Low Aragonite Saturation State Research Articles

Macroalgal cover on coral reefs: Spatial and environmental predictors, and decadal trends in the Great Barrier Reef.

Macroalgae are an important component of coral reef ecosystems. We identified spatial patterns, environmental drivers and long-term trends of total cover of upright fleshy and calcareous coral reef inhabiting macroalgae in the Great Barrier Reef. The spatial study comprised of one-off surveys of 1257 sites (latitude 11-24°S, coastal to offshore, 0-18 m depth), while the temporal trends analysis was based on 26 years of long-term monitoring data from 93 reefs. Environmental predictors were obtained from in situ data and from the coupled hydrodynamic-biochemical model eReefs. Macroalgae dominated the benthos (≥50% cover) on at least one site of 40.4% of surveyed inshore reefs. Spatially, macroalgal cover increased steeply towards the coast, with latitude away from the equator, and towards shallow (≤3 m) depth. Environmental conditions associated with macroalgal dominance were: high tidal range, wave exposure and irradiance, and low aragonite saturation state, Secchi depth, total alkalinity and temperature. Evidence of space competition between macroalgal cover and hard coral cover was restricted to shallow inshore sites. Temporally, macroalgal cover on inshore and mid-shelf reefs showed some fluctuations, but unlike hard corals they showed no systematic trends. Our extensive empirical data may serve to parameterize ecosystem models, and to refine reef condition indices based on macroalgal data for Pacific coral reefs.

Global Synthesis of the Status and Trends of Ocean Acidification Impacts on Shelled Pteropods

The accumulation of anthropogenic CO2 in the ocean has major ecological, socioeconomic, and biogeochemical impacts, with repercussions for the ocean as a critical carbon sink. Ocean acidification (OA) disproportionally affects marine calcifiers, among which pelagic zooplanktonic pteropods play a significant role in carbonate export. The pteropod, due to the susceptibility of its aragonite shell to rapid dissolution, is one of most vulnerable groups and a key indicator for OA regional monitoring, but its regional sensitivities have not yet been extrapolated over global scales. To delineate spatial and temporal changes in pteropod shell dissolution, global OA status and the OA rate of change were evaluated, based on gridded climatologies of observations and using a Regional Ocean Modeling System (ROMS) biogeochemical/ecosystem model. Pteropods dominate in the polar and upwelling regions characterized by low aragonite saturation state and low buffering capacity, where extended pteropod subsurface dissolution is projected. We show that pteropods are most susceptible to OA in the polar regions, subpolar North Pacific, and eastern boundary upwelling system regions, particularly the California and Humboldt Current Systems. Rates of acidification and corresponding increases in pteropod shell dissolution are projected to be the fastest in the North and South Equatorial Currents.

Understanding the Implications of Hydrographic Processes on the Dynamics of the Carbonate System in a Sub-Antarctic Marine-Terminating Glacier-Fjord (53°S)

The biogeochemical dynamics of fjords in the southeastern Pacific Ocean are strongly influenced by hydrological and oceanographic processes occurring at a seasonal scale. In this study, we describe the role of hydrographic forcing on the seasonal variability of the carbonate system of the Sub-Antarctic glacial fjord, Seno Ballena, in the Strait of Magellan (53°S). Biogeochemical variables were measured in 2018 during three seasonal hydrographic cruises (fall, winter and spring) and from a high-frequency pCO2-pH mooring for 10 months at 10 ± 1 m depth in the fjord. The hydrographic data showed that freshwater input from the glacier influenced the adjacent surface layer of the fjord and forced the development of undersaturated CO2 (< 400 μatm) and low aragonite saturation state (ΩAr < 1) water. During spring, the surface water had relatively low pCO2 (mean = 365, range: 167 - 471 μatm), high pH (mean = 8.1 on the total proton concentration scale, range: 8.0 - 8.3), and high ΩAr (mean = 1.6, range: 1.3 - 4.0). Concurrent measurements of phytoplankton biomass and nutrient conditions during spring indicated that the periods of lower pCO2 values corresponded to higher phytoplankton photosynthesis rates, resulting from autochthonous nutrient input and vertical mixing. In contrast, higher values of pCO2 (range: 365 – 433 μatm) and relatively lower values of pHT (range: 8.0 – 8.1) and ΩAr (range: 0.9 – 2.0) were recorded in cold surface waters during winter and fall. The naturally low freshwater carbonate ion concentrations diluted the carbonate ion concentrations in seawater and decreased the calcium carbonate saturation of the fjord. In spring, at 10 m depth, higher primary productivity caused a relative increase in ΩAr and pHT. Assuming global climate change will bring further glacier retreat and ocean acidification, this study represents important advances in our understanding of glacier meltwater processes on CO2 dynamics in glacier–fjord systems.

