Seawater carbonate chemistry and benthic marine community during experiments, 2011

Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Eocene benthic foraminiferal community DSDP of Holes 95-612, 95-613 and 11-108

Benthic foraminiferal biofacies may vary independently of water depth and water mass; however, calibration of biofacies and stratigraphic ranges with independent paleodepth estimates allows reconstruction of age-depth patterns applicable throughout the deep Atlantic (Tjalsma and Lohmann, 1983). We have attempted to test these faunal calibrations in a continental margin setting, reconstructing Eocene benthic foraminiferal distributions along a dip section afforded by the New Jersey Transect (DSDP Sites 612, 108, 613). The following independent estimates of Eocene depths for the transect were obtained by "backtracking," "backstripping," and by assuming increasing depth downdip ("paleoslope"): Site 612, near the middle/lower bathyal boundary (about 1000 m); Site 108, in the middle bathyal zone (about 1600 m); and Site 613, near the lower bathyal/upper abyssal boundary (about 2000 m). Within uncertainties of backtracking (hundreds of meters), these estimates agree with estimates of paleodepth based on comparison of the New Jersey margin biofacies with other backtracked faunas. The stratigraphic ranges of many benthic taxa correspond to those found at other Atlantic DSDP sites. The major biofacies patterns show: (1) a depth dichotomy between an early to middle Eocene Nuttallides truempyidominated biofacies (greater than 2000 m) and a Lenticulina-Osangularia-Alabamina cf. dissonata biofacies (1000- 2000 m); and (2) a difference between a middle and a late Eocene biofacies at Site 612. The faunal boundary at about 2000 m, between bathyal and abyssal zones, occurs not only on the margin, but also throughout the deep Atlantic. The faunal change between the middle and late Eocene at Site 612 was due to a decrease of Lenticulina spp., the local disappearance of N. truempyi, and establishment of a Bulimina alazanensis-Gyroidinoides spp. biofacies. Although this change could be attributed to local paleoceanographic or water-depth changes, we argue that it is the bathyal expression of a global deep-sea benthic foraminiferal change which occurred across the middle/late Eocene boundary.

Neogene benthic foraminiferal community and organic carbon at DSDP Holes 95-612 and 95-613

Quantitative study of benthic foraminifers from the upper Miocene to lower Pliocene section at Site 612 (1404 m present water depth) and the Pliocene section at Site 613 (2323 m present water depth) shows no evidence of widespread downslope transport of shallow-water biofacies or reworking of older material in the greater than 150 µm size fraction. In contrast, upper Miocene sediments from Site 604 (2364 m present water depth) show extensive reworking and downslope transport. At Site 612, benthic foraminifers show a succession from an upper Miocene Bolivina alata-Nonionella sp. biofacies, to an uppermost Miocene Bulimina alazanensis biofacies, to a lower Pliocene Cassidulina reflexa biofacies, to an upper Pliocene Melonis barleeanum-Islandiella laevigata biofacies. Evidence suggests that the Pliocene biofacies are in situ, although they could have been transported downslope from the upper-middle bathyal zone. At Site 613, Uvigerina peregrina dominated the "middle" Pliocene, while Globocassidulina subglobosa was dominant in the early and late Pliocene. High abundances of U. peregrina at Site 613 are associated with high values of sedimentary organic carbon.

The benthic foraminiferal community in a naturally CO2-rich coastal habitat of the southwestern Baltic Sea

