Relative sensitivity of soft-bottom intertidal macrofauna to increased CO2 and experimental stress

We used a controlled CO2 perturbation experiment to test hypotheses about changes in diversity, composition and structure of soft-bottom intertidal macrobenthic assemblages, under realistic and locally relevant scenarios of seawater acidification. Patches of undisturbed sediment were collected from 2 types of intertidal sedimentary habitat in the Ria Formosa coastal lagoon (South Portugal) and exposed to 2 levels of seawater acidification (pH reduced by 0.3 and 0.6 units) and 1 unmanipulated (control) level. After 75 d the assemblages differed significantly between the 2 types of sediment and between field controls and the ex situ treatments, but not among the 3 pH levels tested. The naturally high values of total alkalinity buffered seawater from the changes imposed on carbonate chemistry and may have contributed to offsetting acidification at the local scale. Observed differences on biota were strongly related to the organic matter content and grain-size of the sediments, particularly to the fractions of medium and coarse sand. Soft-bottom intertidal macrofauna was significantly affected by the stress of being held in an artificial environment, but not by CO2-induced seawater acidification. Given the previously observed variations in the sensitivities of marine organisms to seawater acidification, direct extrapolations of the present findings to different regions or other types of assemblages do not seem advisable. However, the contribution of ex situ studies to the assessment of ecosystem-level responses to environmental disturbances could generally be improved by incorporating adequate field controls in the experimental design.

Distribution of planktic foraminifera in surface sediments and sedimentation rates of sediment cores from the Ibero-Moroccan continental rise and slope

Cores from the Atlantic Ibero-Moroccan continental rise and slope contain fine-grained Late Pleistocene and Holocene sediments. These young sediments cover the continental margin in large lensformed litho- and biostratigraphically well-defined units. The total sedimentation rates range from 4 cm/ 1000 yrs. to 27 cm/1000 yrs. off Portugal and from 6 cm/1000 yrs. to 14 cm/1000 yrs. off Morocco. Only a small proportion of these sediments usually consists of sand-sized particles (>0.063 mm) which are mostly dominated by foraminifera. Both planktonic and benthic foraminifera are much more abundant in Late Pleistocene and Holocene samples from the upper slope in comparison to those from the deeper slope and from the abyssal plains. Total sedimentation rates during cool and warm climatic stages are rather similar for both groups of foraminifera. For example, in Late Holocene sediments 7 x 10**3 benthic and 201 x 10**3 planktonic foraminifera (fraction 0.63 -0.20 mm) per 100 cm**2 and 1000 yrs. are preserved in the Tagus, 10-19 X 10**3 benthic and about 1.3 X 10**6 planktonic foraminifera are preserved in the Seine abyssal plain sediments. Values from the upper slope sediments are higher for benthic foraminifera by a factor of 60 off Portugal and 60 to 70 off Morocco. The values for planktonic ones differ by factors of 6-12 and 6 respectively. These numbers seem to reflect differences in production and preservation. Production rates of planktonic foraminifera generally seem to be somewhat higher during Holocene than during Late Pleistocene, and the rates of benthic foraminifera appear rather higher during Late Pleistocene than during Holocene.

Effects of ocean acidification on the swimming ability, development and biochemical responses of sand smelt larvae

Ocean acidification, recognized as a major threat to marine ecosystems, has developed into one of the fastest growing fields of research in marine sciences. Several studies on fish larval stages point to abnormal behaviours, malformations and increased mortality rates as a result of exposure to increased levels of CO2. However, other studies fail to recognize any consequence, suggesting species-specific sensitivity to increased levels of CO2, highlighting the need of further research. In this study we investigated the effects of exposure to elevated pCO2 on behaviour, development, oxidative stress and energy metabolism of sand smelt larvae, Atherina presbyter. Larvae were caught at Arrábida Marine Park (Portugal) and exposed to different pCO2 levels (control: 600 µatm, pH = 8.03; medium: 1000 µatm, pH = 7.85; high: 1800 µatm, pH = 7.64) up to 15 days, after which critical swimming speed (Ucrit), morphometric traits and biochemical biomarkers were determined. Measured biomarkers were related with: 1) oxidative stress-superoxide dismutase and catalase enzyme activities, levels of lipid peroxidation and DNA damage, and levels of superoxide anion production; 2) energy metabolism - total carbohydrate levels, electron transport system activity, lactate dehydrogenase and isocitrate dehydrogenase enzyme activities. Swimming speed was not affected by treatment, but exposure to increasing levels of pCO2 leads to higher energetic costs and morphometric changes, with larger larvae in high pCO2 treatment and smaller larvae in medium pCO2 treatment. The efficient antioxidant response capacity and increase in energetic metabolism only registered at the medium pCO2 treatment may indicate that at higher pCO2 levels the capacity of larvae to restore their internal balance can be impaired. Our findings illustrate the need of using multiple approaches to explore the consequences of future pCO2 levels on organisms.

