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 By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived.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.

Magnetic properties, grain sizes, XRF Scanner Data of sediment core GeoB9307-3

Geochemical and mineralogical proxies for paleoenvironmental conditions have the underlying assumption that climate variations have an impact on terrestrial weathering conditions. Varying properties of terrigenous sediments deposited at sea are therefore often interpreted in terms of paleoenvironmental change. Also in gravity core GeoB9307-3 (18° 33.99' S, 37° 22.89' E), located off the Zambezi River, environmental changes during Heinrich Stadial 1 (HS 1) and the Younger Dryas (YD) are accompanied by changing properties of the terrigenous sediment fraction. Our study focuses on the relationship of variability in the hydrological system and changes in the magnetic properties, major element geochemistry and granulometry of the sediments. We propose that changes in bulk sedimentary properties concur with environmental change, although not as a direct response of climate driven pedogenic processes. Spatial varying rainfall intensities on a sub-basin scale modify sediment export from different parts of the Zambezi River basin. During humid phases, such as HS 1 and the YD, sediment was mainly exported from the coastal areas, while during more arid phases sediments mirror the hinterland soil and lithological properties and are likely derived from the northern Shire sub-basin. We propose that a de-coupling of sedimentological and organic signals with variable discharge and erosional activity can occur.

Lacustrine sediments from two setions in northern Greece

We investigated two lignite quarries in northern Greece for orbital and suborbital climate variability. Sections Lava and Vegora are located at the southern and northern boundaries of the Ptolemais Basin, a northwest southeast elongated intramontane basin that contains Upper Miocene to Lower Pliocene lacustrine sediments. Sediments show cyclic alterations of marl-rich (light), and coal-rich or clay-rich (dark) strata on a decimeter to meter scale. First, we established low-resolution ground-truth stratigraphy based on paleomagnetics and biostratigraphy. Accordingly, the lower 67 m and 65 m that were investigated in both sections Vegora and Lava, respectively, belong to the Upper Miocene and cover a time period of 6.85 to 6.57 and 6.46 to 5.98 Ma at sedimentation rates of roughly 14 and 22 cm/ka. In order to obtain a robust and high-resolution chronology, we then tuned carbonate minima (low L* values; high magnetic susceptibility values) to insolation minima. Besides the known dominance of orbital precession and eccentricity, we detected a robust hemi-precessional cycle in most parameters, most likely indicative for monsoonal influence on climate. Moreover, the insolation-forced time series indicate a number of millennial-scale frequencies that are statistically significant with dominant periods of 1.5-8 kyr. Evolutionary spectral analysis indicates that millennial-scale climate variability documented for the Ptolemais Basin resembles the one that is preserved in ice-core records of Greenland. Most cycles show durations of several tens of thousands of years before they diminish or cease. This is surprising because the generally argued cause for Late Quaternary millennial-scale variability is associated with the presence of large ice sheets, which cannot be the case for the Upper Miocene. Possible explanations maybe a direct response to solar forcing, an influence on the formation of North Atlantic Deep Water through the outflow of high-salinity water, or an atmospheric link to the North Atlantic Oscillation.

Diatom studies in surface and downcore sediments off southern Sumatra and Java

A wealth of sedimentary records aimed at reconstructing late Quaternary changes in productivity and temperature have been devoted to understanding linkages between the Indo-Pacific Warm Pool (IPWP) and other distant oceanic areas. Most of these reconstructions are based, however, on biogeochemical and sedimentological proxies, with comparatively less attention devoted to microfossils. A high-resolution (<1 ka) study of diatom concentrations and the community at site GeoB10038-4, recovered off southern Sumatra (ca. 6°S, 103°E), closely tracks the variations of diatom concentrations in the westernmost IPWP during the last glacial-interglacial cycle. The diatom record provides evidence that diatom paleoproductivity was highest during interglacials, primarily due to the input of lithogenics and nutrients following the rise in sea level after full glacials. In addition, the co-variation of total diatom concentration and Northern Hemisphere forcing for Marine Isotope Stage 5 suggests a direct response of diatom productivity and upwelling intensity to boreal summer insolation. Temporal shifts of the diverse diatom community at site GeoB10038-4 correspond well with the present-day seasonal monsoon pattern and the strengthening and weakening phases of upwelling along the southern coast of Sumatra. Resting spores of Chaetoceros, typical of nutrient-rich waters, were dominant during periods of highest diatom paleoproductivity and responded to the strengthening of the SE monsoon, while diatoms of oligotrophic to mesotrophic waters characterized intermonsoon periods. The close correspondence between the dominance of upwelling diatoms and the boreal summer insolation resembles the present-day dynamics of diatom production. The observed interglacial highs and glacial lows of diatom productivity at site GeoB10038-4 is a unique pattern in the late Quaternary tropics.

