Marine bivalve shell geochemistry and ultrastructure from modern low pH environments: environmental effect versus experimental bias (update)

Bivalve shells can provide excellent archives of past environmental change but have not been used to interpret ocean acidification events. We investigated carbon, oxygen and trace element records from different shell layers in the mussels Mytilus galloprovincialis combined with detailed investigations of the shell ultrastructure. Mussels from the harbour of Ischia (Mediterranean, Italy) were transplanted and grown in water with mean pHT 7.3 and mean pHT 8.1 near CO2 vents on the east coast of the island. Most prominently, the shells recorded the shock of transplantation, both in their shell ultrastructure, textural and geochemical record. Shell calcite, precipitated subsequently under acidified seawater responded to the pH gradient by an in part disturbed ultrastructure. Geochemical data from all test sites show a strong metabolic effect that exceeds the influence of the low-pH environment. These field experiments showed that care is needed when interpreting potential ocean acidification signals because various parameters affect shell chemistry and ultrastructure. Besides metabolic processes, seawater pH, factors such as salinity, water temperature, food availability and population density all affect the biogenic carbonate shell archive.

A short-term in situ CO2 enrichment experiment on Heron Island (GBR)

Ocean acidification poses multiple challenges for coral reefs on molecular to ecological scales, yet previous experimental studies of the impact of projected CO2 concentrations have mostly been done in aquarium systems with corals removed from their natural ecosystem and placed under artificial light and seawater conditions. The Coral–Proto Free Ocean Carbon Enrichment System (CP-FOCE) uses a network of sensors to monitor conditions within each flume and maintain experimental pH as an offset from environmental pH using feedback control on the injection of low pH seawater. Carbonate chemistry conditions maintained in the −0.06 and −0.22 pH offset treatments were significantly different than environmental conditions. The results from this short-term experiment suggest that the CP-FOCE is an important new experimental system to study in situ impacts of ocean acidification on coral reef ecosystems.

Kline D. I., Teneva L., Schneider K., Miard T., Chai A., Marker M., Headley K., Opdyke B., Nash M., Valetich M., Caves J. K., Russell B. D., Connell S. D., Kirkwood B. J., Brewer P., Peltzer E., Silverman J., Caldeira K., Dunbar R. B., Koseff J. R., Monismith S. G., Mitchell B. G., Dove S. & Hoegh-Guldberg O., 2012. A short-term in situ CO2 enrichment experiment on Heron Island (GBR). Scientific Reports 2:413. Article.

Rising CO2 and increased light exposure synergistically reduce marine primary productivity

Carbon dioxide and light are two major prerequisites of photosynthesis. Rising CO₂ levels in oceanic surface waters in combination with ample light supply are therefore often considered stimulatory to marine primary production^{1, 2, 3}. Here we show that the combination of an increase in both CO₂ and light exposure negatively impacts photosynthesis and growth of marine primary producers. When exposed to CO₂ concentrations projected for the end of this century⁴, natural phytoplankton assemblages of the South China Sea responded with decreased primary production and increased light stress at light intensities representative of the upper surface layer. The phytoplankton community shifted away from diatoms, the dominant phytoplankton group during our field campaigns. To examine the underlying mechanisms of the observed responses, we grew diatoms at different CO₂ concentrations and under varying levels (5–100%) of solar radiation experienced by the phytoplankton at different depths of the euphotic zone. Above 22–36% of incident surface irradiance, growth rates in the high-CO₂-grown cells were inversely related to light levels and exhibited reduced thresholds at which light becomes inhibitory. Future shoaling of upper-mixed-layer depths will expose phytoplankton to increased mean light intensities⁵. In combination with rising CO₂ levels, this may cause a widespread decline in marine primary production and a community shift away from diatoms, the main algal group that supports higher trophic levels and carbon export in the ocean.

