Macroalgal communities of gas vents near Ischia island in the Medterranean sea, 2011

There are high levels of uncertainty about how coastal ecosystems will be affected by rapid ocean acidification caused by anthropogenic CO2, due to a lack of data. The few experiments to date have been short-term (< 1 year) and reveal mixed responses depending on the species examined and the culture conditions used. It is difficult to carry out long-term manipulations of CO2 levels, therefore areas with naturally high CO2 levels are being used to help understand which species, habitats and processes are resilient to the effects of ocean acidification, and which are adversely affected. Here we describe the effects of increasing CO2 levels on macroalgal communities along a pH gradient caused by volcanic vents. Macroalgal habitat differed at taxonomic and morphological group levels along a pH gradient. The vast majority of the 101 macroalgal species studied were able to grow with only a 5% decrease in species richness as the mean pH fell from 8.1 to 7.8. However, this small fall in species richness was associated with shifts in community structure as the cover of turf algae decreased disproportionately. Calcitic species were significantly reduced in cover and species richness whereas a few non-calcified species became dominant. At mean pH 6.7, where carbonate saturation levels were < 1, calcareous species were absent and there was a 72% fall in species richness. Under these extremely high CO2 conditions a few species dominated the simplified macroalgal assemblage and a very few exhibited enhanced reproduction, although high CO2 levels seemed to inhibit reproduction in others. Our data show that many macroalgal species are tolerant of long-term elevations in CO2 levels but that macroalgal habitats are altered significantly as pH drops, contributing to a scant but growing body of evidence concerning the long-term effects of CO2 emissions in vegetated marine systems. Further study is now needed to investigate whether the observed response of macroalgal communities can be replicated in different seasons and from a range of geographical regions for incorporation into global modelling studies to predict effects of CO2 emissions on Earth's ecosystems.

Seasonality affects macroalgal community response to increases in pCO2

Ocean acidification is expected to alter marine systems, but there is uncertainty about its effects due to the logistical difficulties of testing its large-scale and long-term effects. Responses of biological communities to increases in carbon dioxide can be assessed at CO2 seeps that cause chronic exposure to lower seawater pH over localised areas of seabed. Shifts in macroalgal communities have been described at temperate and tropical pCO2 seeps, but temporal and spatial replication of these observations is needed to strengthen confidence our predictions, especially because very few studies have been replicated between seasons. Here we describe the seawater chemistry and seasonal variability of macroalgal communities at CO2 seeps off Methana (Aegean Sea). Monitoring from 2011 to 2013 showed that seawater pH decreased to levels predicted for the end of this century at the seep site with no confounding gradients in Total Alkalinity, salinity, temperature or wave exposure. Most nutrient levels were similar along the pH gradient; silicate increased significantly with decreasing pH, but it was not limiting for algal growth at all sites. Metal concentrations in seaweed tissues varied between sites but did not consistently increase with pCO2. Our data on the flora are consistent with results from laboratory experiments and observations at Mediterranean CO2 seep sites in that benthic communities decreased in calcifying algal cover and increased in brown algal cover with increasing pCO2. This differs from the typical macroalgal community response to stress, which is a decrease in perennial brown algae and proliferation of opportunistic green algae. Cystoseira corniculata was more abundant in autumn and Sargassum vulgare in spring, whereas the articulated coralline alga Jania rubens was more abundant at reference sites in autumn. Diversity decreased with increasing CO2 regardless of season. Our results show that benthic community responses to ocean acidification are strongly affected by season.

(Table 1) Presence of sublittoral macroalgae in the investigated profiles in Potter Cove, King George Island, Antarctica

The vegetation of a small fjord and its adjacent open shore was documented by subaquatic video. The distribution of individual species of macroalgae and the composition of assemblages were compared with gradients of light availability, hydrography, slope inclination, substratum, and exposition to turbulence and ice. The sublittoral fringe is usually abraded by winterly ice floes and devoid of large, perennial algae. Below this zone, the upper sublittoral is dominated by Desmarestia menziesii on steep rock faces, where water movements become irregular, or by Ascoseira mirabilis and Palmaria decipiens on weakly inclined slopes with steady rolling water movements. In the central sublittoral above 15 m, where turbulence is still active, Desmarestia anceps is outcompeting all other species on solid substratum, However, the species is not able to persist on loose material under these conditions. Instead, Himantothallus grandifolius may occur. Deeper, where turbulence usually is negligible, Desmarestia anceps also covers loose material. The change of dominance to Himantothallus grandifolius in the deep sublittoral cannot completely be explained at present. Himantothallus grandifolius also prevails in a mixed assemblage under the influence of grounding icebergs. Most of the smaller algae are opportunists with different degrees of tolerance for turbulence, but some apparently need more stable microhabitats and thus are dependent from continuing suppression of competitive large phaeophytes.

Spatial community shift from hard to soft corals in acidified water

Anthropogenic increases in the partial pressure of CO2 (pCO2) cause ocean acidification, declining calcium carbonate saturation states, reduced coral reef calcification and changes in the compositions of marine communities. Most projected community changes due to ocean acidification describe transitions from hard coral to non-calcifying macroalgal communities; other organisms have received less attention, despite the biotic diversity of coral reef communities. We show that the spatial distributions of both hard and soft coral communities in volcanically acidified, semi-enclosed waters off Iwotorishima Island, Japan, are related to pCO2 levels. Hard corals are restricted to non-acidified low- pCO2 (225 µatm) zones, dense populations of the soft coral Sarcophyton elegans dominate medium- pCO2 (831 µatm) zones, and both hard and soft corals are absent from the highest- pCO2 (1,465 µatm) zone. In CO2-enriched culture experiments, high- pCO2 conditions benefited Sarcophyton elegans by enhancing photosynthesis rates and did not affect light calcification, but dark decalcification (negative net calcification) increased with increasing pCO2. These results suggest that reef communities may shift from reef-building hard corals to non-reef-building soft corals under pCO2 levels (550-970 µatm) predicted by the end of this century, and that higher pCO2 levels would challenge the survival of some reef organisms.