Nannofossil assemblages of mid-Pliocene sediments from the equatorial Atlantic

Downcore cyclic variation in high-resolution nannofossil abundance records from mid-Pliocene equatorial Atlantic ODP Sites 662 and 926 demonstrate the direct response by several Pliocene taxa (notably Discoaster, Sphenolithus and Florisphaera profunda) to orbitally forced climatic variation. In particular, these records display strong obliquity and precessional signals reflecting primarily high latitude, Southern hemisphere changes influencing upwelling intensity and local low-latitude, insolation-driven climatic changes (via the productivity and/or turbidity influence of Amazon-sourced terrigenous material) at Sites 622 and 926 respectively. In seasonal studies of coccolithophorid assemblages, only part of the variation observed can be explained by abiotic processes, so it is perhaps not surprising that in this study few Pliocene nannofossil taxa demonstrate significant correlations with each other or with physical environmental parameters. Only some variance in nannofossil abundances can be explained by the primary controls of temperature and productivity. The rest is attributed to nonlinear responses to climatic changes; biotic processes such as grazing, predation, viral infection and competition, and/or, abiotic factors for which there is no readily available proxy (e.g. salinity). The lack of strong, consistent intra- and inter-relationships of the nannoflora and the environment reflects an ecologically complex, differentiated original community producing a complex integrated signal transmitted into the fossil record.

Seawater carbonate chemistry, length and survival of Inland silverside, Menidia beryllina, during experiments, 2012

Absorption of anthropogenic carbon dioxide by the world's oceans is causing mankind's 'other CO2 problem', ocean acidification. Although this process will challenge marine organisms that synthesize calcareous exoskeletons or shells, it is unclear how it will affect internally calcifying organisms, such as marine fish. Adult fish tolerate short-term exposures to CO2 levels that exceed those predicted for the next 300 years (~2,000 ppm), but potential effects of increased CO2 on growth and survival during the early life stages of fish remain poorly understood. Here we show that the exposure of early life stages of a common estuarine fish (Menidia beryllina) to CO2 concentrations expected in the world's oceans later this century caused severely reduced survival and growth rates. When compared with present-day CO2 levels (~400 ppm), exposure of M. beryllina embryos to ~1,000 ppm until one week post-hatch reduced average survival and length by 74% and 18%, respectively. The egg stage was significantly more vulnerable to high CO2-induced mortality than the post-hatch larval stage. These findings challenge the belief that ocean acidification will not affect fish populations, because even small changes in early life survival can generate large fluctuations in adult-fish abundance.

(Table 2) Age interval and accumulation rates of organic and inorganic carbon from ODP Sites 175-1082, 175-1084, 175-1085 and 175-1087

