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Acesta excavata (Fabricius, 1779) is a slow growing bivalve from the Limidae family and is often found associated with cold-water coral reefs along the European continental margin. Here we present the compositional variability of frequently used proxy elemental ratios (Mg/ Ca, Sr/Ca, Na/Ca) measured by laser-ablation mass spectrometry (LA-ICP-MS) and com- pare it to in-situ recorded instrumental seawater parameters such as temperature and salin- ity. Shell Mg/Ca measured in the fibrous calcitic shell section was overall not correlated with seawater temperature or salinity; however, some samples show significant correlations with temperature with a sensitivity that was found to be unusually high in comparison to other marine organisms. Mg/Ca and Sr/Ca measured in the fibrous calcitic shell section display significant negative correlations with the linear extension rate of the shell, which indicates strong vital effects in these bivalves. Multiple linear regression analysis indicates that up to 79% of elemental variability is explicable with temperature and salinity as independent pre- dictor values. Yet, the overall results clearly show that the application of Element/Ca (E/Ca) ratios in these bivalves to reconstruct past changes in temperature and salinity is likely to be complicated due to strong vital effects and the effects of organic material embedded in the shell. Therefore, we suggest to apply additional techniques, such as clumped isotopes, in order to exactly determine and quantify the underlying vital effects and possibly account for these. We found differences in the chemical composition between the two calcitic shell lay- ers that are possibly explainable through differences of the crystal morphology. Sr/Ca ratios also appear to be partly controlled by the amount of magnesium, because the small magne- sium ions bend the crystal lattice which increases the space for strontium incorporation. Oxi- dative cleaning with H2O2 did not significantly change the Mg/Ca and Sr/Ca composition of the shell. Na/Ca ratios decreased after the oxidative cleaning, which is most likely a leaching effect and not caused by the removal of organic matter.
Geochemical investigations on biogenic carbonates are commonly conducted to reconstruct the environmental conditions of the past. However, different carbonate producers incorporate elements to varying degrees, due to biological vital effects. Detecting and quantifying these effects is crucial to produce reliable reconstructions. These paleoreconstructions are of great importance to evaluate the consequences of our recent climate change and identify control mechanisms on the distribution of endangered species such as Desmophyllum pertusum. In chapter three we tested Mg/Ca, Sr/Ca and Na/Ca ratios on this species, among other coldwater scleractinians, to test if they provide reliable proxy information. The results reveal no apparent control of Mg/Ca or Sr/Ca ratios through seawater temperature, salinity or pH. Na/Ca ratios appear to be partly controlled by the seawater temperature, which is also true for other aragonitic organisms such as warm-water corals and the bivalve Mytilus edulis. However, a large variability complicates possible reconstructions by means of Na/Ca. In addition, we explore different models to explain the apparent temperature effect on Na/Ca ratios based on temperature sensitive Na and Ca pumping enzymes.
The bivalve Acesta excavata is commonly found in cold-water coral reefs among the North Atlantic, together with D. pertusum. Multiple linear regression analysis, presented in chapter four, indicates that up to 79% of the elemental variability in Mg/Ca, Sr/Ca and Na/Ca is explainable with temperature and salinity as independent predictor variables. Vital effects, for instance growth rate effects, are evident and make paleoreconstructions not feasible. Furthermore, organic material embedded in the shell, as well as possible stress effects can drastically change the elemental composition. Removal of these organic matrices from bulk samples for LA-ICP-MS (laser ablation inductively coupled mass spectrometer) measurements by means of oxidative cleaning is not possible, but Na/Ca ratios decrease after this cleaning. This is presumably an effect of leaching and not caused by the removal of organic matrices.