It is expected that the calcification of foraminifera will be negatively affected by the ongoing acidification of the oceans. Compared to the open oceans, these organisms are subjected to much more adverse carbonate system conditions in coastal and estuarine environments such as the southwestern Baltic Sea, where benthic foraminifera are abundant. This study documents the seasonal changes of carbonate chemistry and the ensuing response of the foraminiferal community with bi-monthly resolution in Flensburg Fjord. In comparison to the surface pCO2, which is close to equilibrium with the atmosphere, we observed large seasonal fluctuations of pCO2 in the bottom and sediment pore waters. The sediment pore water pCO2 was constantly high during the entire year ranging from 1244 to 3324 µatm. Nevertheless, in contrast to the bottom water, sediment pore water was slightly supersaturated with respect to calcite as a consequence of higher alkalinity (AT) for most of the year. Foraminiferal assemblages were dominated by two calcareous species, Ammonia aomoriensis and Elphidium incertum, and the agglutinated Ammotium cassis. The one-year cycle was characterised by seasonal community shifts. Our results revealed that there is no dynamic response of foraminiferal population density and diversity to elevated sediment pore water pCO2. Surprisingly, the fluctuations of sediment pore water undersaturation (Omega calc) co-vary with the population densities of living Ammonia aomoriensis. Further, we observed that most of the tests of living calcifying foraminifera were intact. Only Ammonia aomorienis showed dissolution and recalcification structures on the tests, especially at undersaturated conditions. Therefore, the benthic community is subjected to high pCO2 and tolerates elevated levels as long as sediment pore water remains supersaturated. Model calculations inferred that increasing atmospheric CO2 concentrations will finally lead to a perennial undersaturation in sediment pore waters. Whereas benthic foraminifera indeed may cope with a high sediment pore water pCO2, the steady undersaturation of sediment pore waters would likely cause a significant higher mortality of the dominating Ammonia aomoriensis. This shift may eventually lead to changes in the benthic foraminiferal communities in Flensburg Fjord, as well as in other regions experiencing naturally undersaturated Omega calc levels.

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.

Structural and functional vulnerability to elevated pCO2 in marine benthic communities

The effect of elevated pCO2/low pH on marine invertebrate benthic biodiversity, community structure and selected functional responses which underpin ecosystem services (such as community production and calcification) was tested in a medium-term (30 days) mesocosm experiment in June 2010. Standardised intertidal macrobenthic communities, collected (50.3567°N, 4.1277°W) using artificial substrate units (ASUs), were exposed to one of seven pH treatments (8.05, 7.8. 7.6, 7.4, 7.2, 6.8 and 6.0). Community net calcification/dissolution rates, as well as changes in biomass, community structure and diversity, were measured at the end of the experimental period. Communities showed significant changes in structure and reduced diversity in response to reduced pH: shifting from a community dominated by calcareous organisms to one dominated by non-calcareous organisms around either pH 7.2 (number of individuals and species) or pH 7.8 (biomass). These results were supported by a reduced total weight of CaCO3 structures in all major taxa at lowered pH and a switch from net calcification to net dissolution around pH 7.4 (Omega calc = 0.78, Omega ara = 0.5). Overall community soft tissue biomass did not change with pH and high mortality was observed only at pH 6.0, although molluscs and arthropods showed significant decreases in soft tissue. This study supports and refines previous findings on how elevated pCO2 can induce changes in marine biodiversity, underlined by differential vulnerability of different phyla. In addition, it shows significant elevated pCO2-/low pH-dependent changes in fundamental community functional responses underpinning changes in ecosystem services.

Benthic foraminifera abundance data of sediment core GeoB7926-2

We present a high resolution, multiproxy study of the relationship between pelagic and benthic environments of a coastal upwelling system in the subtropical NE Atlantic Ocean. Marine sediments corresponding to late MIS3 to the Holocene in the radiocarbon dated core GeoB7926, retrieved off Mauritania (21°N) were analysed to reconstruct productivity in surface waters and its linkage to deep waters during the last 35 ka BP. High latitude cold events and changes in atmospheric and oceanographic dynamics influenced upwelling intensity over this time period. Subsequently, this caused changes in primary productivity off this low-latitude coastal upwelling locality. The benthic foraminiferal fauna displays four main community shifts corresponding to fundamental climatic events, first of all during late MIS3 (35-28 ka BP), secondly from 28 to 19 ka BP (including Heinrich event 2 and the LGM), thirdly within Heinrich event 1, the Bølling Allerød and the Younger Dryas (18-11.5 ka BP) and finally during the Holocene (11.5-0 ka BP). In particular, strong pelagic-benthic coupling is apparent in MIS 3, as demonstrated by increased primary productivity, indicated by moderate DAR and the dominance of benthic foraminiferal species which prefer fresh phytodetritus. A decline in upwelling intensity and nutrient availability follows, which resulted in a proportionately larger amount of older, degraded matter, provoking a shift in the benthic foraminifera fauna composition. This rapid response of the benthic environment continues with a progressive increase in upwelling intensity due to sea level and oceanographic changes and according high surface production during the LGM. During Heinrich event 1 and the Younger Dryas, extreme levels of primary production actually hindered benthic environment through the development of low oxygen conditions. After this period, a final change in benthic foraminiferal community composition occurs which indicates a return to more oxygenated conditions during the Holocene.