the El Masnou coast (Catalonia, NW Mediterranean Sea) was carried out using multibeam bathymetry and GeoPulse seismic data. This environment extends down to 26-30 m water depth, and is defined morphologically by two depositional wedges whose seafloor is affected by erosive furrows, slides, fields of large- and small-scale wavy bedforms, and dredging trenches and pits. Erosive terraces are also identified in the transition domain toward the inner continental shelf. The Holocene stratigraphy of the infralittoral environment is defined by two major seismic sequences (lower and upper), each one formed by internal seismic units. The sequences and units are characterised by downlapping surfaces made up of deposits formed by progradation of coastal lithosomes. The stratigraphy and stratal architecture, displaying a retrogradational arrangement with progradational patterns of minor order, were controlled by different sea-level positions. The stratigraphic division represents the coastal response to the last fourth-order transgressive and highstand conditions, modulated by small-scale sea-level oscillations (≈1-2 m) of fith to sixth order. This study also highlights the advantage of an integrated analysis using acoustic/seismic methods for practical assessment of the anthropogenic effects on infralittoral domains based on the association of marine geological observations.

Tracking Invasion Histories in the Sea: Facing Complex Scenarios Using Multilocus Data

In recent years, new analytical tools have allowed researchers to extract historical information contained in molecular data, which has fundamentally transformed our understanding of processes ruling biological invasions. However, the use of these new analytical tools has been largely restricted to studies of terrestrial organisms despite the growing recognition that the sea contains ecosystems that are amongst the most heavily affected by biological invasions, and that marine invasion histories are often remarkably complex. Here, we studied the routes of invasion and colonisation histories of an invasive marine invertebrate Microcosmus squamiger (Ascidiacea) using microsatellite loci, mitochondrial DNA sequence data and 11 worldwide populations. Discriminant analysis of principal components, clustering methods and approximate Bayesian computation (ABC) methods showed that the most likely source of the introduced populations was a single admixture event that involved populations from two genetically differentiated ancestral regions - the western and eastern coasts of Australia. The ABC analyses revealed that colonisation of the introduced range of M. squamiger consisted of a series of non-independent introductions along the coastlines of Africa, North America and Europe. Furthermore, we inferred that the sequence of colonisation across continents was in line with historical taxonomic records - first the Mediterranean Sea and South Africa from an unsampled ancestral population, followed by sequential introductions in California and, more recently, the NE Atlantic Ocean. We revealed the most likely invasion history for world populations of M. squamiger, which is broadly characterized by the presence of multiple ancestral sources and non-independent introductions within the introduced range. The results presented here illustrate the complexity of marine invasion routes and identify a cause-effect relationship between human-mediated transport and the success of widespread marine non-indigenous species, which benefit from stepping-stone invasions and admixture processes involving different sources for the spread and expansion of their range.

Sponge Mass Mortalities in a Warming Mediterranean Sea: Are Cyanobacteria-Harboring Species Worse Off?

Mass mortality events are increasing dramatically in all coastal marine environments. Determining the underlying causes of mass mortality events has proven difficult in the past because of the lack of prior quantitative data on populations and environmental variables. Four-year surveys of two shallow-water sponge species, Ircinia fasciculata and Sarcotragus spinosulum, were carried out in the western Mediterranean Sea. These surveys provided evidence of two severe sponge die-offs (total mortality ranging from 80 to 95% of specimens) occurring in the summers of 2008 and 2009. These events primarily affected I. fasciculata, which hosts both phototrophic and heterotrophic microsymbionts, while they did not affect S. spinosulum, which harbors only heterotrophic bacteria. We observed a significant positive correlation between the percentage of injured I. fasciculata specimens and exposure time to elevated temperature conditions in all populations, suggesting a key role of temperature in triggering mortality events. A comparative ultrastructural study of injured and healthy I. fasciculata specimens showed that cyanobacteria disappeared from injured specimens, which suggests that cyanobacterial decay could be involved in I. fasciculata mortality. A laboratory experiment confirmed that the cyanobacteria harbored by I. fasciculata displayed a significant reduction in photosynthetic efficiency in the highest temperature treatment. The sponge disease reported here led to a severe decrease in the abundance of the surveyed populations. It represents one of the most dramatic mass mortality events to date in the Mediterranean Sea

Cyanobacterial diversity and a new Acaryochloris-like symbiont from Bahamian sea-squirts

Symbiotic interactions between ascidians (sea-squirts) and microbes are poorly understood. Here we characterized the cyanobacteria in the tissues of 8 distinct didemnid taxa from shallow-water marine habitats in the Bahamas Islands by sequencing a fragment of the cyanobacterial 16S rRNA gene and the entire 16S-23S rRNA internal transcribed spacer region (ITS) and by examining symbiont morphology with transmission electron (TEM) and confocal microscopy (CM). As described previously for other species, Trididemnum spp. mostly contained symbionts associated with the Prochloron-Synechocystis group. However, sequence analysis of the symbionts in Lissoclinum revealed two unique clades. The first contained a novel cyanobacterial clade, while the second clade was closely associated with Acaryochloris marina. CM revealed the presence of chlorophyll d (chl d) and phycobiliproteins (PBPs) within these symbiont cells, as is characteristic of Acaryochloris species. The presence of symbionts was also observed by TEM inside the tunic of both the adult and larvae of L. fragile, indicating vertical transmission to progeny. Based on molecular phylogenetic and microscopic analyses, Candidatus Acaryochloris bahamiensis nov. sp. is proposed for this symbiotic cyanobacterium. Our results support the hypothesis that photosymbiont communities in ascidians are structured by host phylogeny, but in some cases, also by sampling location.