Spatial patterns in aragonite saturation horizon over the northern California shelf

Ocean acidification along the California Shelf is controlled by the upwelling of deep CO2-rich waters. This coastal upwelling is driven by along-shore winds, which vary seasonally. We report aragonite undersaturation based on proxy values derived from temperature, salinity, and dissolved oxygen data and show the relationship between the spatial extent of low aragonite saturation state values and upwelling-favorable along-shore winds. Our data were collected on seasonal hydrographic surveys off northern California that span several years. Transect data show a persistent spatial pattern with shoaling of the aragonite saturation horizon over the shelf (during May and July surveys, mean depth of horizon is less than 20 m within ∼20 km of the coast), and surfacing of the saturation horizon during times of strong upwelling events. Our model shows that a higher proportion of the water column is undersaturated in aragonite when northerly winds are stronger. In addition, we found that pteropod abundance was positively correlated with aragonite saturation state, with lower densities associated with lower saturation values. If upwelling-favorable winds increase with global warming, as anticipated, more shelf water habitat will become undersaturated in aragonite, negatively impacting populations of calcifying organisms like pteropods.

Integrated Assessment of Ocean Acidification Risks to Pteropods in the Northern High Latitudes: Regional Comparison of Exposure, Sensitivity and Adaptive Capacity

Exposure to the impact of ocean acidification (OA) is increasing in high-latitudinal productive habitats. Pelagic calcifying snails (pteropods), a significant component of the diet of economically important fish, are found in high abundance in these regions. Pteropods have thin shells that readily dissolve at low aragonite saturation state (Ωar), making them susceptible to OA. Here, we conducted a first integrated risk assessment for pteropods in the Eastern Pacific subpolar gyre, the Gulf of Alaska (GoA), Bering Sea, and Amundsen Gulf. We determined the risk for pteropod populations by integrating measures of OA exposure, biological sensitivity, and resilience. Exposure was based on physical-chemical hydrographic observations and regional biogeochemical model outputs, delineating seasonal and decadal changes in carbonate chemistry conditions. Biological sensitivity was based on pteropod morphometrics and shell-building processes, including shell dissolution, density and thickness. Resilience and adaptive capacity were based on species diversity and spatial connectivity, derived from the particle tracking modeling. Extensive shell dissolution was found in the central and western part of the subpolar gyre, parts of the Bering Sea, and Amundsen Gulf. We identified two distinct morphotypes:L. helicina helicinaandL. helicina pacifica, with high-spired and flatter shells, respectively. Despite the presence of different morphotypes, genetic analyses based on mitochondrial haplotypes identified a single species, without differentiation between the morphological forms, coinciding with evidence of widespread spatial connectivity. We found that shell morphometric characteristics depends on omega saturation state (Ωar); under Ωardecline, pteropods build flatter and thicker shells, which is indicative of a certain level of phenotypic plasticity. An integrated risk evaluation based on multiple approaches assumes a high risk for pteropod population persistence with intensification of OA in the high latitude eastern North Pacific because of their known vulnerability, along with limited evidence of species diversity despite their connectivity and our current lack of sufficient knowledge of their adaptive capacity. Such a comprehensive understanding would permit improved prediction of ecosystem change relevant to effective fisheries resource management, as well as a more robust foundation for monitoring ecosystem health and investigating OA impacts in high-latitudinal habitats.

Severe biological effects under present-day estuarine acidification in the seasonally variable Salish Sea

Estuaries are recognized as one of the habitats most vulnerable to coastal ocean acidification due to seasonal extremes and prolonged duration of acidified conditions. This is combined with co-occurring environmental stressors such as increased temperature and low dissolved oxygen. Despite this, evidence of biological impacts of ocean acidification in estuarine habitats is largely lacking. By combining physical, biogeochemical, and biological time-series observations over relevant seasonal-to-interannual time scales, this study is the first to describe both the spatial and temporal variation of biological response in the pteropod Limacina helicina to estuarine acidification in association with other stressors. Using clustering and principal component analyses, sampling sites were grouped according to their distribution of physical and biogeochemical variables over space and time. This identified the most exposed habitats and time intervals corresponding to the most severe negative biological impacts across three seasons and three years. We developed a cumulative stress index as a means of integrating spatial-temporal OA variation over the organismal life history. Our findings show that over the 2014–2016 study period, the severity of low aragonite saturation state combined with the duration of exposure contributed to overall cumulative stress and resulted in severe shell dissolution. Seasonally-variable estuaries such as the Salish Sea (Washington, U.S.A.) predispose sensitive organisms to more severe acidified conditions than those of coastal and open-ocean habitats, yet the sensitive organisms persist. We suggest potential environmental factors and compensatory mechanisms that allow pelagic calcifiers to inhabit less favorable habitats and partially offset associated stressors, for instance through food supply, increased temperature, and adaptation of their life history. The novel metric of cumulative stress developed here can be applied to other estuarine environments with similar physical and chemical dynamics, providing a new tool for monitoring biological response in estuaries under pressure from accelerating global change.

