ALBERT

Description: Increased atmospheric CO 2 concentration is leading to changes in the carbonate chemistry and the temperature of the ocean. The impact of these processes on marine organisms will depend on their ability to cope with those changes, particularly the maintenance of calcium carbonate structures. Both a laboratory experiment (long-term exposure to decreased pH and increased temperature) and collections of individuals from natural environments characterized by low pH levels (individuals from intertidal pools and around a CO 2 seep) were here coupled to comprehensively study the impact of near-future conditions of pH and temperature on the mechanical properties of the skeleton of the euechinoid sea urchin Paracentrotus lividus . To assess skeletal mechanical properties, we characterized the fracture force, Young's modulus, second moment of area, material nanohardness, and specific Young's modulus of sea urchin test plates. None of these parameters were significantly affected by low pH and/or increased temperature in the laboratory experiment and by low pH only in the individuals chronically exposed to lowered pH from the CO 2 seeps. In tidal pools, the fracture force was higher and the Young's modulus lower in ambital plates of individuals from the rock pool characterized by the largest pH variations but also a dominance of calcifying algae, which might explain some of the variation. Thus, decreases of pH to levels expected for 2100 did not directly alter the mechanical properties of the test of P. lividus . Since the maintenance of test integrity is a question of survival for sea urchins and since weakened tests would increase the sea urchins' risk of predation, our findings indicate that the decreasing seawater pH and increasing seawater temperature expected for the end of the century should not represent an immediate threat to sea urchins vulnerability.

Description: Most studies assessing the impacts of ocean acidification (OA) on benthic marine invertebrates have used stable mean pH/ p CO 2 levels to highlight variation in the physiological sensitivities in a range of taxa. However, many marine environments experience natural fluctuations in carbonate chemistry, and to date little attempt has been made to understand the effect of naturally fluctuating seawater p CO 2 ( p CO 2 sw ) on the physiological capacity of organisms to maintain acid–base homeostasis. Here, for the first time, we exposed two species of sea urchin with different acid–base tolerances, Paracentrotus lividus and Arbacia lixula , to naturally fluctuating p CO 2 sw conditions at shallow water CO 2 seep systems (Vulcano, Italy) and assessed their acid–base responses. Both sea urchin species experienced fluctuations in extracellular coelomic fluid pH, p CO 2 , and [HCO3–] (pH e , p CO 2 e , and [HCO3–]e , respectively) in line with fluctuations in p CO 2 sw . The less tolerant species, P. lividus, had the greatest capacity for [HCO3–]e buffering in response to acute p CO 2 sw fluctuations, but it also experienced greater extracellular hypercapnia and acidification and was thus unable to fully compensate for acid–base disturbances. Conversely, the more tolerant A. lixula relied on non-bicarbonate protein buffering and greater respiratory control. In the light of these findings, we discuss the possible energetic consequences of increased reliance on bicarbonate buffering activity in P. lividus compared with A. lixula and how these differing physiological responses to acute fluctuations in p CO 2 sw may be as important as chronic responses to mean changes in p CO 2 sw when considering how CO 2 emissions will affect survival and success of marine organisms within naturally assembled systems.

Description: Ideally, networks of marine protected areas should be designed with consideration for future changes. We examine how this could be tackled using the example of cold-water coral reefs which provide a number of ecosystem services but are vulnerable to both managed pressures (e.g. deep-water trawling) and unmanaged pressures (e.g. ocean acidification). We collated data on the known and predicted distribution of Northeast Atlantic coral reefs, their protected areas, and fishing effort. We modelled the effects of ocean acidification on aragonite saturation to examine whether existing protected areas will ensure adequate protection for cold-water coral reefs under four possible future scenarios across two models. The best-case scenario suggests only minor impacts of ocean acidification, and that trawling remains the main threat to these reefs. However, in the worst-case scenario, by 2060, over 85% of these reefs are expected to be exposed to corrosive waters. We argue that unmanaged pressures such as ocean acidification and global warming should be incorporated into marine management decisions, with a focus on the protection of cold-water coral reefs to ensure long-term survival of these habitats. A similar approach could be taken for other iconic marine habitats in the face of climate change.