Deep incision in an Aptian carbonate succession indicates major sea-level fall in the Cretaceous

Long-term relative sea-level cycles (0 5 to 6 Myr) have yet to be fully understood for the Cretaceous. During the Aptian, in the northern Maestrat Basin (Eastern Iberian Peninsula), fault-controlled subsidence created depositional space, but eustasy governed changes in depositional trends. Relative sea-level history was reconstructed by sequence stratigraphic analysis. Two forced regressive stages of relative sea-level were recognized within three depositional sequences. The first stage is late Early Aptian age (intra Dufrenoyia furcata Zone) and is characterized by foreshore to upper shoreface sedimentary wedges, which occur detached from a highstand carbonate platform, and were deposited above basin marls. The amplitude of relative sea-level drop was in the order of tens of metres, with a duration of <1 Myr. The second stage of relative sea-level fall occurred within the Late Aptian and is recorded by an incised valley that, when restored to its pre-contractional attitude, was >2 km wide and cut 115 m down into the underlying Aptian succession. With the subsequent transgression, the incision was back-filled with peritidal to shallow subtidal deposits. The changes in depositional trends, lithofacies evolution and geometric relation of the stratigraphic units characterized are similar to those observed in coeval rocks within the Maestrat Basin, as well as in other correlative basins elsewhere. The pace and magnitude of the two relative sea-level drops identified fall within the glacio-eustatic domain. In the Maestrat Basin, terrestrial palynological studies provide evidence that the late Early and Late Aptian climate was cooler than the earliest part of the Early Aptian and the Albian Stage, which were characterized by warmer environmental conditions. The outcrops documented here are significant because they preserve the results of Aptian long-term sea-level trends that are often only recognizable on larger scales (i.e. seismic) such as for the Arabian Plate.

Deep incision in an Aptian carbonate succession indicates major sea-level fall in the Cretaceous

Long-term relative sea-level cycles (0 5 to 6 Myr) have yet to be fully understood for the Cretaceous. During the Aptian, in the northern Maestrat Basin (Eastern Iberian Peninsula), fault-controlled subsidence created depositional space, but eustasy governed changes in depositional trends. Relative sea-level history was reconstructed by sequence stratigraphic analysis. Two forced regressive stages of relative sea-level were recognized within three depositional sequences. The first stage is late Early Aptian age (intra Dufrenoyia furcata Zone) and is characterized by foreshore to upper shoreface sedimentary wedges, which occur detached from a highstand carbonate platform, and were deposited above basin marls. The amplitude of relative sea-level drop was in the order of tens of metres, with a duration of <1 Myr. The second stage of relative sea-level fall occurred within the Late Aptian and is recorded by an incised valley that, when restored to its pre-contractional attitude, was >2 km wide and cut 115 m down into the underlying Aptian succession. With the subsequent transgression, the incision was back-filled with peritidal to shallow subtidal deposits. The changes in depositional trends, lithofacies evolution and geometric relation of the stratigraphic units characterized are similar to those observed in coeval rocks within the Maestrat Basin, as well as in other correlative basins elsewhere. The pace and magnitude of the two relative sea-level drops identified fall within the glacio-eustatic domain. In the Maestrat Basin, terrestrial palynological studies provide evidence that the late Early and Late Aptian climate was cooler than the earliest part of the Early Aptian and the Albian Stage, which were characterized by warmer environmental conditions. The outcrops documented here are significant because they preserve the results of Aptian long-term sea-level trends that are often only recognizable on larger scales (i.e. seismic) such as for the Arabian Plate.

Vegetative and reproductive morphology of Kallymenia patens (Kallymeniaceae, Rhodophyta) in the Mediterranean Sea

Reproductive morphology of the Mediterranean red alga Kallymenia patens is described for the first time, confirming its position in the genus. K. patens is characterized by a non-procarpic female reproductive apparatus, carpogonial branch systems consisting of supporting cells bearing both three-celled carpogonial branches and subsidiary cells that lack a hypogynous cell and carpogonium; fusion cells develop numerous connecting filaments, and tetrasporangia are scattered over the thallus and are probably cruciately divided. Old fertile spathulate specimens of K. patens are morphologically similar to K. spathulata, but they can be distinguished by the length of spathulated proliferations (up to 0.6 cm and 6 cm, respectively), the length of inner cortical cells (up to 70 and 30 μm, respectively), and the gonimoblast location (in proliferations from the perennial part of the blade and over all the thallus surface, respectively)