Community structure and biodiversity in marine benthic communities during experiments, 2011

A mesocosm experiment was conducted to quantify the effects of reduced pH and elevated temperature on an intact marine invertebrate community. Standardised faunal communities, collected from the extreme low intertidal zone using artificial substrate units, were exposed to one of eight nominal treatments (four pH levels: 8.0, 7.7, 7.3 and 6.7, crossed with two temperature levels: 12 and 16°C). After 60 days exposure communities showed significant changes in structure and lower diversity in response to reduced pH. The response to temperature was more complex. At higher pH levels (8.0 and 7.7) elevated temperature treatments contained higher species abundances and diversity than the lower temperature treatments. In contrast, at lower pH levels (7.3 and 6.7), elevated temperature treatments had lower species abundances and diversity than lower temperature treatments. The species losses responsible for these changes in community structure and diversity were not randomly distributed across the different phyla examined. Molluscs showed the greatest reduction in abundance and diversity in response to low pH and elevated temperature, whilst annelid abundance and diversity was mostly unaffected by low pH and was higher at the elevated temperature. The arthropod response was between these two extremes with moderately reduced abundance and diversity at low pH and elevated temperature. Nematode abundance increased in response to low pH and elevated temperature, probably due to the reduction of ecological constraints, such as predation and competition, caused by a decrease in macrofaunal abundance. This community-based mesocosm study supports previous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical groups. This study also illustrates the importance of considering indirect effects that occur within multispecies assemblages when attempting to predict the consequences of ocean acidification and global warming on marine communities.

Global Ocean Particulate Organic Carbon flux merged with satellite parameters

The efficiency of the biological pump of carbon to the deep ocean depends largely on the biologically mediated export of carbon from the surface ocean and its remineralization with depth. Global satellite studies have primarily focused on chlorophyll concentration and net primary production (NPP) to understand the role of phytoplankton in these processes. Recent satellite retrievals of phytoplankton composition now allow for the size of phytoplankton cells to be considered. Here, we improve understanding of phytoplankton size structure impacts on particle export, remineralization and transfer. Particulate organic carbon (POC) flux observations from sediment traps and 234Th are compiled across the global ocean. Annual climatologies of NPP, percent microplankton, and POC flux at four time series locations and within biogeochemical provinces are constructed, and sinking velocities are calculated to align surface variables with POC flux at depth. Parameters that characterize POC flux vs. depth (export flux ratio, labile fraction, remineralization length scale) are then fit to the aligned dataset. Times of the year dominated by different size compositions are identified and fit separately in regions of the ocean where phytoplankton cell size showed enough dynamic range over the annual cycle. Considering all data together, our findings support the paradigm of high export flux but low transfer efficiency in more productive regions and vice versa for oligotrophic regions. However, when parsing by dominant size class, we find periods dominated by small cells to have both greater export flux and lower transfer efficiency than periods when large cells comprise a greater proportion of the phytoplankton community.

Carbon and deuterium isotopes and C ratios in gas hydrates of ODP Leg 204 sites

Sediments at the southern summit of Hydrate Ridge display two distinct modes of gas hydrate occurrence. The dominant mode is associated with active venting of gas exsolved from the accretionary prism and leads to high concentrations (15%-40% of pore space) of gas hydrate in seafloor or near-surface sediments at and around the topographic summit of southern Hydrate Ridge. These near-surface gas hydrates are mainly composed of previously buried microbial methane but also contain a significant (10%-15%) component of thermogenic hydrocarbons and are overprinted with microbial methane currently being generated in shallow sediments. Focused migration pathways with high gas saturation (>65%) abutting the base of gas hydrate stability create phase equilibrium conditions that permit the flow of a gas phase through the gas hydrate stability zone. Gas seepage at the summit supports rapid growth of gas hydrates and vigorous anaerobic methane oxidation. The other mode of gas hydrate occurs in slope basins and on the saddle north of the southern summit and consists of lower average concentrations (0.5%-5%) at greater depths (30-200 meters below seafloor [mbsf]) resulting from the buildup of in situ-generated dissolved microbial methane that reaches saturation levels with respect to gas hydrate stability at 30-50 mbsf. Net rates of sulfate reduction in the slope basin and ridge saddle sites estimated from curve fitting of concentration gradients are 2-4 mmol/m**3/yr, and integrated net rates are 20-50 mmol/m**2/yr. Modeled microbial methane production rates are initially 1.5 mmol/m**3/yr in sediments just beneath the sulfate reduction zone but rapidly decrease to rates of <0.1 mmol/m**3/yr at depths >100 mbsf. Integrated net rates of methane production in sediments away from the southern summit of Hydrate Ridge are 25-80 mmol/m**2/yr. Anaerobic methane oxidation is minor or absent in cored sediments away from the summit of southern Hydrate Ridge. Ethane-enriched Structure I gas hydrate solids are buried more rapidly than ethane-depleted dissolved gas in the pore water because of advection from compaction. With subsidence beneath the gas hydrate stability zone, the ethane (mainly of low-temperature thermogenic origin) is released back to the dissolved gas-free gas phases and produces a discontinuous decrease in the C1/C2 vs. depth trend. These ethane fractionation effects may be useful to recognize and estimate levels of gas hydrate occurrence in marine sediments.

