Combined effects of coral reef foraminifera Marginopora vertebralis and Heterostegina depressa in a multi-fractorial experiment

Warming and changes in ocean carbonate chemistry alter marine coastal ecosystems at an accelerating pace. The interaction between these stressors has been the subject of recent studies on reef organisms such as corals, bryozoa, molluscs, and crustose coralline algae. Here we investigated the combined effects of elevated sea surface temperatures and pCO2 on two species of photosymbiont-bearing coral reef Foraminifera: Heterostegina depressa (hosting diatoms) and Marginopora vertebralis (hosting dinoflagellates). The effects of single and combined stressors were studied by monitoring survivorship, growth, and physiological parameters, such as respiration, photochemistry (pulse amplitude modulation fluorometry and oxygen production), and chl a content. Specimens were exposed in flow-through aquaria for up to seven weeks to combinations of two pCO2 (~790 and ~490 µatm) and two temperature (28 and 31 °C) regimes. Elevated temperature had negative effects on the physiology of both species. Elevated pCO2 had negative effects on growth and apparent photosynthetic rate in H.depressa but a positive effect on effective quantum yield. With increasing pCO2, chl a content decreased in H. depressa and increased in M. vertebralis. The strongest stress responses were observed when the two stressors acted in combination. An interaction term was statistically significant in half of the measured parameters. Further exploration revealed that 75 % of these cases showed a synergistic (= larger than additive) interaction between the two stressors. These results indicate that negative physiological effects on photosymbiont-bearing coral reef Foraminifera are likely to be stronger under simultaneous acidification and temperature rise than what would be expected from the effect of each of the stressors individually.

Late pleistocene and holocene sedimentation in the central and eastern Persian Gulf

The sandfraction of the sediment was analysed in five cores, taken from 65 m water depth in the central and eastern part of the Persian Gulf. The holocene marls are underlayn by aragonite muds, which are probably 10-11,000 years old. 1. The cores could be subdivided into coarse grained and fine grained layers. Sorting is demonstrated by the following criteria: With increasing median values of the sandfraction - the fine grained fraction decreases within each core; - the median of each biogenic component, benthonic as well as planktonic, increases; - the median of the relict sediment, which in core 1179 was carried upward into the marl by bioturbation, increases; - the percentages of pelecypods, gastropods, decapods and serpulid worms in the sandfraction increase, the percentages of foraminifera and ostracods decrease; - the ratios of pelecypods to foraminifera and of decapods to ostracods increase; - the ratios of benthonic molluscs to planktonic molluscs (pteropods) and of benthonic foraminifera to planktonic foraminifera increase (except in core 1056 and 1179); - the ratio of planktonic molluscs (pteropods) to planktonic foraminifera increases; - the globigerinas without orbulinas increase, the orbulinas decrease in core 1056. Different settling velocities of these biogenic particles help in better understanding the results : the settling velocities, hence the equivalent hydrodynamic diameters, of orbulinas are smaller than those of other globigerinas, those of planktonic foraminifera are smaller than those of planktonic molluscs, those of planktonic molluscs are smaller than those of benthonic molluscs, those of pelecypods are smaller than those of gastropods. Bioturbation could not entirely distroy this "grain-size-stratification". Sorting has been stronger in the coarse layers than in the finer ones. As a cause variations in the supply of terrigenous material at constant strength of tidal currents is suggested. When much terrigenous material is supplied (large contents of fine grained fraction) the sedimentation rates are high: the respective sediment surface is soon covered and removed from the influence of tidal currents. When, however, the supply of terrigenous material is small, more sandy material is taken away in all locations within the influence of terrigenous supply. Thus the biogenic particles in the sediment do not only reflect the organic production, but also the influence of currents. 2. There is no parameter present in all cores that is independently variable from grain size and can be used for stratigraphic correlation. The two cores from the Strait of Hormus were correlated by their sequences of coarse and fine grained layers. 3. The sedimentation rates of terrigenous material, of total planktonic and benthonic organisms and of molluscs, foraminifera, echinoids and ophiuroids are shown in table 1 (total sediment 6.3-75.5 cm/1000 yr, biogenic carbonate 1.9-3.6 cm/1000 yr). The sedimentation rates of benthonic organisms are nearly the same in the cores of the Strait of Hormus, whereas near the Central Swell they are smaller. In the upper parts of the two cores of the Strait of Hormus sedimentation rates are higher than in the deeper parts, where higher median values point to stronger reworking. 4. The sequence of coarse and fine grained intervals in the two cores of the Hormus Strait, attributed to variations in climate, as well as the increase of terrigenous supply from the deeper to the upper parts of the cores, agrees with the descriptions in the literature of the post Pleistocene climate as becoming more humid. The rise of sea level is sedimentologically not measurable in the marly sediments - except perhaps for the higher content of echinoids in the lower part of core 1056. These may be attributed to the influence of a migrating wave-base. 5. The late Pleistocene aragonite mud is very fine grained (> 50%< 2 p) and poor in fossils (0.5-1.8%) biogenic particles of total sediment. The sand fraction consists almost entirely of white clumps, c. 0.1 mm in diameter (1177), composed of aragonite needles and of detrital minerals with the same size (1201). The argonite mud was probably not formed in situ, because the water depth at time of formation was at most 35 m at least 12 m. The sorting of the sediment (predominance of the fine grained sand), the absence of larger biogenic components and of pellets, c. 0.2-0.5 mm in diameter, which are typical for Recent and Pleistocene locations of aragonite formation, as well as the sedimentological conditions near the sampling points, indicate rather a transport of aragonite mud from an area of formation in very shallow waters. Sorting as well as lenticular fabric in core 1201 point to sedimentation within the influence of currents. During alternating sedimentation - and reworking processes the aragonitic matrix was separated from the silt - and sand-sized minerals. The lenses grade into touches because of bioturbation. 6. In core 1056 D2 from Hormus Bay the percentages of organic carbon, total nitrogen and total carbonate were determined. With increasing amounts of smaller grain sizes the content of organic matter increases, whereas the amount of carbonate decreases. The amounts of organic carbon and of nitrogen decrease with increasing depth, probably due to early-diagenetic decomposition processes. Most of the total nitrogen is of organic origin, only about 10% may well be inorganically fixed as ammonium-nitrogen. In the upper part of the core the C/N-ratio increases with increasing depth. This may be connected with a stronger decomposition of nitrogen-containing organic compounds. The general decrease of the C/N-ratios in the lower part of the core may be explained by the relative increase of inorganically fixed ammonium-nitrogen with decreasing content of organic matter.