The Ocean Drilling Program Leg 175 recovered a unique series of stratigraphically continuous sedimentary sections along the SW African margin, an area which is presently affected by active coastal upwelling. The accumulation rates of organic and inorganic carbon are a major component of this record. Four Leg 175 sites (1082, 1084, 1085, 1087) are chosen as part of a latitudinal transect from the present northern to southern boundaries of the Benguela Current upwelling system, to decipher the Pliocene-Pleistocene history of biogenic production and its relationship with global and local changes in oceanic circulation and climate. The pattern of CaCO3 and Corg mass accumulation rates (MARs) over 0.25-Myr intervals indicates that the evolution of carbon burial is highly variable between the northern and the southern Benguela regions, as well as between sites that have similar hydrological conditions. This, as well as the presence over most locations of high-amplitude, rapid changes of carbon burial, reflect the partitioning of biogenic production and patterns of sedimentation into local compartments over the Benguela margin. The combined mapping of CaCO3 and Corg MARs at the study locations suggests four distinct evolutionary periods, which are essentially linked with major steps in global climate change: the early Pliocene, the mid-Pliocene warm event, a late Pliocene intensification of northern hemisphere glaciation and the Pleistocene. The early Pliocene spatially heterogeneous patterns of carbon burial are thought to reflect the occurrence of mass-gravitational movements over the Benguela slope which resulted in disruption of the recorded biogenic production. This was followed (3.5-3 Ma) by an episode of peak carbonate accumulation over the whole margin and, subsequently, by the onset of Benguela provincialism into a northern and a southern sedimentary regime near 2 Ma. This mid and late Pliocene evolution is interpreted as a direct response to changes in the ventilation of bottom and intermediate waters, as well as to dynamics of the subtropical gyral circulation and associated wind stress.

Susceptibility measurements at Gubbio sections

The complex interplay between extraterrestrial events and earth-bound processes that triggered one of the greatest biological crises of the Phanerozoic requires a high resolution timescale. Detailed magnetic susceptibility measurements at the Contessa Highway and Bottaccione sections (Italy) span the Cretaceous-Paleogene boundary and reveal clear orbital signatures in the sedimentary record. Identification of precession and 405 kyr eccentricity cycles allows an estimate of 324+/-40 kyr for the duration of the Maastrichtian part of Chron C29r. We present in the same high resolution time frame sites in Spain and the North and South Atlantic and bio-horizons, biotic changes, stable isotopic excursions and the decrease in Osmium isotopes recorded in these sections. The onset of 187 Os/ 188 Os decrease coincides with the d13 C negative excursion K-PgE1, thus suggesting a first pulse in Deccan volcanism at 66.64 Ma. The K-PgE3 d13 C negative excursion is possibly the expression of a second pulse at 66.26 Ma. Late Maastrichtian d13 C negative excursions are of low intensity and span durations of one to two eccentricity cycles, whereas early Danian excursions are brief (about 30 kyr) and acute. In Biotic response to late Maastrichtian perturbations occurred with a delay of ca. 200 kyr after the beginning of K-PgE1 shortly before K-PgE3. The biotic perturbation could be thus either a delayed response to K-PgE1, or a direct response to K-PgE3, and possibly, a threshold response to the stepwise buildup of CO2 atmospheric injections. No delay is evident in response to early Danian hyperthermal events. These differences suggest that short-lived, volcanically-derived environmental perturbations were buffered within the stable late Maastrichtian oceanic realm whereas they were amplified by the more sensitive and highly disturbed early Danian oceanic ecosystem.

Holocene experiment with coupled atmosphere-ocean-model ECHAM5/MPI-OM

Changes in the Earth's orbit lead to changes in the seasonal and meridional distribution of insolation. We quantify the influence of orbitally induced changes on the seasonal temperature cycle in a transient simulation of the last 6000 years - from the mid-Holocene to today - using a coupled atmosphere-ocean general circulation model (ECHAM5/MPI-OM) including a land surface model (JSBACH). The seasonal temperature cycle responds directly to the insolation changes almost everywhere. In the Northern Hemisphere, its amplitude decreases according to an increase in winter insolation and a decrease in summer insolation. In the Southern Hemisphere, the opposite is true. Over the Arctic Ocean, decreasing summer insolation leads to an increase in sea-ice cover. The insulating effect of sea ice between the ocean and the atmosphere leads to decreasing heat flux and favors more "continental" conditions over the Arctic Ocean in winter, resulting in strongly decreasing temperatures. Consequently, there are two competing effects: the direct response to insolation changes and a sea-ice insulation effect. The sea-ice insulation effect is stronger, and thus an increase in the amplitude of the seasonal temperature cycle over the Arctic Ocean occurs. This increase is strongest over the Barents Shelf and influences the temperature response over northern Europe. We compare our modeled seasonal temperatures over Europe to paleo reconstructions. We find better agreements in winter temperatures than in summer temperatures and better agreements in northern Europe than in southern Europe, since the model does not reproduce the southern European Holocene summer cooling inferred from the paleo reconstructions. The temperature reconstructions for northern Europe support the notion of the influence of the sea-ice insulation effect on the evolution of the seasonal temperature cycle.