Oxygen and carbon isotopes from benthic and planktonic foraminifers at DSDP Leg 74 Holes

Oxygen and carbon isotope measurements have been made in picked planktonic and benthonic foraminifers from the five sites drilled on Leg 74, covering the whole Cenozoic. For the Neogene, the coverage gives good information on the development of the vertical temperature structure of Atlantic deep water. For the Paleogene, vertical gradients were weak and it is possible to combine data from different sites to obtain a very detailed record of both the temperature and carbon isotope history of Atlantic deep waters.

Tab. 1: Chupa Inlet, Kandalaksha Bay of the White Sea. Population density of the dominant boreal copepod species and daily pellet carbon flux

Data on the zooplankton community structure, gut evacuation rate and carbon content of zooplankton faecal pellets were used for assessing the contribution of zooplankton to vertical carbon fluxes in the White and Kara Seas. The results revealed strong regional and seasonal variations of pellet carbon input related to differences in structure and dynamics of the zooplankton communities in the regions studied. In the deep regions of the White Sea, maximum daily pellet carbon flux from the 0-50 m layer was observed in the spring. It reached 98 mg Corg m-2 day-1 and coincided with a strong predominance of the large arctic herbivorous copepod Calanus glacialis in the surface layers. In summer and fall, it decreased by 1 to 2 orders of magnitude due to migration of this copepod to its overwintering depths. In contrast, in the shallow coastal regions, the pellet production was low in spring, gradually increased during summer and reached its maximum of 138 mg Corg m-2 day-1 by late summer to beginning of autumn. Such a seasonal pattern was in accordance with the seasonal variation of abundance of major pellet producers, the small boreal copepods Acartia bifilosa, Centropages hamatus, and Temora longicornis. In the estuarine zone of the Kara Sea, the pellet flux was mostly formed by pellets of brackish-water omnivorous copepods. It varied from 35 mg Corg m-2 day-1 in 1997 to 96 mg Corg m-2 day-1 in 1999. In the central Kara Sea with its typical marine community, the daily flux reached 125 mg Corg m-2 day-1 in summer. The results of our calculations indicate that both in the White and Kara seas zooplankton pellet carbon contributes up to 30 % to the total carbon flux during particular seasons.

Carbon cycling in the northern Humboldt Current off Chile

The structure of the zooplankton foodweb and their dominant carbon fluxes were studied in the upwelling system off northern Chile (Mejillones Bay; 23°S) between October 2000 and December 2002. High primary production (PP) rates (18 gC/m**2 d) were mostly due to the net-phytoplankton size fraction (>23 µm). High PP has been traditionally associated with the wind-driven upwelling fertilizing effect of equatorial subsurface waters, which favour development of a short food chain dominated by a few small clupeiform fish species. The objective of the present work was to study the trophic carbon flow through the first step of this 'classical chain' (from phytoplankton to primary consumers such as copepods and euphausiids) and the carbon flow towards the gelatinous web composed of both filter-feeding and carnivorous zooplankton. To accomplish this objective, feeding experiments with copepods, appendicularians, ctenophores, and chaetognaths were conducted using naturally occurring plankton prey assemblages. Throughout the study, the total carbon ingestion rates showed that the dominant appendicularian species and small copepods consumed an average of 7 and 5 µgC/ind d, respectively. In addition, copepods ingested particles mainly in the size range of nano- and microplankton, whereas appendicularians ingested in the range of pico- and nanoplankton. Small copepods and appendicularians removed a small fraction of total daily PP (range 6-11%). However, when the pico- + nanoplankton fractions were the major contributors to total PP (oligotrophic conditions), grazing by small copepods increased markedly to 86% of total PP. Under these more oligotrophic conditions, the euphausiids grazing increased as well, but only reached values lower than 5% of total PP. During this study, chaetognaths and ctenophores ingested an average of 1 and 14 copepods/ind d, respectively. In terms of biomass consumed, the potential impact of carnivorous gelatinous zooplankton on the small-size copepod community (preferred prey) was important (2-12% of biomass removed daily). However, their impact produced more significant results on copepod abundance (up to 33%), which suggests that carnivorous gelatinous zooplankton may even modulate (control) the abundance of some species as well as the size structure of the copepod community.

