Could the acid–base status of Antarctic sea urchins indicate a better‐than‐expected resilience to near‐future ocean acidification?

Increasing atmospheric carbon dioxide concentration alters the chemistry of the oceans towards more acidic conditions. Polar oceans are particularly affected due to their low temperature, low carbonate content and mixing patterns, for instance upwellings. Calcifying organisms are expected to be highly impacted by the decrease in the oceans' pH and carbonate ions concentration. In particular, sea urchins, members of the phylum Echinodermata, are hypothesized to be at risk due to their high‐magnesium calcite skeleton. However, tolerance to ocean acidification in metazoans is first linked to acid–base regulation capacities of the extracellular fluids. No information on this is available to date for Antarctic echinoderms and inference from temperate and tropical studies needs support. In this study, we investigated the acid–base status of 9 species of sea urchins (3 cidaroids, 2 regular euechinoids and 4 irregular echinoids). It appears that Antarctic regular euechinoids seem equipped with similar acid–base regulation systems as tropical and temperate regular euechinoids but could rely on more passive ion transfer systems, minimizing energy requirements. Cidaroids have an acid–base status similar to that of tropical cidaroids. Therefore Antarctic cidaroids will most probably not be affected by decreasing seawater pH, the pH drop linked to ocean acidification being negligible in comparison of the naturally low pH of the coelomic fluid. Irregular echinoids might not suffer from reduced seawater pH if acidosis of the coelomic fluid pH does not occur but more data on their acid–base regulation are needed. Combining these results with the resilience of Antarctic sea urchin larvae strongly suggests that these organisms might not be the expected victims of ocean acidification. However, data on the impact of other global stressors such as temperature and of the combination of the different stressors needs to be acquired to assess the sensitivity of these organisms to global change.     

Anthropogenic changes to seawater buffer capacity combined with natural reef metabolism induce extreme future coral reef CO2 conditions

Ocean acidification, via an anthropogenic increase in seawater carbon dioxide (CO₂), is potentially a major threat to coral reefs and other marine ecosystems. However, our understanding of how natural short‐term diurnal CO₂ variability in coral reefs influences longer term anthropogenic ocean acidification remains unclear. Here, we combine observed natural carbonate chemistry variability with future carbonate chemistry predictions for a coral reef flat in the Great Barrier Reef based on the RCP8.5 CO₂ emissions scenario. Rather than observing a linear increase in reef flat partial pressure of CO₂ (pCO₂) in concert with rising atmospheric concentrations, the inclusion of in situ diurnal variability results in a highly nonlinear threefold amplification of the pCO₂ signal by the end of the century. This significant nonlinear amplification of diurnal pCO₂ variability occurs as a result of combining natural diurnal biological CO₂ metabolism with long‐term decreases in seawater buffer capacity, which occurs via increasing anthropogenic CO₂ absorption by the ocean. Under the same benthic community composition, the amplification in the variability in pCO₂ is likely to lead to exposure to mean maximum daily pCO₂ levels of ca. 2100 μatm, with corrosive conditions with respect to aragonite by end‐century at our study site. Minimum pCO₂ levels will become lower relative to the mean offshore value (ca. threefold increase in the difference between offshore and minimum reef flat pCO₂) by end‐century, leading to a further increase in the pCO₂ range that organisms are exposed to. The biological consequences of short‐term exposure to these extreme CO₂ conditions, coupled with elevated long‐term mean CO₂ conditions are currently unknown and future laboratory experiments will need to incorporate natural variability to test this. The amplification of pCO₂ that we describe here is not unique to our study location, but will occur in all shallow coastal environments where high biological productivity drives large natural variability in carbonate chemistry.     

Post‐glacial colonization routes coincide with a life‐history breakpoint along a latitudinal gradient

Although adaptive divergence along environmental gradients has repeatedly been demonstrated, the role of post‐glacial colonization routes in determining phenotypic variation along gradients has received little attention. Here, we used a hierarchical Q_ST–F_ST approach to separate the roles of adaptive and neutral processes in shaping phenotypic variation in moor frog (Rana arvalis) larval life histories along a 1,700 km latitudinal gradient across northern Europe. This species has colonized Scandinavia via two routes with a contact zone in northern Sweden. By using neutral SNP and common garden phenotypic data from 13 populations at two temperatures, we showed that most of the variation along the gradient occurred between the two colonizing lineages. We found little phenotypic divergence within the lineages; however, all phenotypic traits were strongly diverged between the southern and northern colonization routes, with higher growth and development rates and larger body size in the north. The Q_ST estimates between the colonization routes were four times higher than F_ST, indicating a prominent role for natural selection. Q_ST within the colonization routes did not generally differ from F_ST, but we found temperature‐dependent adaptive divergence close to the contact zone. These results indicate that lineage‐specific variation can account for much of the adaptive divergence along a latitudinal gradient.     