Description: Ideally, networks of marine protected areas should be designed with consideration for future changes. We examine how this could be tackled using the example of cold-water coral reefs which provide a number of ecosystem services but are vulnerable to both managed pressures (e.g. deep-water trawling) and unmanaged pressures (e.g. ocean acidification). We collated data on the known and predicted distribution of Northeast Atlantic coral reefs, their protected areas, and fishing effort. We modelled the effects of ocean acidification on aragonite saturation to examine whether existing protected areas will ensure adequate protection for cold-water coral reefs under four possible future scenarios across two models. The best-case scenario suggests only minor impacts of ocean acidification, and that trawling remains the main threat to these reefs. However, in the worst-case scenario, by 2060, over 85% of these reefs are expected to be exposed to corrosive waters. We argue that unmanaged pressures such as ocean acidification and global warming should be incorporated into marine management decisions, with a focus on the protection of cold-water coral reefs to ensure long-term survival of these habitats. A similar approach could be taken for other iconic marine habitats in the face of climate change.

Description: 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 (from the Mediterranean) and M. edulis (from the Wadden Sea) 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 of Ischia. The shells of transplanted mussels were compared with M. edulis collected at pH ~8.2 from Sylt (German Wadden Sea). 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.

Description: Here we show the use of the 210Pb-226Ra excess method to determine the growth rate of corals from one of the world's largest known cold-water coral reef, the Røst Reef off Norway. Two large branching framework-forming cold-water coral specimens, one Lophelia pertusa and one Madrepora oculata were collected alive at 350 m water depth from the Røst Reef at ~67° N and ~9° E. Pb and Ra isotopes were measured along the major growth axis of both specimens using low level alpha and gamma spectrometry and the corals trace element compositions were studied using ICP-QMS. Due to the different chemical behaviors of Pb and Ra in the marine environment, 210Pb and 226Ra were not incorporated the same way into the aragonite skeleton of those two cold-water corals. Thus to assess of the growth rates of both specimens we have here taken in consideration the exponential decrease of initially incorporated 210Pb as well as the ingrowth of 210Pb from the decay of 226Ra. Moreover a~post-depositional 210Pb incorporation is found in relation to the Mn-Fe coatings that could not be entirely removed from the oldest parts of the skeletons. The 226Ra activities in both corals were fairly constant, then assuming constant uptake of 210Pb through time the 210Pb-226Ra chronology can be applied to calculate linear growth rate. The 45.5 cm long branch of M. oculata reveals an age of 31 yr and a~linear growth rate of 14.4 ± 1.1 mm yr−1, i.e. 2.6 polyps per year. However, a correction regarding a remaining post-depositional Mn-Fe oxide coating is needed for the base of the specimen. The corrected age tend to confirm the radiocarbon derived basal age of 40 yr (using 14C bomb peak) with a mean growth rate of 2 polyps yr−1. This rate is similar to the one obtained in Aquaria experiments under optimal growth conditions. For the 80 cm-long specimen of L. pertusa a remaining contamination of metal-oxides is observed for the middle and basal part of the coral skeleton, inhibiting similar accurate age and growth rate estimates. However, the youngest branch was free of Mn enrichment and this 15 cm section reveals a growth rate of 8 mm yr−1 (~1 polyp every two to three years). However, the 210Pb growth rate estimate is within the lowermost ranges of previous growth rate estimates and may thus reflect that the coral was not developing at optimal growth conditions. Overall, 210Pb-226Ra dating can be successfully applied to determine the age and growth rate of framework-forming cold-water corals, however, removal of post-depositional Mn-Fe oxide deposits is a prerequisite. If successful, large branching M. oculata and L. pertusa coral skeletons provide unique oceanographic archive for studies of intermediate water environmentals with an up to annual time resolution and spanning over many decades.