Nighttime dissolution in a temperate coastal ocean ecosystem increases under acidification

Anthropogenic emissions of carbon dioxide (CO2) are causing ocean acidification, lowering seawater aragonite (CaCO3) saturation state (Omega arag), with potentially substantial impacts on marine ecosystems over the 21st Century. Calcifying organisms have exhibited reduced calcification under lower saturation state conditions in aquaria. However, the in situ sensitivity of calcifying ecosystems to future ocean acidification remains unknown. Here we assess the community level sensitivity of calcification to local CO2-induced acidification caused by natural respiration in an unperturbed, biodiverse, temperate intertidal ecosystem. We find that on hourly timescales nighttime community calcification is strongly influenced by Omega arag, with greater net calcium carbonate dissolution under more acidic conditions. Daytime calcification however, is not detectably affected by Omega arag. If the short-term sensitivity of community calcification to Omega arag is representative of the long-term sensitivity to ocean acidification, nighttime dissolution in these intertidal ecosystems could more than double by 2050, with significant ecological and economic consequences.

Fertilisation and larval development in an Antarctic bivalve, Laternula elliptica, under reduced pH and elevated temperatures

Elevated temperatures associated with ocean warming and acidification can influence development and, ultimately, success of larval molluscs. The effect of projected oceanic changes on fertilisation and larval development in an Antarctic bivalve, Laternula elliptica, was investigated through successive larval stages at ambient temperature and pH conditions (-1.6°C and pH 7.98) and conditions representative of projections through to 2100 (-0.5°C to +0.4°C and pH 7.80 to pH 7.65). Where significant effects were detected, increased temperature had a consistently positive influence on larval development, regardless of pH level, while effects of reduced pH varied with larval stage and incubation temperature. Fertilisation was high and largely independent of stressors, with no loss of gamete viability. Mortality was unaffected at all development stages under experimental conditions. Elevated temperatures reduced occurrences of abnormalities in D-larvae and accelerated larval development through late veliger and D-larval stages, with D-larvae occurring 5 d sooner at 0.4°C compared to ambient temperature. Reduced pH did not affect occurrences of abnormalities in larvae, but it slowed the development of calcifying stages. More work is required to investigate the effects of developmental delays of the magnitude seen here in order to better determine the ecological relevance of these changes on longer term larval and juvenile success.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Relative contents of foraminifera species in bottom sediments of northern and southern seas

