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Seawater carbonate chemistry and in situ and laboratory measurements of calcification

Ocean acidification (OA) is generally assumed to negatively impact calcification rates of marine organisms. At a local scale however, biological activity of macrophytes may generate pH fluctuations with rates of change that are orders of magnitude larger than the long-term trend predicted for the open ocean. These fluctuations may in turn impact benthic calcifiers in the vicinity. Combining laboratory, mesocosm and field studies, such interactions between OA, the brown alga Fucus vesiculosus, the sea grass Zostera marina and the blue mussel Mytilus edulis were investigated at spatial scales from decimetres to 100s of meters in the western Baltic. Macrophytes increased the overall mean pH of the habitat by up to 0.3 units relative to macrophyte- free, but otherwise similar, habitats and imposed diurnal pH fluctuations with amplitudes ranging from 0.3 to more than 1 pH unit. These amplitudes and their impact on mussel calcification tended to increase with increasing macrophyte biomass to bulk water ratio. At the laboratory and mesocosm scales, biogenic pH fluc- tuations allowed mussels to maintain calcification even under acidified conditions by shifting most of their calcification activity into the daytime when biogenic fluctuations caused by macrophyte activity offered temporal refuge from OA stress. In natural habitats with a low biomass to water body ratio, the impact of biogenic pH fluctuations on mean calcification rates of M. edulis was less pronounced. Thus, in dense algae or seagrass habitats, macrophytes may mitigate OA impact on mussel calcification by raising mean pH and providing temporal refuge from acidification stress.

Seawater carbonate chemistry and benthic foraminifera Ammonia sp. mass, size, and growth rate during experiments, 2013

About 30% of the anthropogenically released CO2 is taken up by the oceans; such uptake causes surface ocean pH to decrease and is commonly referred to as ocean acidification (OA). Foraminifera are one of the most abundant groups of marine calcifiers, estimated to precipitate ca. 50 % of biogenic calcium carbonate in the open oceans. We have compiled the state of the art literature on OA effects on foraminifera, because the majority of OA research on this group was published within the last three years. Disparate responses of this important group of marine calcifiers to OA were reported, highlighting the importance of a process-based understanding of OA effects on foraminifera. We cultured the benthic foraminifer Ammonia sp. under a range of carbonate chemistry manipulation treatments to identify the parameter of the carbonate system causing the observed effects. This parameter identification is the first step towards a process-based understanding. We argue that CO3 is the parameter affecting foraminiferal size-normalized weights (SNWs) and growth rates. Based on the presented data, we can confirm the strong potential of Ammonia sp. foraminiferal SNW as a CO3 proxy.

Schwerpunktprogramm (SPP) 527: Bereich Infrastruktur - International Ocean Discovery Program, Teilprojekt: Terrestrische Ökosystem- und Klimadynamik in SE-Afrika für den Zeitraum der Entwicklung, des Vorkommens und des Aussterbens archaischer Hominiden (4-2 Ma) basierend auf Kernmaterial von IODP-Expedition 361

Das beantragte Projekt hat zum Ziel, die terrestrische Ökosystem- und Klimadynamik - und damit die naturräumlichen Rahmenbedingungen für die Evolution früher Hominiden - in SE-Afrika während des 'mittleren' Pliozäns und frühen Pleistozäns zu rekonstruieren. Um dieses Ziel zu erreichen, soll an Hand von Kernmaterial von IODP Expedition 361 ('Southern African Climates') eine Land/Meer-Korrelation vor SE-Afrika erarbeitet werden; diese wird die erste kontinuierliche Rekonstruktion der terrestrischen Ökosystem- und Klimaänderungen in SE-Afrika während des 'mittleren' Pliozäns bis frühen Pleistozäns liefern. Methodisch basiert das Projekt auf einem integrierten Ansatz, der palynologische (Pollen und Sporen) und elementgeochemische (XRF-Scanning) Analysen vereint und auf den Splice von IODP-Site U1478 (Straße von Mosambik) angewendet werden soll. Eine präzise Alterskontrolle wird durch die hochauflösende Benthos-Sauerstoffisotopenstratigraphie ermöglicht, die aktuell für Site U1478 erarbeitet wird. Site U1478 ist für die hier vorgeschlagenen Untersuchungen aus einer Reihe von Gründen ideal geeignet: (i) er stellt ein stratigraphisch außergewöhnlich vollständiges Archiv dar und verfügt dabei über hohe Sedimentationsraten; (ii) seine proximale Lage in Bezug auf das Limpopo-Delta gewährleistet einen hohen Anteil terrigenen Inputs im Kernmaterial; (iii) die Ursprungsregion dieser terrigenen Komponenten lässt sich hervorragend eingrenzen; (iv) er ist gegenüber terrestrischen Klimaänderungen hoch empfindlich, wie frühere Studien an nahe gelegenen Kurzkernen belegen; (v) die für das vorgeschlagene Projekt durchgeführten Pilotstudien an Kernfänger-Material belegen, dass seine Sedimente in Bezug auf Pollen und Sporen extrem produktiv sind; er liegt in einer proximal Position hinsichtlich der paläoanthropologischen 'Cradle of Humankind'-Fundstätten in Südafrika. Unter Berücksichtigung des gegenwärtigen Forschungsstandes zur Evolution archaischer Hominiden (insbesondere Australopithecus africanus) fokussiert das Projekt auf den Zeitraum zwischen 4 und 2 Ma; kritische Intervalle der Evolution archaischer Hominiden sollen in besonders hoher zeitlicher Auflösung analysiert werden. Die Integration der palynologischen und elementgeochemischen Proxy-Daten wird detaillierte Aussagen zum Charakter und Zeitpunkt wie auch zur Stärke und Geschwindigkeit der Ökosystem- und Klimavariabilität im Einzugsgebiet des Limpopo River und damit in der 'Cradle of Humankind'-Region erlauben. Dadurch wird nicht nur die Klärung der Frage möglich, inwiefern Intervalle mit besonders ausgeprägtem Umweltwandel tatsächlich mit Schritten in der Hominiden-Evolution einhergehen, sondern es lassen sich auch die einzelnen Komponenten dieses Umweltwandels identifizieren. Diese Informationen können neues Licht auf die aktuelle Diskussion um potenzielle kausale Zusammenhänge zwischen Umwelt-'Forcing' und menschlicher Evolution werfen.