Comparing Subsurface Seasonal Deoxygenation and Acidification in the Yellow Sea and Northern East China Sea Along the North-to-South Latitude Gradient

To better understand the relationship between subsurface seasonal deoxygenation and acidification in the Yellow Sea and northern East China Sea (ECS), we examined carbonate system parameters and dissolved oxygen (DO) of seven field surveys conducted in 2017–2018, spanning all four seasons. Low pHT values of 7.71‒7.80 and critically low aragonite saturation state (Ωarag) values of 1.07‒1.40 along with undersaturated DO of mostly higher than 150 μmol O2 kg–1 occurred in the Yellow Sea Cold Water Mass area in summer and autumn, while hypoxic DO values of 49‒63 μmol O2 kg–1 and extremely low pHT values of 7.68‒7.74 as well as critically low Ωarag values of 1.21‒1.39 were observed in the northern ECS in July 2018. At the beginning of warm-season stratification formation, the cold Yellow Sea waters had much higher DO but lower Ωarag values than those in relatively warmer ECS waters, while yearly initial pHT values rarely exhibited differences between the two coastal seas. During warm seasons, the central Yellow Sea accumulated respiration products beneath the thermocline in summer and autumn, while the northern ECS bottom waters preserved them only in summer. This study highlights fundamental roles of wintertime carbon dioxide (CO2) solubility along a north-to-south latitude gradient in the coastal acidification development. In comparison with the relatively low-latitude northern ECS subject to seasonal hypoxia, relatively high-latitude Yellow Sea exhibits higher CO2 solubility in winter and longer respiration-product accumulations in warm seasons, leading to lower Ωarag in the central Yellow Sea than those in the northern ECS. However, the present-day central Yellow Sea is free from hypoxia.

Uptake of groundwater nitrogen by a near-shore coral reef community on Bermuda

Nutrient enrichment can slow growth, enhance bioerosion rates, and intensify algal competition for reef-building corals. In areas of high human population density and/or limited waste management, submarine groundwater discharge can transfer anthropogenic nutrients from polluted groundwater to coastal reefs. In this case study, we investigate the impact of submarine groundwater discharge on a near-shore reef in Bermuda, where over 60% of sewage generated by the island’s 64,000 residents enters the groundwater through untreated cesspits. Temperature, salinity, pH, and alkalinity were monitored at a groundwater discharge vent, three locations across the adjacent coral reef (0–30 m from shore), and a comparison patch reef site 2 km from shore. Groundwater discharge was characterized by low salinity, low aragonite saturation state (Ωar), high alkalinity, elevated nitrate + nitrite (NO3– + NO2–; hereafter, “NO3–”) concentrations (> 400 µM), and an elevated 15N/14N ratio of NO3– (δ15N = 10.9 ± 0.02‰ vs. air, mean ± SD). Rainfall and tidal cycles strongly impacted groundwater discharge, with maximum discharge during low tide. NO3– concentrations on the near-shore reef averaged 4 µM, ten times higher than that found at the control site 2 km away, and elevated NO3– δ15N at the near-shore reef indicated sewage-contaminated groundwater as a significant nitrogen source. Tissue δ15N of Porites astreoides, a dominant reef-building coral, was elevated by ~ 3‰ on the near-shore reef compared to the control site, indicating that corals across the near-shore reef were assimilating groundwater-derived nitrogen. In addition, coral skeletal density and calcification rates across the near-shore reef were inversely correlated with NO3– concentration and δ15N, indicating a negative coral health response to groundwater-borne nutrient inputs. P. astreoides bioerosion rates, in contrast, did not show an effect from the groundwater input.

