Carbonate chemistry, temperature and salinity of Lady Elliot Island Reef 2009-2010

Ocean acidification leads to changes in marine carbonate chemistry that are predicted to cause a decline in future coral reef calcification. Several laboratory and mesocosm experiments have described calcification responses of species and communities to increasing CO2. The few in situ studies on natural coral reefs that have been carried out to date have shown a direct relationship between aragonite saturation state (Omega arag) and net community calcification (Gnet). However, these studies have been performed over a limited range of Omega arag values, where extrapolation outside the observational range is required to predict future changes in coral reef calcification. We measured extreme diurnal variability in carbonate chemistry within a reef flat in the southern Great Barrier Reef, Australia. Omega arag varied between 1.1 and 6.5, thus exceeding the magnitude of change expected this century in open ocean subtropical/tropical waters. The observed variability comes about through biological activity on the reef, where changes to the carbonate chemistry are enhanced at low tide when reef flat waters are isolated from open ocean water. We define a relationship between net community calcification and Omega arag, using our in situ measurements. We find net community calcification to be linearly related to Omega arag, while temperature and nutrients had no significant effect on Gnet. Using our relationship between Gnet and Omega arag, we predict that net community calcification will decline by 55% of its preindustrial value by the end of the century. It is not known at this stage whether exposure to large variability in carbonate chemistry will make reef flat organisms more or less vulnerable to the non-calcifying physiological effects of increasing ocean CO2 and future laboratory studies will need to incorporate this natural variability to address this question.

(Table 1) Stage of the lemming cycle and nesting parameters of the greater snow geese between 1996-2005, Bylot Island

Resource pulses are common in various ecosystems and often have large impacts on ecosystem functioning. Many animals hoard food during resource pulses, yet how this behaviour affects pulse diffusion through trophic levels is poorly known because of a lack of individual-based studies. Our objective was to examine how the hoarding behaviour of arctic foxes (Alopex lagopus) preying on a seasonal pulsed resource (goose eggs) was affected by annual and seasonal changes in resource availability. We monitored foraging behaviour of foxes in a greater snow goose (Chen caerulescens atlanticus) colony during 8 nesting seasons that covered 2 lemming cycles. The number of goose eggs taken and cached per hour by foxes declined 6-fold from laying to hatching, while the proportion of eggs cached remained constant. In contrast, the proportion of eggs cached by foxes fluctuated in response to the annual lemming cycle independently of the seasonal pulse of goose eggs. Foxes cached the majority of eggs taken (> 90%) when lemming abundance was high or moderate but only 40% during the low phase of the cycle. This likely occurred because foxes consumed a greater proportion of goose eggs to fulfill their energy requirement at low lemming abundance. Our study clearly illustrates a behavioural mechanism that extends the energetic benefits of a resource pulse. The hoarding behaviour of the main predator enhances the allochthonous nutrients input brought by migrating birds from the south into the arctic terrestrial ecosystem. This could increase average predator density and promote indirect interactions among prey.

Anemone abundance and productivity at North Vulcano Island in May 2011

Increased seawater pCO2, and in turn 'ocean acidification' (OA), is predicted to profoundly impact marine ecosystem diversity and function this century. Much research has already focussed on calcifying reef-forming corals (Class: Anthozoa) that appear particularly susceptible to OA via reduced net calcification. However, here we show that OA-like conditions can simultaneously enhance the ecological success of non-calcifying anthozoans, which not only play key ecological and biogeochemical roles in present day benthic ecosystems but also represent a model organism should calcifying anthozoans exist as less calcified (soft-bodied) forms in future oceans. Increased growth (abundance and size) of the sea anemone (Anemonia viridis) population was observed along a natural CO2 gradient at Vulcano, Italy. Both gross photosynthesis (PG) and respiration (R) increased with pCO2 indicating that the increased growth was, at least in part, fuelled by bottom up (CO2 stimulation) of metabolism. The increase of PG outweighed that of R and the genetic identity of the symbiotic microalgae (Symbiodinium spp.) remained unchanged (type A19) suggesting proximity to the vent site relieved CO2 limitation of the anemones' symbiotic microalgal population. Our observations of enhanced productivity with pCO2, which are consistent with previous reports for some calcifying corals, convey an increase in fitness that may enable non-calcifying anthozoans to thrive in future environments, i.e. higher seawater pCO2. Understanding how CO2-enhanced productivity of non- (and less-) calcifying anthozoans applies more widely to tropical ecosystems is a priority where such organisms can dominate benthic ecosystems, in particular following localized anthropogenic stress.