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO2 by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO2 ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.

Direct effects of increased CO2 concentrations in seawater on the net primary production of charophytes in a shallow, coastal, brackish-water ecosystem

Charophytes are found in fresh and brackish waters across the globe and play key roles in coastal ecosystems. However, their response to increasing CO2 is not well understood. The aim of the study was to detect the effects of elevated CO2 on the physiology of charophyte species growing in the brackish Baltic Sea by measuring net primary production. Mesocosm experiments were conducted in the Kõiguste Bay (N Gulf of Riga) during the field season of 2012. Separate mesocosms were maintained at different pCO2 levels: 2000, 1000 and 200 µatm. The experiments were carried out with three species of charophytes: Chara aspera, C. tomentosa and C. horrida. The short-term photosynthetic responses of charophytes to different treatments were measured by the oxygen method. The results show that elevated CO2 levels in brackish water may enhance the photosynthetic activity of charophyte species and suggest that increasing CO2 in the Baltic Sea could have implications for interspecific competition and community structure in a future high CO2 world.

Small Changes in pH Have Direct Effects on Marine Bacterial Community Composition: A Microcosm Approach

As the atmospheric CO2 concentration rises, more CO2 will dissolve in the oceans, leading to a reduction in pH. Effects of ocean acidification on bacterial communities have mainly been studied in biologically complex systems, in which indirect effects, mediated through food web interactions, come into play. These approaches come close to nature but suffer from low replication and neglect seasonality. To comprehensively investigate direct pH effects, we conducted highly-replicated laboratory acidification experiments with the natural bacterial community from Helgoland Roads (North Sea). Seasonal variability was accounted for by repeating the experiment four times (spring, summer, autumn, winter). Three dilution approaches were used to select for different ecological strategies, i.e. fast-growing or low-nutrient adapted bacteria. The pH levels investigated were in situ seawater pH (8.15-8.22), pH 7.82 and pH 7.67, representing the present-day situation and two acidification scenarios projected for the North Sea for the year 2100. In all seasons, both automated ribosomal intergenic spacer analysis and 16S ribosomal amplicon pyrosequencing revealed pH-dependent community shifts for two of the dilution approaches. Bacteria susceptible to changes in pH were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. Their specific response to reduced pH was often context-dependent. Bacterial abundance was not influenced by pH. Our findings suggest that already moderate changes in pH have the potential to cause compositional shifts, depending on the community assembly and environmental factors. By identifying pH-susceptible groups, this study provides insights for more directed, in-depth community analyses in large-scale and long-term experiments.

Response of A. aurita ephyrae to three stressors: high-temperature, high-CO2 and low-oxygen

Global change is affecting marine ecosystems through a combination of different stressors such as warming, ocean acidification and oxygen depletion. Very little is known about the interactions among these factors, especially with respect to gelatinous zooplankton. Therefore, in this study we investigated the direct effects of pH, temperature and oxygen availability on the moon jellyfish Aurelia aurita, concentrating on the ephyral life stage. Starved one-day-old ephyrae were exposed to a range of pCO2 (400-4000 ppm) and three different dissolved oxygen levels (from saturated to hypoxic conditions), in two different temperatures (5 and 15 °C) for 7 days. Carbon content and swimming activity were analysed at the end of the incubation period, and mortality noted. General linearized models were fitted through the data, with the best fitting models including two- and three-way interactions between pCO2, temperature and oxygen concentration. The combined effect of the stressors was small but significant, with the clearest negative effect on growth caused by the combination of all three stressors present (high temperature, high CO2, low oxygen). We conclude that A. aurita ephyrae are robust and that they are not likely to suffer from these environmental stressors in a near future.