Effects of ocean warming and acidification on survival,growth and skeletal development in the early benthic juvenile sea urchin (Heliocidaris erythrogramma)

Co‐occurring ocean warming, acidification and reduced carbonate mineral saturation have significant impacts on marine biota, especially calcifying organisms. The effects of these stressors on development and calcification in newly metamorphosed juveniles (ca. 0.5 mm test diameter) of the intertidal sea urchin Heliocidaris erythrogramma, an ecologically important species in temperate Australia, were investigated in context with present and projected future conditions. Habitat temperature and pH/pCO₂ were documented to place experiments in a biologically and ecologically relevant context. These parameters fluctuated diurnally up to 10 °C and 0.45 pH units. The juveniles were exposed to three temperature (21, 23 and 25 °C) and four pH (8.1, 7.8, 7.6 and 7.4) treatments in all combinations, representing ambient sea surface conditions (21 °C, pH 8.1; pCO₂ 397; Ω_Ca 4.7; Ω_Ar 3.1), near‐future projected change (+2–4 °C, ?0.3–0.5 pH units; pCO₂ 400–1820; Ω_Ca 5.0–1.6; Ω_Ar 3.3–1.1), and extreme conditions experienced at low tide (+4 °C, ?0.3–0.7 pH units; pCO₂ 2850–2967; Ω_Ca 1.1–1.0; Ω_Ar 0.7–0.6). The lowest pH treatment (pH 7.4) was used to assess tolerance levels. Juvenile survival and test growth were resilient to current and near‐future warming and acidification. Spine development, however, was negatively affected by near‐future increased temperature (+2–4 °C) and extreme acidification (pH 7.4), with a complex interaction between stressors. Near‐future warming was the more significant stressor. Spine tips were dissolved in the pH 7.4 treatments. Adaptation to fluctuating temperature‐pH conditions in the intertidal may convey resilience to juvenile H. erythrogramma to changing ocean conditions, however, ocean warming and acidification may shift baseline intertidal temperature and pH/pCO₂ to levels that exceed tolerance limits.     

Elevated pCO2 increases sperm limitation and risk of polyspermy in the red sea urchin Strongylocentrotus franciscanus

Anthropogenic carbon dioxide (CO₂) emissions and the resultant acidification of surface ocean waters are predicted to have far‐reaching consequences for biological processes in the marine environment. For example, because changes in pH and pCO₂ can alter sperm performance, ocean acidification may be accompanied by reductions in the success of fertilization in marine broadcast spawners. Several studies have attempted to determine the effects of elevated pCO₂ on marine invertebrate fertilization success, albeit with differing results. These conflicts may stem from the use of inappropriate sperm–egg contact times and, in several cases, the lack of measurements over a range of sperm concentrations extending from sperm‐limited conditions to polyspermy scenarios. In our study, we used biologically realistic sperm–egg contact times and a full range of sperm concentrations to assess the effect of elevated pCO₂ on fertilization in the broadcast spawning sea urchin, Strongylocentrotus franciscanus. Fertilization experiments were carried out in seawater bubbled with CO₂ to 400 (control), 800, and 1800 ppm. Using a fertilization kinetics model, we estimate that elevated pCO₂ levels both increased sperm limitation and reduced the efficiency of fast blocks to polyspermy. Thus, elevated pCO₂ decreased the range of sperm concentrations over which high fertilization success was likely. Given the inherent difficulties in achieving high fertilization success in broadcast spawners, raised pCO₂ levels are likely to exacerbate low fertilization success in low‐density populations or in areas with high water turbulence.     

