Increasing atmospheric CO2 concentrations are expected to impact pelagic ecosystem functioning in the near future by driving ocean warming and acidification. While numerous studies have investigated impacts of rising temperature and seawater acidification on planktonic organisms separately, little is presently known on their combined effects. To test for possible synergistic effects we exposed two coccolithophore species, Emiliania huxleyi and Gephyrocapsa oceanica, to a CO2 gradient ranging from ∼0.5–250 µmol kg−1 (i.e. ∼20–6000 µatm pCO2) at three different temperatures (i.e. 10, 15, 20°C for E. huxleyi and 15, 20, 25°C for G. oceanica). Both species showed CO2-dependent optimum-curve responses for growth, photosynthesis and calcification rates at all temperatures. Increased temperature generally enhanced growth and production rates and modified sensitivities of metabolic processes to increasing CO2. CO2 optimum concentrations for growth, calcification, and organic carbon fixation rates were only marginally influenced from low to intermediate temperatures. However, there was a clear optimum shift towards higher CO2 concentrations from intermediate to high temperatures in both species. Our results demonstrate that the CO2 concentration where optimum growth, calcification and carbon fixation rates occur is modulated by temperature. Thus, the response of a coccolithophore strain to ocean acidification at a given temperature can be negative, neutral or positive depending on that strain's temperature optimum. This emphasizes that the cellular responses of coccolithophores to ocean acidification can only be judged accurately when interpreted in the proper eco-physiological context of a given strain or species. Addressing the synergistic effects of changing carbonate chemistry and temperature is an essential step when assessing the success of coccolithophores in the future ocean.

Thousands of moribund thaliacean carcasses (Pyrosoma atlanticum) were deposited between February and March 2006 at the seafloor in the Ivory Coast area (West Africa). Remotely operated vehicle surveys were conducted in a continuous depth gradient between 20 and 1275 m along an oil pipeline. Video and still photography were used to estimate the carcass distribution, density, and size on the seabed, as well as recording the local megafauna interactions with the gelatinous material. Large patches of dead pyrosomids covered extensive areas on the continental slope, whereas minor aggregations were found on the shelf. The carcasses were in many instances trapped along the pipelines, accumulating extensively in troughs and furrows in the slope, and especially in soft sediment channels. Pyrosoma atlanticum dried samples were used to calculate the carbon content, enabling the extrapolation to the densities and sizes recorded in the video surveys. The average standing stock of organic carbon associated with the carcasses was .5 g C m 22 in the whole slope and canyon, with values as high as 22 g C m 22 in certain areas. Eight megafaunal species from three different phyla were observed 63 times directly feeding on the decomposing carcasses. The gelatinous carbon may have contributed substantially to the detrital food web including microbes at the seabed, and certainly to the diet of larger benthic organisms. The organism-level carbon measurements and documented fate of pyrosomid organic carbon is new evidence of the importance of gelatinous material in large-scale processes and elemental cycling.

In situ coral calcification was primarily controlled by temperature and relatively insensitive to seawater CO2 chemistry.

Sinking of gelatinous zooplankton biomass is an important component of the biological pump removing carbon from the upper ocean. The export efficiency, e.g., how much biomass reaches the ocean interior sequestering carbon, is poorly known because of the absence of reliable sinking speed data. We measured sinking rates of gelatinous particulate organic matter (jelly-POM) from different species of scyphozoans, ctenophores, thaliaceans, and pteropods, both in the field and in the laboratory in vertical columns filled with seawater using high-quality video. Using these data, we determined taxon-specific jelly-POM export efficiencies using equations that integrate biomass decay rate, seawater temperature, and sinking speed. Two depth scenarios in several environments were considered, with jelly-POM sinking from 200 and 600 m in temperate, tropical, and polar regions. Jelly-POM sank on average between 850 and 1500 m d 21 (salps: 800-1200 m d 21 ; ctenophores: 1200-1500 m d 21 ; scyphozoans: 1000-1100 m d 21 ; pyrosomes: 1300 m d 21 ). High latitudes represent a fast-sinking and low-remineralization corridor, regardless of species. In tropical and temperate regions, significant decomposition takes place above 1500 m unless jelly-POM sinks below the permanent thermocline. Sinking jelly-POM sequesters carbon to the deep ocean faster than anticipated, and should be incorporated into biogeochemical and modeling studies to provide more realistic quantification of export via the biological carbon pump worldwide.