Effects of ocean acidification on population dynamics and community structure of crustose coralline algae

Calcification and growth of crustose coralline algae (CCA) are affected by elevated seawater pCO2 and associated changes in carbonate chemistry. However, the effects of ocean acidification (OA) on population and community-level responses of CCA have barely been investigated. We explored changes in community structure and population dynamics (size structure and reproduction) of CCA in response to OA. Recruited from an experimental flow-through system, CCA settled onto the walls of plastic aquaria and developed under exposure to one of three pCO2 treatments (control [present day, 389±6 ppm CO2], medium [753±11 ppm], and high [1267±19 ppm]). Elevated pCO2 reduced total CCA abundance and affected community structure, in particular the density of the dominant species Pneophyllum sp. and Porolithon onkodes. Meanwhile, the relative abundance of P. onkodes declined from 24% under control CO2 to 8.3% in high CO2 (65% change), while the relative abundance of Pneophyllum sp. remained constant. Population size structure of P. onkodes differed significantly across treatments, with fewer larger individuals under high CO2. In contrast, the population size structure and number of reproductive structures (conceptacles) per crust of Pneophyllum sp. was similar across treatments. The difference in the magnitude of the response of species abundance and population size structure between species may have the potential to induce species composition changes in the future. These results demonstrate that the impacts of OA on key coral reef builders go beyond declines in calcification and growth, and suggest important changes to aspects of population dynamics and community ecology.

Elevated CO2 levels do not affect the shell structure of the bivalve Arctica islandica from the western Baltic

Shells of the bivalve Arctica islandica are used to reconstruct paleo-environmental conditions (e.g. temperature) via biogeochemical proxies, i.e. biogenic components that are related closely to environmental parameters at the time of shell formation. Several studies have shown that proxies like element and isotope-ratios can be affected by shell growth and microstructure. Thus it is essential to evaluate the impact of changing environmental parameters such as high pCO2 and consequent changes in carbonate chemistry on shell properties to validate these biogeochemical proxies for a wider range of environmental conditions. Growth experiments with Arctica islandica from the Western Baltic Sea kept under different pCO2 levels (from 380 to 1120 µatm) indicate no affect of elevated pCO2 on shell growth or crystal microstructure, indicating that A. islandica shows an adaptation to a wider range of pCO2 levels than reported for other species. Accordingly, proxy information derived from A. islandica shells of this region contains no pCO2 related bias.