Soil‐surface CO2 efflux and its spatial and temporal variations in a young ponderosa pine plantation in northern California

Soil‐surface CO₂ efflux and its spatial and temporal variations were examined in an 8‐y‐old ponderosa pine plantation in the Sierra Nevada Mountains in California from June 1998 to August 1999. Continuous measurements of soil CO₂ efflux, soil temperatures and moisture were conducted on two 20 × 20 m sampling plots. Microbial biomass, fine root biomass, and the physical and chemical properties of the soil were also measured at each of the 18 sampling locations on the plots. It was found that the mean soil CO₂ efflux in the plantation was 4.43 µmol m^?2 s^?1 in the growing season and 3.12 µmol m^?2 s^?1 in the nongrowing season. These values are in the upper part of the range of published soil‐surface CO₂ efflux data. The annual maximum and minimum CO₂ efflux were 5.87 and 1.67 µmol m^?2 s^?1, respectively, with the maximum occurring between the end of May and early June and the minimum in December. The diurnal fluctuation of CO₂ efflux was relatively small (< 20%) with the minimum appearing around 09.00 hours and the maximum around 14.00 hours. Using daytime measurements of soil CO₂ efflux tends to overestimate the daily mean soil CO₂ efflux by 4–6%. The measurements taken between 09.00 and 11.00 hours (local time) seem to better represent the daily mean with a reduced sampling error of 0.9–1.5%. The spatial variation of soil CO₂ efflux among the 18 sampling points was high, with a coefficient of variation of approximately 30%. Most (84%) of the spatial variation was explained by fine root biomass, microbial biomass, and soil physical and chemical properties. Although soil temperature and moisture explained most of the temporal variations (76–95%) of soil CO₂ efflux, the two variables together explained less than 34% of the spatial variation. Microbial biomass, fine root biomass, soil nitrogen content, organic matter content, and magnesium content were significantly and positively correlated with soil CO₂ efflux, whereas bulk density and pH value were negatively correlated with CO₂ efflux. The relationship between soil CO₂ efflux and soil temperature was significantly controlled by soil moisture with a Q₁₀ value of 1.4 when soil moisture was <14% and 1.8 when soil moisture was >14%. Understanding the spatial and temporal variations is essential to accurately assessment of carbon budget at whole ecosystem and landscape scales. Thus, this study bears important implications for the study of large‐scale ecosystem dynamics, particularly in response to climatic variations and management regimes.     

Modified future diurnal variability of the global surface ocean CO2 system

Our understanding of how increasing atmospheric CO₂ and climate change influences the marine CO₂ system and in turn ecosystems has increasingly focused on perturbations to carbonate chemistry variability. This variability can affect ocean-climate feedbacks and has been shown to influence marine ecosystems. The seasonal variability of the ocean CO₂ system has already changed, with enhanced seasonal variations in the surface ocean pCO₂ over recent decades and further amplification projected by models over the 21st century. Mesocosm studies and CO₂ vent sites indicate that diurnal variability of the CO₂ system, the amplitude of which in extreme events can exceed that of mean seasonal variability, is also likely to be altered by climate change. Here, we modified a global ocean biogeochemical model to resolve physically and biologically driven diurnal variability of the ocean CO₂ system. Forcing the model with 3-h atmospheric outputs derived from an Earth system model, we explore how surface ocean diurnal variability responds to historical changes and project how it changes under two contrasting 21st-century emission scenarios. Compared to preindustrial values, the global mean diurnal amplitude of pCO₂ increases by 4.8 μatm (+226%) in the high-emission scenario but only 1.2 μatm (+55%) in the high-mitigation scenario. The probability of extreme diurnal amplitudes of pCO₂ and [H⁺] is also affected, with 30- to 60-fold increases relative to the preindustrial under high 21st-century emissions. The main driver of heightened pCO₂ diurnal variability is the enhanced sensitivity of pCO₂ to changes in temperature as the ocean absorbs atmospheric CO₂. Our projections suggest that organisms in the future ocean will be exposed to enhanced diurnal variability in pCO₂ and [H⁺], with likely increases in the associated metabolic cost that such variability imposes.     

Food availability outweighs ocean acidification effects in juvenile Mytilus edulis: laboratory and field experiments

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 out‐compete 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.     

Does elevated pCO2 affect reef octocorals?