(Table 1) Age control points and of sediment core MD01-2444

The marine isotopic stage 3 (MIS3) at Ocean Drilling Program (ODP) Site 1060 (Gulf Stream) shows both sharp onset and end of interstadials, the existence of very short lived warm events during stadials, and points to differences in detail between the sea surface temperature (SST) record from the western North Atlantic and the atmospheric temperature record inferred from d18O in Greenland ice. Investigating MIS3 and obtaining comparable data from other locations appears crucial. The eastern Atlantic provides well-documented records of climate changes. We have selected a core from off Portugal and use it to examine Dansgaard/Oeschger events (D/O) at centennial-scale resolution (139 years on average between two data points). We have obtained a faunal data set for core MD01-2444, 37°N, 10°W, 2600 m water depth and use a group of species (Globigerina bulloides + Globigerinita glutinata) as a proxy of upwelling intensity driven by trade winds intensity changes. We tentatively relate the variation of this group to a North Atlantic Oscillation-like phenomenon (NAO) off Portugal. We observe that it resembles the rainfall index in the Caribbean as recorded at ODP Site 1002 (Cariaco Basin) which traces the Intertropical Convergence Zone (ITCZ) location through changes of terrigenous inputs. The driest intervals (ITZC to the south) at Site 1002 correspond to intervals of increased upwelling in MD01-2444 as well as the driest periods identified during stadials on similar cores in the area. Because the ITZC to the south is consistent with an El Niño-Southern Oscillation (ENSO+) situation, our study suggests a positive correlation between ENSO-like conditions and NAO-like conditions at a millennial timescale. During interstadial intervals when increased wetness over Cariaco is recorded (ITCZ to the north) and the upwelling in MD01-2444 is decreased, we see from both SSTs and faunal tropical indicators that MD01-2444 site is warm. In addition, interstadials are equally warm through each so-called Bond cycle. This contrasts with the Greenland Ice Core Project (GRIP) record where interstadial peaks are successively cooler through each Bond cycle. This record confirms a link between tropical climate linked to ITCZ position and the climate of southern Europe at millennial timescales, in spite of showing a very good correlation with polar latitudes (GRIP) through d18O on Globigerina bulloides. In addition, because the warmest SSTs and the d18O on G. bulloides are so remarkably different, our work points to changes in seasonality as a strong control over the climatic pattern of the North Atlantic area and the marked influence of winter conditions.

Relative abundance of pteropod species of the plankton pump samples in the surface water of the eastern North Atlantic Ocean (Table 4)

Plankton pump samples and plankton tows (size fractions between 0.04 mm and 1.01 mm) from the eastern North Atlantic Ocean contain the following shell- and skeleton-producing planktonic and nektonic organisms, which can be fossilized in the sediments: diatoms, radiolarians, foraminifers, pteropods, heteropods, larvae of benthic gastropods and bivalves, ostracods, and fish. The abundance of these components has been mapped quantitatively in the eastern North Atlantic surface waters in October - December 1971. More ash (after ignition of the organic matter, consisting mostly of these components) per cubic meter of water is found close to land masses (continents and islands) and above shallow submarine elevations than in the open ocean. Preferred biotops of planktonic diatoms in the region described are temperate shallow water and tropical coastal upwelling areas. Radiolarians rarely occur close to the continent, but are abundant in pelagic warm water masses, even near islands. Foraminifers are similar to the radiolarians, rarer in the coastal water mass of the continent than in the open ocean or off oceanic islands. Their abundance is highest outside the upwelling area off NW Africa. Molluscs generally outnumber planktonic foraminifers, implying that the carbonate cycle of the ocean might be influenced considerably by these animals. The molluscs include heteropods, pteropods, and larvae of benthic bivalves and gastropods. Larvae of benthic molluscs occur more frequently close to continental and island margins and above submarine shoals (in this case mostly guyots) than in the open ocean. Their size increases, but they decrease in number with increasing distance from their area of origin. Ostracods and fish have only been found in small numbers concentrated off NW Africa. All of the above-mentioned components occur in higher abundances in the surface water than in subsurface waters. They are closely related to the hydrography of the sampled water masses (here defined through temperature measurements). Relatively warm water masses of the southeastern branches of the Gulf Stream system transport subtropical and southern temperate species to the Bay of Biscay, relatively cool water masses of the Portugal and Canary Currents carry transitional faunal elements along the NW African coast southwards to tropical regions. These mix in the northwest African upwelling area with tropical faunal elements which are generally assumed to live in the subsurface water masses and which probably have been transported northwards to this area by a subsurface counter current. The faunas typical for tropical surface water masses are not only reduced due to the tongue of cool water extending southwards along the coast, but they are also removed from the coastal zone by the upwelling subsurface water masses carrying their own shell and skeleton assemblages. Tropical water masses contain much more shelland skeleton-producing plankters than subtropical and temperate ones. The climatic conditions found at different latitudes control the development and intensity of a separate continental coastal water mass with its own plankton assemblages. Extent of this water mass and steepness of gradients between the pelagic and coastal environment limit the occurrence of pelagic plankton close to the continental coast. A similar water mass in only weakly developed off oceanic islands.