Stable carbon and oxygen isotope ratios of carbonates of ODP Site 103-639

A Tithonian sequence of shallow-water limestones, intercalated with siliciclastics and overlain by dolomite, was recovered during drilling at ODP Site 639 on the edge of a tilted fault block. The carbonates were strongly affected by fracturing, dolomitization, dedolomitization, and compaction. The chronology and nature of the fractures, fracture infilling, and diagenesis of the host rock are established and correlated for both the limestone and the dolomite. A first phase of dolomitization affected limestone that was already, at least partially, indurated. In the limestone unit, fractures were filled by calcite and dolomite; most of the dolomite was recrystallized into calcite, except for the upper part. In the dolomitic unit, the first-formed dolomite was progressively recrystallized into saddle dolomite, as fractures were simultaneously activated. The dolomitic textures become less magnesian (the molar ratio mMg/mCa goes from 1.04-0.98 to 0.80), and the d18O (PDB) ranges from -10 per mil to -8 per mil. The varying pores and fissures are either cemented by a calcic saddle dolomite (mMg/mCa ranging from 0.95 to 0.80) or filled with diverse internal sediments of detrital calcic dolomite, consisting of detrital dolomite silt (d18O from -9 per mil to -7 per mil) and laminated yellow filling (with different d18O values that range from -4 per mil to +3 per mil). These internal sediments clearly contain elements of the host rock and fragments of saddle crystals. They are covered by marls with calpionellids of early Valanginian age, which permits dating of most of the diagenetic phases as pre-Valanginian. The dolomitization appears to be related to fracturing resulting from extensional tectonics; it is also partially related to an erosional episode. Two models of dolomitization can be proposed from the petrographic characteristics and isotopic data. Early replacement of aragonite bioclasts by sparite, dissolution linked to dolomitization, and negative d18O values of dolomite suggest a freshwater influence and 'mixing zone' model. On the other hand, the significant presence of saddle dolomite and repeated negative d18O values suggest a temperature effect; because we can dismiss deep burial, hydrothermal formation of dolomite would be the most probable model. For both of these hypotheses, the vadose filling of cavities and fractures by silt suggests emersion, and the different, and even positive, d18O values of the last-formed yellow internal sediment could suggest dolomitization of the top of the sequence under saline to hypersaline conditions. Fracturing resulting in the reopening of porosity and the draining of dolomitizing fluids was linked to extensional tectonics prior to the tilting of the block. These features indicate an earlier beginning to the rifting of the Iberian margin than previously known. Dolomitization, emersion, and erosion correspond to eustatic sea-level lowering at the Berriasian/Valanginian boundary. Diagenesis, rather than sedimentation, seems to mark this global event and to provide a record of the regional tectonic history.

Freshwater mesocosm experiment 2012 in Vancouver, Canada on effects of increased temperature and food chain length on oxygen fluxes and plankton community structure

Over broad thermal gradients, the effect of temperature on aerobic respiration and photosynthesis rates explains variation in community structure and function. Yet for local communities, temperature dependent trophic interactions may dominate effects of warming. We tested the hypothesis that food chain length modifies the temperature-dependence of ecosystem fluxes and community structure. In a multi-generation aquatic food web experiment, increasing temperature strengthened a trophic cascade, altering the effect of temperature on estimated mass-corrected ecosystem fluxes. Compared to consumer-free and 3-level food chains, grazer-algae (2-level) food chains responded most strongly to the temperature gradient. Temperature altered community structure, shifting species composition and reducing zooplankton density and body size. Still, food chain length did not alter the temperature dependence of net ecosystem fluxes. We conclude that locally, food chain length interacts with temperature to modify community structure, but only temperature, not food chain length influenced net ecosystem fluxes.

Phytoplankton responses and associated carbon cycling during shipboard carbonate chemistry manipulation experiments conducted around Northwest European shelf seas

The ongoing oceanic uptake of anthropogenic carbon dioxide (CO2) is significantly altering the carbonate chemistry of seawater, a phenomenon referred to as ocean acidification. Experimental manipulations have been increasingly used to gauge how continued ocean acidification will potentially impact marine ecosystems and their associated biogeochemical cycles in the future; however, results amongst studies, particularly when performed on natural communities, are highly variable, which may reflect community/environment-specific responses or inconsistencies in experimental approach. To investigate the potential for identification of more generic responses and greater experimentally reproducibility, we devised and implemented a series (n = 8) of short-term (2-4 days) multi-level (>=4 conditions) carbonate chemistry/nutrient manipulation experiments on a range of natural microbial communities sampled in Northwest European shelf seas. Carbonate chemistry manipulations and resulting biological responses were found to be highly reproducible within individual experiments and to a lesser extent between geographically separated experiments. Statistically robust reproducible physiological responses of phytoplankton to increasing pCO2, characterised by a suppression of net growth for small-sized cells (<10 µm), were observed in the majority of the experiments, irrespective of natural or manipulated nutrient status. Remaining between-experiment variability was potentially linked to initial community structure and/or other site-specific environmental factors. Analysis of carbon cycling within the experiments revealed the expected increased sensitivity of carbonate chemistry to biological processes at higher pCO2 and hence lower buffer capacity. The results thus emphasise how biogeochemical feedbacks may be altered in the future ocean.