Correlation of paleoceanographic events in several key regions of the World Ocean: North Atlantic, Antarctic, West Arctic Seas, North Pacific and tropical Indo-Pacific has been carried out for the last 135 ka based on micropaleontological, stable isotope, geochronological (AMS-14C) and other data. It has been shown that the global thermohaline circulation controls remote climatic teleconnections on millennial-scale and partly on centennial-scale, while short-term climate changes are mainly transferred by the atmosphere. The basic information is given about the recent thermohaline circulation and stages of its development during Neogene.

Seawater carbonate chemistry and biological processes during experiments with postlarvae of barnacle of Semibalanus balanoides and Elminius modestus, 2010

Ocean acidification and global warming are occurring concomitantly, yet few studies have investigated how organisms will respond to increases in both temperature and CO2. Intertidal microcosms were used to examine growth, shell mineralogy and survival of two intertidal barnacle post-larvae, Semibalanus balanoides and Elminius modestus, at two temperatures (14 and 19°C) and two CO2 concentrations (380 and 1,000 ppm), fed with a mixed diatom-flagellate diet at 15,000 cells ml-1 with flow rate of 10 ml-1 min-1. Control growth rates, using operculum diameter, were 14 ± 8 µm day-1 and 6 ± 2 µm day-1 for S. balanoides and E. modestus, respectively. Subtle, but significant decreases in E. modestus growth rate were observed in high CO2 but there were no impacts on shell calcium content and survival by either elevated temperature or CO2. S. balanoides exhibited no clear alterations in growth rate but did show a large reduction in shell calcium content and survival under elevated temperature and CO2. These results suggest that a decrease by 0.4 pH(NBS) units alone would not be sufficient to directly impact the survival of barnacles during the first month post-settlement. However, in conjunction with a 4-5°C increase in temperature, it appears that significant changes to the biology of these organisms will ensue.

Seawater carbonate chemistry and energy status in the periwinkle Littorina littorea during experiments, 2011

In the future, marine organisms will face the challenge of coping with multiple environmental changes associated with increased levels of atmospheric Pco2, such as ocean warming and acidification. To predict how organisms may or may not meet these challenges, an in-depth understanding of the physiological and biochemical mechanisms underpinning organismal responses to climate change is needed. Here, we investigate the effects of elevated Pco2 and temperature on the whole-organism and cellular physiology of the periwinkle Littorina littorea. Metabolic rates (measured as respiration rates), adenylate energy nucleotide concentrations and indexes, and end-product metabolite concentrations were measured. Compared with values for control conditions, snails decreased their respiration rate by 31% in response to elevated Pco2 and by 15% in response to a combination of increased Pco2 and temperature. Decreased respiration rates were associated with metabolic reduction and an increase in end-product metabolites in acidified treatments, indicating an increased reliance on anaerobic metabolism. There was also an interactive effect of elevated Pco2 and temperature on total adenylate nucleotides, which was apparently compensated for by the maintenance of adenylate energy charge via AMP deaminase activity. Our findings suggest that marine intertidal organisms are likely to exhibit complex physiological responses to future environmental drivers, with likely negative effects on growth, population dynamics, and, ultimately, ecosystem processes.