Description: There is concern that the use of natural volcanic CO2 vents as analogs for studies of the impacts of ocean acidification on marine organisms are biased due to physiochemical influences other than seawater pH alone. One issue that has been raised is whether potentially harmful trace elements in sediments that are rendered more soluble and labile in low pH environments are made more bioavailable, and sequestered in the local flora and fauna at harmful levels. In order to evaluate this hypothesis, we analyzed the concentrations of trace elements in shells (an established proxy for tissues) of four species of gastropods (two limpets, a topshell and a whelk) collected from three sites in Levante Bay, Vulcano Island. Each sampling site increased in distance from the primary CO2 vent and thus represented low, moderate, and ambient seawater pH conditions. Concentrations of As, Cd, Co, Cr, Hg, Mo, Ni, Pb, and V measured in shells using ICP-OES were below detection thresholds for all four gastropod species at all three sites. However, there were measurable concentrations of Sr, Mn, and U in the shells of the limpets Patella caerulea, P. rustica, and the snail Osilinus turbinatus, and similarly, Sr, Mn, U, and also Zn in the shells of the whelk Hexaplex trunculus. Levels of these elements were within the ranges measured in gastropod shells in non-polluted environments, and with the exception of U in the shells of P. caerulea, where the concentration was significantly lower at the collecting site closest to the vent (low pH site), there were no site-specific spatial differences in concentrations for any of the trace elements in shells. Thus trace element enhancement in sediments in low-pH environments was not reflected in greater bioaccumulations of potentially harmful elements in the shells of common gastropods.

Description: 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.

Description: Here we show the use of the 210Pb-226Ra excess method to determine the growth rate of two corals from the world's largest known cold-water coral reef, Røst Reef, north of the Arctic circle off Norway. Colonies of each of the two species that build the reef, Lophelia pertusa and Madrepora oculata, were collected alive at 350 m depth using a submersible. Pb and Ra isotopes were measured along the major growth axis of both specimens using low level alpha and gamma spectrometry and trace element compositions were studied. 210Pb and 226Ra differ in the way they are incorporated into coral skeletons. Hence, to assess growth rates, we considered the exponential decrease of initially incorporated 210Pb, as well as the increase in 210Pb from the decay of 226Ra and contamination with 210Pb associated with Mn-Fe coatings that we were unable to remove completely from the oldest parts of the skeletons. 226Ra activity was similar in both coral species, so, assuming constant uptake of 210Pb through time, we used the 210Pb-226Ra chronology to calculate growth rates. The 45.5 cm long branch of M. oculata was 31 yr with an average linear growth rate of 14.4 ± 1.1 mm yr−1 (2.6 polyps per year). Despite cleaning, a correction for Mn-Fe oxide contamination was required for the oldest part of the colony; this correction corroborated our radiocarbon date of 40 yr and a mean growth rate of 2 polyps yr−1. This rate is similar to the one obtained in aquarium experiments under optimal growth conditions. For the 80 cm-long L. pertusa colony, metal-oxide contamination remained in both the middle and basal part of the coral skeleton despite cleaning, inhibiting similar age and growth rate estimates. The youngest part of the colony was free of metal oxides and this 15 cm section had an estimated a growth rate of 8 mm yr−1, with high uncertainty (~1 polyp every two to three years). We are less certain of this 210Pb growth rate estimate which is within the lowermost ranges of previous growth rate estimates. We show that 210Pb-226Ra dating can be successfully applied to determine the age and growth rate of framework-forming cold-water corals if Mn-Fe oxide deposits can be removed. Where metal oxides can be removed, large M. oculata and L. pertusa skeletons provide archives for studies of intermediate water masses with an up to annual time resolution and spanning over many decades.