Reconstructing Aragonite Saturation State Based on an Empirical Relationship for Northern California

Ocean acidification is a global phenomenon with highly regional spatial and temporal patterns. In order to address the challenges of future ocean acidification at a regional scale, it is necessary to increase the resolution of spatial and temporal monitoring of the inorganic carbon system beyond what is currently available. One approach is to develop empirical regional models that enable aragonite saturation state to be estimated from existing hydrographic measurements, for which greater spatial coverage and longer time series exist in addition to higher spatial and temporal resolution. We present such a relationship for aragonite saturation state for waters off Northern California based on in situ bottle sampling and instrumental measurements of temperature, salinity, and dissolved oxygen. Application of this relationship to existing datasets (5 to 200 m depth) demonstrates both seasonal and interannual variability in aragonite saturation state. We document a deeper aragonite saturation horizon and higher near surface aragonite saturation state in the summers of 2014 and 2015 (compared with 2010–2013), associated with anomalous warm conditions and decadal scale oscillations. Application of this model to time series data reiterates the direct association between low aragonite saturation state and upwelled waters and highlights the extent to which benthic communities on the Northern California shelf are already exposed to aragonite undersaturated waters.

Export of calcium carbonate corrosive waters from the East Siberian Sea

Abstract. The Siberian shelf seas are areas of extensive biogeochemical transformation of organic matter, both of marine and terrestrial origin. This in combination with brine production from sea ice formation results in a cold bottom water of relative high salinity and partial pressure of carbon dioxide (pCO2). Data from the SWERUS-C3 expedition compiled on the icebreaker Oden in July to September 2014 show the distribution of such waters at the outer shelf, as well as their export into the deep central Arctic basins. Very high pCO2 water, up to ∼ 1000 µatm, was observed associated with high nutrients and low oxygen concentrations. Consequently, this water had low saturation state with respect to calcium carbonate down to less than 0.8 for calcite and 0.5 for aragonite. Waters undersaturated in aragonite were also observed in the surface in waters at equilibrium with atmospheric CO2; however, at these conditions the cause of under-saturation was low salinity from river runoff and/or sea ice melt. The calcium carbonate corrosive water was observed all along the continental margin and well out into the deep Makarov and Canada basins at a depth from about 50 m depth in the west to about 150 m in the east. These waters of low aragonite saturation state are traced in historic data to the Canada Basin and in the waters flowing out of the Arctic Ocean north of Greenland and in the western Fram Strait, thus potentially impacting the marine life in the North Atlantic Ocean.

Chemical and biological impacts of ocean acidification along the west coast of North America

The continental shelf region off the west coast of North America is seasonally exposed to water with a low aragonite saturation state by coastal upwelling of CO2-rich waters. To date, the spatial and temporal distribution of anthropogenic CO2 (Canth) within the CO2-rich waters is largely unknown. Here we adapt the multiple linear regression approach to utilize the GO-SHIP Repeat Hydrography data from the northeast Pacific to establish an annually updated relationship between Canth and potential density. This relationship was then used with the NOAA Ocean Acidification Program West Coast Ocean Acidification (WCOA) cruise data sets from 2007, 2011, 2012, and 2013 to determine the spatial variations of Canth in the upwelled water. Our results show large spatial differences in Canth in surface waters along the coast, with the lowest values (37–55 μmol kg−1) in strong upwelling regions off southern Oregon and northern California and higher values (51–63 μmol kg−1) to the north and south of this region. Coastal dissolved inorganic carbon concentrations are also elevated due to a natural remineralized component (Cbio), which represents carbon accumulated through net respiration in the seawater that has not yet degassed to the atmosphere. Average surface Canth is almost twice the surface remineralized component. In contrast, Canth is only about one third and one fifth of the remineralized component at 50 m and 100 m depth, respectively. Uptake of Canth has caused the aragonite saturation horizon to shoal by approximately 30–50 m since the preindustrial period so that undersaturated waters are well within the regions of the continental shelf that affect the shell dissolution of living pteropods. Our data show that the most severe biological impacts occur in the nearshore waters, where corrosive waters are closest to the surface. Since the pre-industrial times, pteropod shell dissolution has, on average, increased approximately 19–26% in both nearshore and offshore waters.

Shell condition and survival of Puget Sound pteropods are impaired by ocean acidification conditions.