Ocean acidification and the loss of phenolic substances in marine plants

Rising atmospheric CO₂ often triggers the production of plant phenolics, including many that serve as herbivore deterrents, digestion reducers, antimicrobials, or ultraviolet sunscreens. Such responses are predicted by popular models of plant defense, especially resource availability models which link carbon availability to phenolic biosynthesis. CO₂ availability is also increasing in the oceans, where anthropogenic emissions cause ocean acidification, decreasing seawater pH and shifting the carbonate system towards further CO₂ enrichment. Such conditions tend to increase seagrass productivity but may also increase rates of grazing on these marine plants. Here we show that high CO₂ / low pH conditions of OA decrease, rather than increase, concentrations of phenolic protective substances in seagrasses and eurysaline marine plants. We observed a loss of simple and polymeric phenolics in the seagrass Cymodocea nodosa near a volcanic CO₂ vent on the Island of Vulcano, Italy, where pH values decreased from 8.1 to 7.3 and pCO₂ concentrations increased ten-fold. We observed similar responses in two estuarine species, Ruppia maritima and Potamogeton perfoliatus, in in situ Free-Ocean-Carbon-Enrichment experiments conducted in tributaries of the Chesapeake Bay, USA. These responses are strikingly different than those exhibited by terrestrial plants. The loss of phenolic substances may explain the higher-than-usual rates of grazing observed near undersea CO₂ vents and suggests that ocean acidification may alter coastal carbon fluxes by affecting rates of decomposition, grazing, and disease. Our observations temper recent predictions that seagrasses would necessarily be “winners” in a high CO₂ world.

Temperate and tropical brown macroalgae thrive, despite decalcification, along natural CO2 gradients

Predicting the impacts of ocean acidification on coastal ecosystems requires an understanding of the effects on macroalgae and their grazers, as these underpin the ecology of rocky shores. Whilst calcified coralline algae (Rhodophyta) appear to be especially vulnerable to ocean acidification, there is a lack of information concerning calcified brown algae (Phaeophyta), which are not obligate calcifiers but are still important producers of calcium carbonate and organic matter in shallow coastal waters. Here we compare ecological shifts in sub-tidal rocky shore systems along CO₂ gradients created by volcanic seeps in the Mediterranean and Papua New Guinea, focussing on abundant macroalgae and grazing sea urchins. In both the temperate and tropical systems the abundances of grazing sea urchins declined dramatically along CO₂ gradients. Temperate and tropical species of the calcifying macroalgal genus Padina (Dictyoaceae, Phaeophyta) showed reductions in CaCO₃ content with CO₂ enrichment. In contrast to other studies of calcified macroalgae, however, we observed an increase in the abundance of Padina spp. in acidified conditions. Reduced sea urchin grazing pressure and significant increases in photosynthetic rates may explain the unexpected success of decalcified Padina spp. at elevated levels of CO₂. This is the first study to provide a comparison of ecological changes along CO₂ gradients between temperate and tropical rocky shores. The similarities we found in the responses of Padina spp. and sea urchin abundance at several vent systems increases confidence in predictions of the ecological impacts of ocean acidification over a large geographical range.

Storm-induced upwelling of high pCO2 waters onto the continental shelf of the western Arctic Ocean and implications for carbonate mineral saturation states

The carbon system of the western Arctic Ocean is undergoing a rapid transition as sea ice extent and thickness decline. These processes are dynamically forcing the region, with unknown consequences for CO2 fluxes and carbonate mineral saturation states, particularly in the coastal regions where sensitive ecosystems are already under threat from multiple stressors. In October 2011, persistent wind-driven upwelling occurred in open water along the continental shelf of the Beaufort Sea in the western Arctic Ocean. During this time, cold (<−1.2°C), salty (>32.4) halocline water—supersaturated with respect to atmospheric CO2 (pCO2 > 550 μatm) and undersaturated in aragonite (Ωaragonite < 1.0) was transported onto the Beaufort shelf. A single 10-day event led to the outgassing of 0.18–0.54 Tg-C and caused aragonite undersaturations throughout the water column over the shelf. If we assume a conservative estimate of four such upwelling events each year, then the annual flux to the atmosphere would be 0.72–2.16 Tg-C, which is approximately the total annual sink of CO2 in the Beaufort Sea from primary production. Although a natural process, these upwelling events have likely been exacerbated in recent years by declining sea ice cover and changing atmospheric conditions in the region, and could have significant impacts on regional carbon budgets. As sea ice retreat continues and storms increase in frequency and intensity, further outgassing events and the expansion of waters that are undersaturated in carbonate minerals over the shelf are probable.

Assembly rules of reef corals are flexible along a steep climatic gradient

Coral reefs, one of the world’s most complex and vulnerable ecosystems, face an uncertain future in coming decades as they continue to respond to anthropogenic climate change, overfishing, pollution, and other human impacts [[1] and [2]]. Traditionally, marine macroecology is based on presence/absence data from taxonomic checklists or geographic ranges, providing a qualitative overview of spatial shifts in species richness that treats rare and common species equally [[3] and [4]]. As a consequence, regional and long-term shifts in relative abundances of individual taxa are poorly understood. Here we apply a more rigorous quantitative approach to examine large-scale spatial variation in the species composition and abundance of corals on midshelf reefs along the length of Australia’s Great Barrier Reef, a biogeographic region where species richness is high and relatively homogeneous [5]. We demonstrate that important functional components of coral assemblages “sample” space differently at 132 sites separated by up to 1740 km, leading to complex latitudinal shifts in patterns of absolute and relative abundance. The flexibility in community composition that we document along latitudinal environmental gradients indicates that climate change is likely to result in a reassortment of coral reef taxa rather than wholesale loss of entire reef ecosystems.