(Table 2) Carbon and nitrogen isotopic ratios in blood sampled from arctic foxes (Vulpes lagopus) on Bylot Island between 2003-2008

Inter-individual variation in diet within generalist animal populations is thought to be a widespread phenomenon but its potential causes are poorly known. Inter-individual variation can be amplified by the availability and use of allochthonous resources, i.e., resources coming from spatially distinct ecosystems. Using a wild population of arctic fox as a study model, we tested hypotheses that could explain variation in both population and individual isotopic niches, used here as proxy for the trophic niche. The arctic fox is an opportunistic forager, dwelling in terrestrial and marine environments characterized by strong spatial (arctic-nesting birds) and temporal (cyclic lemmings) fluctuations in resource abundance. First, we tested the hypothesis that generalist foraging habits, in association with temporal variation in prey accessibility, should induce temporal changes in isotopic niche width and diet. Second, we investigated whether within-population variation in the isotopic niche could be explained by individual characteristics (sex and breeding status) and environmental factors (spatiotemporal variation in prey availability). We addressed these questions using isotopic analysis and Bayesian mixing models in conjunction with linear mixed-effects models. We found that: i) arctic fox populations can simultaneously undergo short-term (i.e., within a few months) reduction in both isotopic niche width and inter-individual variability in isotopic ratios, ii) individual isotopic ratios were higher and more representative of a marine-based diet for non-breeding than breeding foxes early in spring, and iii) lemming population cycles did not appear to directly influence the diet of individual foxes after taking their breeding status into account. However, lemming abundance was correlated to proportion of breeding foxes, and could thus indirectly affect the diet at the population scale.

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.

Seawater carbonate chemistry, community calcification and photosynthesis during experiments with coral reefs at Shiraho reef, Ishigaki Island, southwest Japan, 2009

Seven coral reef communities were defined on Shiraho fringing reef, Ishigaki Island, Japan. Net photosynthesis and calcification rates were measured by in situ incubations at 10 sites that included six of the defined communities, and which occupied most of the area on the reef flat and slope. Net photosynthesis on the reef flat was positive overall, but the reef flat acts as a source for atmospheric CO2, because the measured calcification/photosynthesis ratio of 2.5 is greater than the critical ratio of 1.67. Net photosynthesis on the reef slope was negative. Almost all excess organic production from the reef flat is expected to be effused to the outer reef and consumed by the communities there. Therefore, the total net organic production of the whole reef system is probably almost zero and the whole reef system also acts as a source for atmospheric CO2. Net calcification rates of the reef slope corals were much lower than those of the branching corals. The accumulation rate of the former was approximately 0.5 m kyr?1 and of the latter was ~0.7-5 m kyr?1. Consequently, reef slope corals could not grow fast enough to keep up with or catch up to rising sea levels during the Holocene. On the other hand, the branching corals grow fast enough to keep up with this rising sea level. Therefore, a transition between early Holocene and present-day reef communities is expected. Branching coral communities would have dominated while reef growth kept pace with sea level rise, and the reef was constructed with a branching coral framework. Then, the outside of this framework was covered and built up by reef slope corals and present-day reefs were constructed.

Seawater carbonate chemistry and biological processes of coral Acropora muricata during experiments and observations in La Saline fringing reef, La Reunion Island, western Indian Ocean, 2011