Raw data of physical oceanography during RV HEINCKE cruise HE670

Raw physical oceanography data was acquired by a ship-based Seabird SBE911plus CTD-Rosette system onboard RV HEINCKE. The CTD was equipped with duplicate sensors for temperature (SBE3plus), conductivity (SBE4) and oxygen (SBE43) as well as one sensor for pressure (SBE9plus). Additional sensors such as a WET Labs C-Star transmissometer, a WET Labs ECO-AFL fluorometer (FLRTD) and an altimeter (Teledyne Benthos PSA-916) were mounted to the CTD. The data was recorded using pre-cruise calibration coefficients. No correction, post-cruise calibration or quality control was applied. Processed profile data are available via the link below (Other version).

Physical oceanography during RV HEINCKE cruise HE672

Conductivity-temperature-depth profiles were measured using a Seabird SBE 911plus CTD during RV HEINCKE cruise HE672. The CTD was equipped with duplicate sensors for temperature (SBE3plus), conductivity (SBE4) and oxygen (SBE43) as well as one sensor for pressure (SBE9plus). Additional sensors such as a WET Labs C-Star transmissometer, a WET Labs ECO-AFL fluorometer and an altimeter (PSA-916 Teledyne (Benthos)) were mounted to the CTD. Temperature, conductivity and oxygen sensors are calibrated by the manufacturer once a year before being mounted in January. They are used throughout the year and no post-cruise or in-situ calibration is applied. All other sensors are calibrated irregularly. Data were connected to the station book of the specific cruise as available in the DSHIP database. Processing of the data including removal of obvious outliers followed the procedures described in CTD Processing Logbook of RV HEINCKE (hdl:10013/epic.47427). The processing report for this dataset is linked below.

Physical oceanography during RV HEINCKE cruise HE655/2

Conductivity-temperature-depth profiles were measured using a Seabird SBE 911plus CTD during RV HEINCKE cruise HE655/2. The CTD was equipped with duplicate sensors for temperature (SBE3plus), conductivity (SBE4) and oxygen (SBE43) as well as one sensor for pressure (SBE9plus). Additional sensors such as a WET Labs C-Star transmissometer, a WET Labs ECO-AFL fluorometer and an altimeter (PSA-916 Teledyne (Benthos)) were mounted to the CTD. Temperature, conductivity and oxygen sensors are calibrated by the manufacturer once a year before being mounted in January. They are used throughout the year and no post-cruise or in-situ calibration is applied. All other sensors are calibrated irregularly. Data were connected to the station book of the specific cruise as available in the DSHIP database. Processing of the data including removal of obvious outliers followed the procedures described in CTD Processing Logbook of RV HEINCKE (hdl:10013/epic.47427). The processing report for this dataset is linked below.