Biodiversity stability of shallow marine benthos in Strait of Georgia, British Columbia, Canada through climate regimes, overfishing and ocean acidification

The highest human population density in British Columbia, Canada is situated around the shores of the Strait of Georgia, where current government policy is focusing early efforts toward achieving ecosystem-based management of marine resources. Climate regime shifts are acknowledged to have affected commercial fishery production in southern British Columbia (McFarlane et al., 2000), and overfishing is well documented in the Strait of Georgia region for a variety of important species, to the extent that Rockfish Conservation Areas have been created (Marliave & Challenger, 2009). As CO2 levels rise in the atmosphere, the oceans become progressively more acidic. While ocean acidification is predicted to be a great threat to marine ecosystems, little is known about its ecosystem impacts. Few taxpayer-funded studies have committed to long-term monitoring of full ecosystem biodiversity. This document presents results of over forty years of private taxonomic monitoring of shallow seafloors in the region centering on the Strait of Georgia.
Also presented are records of ambient ocean acidity levels (pH), documented continuously by the Vancouver Aquarium through the same time period. Biodiversity data are summarized in ways that enable visualization of possible relationships to climate regimes and ocean acidification. This work does not attempt statistical analyses, in the hope that the data trends can be incorporated into future models.

Coastal acidification in summer bottom oxygen-depleted waters in northwestern-northern Bohai Sea from June to August in 2011

Dissolved oxygen (DO) and pH in the central part of the Bohai Sea were surveyed in late June and late August, 2011. During the June cruise, the bottom DO was in the range of 215–290 μmol-O2 kg−1 (i.e. 85%–115% of the saturation level), and the bottom pH was in the range of 7.82–8.04 on the total-hydrogen-ion scale. In August, however, both the bottom DO and the pH had significantly declined in the northwestern-northern near-shore areas, where the water depth was no more than 35 m. The lowest bottom DO was 100–110 μmol-O2 kg−1 (only 44%–47% of the June DO values) in the northern near-shore area, where the bottom pH was 7.64–7.68 on the total-hydrogen-ion scale (0.16–0.20 units lower than the June pH value). The largest decreases in DO and in pH were observed in the northwestern near-shore bottom waters, corresponding to declines of 170 μmol-O2 kg−1 (as high as 59% of the June DO value) and 0.29 pH units, respectively. The greatest pH decline of 0.29 pH units meant that the total-hydrogen-ion concentration doubled in the bottom waters from June to August. Based on field measurements of bottom DO/pH combined with a simplified model simulation, we suggest that respiration/remineralization-derived CO2 increased the acidity in the bottom oxygen-depleted waters of northwestern-northern near-shore areas in the Bohai Sea as a result of coastal red tides and/or marine aquaculture. This aquatic chemistry is suggested to be partially responsible for scallop-breeding failures in the northwestern Bohai Sea in summer 2011.

Decreasing pH trend estimated from 35-yr time series of carbonate parameters in the Pacific sector of the Southern Ocean in summer

The Southern Ocean is an important region for investigation because it has a major effect on global air-to-sea CO2 fluxes and because of the ocean acidification resulting from the uptake of anthropogenic carbon, leading to serious consequences for marine ecosystems in the near future. We estimated long-term trends of ocean acidification in surface waters of the Pacific sector of the Southern Ocean, based on the summer observational records of oceanic CO2 partial pressure and related surface properties during 1969–2003. The computed pH time series exhibited substantial decreasing trends in the extensive region from the subtropical to polar zones. The mean rates of pH decrease over the 35-year period were 0.0011 to 0.0013 yr–1 in the zones north of the Polar Front and were larger in the polar zone (0.0020 yr–1). The contribution of trends in sea surface temperature to the trends of pH decrease was small in all zones. The high rate of pH decrease in the polar zone was attributed to the supply of dissolved inorganic carbon from lower layers, enhanced by intensified wind stress and superimposed onto the accumulation of anthropogenic CO2. A preliminary evaluation of thermodynamic changes in the upper carbonate system, using observational results, projected that the polar zone south of the Polar Front would be undersaturated with respect to aragonite in summer after 80 years.