The effect of decreasing aragonite saturation state (Omega Arag) of seawater (elevated pCO2) on calcification rates of Acropora muricata was studied using nubbins prepared from parent colonies located at two sites of La Saline reef (La Réunion Island, western Indian Ocean): a back-reef site (BR) affected by nutrient-enriched groundwater discharge (mainly nitrate), and a reef flat site (RF) with low terrigenous inputs. Protein and chlorophyll a content of the nubbins, as well as zooxanthellae abundance, were lower at RF than BR. Nubbins were incubated at ~27°C over 2 h under sunlight, in filtered seawater manipulated to get differing initial pCO2 (1,440-340 µatm), Omega Arag (1.4-4.0), and dissolved inorganic carbon (DIC) concentrations (2,100-1,850 µmol/kg). Increasing DIC concentrations at constant total alkalinity (AT) resulted in a decrease in Omega Arag and an increase in pCO2. AT at the beginning of the incubations was kept at a natural level of 2,193 ± 6 µmol/kg (mean ± SD). Net photosynthesis (NP) and calcification were calculated from changes in pH and AT during the incubations. Calcification decrease in response to doubling pCO2 relative to preindustrial level was 22% for RF nubbins. When normalized to surface area of the nubbins, (1) NP and calcification were higher at BR than RF, (2) NP increased in high pCO2 treatments at BR compared to low pCO2 treatments, and (3) calcification was not related to Omega Arag at BR. When normalized to NP, calcification was linearly related to Omega Arag at both sites, and the slopes of the relationships were not significantly different. The increase in NP at BR in the high pCO2 treatments may have increased calcification and thus masked the negative effect of low Omega Arag on calcification. Removing the effect of NP variations at BR showed that calcification declined in a similar manner with decreased Omega Arag (increased pCO2) whatever the nutrient loading.

Seawater carbonate chemistry and community calcification during Shiraha coral reef (Ishigaki Island, Japan) studies, 1995

Only about half of all the CO_2 that has been produced by the burning of fossil fuels now remains in the atmosphere. The CO_2 "missing" from the atmosphere is the subject of an important debate. It was thought that the great majority of the missing CO_2 has invaded the ocean, for this system naturally acts as a giant chemical regulator of the atmosphere. Although it is clear that ocean processes have a major role in the regulation of the carbon dioxide content of the atmosphere through air-sea exchange processes, recent studies of the oceanic carbon cycle and air-sea interaction indicate that oceanic carbon is in a quasi-steady state via the system of biological and physical processes in the ocean interior. It is difficult to determine whether the ocean has the capacity to take up the increasing air-born CO_2 released by human activities over the past five or six decades. To understand this enigma, we need a better understanding of the natural variability of the oceanic carbon cycle.

Wet tundra polygon centers and distance between polygon centroids on Samoylov Island, Lena Delta, Siberia, Russia, with links to shape files

Subgrid processes occur in various ecosystems and landscapes but, because of their small scale, they are not represented or poorly parameterized in climate models. These local heterogeneities are often important or even fundamental for energy and carbon balances. This is especially true for northern peatlands and in particular for the polygonal tundra, where methane emissions are strongly influenced by spatial soil heterogeneities. We present a stochastic model for the surface topography of polygonal tundra using Poisson-Voronoi diagrams and we compare the results with available recent field studies. We analyze seasonal dynamics of water table variations and the landscape response under different scenarios of precipitation income. We upscale methane fluxes by using a simple idealized model for methane emission. Hydraulic interconnectivities and large-scale drainage may also be investigated through percolation properties and thresholds in the Voronoi graph. The model captures the main statistical characteristics of the landscape topography, such as polygon area and surface properties as well as the water balance. This approach enables us to statistically relate large-scale properties of the system to the main small-scale processes within the single polygons.

Seawater conditions during the experiment in May 2011 at the sampling sites off Vulcano Island

The impacts of ocean acidification on coastal biofilms are poorly understood. Carbon dioxide vent areas provide an opportunity to make predictions about the impacts of ocean acidification. We compared biofilms that colonised glass slides in areas exposed to ambient and elevated levels of pCO2 along a coastal pH gradient, with biofilms grown at ambient and reduced light levels. Biofilm production was highest under ambient light levels, but under both light regimes biofilm production was enhanced in seawater with high pCO2. Uronic acids are a component of biofilms and increased significantly with high pCO2. Bacteria and Eukarya denaturing gradient gel electrophoresis profile analysis showed clear differences in the structures of ambient and reduced light biofilm communities, and biofilms grown at high pCO2 compared with ambient conditions. This study characterises biofilm response to natural seabed CO2 seeps and provides a baseline understanding of how coastal ecosystems may respond to increased pCO2 levels.