Critical bed shear stress and threshold of motion of maërl biogenic gravel

Critical bed shear stress for incipient motion has been determined for biogenic free-living coralline algae known as maërl. Maërl from three different sedimentary environments (beach, intertidal, and open marine) in Galway Bay, west of Ireland have been analysed in a rotating annular flume and linear flume. Velocity profile measurements of the benthic boundary layer, using an Acoustic Doppler Velocimeter, have been obtained in four different velocity experiments. The bed shear stress has been determined using three methods: Law of the Wall, Turbulent Kinetic Energy and Reynolds Stress. The critical Shields parameter has been estimated as a non-dimensional mobility number and the results have been compared with the Shields curve for natural sand. Maërl particles fall below this curve because its greater angularity allows grains to be mobilised easier than hydraulically equivalent particles. From previous work, the relationship between grain shape and the settling velocity of maërl suggests that the roughness is greatest for intertidal maërl particles. During critical shear stress determinations, beds of such rough particles exhibited the greatest critical shear stress probably because the particle thalli interlocked and resisted entrainment. The Turbulent Kinetic Energy methodology gives the most consistent results, agreeing with previous comparative studies. Rarely-documented maërl megaripples were observed in the rotating annular flume and are hypothesised to form at velocities ~10 cm s-1 higher than the critical threshold velocity, where tidal currents, oscillatory flow or combined-wave current interaction results in the preferential transport of maërl. A determination of the critical bed shear stress of maërl allows its mobility and rate of erosion and deposition to be evaluated spatially in subsequent applications to biological conservation management.

Pollen and spore counts of sediment core GeoB12624

In tropical eastern Africa, vegetation distribution is largely controlled by regional hydrology, which has varied over the past 20 000 years. Therefore, accurate reconstructions of past vegetation and hydrological changes are crucial for a better understanding of climate variability in the tropical southeastern African region. We present high-resolution pollen records from a marine sediment core recovered offshore of the Rufiji River delta. Our data document significant shifts in pollen assemblages during the last deglaciation, identifying, through changes in both upland and lowland vegetation, specific responses of plant communities to atmospheric (precipitation) and coastal (coastal dynamics and sea-level changes) alterations. Specifically, arid conditions reflected by a maximum pollen representation of dry and open vegetation occurred during the Northern Hemisphere cold Heinrich event 1 (H1), suggesting that the expansion of drier upland vegetation was synchronous with cold Northern Hemisphere conditions. This arid period is followed by an interval in which forest and humid woodlands expanded, indicating a hydrologic shift towards more humid conditions. Droughts during H1 and the shift to humid conditions around 14.8 kyr BP in the uplands are consistent with latitudinal shifts of the intertropical convergence zone (ITCZ) driven by high-latitude Northern Hemisphere climatic fluctuations. Additionally, our results show that the lowland vegetation, consisting of well-developed salt marshes and mangroves in a successional pattern typical for vegetation occurring in intertidal habitats, has responded mainly to local coastal dynamics related to marine inundation frequencies and soil salinity in the Rufiji Delta as well as to the local moisture availability. Lowland vegetation shows a substantial expansion of mangrove trees after ~ 14.8 kyr BP, suggesting an increased moisture availability and river runoff in the coastal area. The results of this study highlight the decoupled climatic and environmental processes to which the vegetation in the uplands and the Rufiji Delta has responded during the last deglaciation.

Seawater carbonate chemistry, physiology and skeletal mineralogy of coralline alga Corallina elongata in a laboratory experiment

Marine organisms inhabiting environments where pCO2/pH varies naturally are suggested to be relatively resilient to future ocean acidification. To test this hypothesis, the effect of elevated pCO2 was investigated in the articulated coralline red alga Corallina elongata from an intertidal rock pool on the north coast of Brittany (France), where pCO2 naturally varied daily between 70 and 1000 µatm. Metabolism was measured on algae in the laboratory after they had been grown for 3 weeks at pCO2 concentrations of 380, 550, 750 and 1000 µatm. Net and gross primary production, respiration and calcification rates were assessed by measurements of oxygen and total alkalinity fluxes using incubation chambers in the light and dark. Calcite mol % Mg/Ca (mMg/Ca) was analysed in the tips, branches and basal parts of the fronds, as well as in new skeletal structures produced by the algae in the different pCO2 treatments. Respiration, gross primary production and calcification in light and dark were not significantly affected by increased pCO2. Algae grown under elevated pCO2 (550, 750 and 1000 µatm) formed fewer new structures and produced calcite with a lower mMg/Ca ratio relative to those grown under 380 µatm. This study supports the assumption that C. elongata from a tidal pool, where pCO2 fluctuates over diel and seasonal cycles, is relatively robust to elevated pCO2 compared to other recently investigated coralline algae.