A comprehensive analysis of tidal notches in the Mediterranean Sea

Recent works (Evelpidou et al., 2012) suggest that the modern tidal notch is disappearing worldwide due sea level rise over the last century. In order to assess this hypothesis, we measured modern tidal notches in several of sites along the Mediterranean coasts. We report observations on tidal notches cut along carbonate coasts from 73 sites from Italy, France, Croatia, Montenegro, Greece, Malta and Spain, plus additional observations carried outside the Mediterranean. At each site, we measured notch width and depth, and we described the characteristics of the biological rim at the base of the notch. We correlated these parameters with wave energy, tide gauge datasets and rock lithology. Our results suggest that, considering 'the development of tidal notches the consequence of midlittoral bioerosion' (as done in Evelpidou et al., 2012) is a simplification that can lead to misleading results, such as stating that notches are disappearing. Important roles in notch formation can be also played by wave action, rate of karst dissolution, salt weathering and wetting and drying cycles. Of course notch formation can be augmented and favoured also by bioerosion which can, in particular cases, be the main process of notch formation and development. Our dataset shows that notches are carved by an ensemble rather than by a single process, both today and in the past, and that it is difficult, if not impossible, to disentangle them and establish which one is prevailing. We therefore show that tidal notches are still forming, challenging the hypothesis that sea level rise has drowned them.

(Appendix A) Relative abundance of Emiliania huxleyi in sediment cores of the North Atlantic

The coccolithophore species Emiliania huxleyi is characterized by a wide range of sizes, which can be easily distinguished in the light microscope. In this study we have quantified the abundance of large (coccoliths > 4 µm in maximum length) E. huxleyi specimens during the last 25 kyr in sedimentary records from eleven cores and drill sites in the NE Atlantic and W Mediterranean Sea, to prove its usefulness in the reconstruction of water mass dynamics and biostratigraphic potential. During the Last Glacial Maximum this large form, a cold-water indicator, was common in the NE Atlantic and Mediterranean, and its regional variation in abundance indicates a displacement of the climatic zones southwards in agreement with the development of ice sheets and sea ice in the Northern Hemisphere during this period. On the other hand, the gradient between northern and southern surface water masses in the Subtropical Gyre appears to have been more pronounced than at present, while the Portugal and Canary Currents were more intense. In the western Mediterranean basin temperatures were cooler than in the adjacent Atlantic, provoking a quasi-endemism of these specimens until the end of Heinrich Event 1. This may have been due to a restriction in the communication between the Atlantic and Mediterranean through the Strait of Gibraltar, the arrival of cold surface water and the amplification of cooling after the development of ice sheets in the Northern Hemisphere. During the deglaciation, large E. huxleyi specimens decreased in abundance at medium and low latitudes, but were still numerous close to the Subarctic region during the Holocene. In transitional waters this decrease to present day abundances occurred after Termination Ib. The abrupt change in abundance of this large E. huxleyi form is proposed as a new biostratigraphic event to characterize the Holocene in mid- to low-latitude water masses in the North Atlantic, although this horizon seems to be diachronous by 5 kyr from tropical to subarctic regions, in agreement with the gradual onset of warm conditions.

Distribution of the sand-ascidian Seriocarpa rhizoides in surface sediments of Josephine-Bank, North Atlantic (Table 4)

Seriocarpa rhizoides Diehl 1969 was collected in abundance from the calcareous sand of the Josephine Bank (between Portugal and Madeira) during the "Meteor" seamount cruises in 1967. Attachment in this loose soft substratum is effected by fine anchoring strands of the tests. Two irregular series of small polycarp-like hermaphrodite bodies which are embedded in a connective tissue lie directly below the endostyle, forming a tubular compound gonad, but without common ducts. The intermediate nature of the reproductive system with respect to arrangement and structure increases our knowledge about the polygenetic relations of the stylid-genera. Some of the hitherto known ecological facts point to the presumed "seamounts effect" on this species.

Theoretical model simulations for variations in thermal reaction norms

Thermal reaction norms for growth rates of six Emiliania huxleyi isolates originating from the central Atlantic (Azores, Portugal) and five isolates from the coastal North Atlantic (Bergen, Norway) were assessed. We used the template mode of variation model to decompose variations in growth rates into modes of biological interest: vertical shift, horizontal shift, and generalist-specialist variation. In line with the actual habitat conditions, isolates from Bergen (Bergen population) grew well at lower temperatures, and isolates from the Azores (Azores population) performed better at higher temperatures. The optimum growth temperature of the Azores population was significantly higher than that of the Bergen population. Neutral genetic differentiation was found between populations by microsatellite analysis. These findings indicate that E. huxleyi populations are adapted to local temperature regimes. Next to between-population variation, we also found variation within populations. Genotype-by-environment interactions resulted in the most pronounced phenotypic differences when isolates were exposed to temperatures outside the range they naturally encounter. Variation in thermal reaction norms between and within populations emphasizes the importance of using more than one isolate when studying the consequences of global change on marine phytoplankton. Phenotypic plasticity and standing genetic variation will be important in determining the potential of natural E. huxleyi populations to cope with global climate change.