Interesting biogeochemical relations were found in the parasitic foraminifera H. sarcophaga. We report Mg/Ca, Sr/Ca, Na/Ca and Mn/Ca ratios measured in H. sarcophaga from two different host species (A. excavata and D. pertusum) in chapter five. Sr/Ca ratios are significantly higher in foraminifera that lived on D. pertusum. This could indicate that dissolved host material is utilized in shell calcification of H. sarcophaga, given the naturally higher strontium concentration in the aragonite of D. pertusum. Mn/Ca ratios are highest in foraminifera that lived on A. excavata but did not fully penetrate the host’s shell. Most likely, this represents a juvenile stadium of the foraminifera during which it feeds on the organic
periostracum of the bivalve, which is enriched in Mn and Fe. The isotopic compositions are similarly affected, both δ18O and δ13C values are significantly lower in foraminifera that lived 23on D. pertusum compared to specimen that lived on A. excavata. Again, this might represent the uptake of dissolved host material or different pH regimes in the calcifying fluid of the hosts (bivalve < 8, coral > 8) that control the extent of hydration/hydroxylation reactions. Temperature reconstructions are possible using stable oxygen isotopes on this foraminifera species; however, the results are only reliable if the foraminifera lived on A. excavata. Samples of H. sarcophaga from D. pertusum would lead to overestimations of the seawater temperature due to the lower δ18O values.
Apart from biological vital effects, storage and preservation methods can significantly change the geochemical composition of different marine biogenic carbonates. In chapter six this is presented on the example of ethanol preservation, a common technique to allow extended storage of biogenic samples. The investigation reveals a significant decrease of Mg/Ca and Na/Ca ratios even after only 45 days storage in ultrapure ethanol. Sr/Ca ratios on the other hand are not influenced.
Besides temperature, salinity and pH further environmental parameters are important such as nutrient availability, especially for the distribution of cold-water corals. In chapter seven we extend the investigations on A. excavata by including the elemental ratios Ba/Ca, Mn/Ca and P/Ca. We expected P/Ca to be helpful in the otherwise difficult process of dentifying growth increments. Based on our observations we had to refute this theory. P/Ca ratios are not systematically enriched in the vicinity of growth lines. Instead, we found a regular sequence of peaks of Ba/Ca, P/Ca and Mn/Ca. This sequence as well as the peaks in general are potentially caused by equential blooms of different algae, diatoms and other planktonic organisms ...
Here we present a comprehensive attempt to correlate aragonitic Na∕Ca ratios from Desmophyllum pertusum (formerly known as Lophelia pertusa), Madrepora oculata and a caryophylliid cold-water coral (CWC) species with different seawater parameters such as temperature, salinity and pH. Living CWC specimens were collected from 16 different locations and analyzed for their Na∕Ca ratios using solution-based inductively coupled plasma-optical emission spectrometry (ICP-OES) measurements.
The results reveal no apparent correlation with salinity (30.1–40.57 g kg−1) but a significant inverse correlation with temperature (−0.31±0.04 mmolmol−1∘C−1). Other marine aragonitic organisms such as Mytilus edulis (inner aragonitic shell portion) and Porites sp. exhibit similar results highlighting the consistency of the calculated CWC regressions. Corresponding Na∕Mg ratios show a similar temperature sensitivity to Na∕Ca ratios, but the combination of two ratios appears to reduce the impact of vital effects and domain-dependent geochemical variation. The high degree of scatter and elemental heterogeneities between the different skeletal features in both Na∕Ca and Na∕Mg, however, limit the use of these ratios as a proxy and/or make a high number of samples necessary. Additionally, we explore two models to explain the observed temperature sensitivity of Na∕Ca ratios for an open and semi-enclosed calcifying space based on temperature-sensitive Na- and Ca-pumping enzymes and transport proteins that change the composition of the calcifying fluid and consequently the skeletal Na∕Ca ratio.
Here we present a comprehensive attempt to correlate aragonitic Na / Ca ratios from Lophelia pertusa, Madrepora oculata and a caryophylliid cold-water coral (CWC) species with different seawater parameters such as temperature, salinity and pH. Living CWC specimens were collected from 16 different locations and analyzed for their Na / Ca content using solution-based inductively coupled plasma-optical emission spectrometry (ICP-OES) measurements. The results reveal no apparent correlation with salinity (30.1–40.57 g/kg) but a significant inverse correlation with temperature (−0.31 mmol/mol/°C). Other marine aragonitic organisms such as Mytilus edulis (inner aragonitic shell portion) and Porites sp. exhibit similar results highlighting the consistency of the calculated CWC regressions. Corresponding Na / Mg ratios show a similar temperature sensitivity to Na / Ca ratios, but the combination of two ratios appear to reduce the impact of vital effects and domain-dependent geochemical variation. The high degree of scatter and elemental heterogeneities between the different skeletal features in both Na / Ca and Na / Mg however limit the use of these ratios as a proxy and/or make a high number of samples necessary. Additionally, we explore two models to explain the observed temperature sensitivity of Na / Ca ratios for an open and semi-enclosed calcifying space based on temperature sensitive Na and Ca pumping enzymes and transport proteins that change the composition of the calcifying fluid and consequently the skeletal Na / Ca ratio.
