Small Changes in pH Have Direct Effects on Marine Bacterial Community Composition: A Microcosm Approach

As the atmospheric CO2 concentration rises, more CO2 will dissolve in the oceans, leading to a reduction in pH. Effects of ocean acidification on bacterial communities have mainly been studied in biologically complex systems, in which indirect effects, mediated through food web interactions, come into play. These approaches come close to nature but suffer from low replication and neglect seasonality. To comprehensively investigate direct pH effects, we conducted highly-replicated laboratory acidification experiments with the natural bacterial community from Helgoland Roads (North Sea). Seasonal variability was accounted for by repeating the experiment four times (spring, summer, autumn, winter). Three dilution approaches were used to select for different ecological strategies, i.e. fast-growing or low-nutrient adapted bacteria. The pH levels investigated were in situ seawater pH (8.15-8.22), pH 7.82 and pH 7.67, representing the present-day situation and two acidification scenarios projected for the North Sea for the year 2100. In all seasons, both automated ribosomal intergenic spacer analysis and 16S ribosomal amplicon pyrosequencing revealed pH-dependent community shifts for two of the dilution approaches. Bacteria susceptible to changes in pH were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. Their specific response to reduced pH was often context-dependent. Bacterial abundance was not influenced by pH. Our findings suggest that already moderate changes in pH have the potential to cause compositional shifts, depending on the community assembly and environmental factors. By identifying pH-susceptible groups, this study provides insights for more directed, in-depth community analyses in large-scale and long-term experiments.

Phytoplankton abundance measured on water samples from rivers of the Lena Delta in 2010

The Lena Delta in Northern Siberia is one of the largest river deltas in the world. During peak discharge, after the ice melt in spring, it delivers between 60-8000 m**3/s of water and sediment into the Arctic Ocean. The Lena Delta and the Laptev Sea coast also constitute a continuous permafrost region. Ongoing climate change, which is particularly pronounced in the Arctic, is leading to increased rates of permafrost thaw. This has already profoundly altered the discharge rates of the Lena River. But the chemistry of the river waters which are discharged into the coastal Laptev Sea have also been hypothesized to undergo considerable compositional changes, e.g. by increasing concentrations of inorganic nutrients such as dissolved organic carbon (DOC) and methane. These physical and chemical changes will also affect the composition of the phytoplankton communities. However, before potential consequences of climate change for coastal arctic phytoplankton communities can be judged, the inherent status of the diversity and food web interactions within the delta have to be established. In 2010, as part of the AWI Lena Delta programme, the phyto- and microzooplankton community in three river channels of the delta (Trofimov, Bykov and Olenek) as well as four coastal transects were investigated to capture the typical river phytoplankton communities and the transitional zone of brackish/marine conditions. Most CTD profiles from 23 coastal stations showed very strong stratification. The only exception to this was a small, shallow and mixed area running from the outflow of Bykov channel in a northerly direction parallel to the shore. Of the five stations in this area, three had a salinity of close to zero. Two further stations had salinities of around 2 and 5 throughout the water column. In the remaining transects, on the other hand, salinities varied between 5 and 30 with depth. Phytoplankton counts from the outflow from the Lena were dominated by diatoms (Aulacoseira species) cyanobacteria (Aphanizomenon, Pseudanabaena) and chlorophytes. In contrast, in the stratified stations the plankton was mostly dominated by dinoflagellates, ciliates and nanoflagellates, with only an insignificant diatom component from the genera Chaetoceros and Thalassiosira (brackish as opposed to freshwater species). Ciliate abundance was significantly coupled with the abundance of total flagellates. A pronounced partitioning in the phytoplankton community was also discernible with depth, with a different community composition and abundance above and below the thermocline in the stratified sites. This work is a first analysis of the phytoplankton community structure in the region where Lena River discharge enters the Laptev Sea.

Particle flux measured on deep sea sediment trap VP-2_trap (Appendix A2.7)

A 17 month record of vertical particle flux of dry weight, carbonate and organic carbon were 25.8, 9.4 and 2.4g/m**2/y, respectively. Parallel to trap deployments, pelagic system structure was recorded with high vertical and temporal resolution. Within a distinct seasonal cycle of vertical particle flux, zooplankton faecal pellets of various sizes, shapes and contents were collected by the traps in different proportions and quantities throughout the year (range: 0-4,500 10**3/m**2/d). The remains of different groups of organisms showed distinct seasonal variations in abundance. In early summer there was a small maximum in the diatom flux and this was followed by pulses of tinntinids, radiolarians, foraminiferans and pteropods between July and November. Food web interactions in the water column were important in controlling the quality and quantity of sinking materials. For example, changes in the population structure of dominant herbivores, the break-down of regenerating summer populations of microflagellates and protozooplankton and the collapse of a pteropod dominated community, each resulted in marked sedimentation pulses. These data from the Norwegian Sea indicate those mechanisms which either accelerate or counteract loss of material via sedimentation. These involve variations in the structure of the pelagic system and they operatè on long (e.g. annual plankton succession) and short (e.g. the end of new production, sporadic grazing of swarm feeders) time scales. Connecting investigation of the water column with a high resolution in time in parallel with drifting sediment trap deployments and shipboard experiments with the dominant zooplankters is a promising approach for giving a better understanding of both the origin and the fate of material sinking to the sea floor.