Abstract. The contribution of carbonate-producing benthic organisms to the global marine carbon budget has been overlooked, the prevailing view being that calcium carbonate (CaCO 3 ) is predominantly produced and exported by marine plankton in the ''biological pump.'' Here, we provide the first estimation of the global contribution of echinoderms to the marine inorganic and organic carbon cycle, based on organism-level measurements from species of the five echinoderm classes. Echinoderms' global CaCO 3 contribution amounts to ;0.861 Pg CaCO 3 /yr (0.102 Pg C/yr of inorganic carbon) as a production rate, and ;2.11 Pg CaCO 3 (0.25 Pg C of inorganic carbon) as a standing stock from the shelves, slopes, and abyssal depths. Echinoderm inorganic carbon production (0.102 Pg C/yr) is less than the global pelagic production (0.4-1.8 Pg C/yr) and similar to the estimates for carbonate shelves globally (0.024-0.120 Pg C/yr Áyr À1 as inorganic carbon) for the slope on a global scale. The biogeography of the CaCO 3 standing stocks of echinoderms showed strong latitudinal variability. More than 80% of the global CaCO 3 production from echinoderms occurs between 0 and 800 m, with the highest contribution attributed to the shelf and upper slope. We provide a global distribution of echinoderm populations in the context of global calcite saturation horizons, since undersaturated waters with respect to mineral phases are surfacing. This shallowing is a direct consequence of ocean acidification, and in some places it may reach the shelf and upper slope permanently, where the highest CaCO 3 standing stocks from echinoderms originate. These organism-level data contribute substantially to the assessment of global carbonate inventories, which at present are poorly estimated. Additionally, it is desirable to include these benthic compartments in coupled global biogeochemical models representing the ''biological pump'' and its feedbacks, since at present all efforts have focused on pelagic processes, dominated by coccolithophores. The omission of the benthic processes from modeling will only diminish the understanding of elemental fluxes at large scales and any future prediction of climate change scenarios.

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Gelatinous zooplankton (Cnidaria, Ctenophora, and Urochordata, namely, Thaliacea) are ubiquitous members of plankton communities linking primary production to higher trophic levels and the deep ocean by serving as food and transferring “jelly‐carbon” (jelly‐C) upon bloom collapse. Global biomass within the upper 200 m reaches 0.038 Pg C, which, with a 2–12 months life span, serves as the lower limit for annual jelly‐C production. Using over 90,000 data points from 1934 to 2011 from the Jellyfish Database Initiative as an indication of global biomass (JeDI: http://jedi.nceas.ucsb.edu, http://www.bco-dmo.org/dataset/526852), upper ocean jelly‐C biomass and production estimates, organism vertical migration, jelly‐C sinking rates, and water column temperature profiles from GLODAPv2, we quantitatively estimate jelly‐C transfer efficiency based on Longhurst Provinces. From the upper 200 m production estimate of 0.038 Pg C year−1, 59–72% reaches 500 m, 46–54% reaches 1,000 m, 43–48% reaches 2,000 m, 32–40% reaches 3,000 m, and 25–33% reaches 4,500 m. This translates into ~0.03, 0.02, 0.01, and 0.01 Pg C year−1, transferred down to 500, 1,000, 2,000, and 4,500 m, respectively. Jelly‐C fluxes and transfer efficiencies can occasionally exceed phytodetrital‐based sediment trap estimates in localized open ocean and continental shelves areas under large gelatinous blooms or jelly‐C mass deposition events, but this remains ephemeral and transient in nature. This transfer of fast and permanently exported carbon reaching the ocean interior via jelly‐C constitutes an important component of the global biological soft‐tissue pump, and should be addressed in ocean biogeochemical models, in particular, at the local and regional scale.