Experiment: Elevated CO2 levels affect the activity of nitrate reductase and carbonic anhydrase in the calcifying rhodophyte Corallina officinalis

The concentration of CO2 in global surface ocean waters is increasing due to rising atmospheric CO2 emissions, resulting in lower pH and a lower saturation state of carbonate ions. Such changes in seawater chemistry are expected to impact calcification in calcifying marine organisms. However, other physiological processes related to calcification might also be affected, including enzyme activity. In a mesocosm experiment, macroalgal communities were exposed to three CO2 concentrations (380, 665, and 1486 µatm) to determine how the activity of two enzymes related to inorganic carbon uptake and nutrient assimilation in Corallina officinalis, an abundant calcifying rhodophyte, will be affected by elevated CO2 concentrations. The activity of external carbonic anhydrase, an important enzyme functioning in macroalgal carbon-concentrating mechanisms, was inversely related to CO2 concentration after long-term exposure (12 weeks). Nitrate reductase, the enzyme responsible for reduction of nitrate to nitrite, was stimulated by CO2 and was highest in algae grown at 665 µatm CO2. Nitrate and phosphate uptake rates were inversely related to CO2, while ammonium uptake was unaffected, and the percentage of inorganic carbon in the algal skeleton decreased with increasing CO2. The results indicate that the processes of inorganic carbon and nutrient uptake and assimilation are affected by elevated CO2 due to changes in enzyme activity, which change the energy balance and physiological status of C. officinalis, therefore affecting its competitive interactions with other macroalgae. The ecological implications of the physiological changes in C. officinalis in response to elevated CO2 are discussed.

Biogenic and environmental fluctuations of temperature and pH in a novel mesocosm concept ("Kiel Outdoor Benthocosms") April 2013 to September 2014

The Kiel Outdoor Benthocosm infrastructure (Kiel, Germany,N 54°19.8'; E 010°09.0') allows combining natural in-situ fluctuations on all environmental variables with the controlled manipulation of treatment factors. The environmental fluctuations are admitted by a continuous flow-through of fjord water. The treatment is applied by delta-treatments which shift the mean of target variables (temperature and pH in this case) while maintaining the frequency and amplitude of natural fluctuations. The data presented here show the treatment levels and the continuously logged temperature and pH conditions in the experimental tanks. The dynamics of temperature and pH are driven by (i) in situ variability, (ii) the treatments imposed and (iii) the biology of the biota in the tanks. These contained macroalgal communities with associated mesograzers, mussels, and sea stars. The data set comprised 5 experimental runs: spring experiment (4.4.-19.6.2013), summer experiment 1 (4.7.-17.9.2013), autumn experiment (10.10-17.12.2013), winter experiment (16.1. - 1.4.2014), summer experiment 2 (10.7. - 26.9.2014).

Species name, classification (WORMS), abundance and biomass of the megabenthic communities of the Algarve

The present study analysed the megabenthic diversity in subtidal soft bottoms and assessed the main environmental drivers of megabenthic community organisation along the Algarve coast (southern Portugal). We tested the hypothesis that megabenthic communities respond to the same environmental drivers than macrofauna. We found that similar to macrofauna, megafaunal communities were organised in relation to the depth of closure, light reaching the bottom, and the hydrodynamic conditions related with exposure within the shallower areas. The influence of the main river outflow prevailed over other drivers, but only up to 9 m depth. We found that seven different spatial units should be considered, each characterised by different indicator species. Additionally, among a total of 412 taxa collected between 4 and 50 m depth, we provide the characteristics of the 64 commonest species in terms of occurrence, frequency, distribution, abundance, bathymetric and sedimentary preferences, which constitutes most valuable information for ecosystem modelling. Megabenthic alpha diversity decreased with depth, contrary to evenness and was higher in the proximity of the river Guadiana and in highly exposed shores. We conclude that the megafauna, which is significantly quicker to collect and analyse, can provide an accurate alternative to macrofauna sampling, as their communities are shaped by the same drivers.

Seawater carbonate chemistry, thickness and carbonate elemental composition of the test of juvenile sea urchins in a laboratory experiment

Continuous anthropogenic CO2 emissions to the atmosphere and uptake by the oceans will cause a reduction of seawater pH and saturation state (Omega) of CaCO3 minerals from which marine calcifiers build their shells and skeletons. Sea urchins use the most soluble form of calcium carbonate, high-magnesium calcite, to build their skeleton, spines and grazing apparatus. In order to highlight the effects of increased pCO2 on the test thickness and carbonate elemental composition of juvenile sea urchins and potential differences in their responses linked to the diet, we performed a laboratory experiment on juvenile Paracentrotus lividus, grazing on calcifying (Corallina elongata) and non-calcifying (Cystoseira amentacea, Dictyota dichotoma) macroalgae, under different pH (corresponding to pCO2 values of 390, 550, 750 and 1000 µatm). Results highlighted the importance of the diet in determining sea urchin size irrespectively of the pCO2 level, and the relevance of macroalgal diet in modulating urchin Mg/Ca ratio. The present study provides relevant clues both in terms of the mechanism of mineral incorporation and in terms of bottom-up processes (algal diet) affecting top-down ones (fish predation) in rocky subtidal communities

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.