We tested whether the thecosome pteropod Limacina helicina from Puget Sound, an urbanized estuary in the northwest continental US, experiences shell dissolution and altered mortality rates when exposed to the high CO2, low aragonite saturation state (Ωa) conditions that occur in Puget Sound and the northeast Pacific Ocean. Five, week-long experiments were conducted in which we incubated pteropods collected from Puget Sound in four carbon chemistry conditions: current summer surface (∼460–500 µatm CO2, Ωa≈1.59), current deep water or surface conditions during upwelling (∼760 and ∼1600–1700 µatm CO2, Ωa≈1.17 and 0.56), and future deep water or surface conditions during upwelling (∼2800–3400 µatm CO2, Ωa≈0.28). We measured shell condition using a scoring regime of five shell characteristics that capture different aspects of shell dissolution. We characterized carbon chemistry conditions in statistical analyses with Ωa, and conducted analyses considering Ωa both as a continuous dataset and as discrete treatments. Shell dissolution increased linearly as aragonite saturation state decreased. Discrete treatment comparisons indicate that shell dissolution was greater in undersaturated treatments compared to oversaturated treatments. Survival increased linearly with aragonite saturation state, though discrete treatment comparisons indicated that survival was similar in all but the lowest saturation state treatment. These results indicate that, under starvation conditions, pteropod survival may not be greatly affected by current and expected near-future aragonite saturation state in the NE Pacific, but shell dissolution may. Given that subsurface waters in Puget Sound’s main basin are undersaturated with respect to aragonite in the winter and can be undersaturated in the summer, the condition and persistence of the species in this estuary warrants further study.

The intensity, duration, and severity of low aragonite saturation state events on the California continental shelf

Ocean acidification will progress in an episodic manner, yet this has rarely been considered. Here, we investigate how the intensity, duration, and severity of episodic low aragonite saturation state events along the California continental shelf have changed since ~1750, and how they might change up to 2050 under the A2 scenario from the Special Report on Emissions Scenarios by the Intergovernmental Panel on Climate Change. Our model‐based results suggest that between 1750 and 2010, aragonite undersaturation events along the shelf have quadrupled in their number and lengthened in duration, but that even larger changes are bound to occur within the next 20 to 40 years. Undersaturation will become very likely the norm near the seafloor by 2030, and if atmospheric CO2 increases beyond ~500 ppm, this layer will become permanently undersaturated. Combined with a fourfold increase in intensity, the resulting increase in severity of low aragonite saturation state events will substantially affect the viability of calcifying organisms and will alter ecosystem structure.

Aragonite saturation state dynamics in a coastal upwelling zone

AbstractCoastal upwelling zones may be at enhanced risk from ocean acidification as upwelling brings low aragonite saturation state (ΩAr) waters to the surface that are further suppressed by anthropogenic CO2. ΩAr was calculated with pH, pCO2, and salinity‐derived alkalinity time series data from autonomous pH and pCO2 instruments moored on the Oregon shelf and shelf break during different seasons from 2007 to 2011. Surface ΩAr values ranged between 0.66 ± 0.04 and 3.9 ± 0.04 compared to an estimated pre‐industrial range of 1.0 ± 0.1 to 4.7 ± 0.1. Upwelling of high‐CO2 water and subsequent removal of CO2 by phytoplankton imparts a dynamic range to ΩAr from ~1.0 to ~4.0 between spring and autumn. Freshwater input also suppresses saturation states during the spring. Winter ΩAr is less variable than during other seasons and is controlled primarily by mixing of the water column.

Response of the Arctic Pteropod Limacina helicina to Projected Future Environmental Conditions

Thecosome pteropods (pelagic mollusks) can play a key role in the food web of various marine ecosystems. They are a food source for zooplankton or higher predators such as fishes, whales and birds that is particularly important in high latitude areas. Since they harbor a highly soluble aragonitic shell, they could be very sensitive to ocean acidification driven by the increase of anthropogenic CO2 emissions. The effect of changes in the seawater chemistry was investigated on Limacina helicina, a key species of Arctic pelagic ecosystems. Individuals were kept in the laboratory under controlled pCO2 levels of 280, 380, 550, 760 and 1020 µatm and at control (0°C) and elevated (4°C) temperatures. The respiration rate was unaffected by pCO2 at control temperature, but significantly increased as a function of the pCO2 level at elevated temperature. pCO2 had no effect on the gut clearance rate at either temperature. Precipitation of CaCO3, measured as the incorporation of 45Ca, significantly declined as a function of pCO2 at both temperatures. The decrease in calcium carbonate precipitation was highly correlated to the aragonite saturation state. Even though this study demonstrates that pteropods are able to precipitate calcium carbonate at low aragonite saturation state, the results support the current concern for the future of Arctic pteropods, as the production of their shell appears to be very sensitive to decreased pH. A decline of pteropod populations would likely cause dramatic changes to various pelagic ecosystems.