Hyrrokkin sarcophaga is a parasitic foraminifera that is commonly found in cold-water coral reefs where it infests the file clam Acesta excavata and the scleractinian coral Desmophyllum pertusum (formerly known as Lophelia pertusa). Here, we present measurements of the trace element and isotopic composition of these parasitic foraminifera, analyzed by inductively coupled optical emission spectrometry (ICP-OES), electron probe microanalysis (EPMA) and mass spectrometry (gas-source MS and inductively-coupled-plasma MS). Our results reveal that the geochemical signature of H. sarcophaga depends on the host organism it infests. Sr / Ca ratios are 1.1 mmol mol−1 higher in H. sarcophaga that infest D. pertusum, which could be an indication that dissolved host carbonate material is utilized in shell calcification, given that the aragonite of D. pertusum has a naturally higher Sr concentration compared to the calcite of A. excavata. Similarly, we measure 3.1 ‰ lower δ13C and 0.25 ‰ lower δ18O values in H. sarcophaga that lived on D. pertusum, which might be caused by the direct uptake of the host's carbonate material with a more negative isotopic composition or different pH regimes in these foraminifera (pH can exert a control on the extent of CO2 hydration/hydroxylation) due to the uptake of body fluids of the host. We also observe higher Mn / Ca ratios in foraminifera that lived on A. excavata but did not penetrate the host shell compared to specimen that penetrated the shell, which could be interpreted as a change in food source, changes in the calcification rate, Rayleigh fractionation or changing oxygen conditions. While our measurements provide an interesting insight into the calcification process of this unusual foraminifera, these data also indicate that the geochemistry of this parasitic foraminifera is unlikely to be a reliable indicator of paleoenvironmental conditions using Sr / Ca, Mn / Ca, δ18O or δ13C unless the host organism is known and its geochemical composition can be accounted for.
Hyrrokkin sarcophaga is a parasitic foraminifer that is commonly found in cold-water coral reefs where it infests the file clam Acesta excavata and the scleractinian coral Lophelia pertusa. Here, we present measurements of the elemental and isotopic composition of this parasitic foraminifer for the first time, analyzed by inductively coupled optical emission spectrometry (ICP-OES), electron probe micro analysis (EPMA) and mass spectrometry (MS). Our results reveal that the geochemical signature of H. sarcophaga depends on the host organism it infests. Sr/Ca ratios are 1.1 mmol mol-1 higher in H. sarcophaga that infest L. pertusa, which could be an indication that dissolved host carbonate material is utilised in shell calcification, given that the aragonite of L. pertusa has a naturally higher Sr concentration compared to the calcite of A. excavata.Similarly, we measure 3.1 ‰ lower δ13C and 0.25 ‰ lower δ18O values in H. sarcophaga that lived on20 L. pertusa, which might be caused by the direct uptake of the host’s carbonate material with a more negative isotopic composition or different pH regimes in these foraminifera (pH can exert a control on the extent of CO2 hydration/hydroxylation) due to the uptake of body fluids of the host. We also observe higher Mn/Ca ratios in foraminifers that lived on A. excavata but did not penetrate the host shell compared to specimen that penetrated the shell, which could be interpreted as a change in food source, changes in the calcification rate, Rayleigh fractionation or changing oxygen conditions. While our measurements provide an interesting insight into the calcification process of this unusual foraminifer, these data also indicate that the geochemistry of this parasitic foraminifer is unlikely to be a reliable indicator of paleoenvironmental conditions using Sr/Ca, Mn/Ca, δ18O or δ13C unless the host organism is known and its geochemical composition can be accounted for.