Data to investigate the role of Pomatoschistus microps in intertidal food webs, sampled off the island of Sylt

Ecological network analysis (ENA) was used to study the effects of Pomatoschistus microps on energy transport through the food web, its impact on other compartments and its possible role as a keystone species in the trophic webs of an Arenicola tidal flat ecosystem and a sparse Zostera noltii bed ecosystem. Three ENA models were constructed: (a) model 1 contains data of the original food web from prior research in the investigated area by Baird et al. (2007), (b) an updated model 2 which included biomass and diet data of P. microps from recent sampling, and (c) model 3 simulating a food web without P. microps. A comparison of energy transport between the different models revealed that more energy is transported from lower trophic levels up the food chain, in the presence of P. microps (models 1 and 2) than in its absence (model 3). Calculations of the keystone index (KSi) revealed the high overall impact (measured as eps_i) of this fish species on food webs. In model 1, P. microps was assigned a low KSi in the Arenicola flat and in the sparse Z. noltii bed. Calculations in model 2 ranked P. microps first for keystoneness and eps_i in both communities, the Arenicola flat and the sparse Z. noltii bed. Taken together, our results give insight into the role of P. microps when considering a whole food web and reveal direct and indirect trophic interactions of this small-sized fish species. These results might illustrate the impact and importance of abundant, widespread species in food webs and facilitate further investigations.

(Table A1) Species list of the high Antarctic Weddell Sea food web

Human-induced habitat destruction, overexploitation, introduction of alien species and climate change are causing species to go extinct at unprecedented rates, from local to global scales. There are growing concerns that these kinds of disturbances alter important functions of ecosystems. Our current understanding is that key parameters of a community (e.g. its functional diversity, species composition, and presence/absence of vulnerable species) reflect an ecological network's ability to resist or rebound from change in response to pressures and disturbances, such as species loss. If the food web structure is relatively simple, we can analyse the roles of different species interactions in determining how environmental impacts translate into species loss. However, when ecosystems harbour species-rich communities, as is the case in most natural systems, then the complex network of ecological interactions makes it a far more challenging task to perceive how species' functional roles influence the consequences of species loss. One approach to deal with such complexity is to focus on the functional traits of species in order to identify their respective roles: for instance, large species seem to be more susceptible to extinction than smaller species. Here, we introduce and analyse the marine food web from the high Antarctic Weddell Sea Shelf to illustrate the role of species traits in relation to network robustness of this complex food web. Our approach was threefold: firstly, we applied a new classification system to all species, grouping them by traits other than body size; secondly, we tested the relationship between body size and food web parameters within and across these groups and finally, we calculated food web robustness. We addressed questions regarding (i) patterns of species functional/trophic roles, (ii) relationships between species functional roles and body size and (iii) the role of species body size in terms of network robustness. Our results show that when analyzing relationships between trophic structure, body size and network structure, the diversity of predatory species types needs to be considered in future studies.

Experiment: Food availability outweighs ocean acidification effects in juvenile Mytilus edulis

Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless in many coastal areas high pCO2 variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12-20 g kg-1) and CO2 enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO2 variability in this habitat and the combined effects of elevated pCO2 and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO2 up to 3350 µatm. Kiel Fjord was characterized by strong seasonal pCO2 variability. During summer, maximal pCO2 values of 2500 µatm were observed at the surface and >3000 µatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO2 inner fjord field station (mean pCO2 ca. 1000 µatm) in comparison to a low pCO2 outer fjord station (ca. 600 µatm). In addition, mussels were able to outcompete the barnacle Amphibalanus improvisus at the high pCO2 site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO2. At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO2 when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.

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.

Nitrogen fuelling of the pelagic food web of the Tropical Atlantic

We estimated the relative contribution of atmospheric Nitrogen (N) input (wet and dry deposition and N fixation) to the epipelagic food web by measuring N isotopes of different functional groups of epipelagic zooplankton along 23°W (17°N-4°S) and 18°N (20-24°W) in the Eastern Tropical Atlantic. Results were related to water column observations of nutrient distribution and vertical diffusive flux as well as colony abundance of Trichodesmium obtained with an Underwater Vision Profiler (UVP5). The thickness and depth of the nitracline and phosphocline proved to be significant predictors of zooplankton stable N isotope values. Atmospheric N input was highest (61% of total N) in the strongly stratified and oligotrophic region between 3 and 7°N, which featured very high depth-integrated Trichodesmium abundance (up to 9.4×104 colonies m-2), strong thermohaline stratification and low zooplankton delta15N (~2 per mil). Relative atmospheric N input was lowest south of the equatorial upwelling between 3 and 5°S (27%). Values in the Guinea Dome region and north of Cape Verde ranged between 45 and 50%, respectively. The microstructure-derived estimate of the vertical diffusive N flux in the equatorial region was about one order of magnitude higher than in any other area (approximately 8 mmol m-2 d 1). At the same time, this region received considerable atmospheric N input (35% of total). In general, zooplankton delta15N and Trichodesmium abundance were closely correlated, indicating that N fixation is the major source of atmospheric N input. Although Trichodesmium is not the only N fixing organism, its abundance can be used with high confidence to estimate the relative atmospheric N input in the tropical Atlantic (r2 = 0.95). Estimates of absolute N fixation rates are two- to tenfold higher than incubation-derived rates reported for the same regions. Our approach integrates over large spatial and temporal scales and also quantifies fixed N released as dissolved inorganic and organic N. In a global analysis, it may thus help to close the gap in oceanic N budgets.