Physical oceanography during RV HEINCKE cruise HE660

Conductivity-temperature-depth profiles were measured using a Seabird SBE 911plus CTD during RV HEINCKE cruise HE660. The CTD was equipped with duplicate sensors for temperature (SBE3plus), conductivity (SBE4) and oxygen (SBE43) as well as one sensor for pressure (SBE9plus). Additional sensors such as a WET Labs C-Star transmissometer, a WET Labs ECO-AFL fluorometer and an altimeter (PSA-916 Teledyne (Benthos)) were mounted to the CTD. Temperature, conductivity and oxygen sensors are calibrated by the manufacturer once a year before being mounted in January. They are used throughout the year and no post-cruise or in-situ calibration is applied. All other sensors are calibrated irregularly. Data were connected to the station book of the specific cruise as available in the DSHIP database. Processing of the data including removal of obvious outliers followed the procedures described in CTD Processing Logbook of RV HEINCKE (hdl:10013/epic.47427). The processing report for this dataset is linked below.

Seawater carbonate chemistry and biometry and dissolution features of the benthic foraminifer Ammonia aomoriensis in a laboratory experiment

Culturing experiments were performed with the benthic foraminifer Ammonia aomoriensis from Flensburg Fjord, western Baltic Sea. The experiments simulated a projected rise in atmospheric CO2 concentrations. We exposed specimens to 5 seawater pCO2 levels ranging from 618 µatm (pH 7.9) to 3130 µatm (pH 7.2) for 6 wk. Growth rates and mortality differed significantly among pCO2 treatments. The highest increase of mean test diameter (19%) was observed at 618 µatm. At partial pressures >1829 µatm, the mean test diameter was observed to decrease, by up to 22% at 3130 µatm. At pCO2 levels of 618 and 751 µatm, A. aomoriensis tests were found intact after the experiment. The outer chambers of specimens incubated at 929 and 1829 µatm were severely damaged by corrosion. Visual inspection of specimens incubated at 3130 µatm revealed wall dissolution of all outer chambers, only their inner organic lining stayed intact. Our results demonstrate that pCO2 values of >=929 µatm in Baltic Sea waters cause reduced growth of A. aomoriensis and lead to shell dissolution. The bottom waters in Flensburg Fjord and adjacent areas regularly experience pCO2 levels in this range during summer and fall. Increasing atmospheric CO2 concentrations are likely to extend and intensify these periods of undersaturation. This may eventually slow down calcification in A. aomoriensis to the extent that net carbonate precipitation terminates. The possible disappearance of this species from the Baltic Sea and other areas prone to seasonal undersaturation would likely cause significant shifts in shallow-water benthic ecosystems in the near future.

Seawater carbonate chemistry and growth and carbon metabolism of the seaweed Fucus vesiculosus in the western Baltic Sea

Warming and acidification of the oceans as a consequence of increasing CO2-concentrations occur at large scales. Numerous studies have shown the impact of single stressors on individual species. However, studies on the combined effect of multiple stressors on a multi-species assemblage, which is ecologically much more realistic and relevant, are still scarce. Therefore, we orthogonally crossed the two factors warming and acidification in mesocosm experiments and studied their single and combined impact on the brown alga Fucus vesiculosus associated with its natural community (epiphytes and mesograzers) in the Baltic Sea in all seasons (from April 2013 to April 2014). We superimposed our treatment factors onto the natural fluctuations of all environmental variables present in the Benthocosms in so-called delta-treatments. Thereby we compared the physiological responses of F. vesiculosus (growth and metabolites) to the single and combined effects of natural Kiel Fjord temperatures and pCO2 conditions with a 5 °C temperature increase and/or pCO2 increase treatment (1100 ppm in the headspace above the mesocosms). Responses were also related to the factor photoperiod which changes over the course of the year. Our results demonstrate complex seasonal pattern. Elevated pCO2 positively affected growth of F. vesiculosus alone and/or interactively with warming. The response direction (additive, synergistic or antagonistic), however, depended on season and daylength. The effects were most obvious when plants were actively growing during spring and early summer. Our study revealed for the first time that it is crucial to always consider the impact of variable environmental conditions throughout all seasons. In summary, our study indicates that in future F. vesiculosus will be more affected by detrimental summer heat-waves than by ocean acidification although the latter consequently enhances growth throughout the year. The mainly negative influence of rising temperatures on the physiology of this keystone macroalga may alter and/or hamper its ecological functions in the shallow coastal ecosystem of the Baltic Sea.

Response of benthic foraminifera to ocean acidification in their natural sediment environment: a long-term culturing experiment

Calcifying foraminifera are expected to be endangered by ocean acidification; however, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment with pCO2-enriched seawater of 430, 907, 1865 and 3247 µatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Omega calc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size-frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 µm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 µm (pCO2 from 430 to 1865 µatm) to 258 µm (pCO2 3247 µatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g/m**2/a, whereas dead empty tests accumulated a rate of 0.27 g /m**2/a. Although Omega calc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 µatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

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