Increasing anthropogenic pCO₂ alters seawater chemistry, with potentially severe consequences for coral reef growth and health. Octocorals are the second most important faunistic component in many reefs, often occupying 50% or more of the available substrate. Three species of octocorals from two families were studied in Eilat (Gulf of Aqaba), comprising the zooxanthellate Ovabunda macrospiculata and Heteroxenia fuscescens (family Xeniidae), and Sarcophyton sp. (family Alcyoniidae). They were maintained under normal (8.2) and reduced (7.6 and 7.3) pH conditions for up to 5 months. Their biolological features, including protein concentration, polyp weight, density of zooxanthellae, and their chlorophyll concentration per cell, as well as polyp pulsation rate, were examined under conditions more acidic than normal, in order to test the hypothesis that rising pCO₂ would affect octocorals. The results indicate no statistically significant difference between the octocorals exposed to reduced pH values compared to the control. It is therefore suggested that the octocorals' tissue may act as a protective barrier against adverse pH conditions, thus maintaining them unharmed at high levels of pCO₂.     

Effects of pCO2 and iron on the elemental composition and cell geometry of the marine diatom Pseudo‐nitzschia pseudodelicatissima (Bacillariophyceae)1

Partial pressure of CO₂ (pCO₂) and iron availability in seawater show corresponding changes due to biological and anthropogenic activities. The simultaneous change in these factors precludes an understanding of their independent effects on the ecophysiology of phytoplankton. In addition, there is a lack of data regarding the interactive effects of these factors on phytoplankton cellular stoichiometry, which is a key driving factor for the biogeochemical cycling of oceanic nutrients. Here, we investigated the effects of pCO₂ and iron availability on the elemental composition (C, N, P, and Si) of the diatom Pseudo‐nitzschia pseudodelicatissima (Hasle) Hasle by dilute batch cultures under 4 pCO₂ (~200, ~380, ~600, and ~800 μatm) and five dissolved inorganic iron (Fe′; ~5, ~10, ~20, ~50, and ~100 pmol · L^?1) conditions. Our experimental procedure successfully overcame the problems associated with simultaneous changes in pCO₂ and Fe′ by independently manipulating carbonate chemistry and iron speciation, which allowed us to evaluate the individual effects of pCO₂ and iron availability. We found that the C:N ratio decreased significantly only with an increase in Fe′, whereas the C:P ratio increased significantly only with an increase in pCO₂. Both Si:C and Si:N ratios decreased with increasing pCO₂ and Fe′. Our results indicate that changes in pCO₂ and iron availability could influence the biogeochemical cycling of nutrients in future oceans with high‐ CO₂ levels, and, similarly, during the time course of phytoplankton blooms. Moreover, pCO₂ and iron availability may also have affected oceanic nutrient biogeochemistry in the past, as these conditions have changed markedly over the Earth's history.     

Local adaptation with high gene flow: temperature parameters drive adaptation to altitude in the common frog (Rana temporaria)

Both environmental and genetic influences can result in phenotypic variation. Quantifying the relative contributions of local adaptation and phenotypic plasticity to phenotypes is key to understanding the effect of environmental variation on populations. Identifying the selective pressures that drive divergence is an important, but often lacking, next step. High gene flow between high‐ and low‐altitude common frog (Rana temporaria) breeding sites has previously been demonstrated in Scotland. The aim of this study was to assess whether local adaptation occurs in the face of high gene flow and to identify potential environmental selection pressures that drive adaptation. Phenotypic variation in larval traits was quantified in R. temporaria from paired high‐ and low‐altitude sites using three common temperature treatments. Local adaptation was assessed using Q_ST–F_ST analyses, and quantitative phenotypic divergence was related to environmental parameters using Mantel tests. Although evidence of local adaptation was found for all traits measured, only variation in larval period and growth rate was consistent with adaptation to altitude. Moreover, this was only evident in the three mountains with the highest high‐altitude sites. This variation was correlated with mean summer and winter temperatures, suggesting that temperature parameters are potentially strong selective pressures maintaining local adaptation, despite high gene flow.     

Combined effects of ocean acidification and solar UV radiation on photosynthesis,growth, pigmentation and calcification of the coralline alga Corallina sessilis (Rhodophyta)

Previous studies have shown that increasing atmospheric CO₂ concentrations affect calcification in some planktonic and macroalgal calcifiers due to the changed carbonate chemistry of seawater. However, little is known regarding how calcifying algae respond to solar UV radiation (UVR, UVA+UVB, 280–400 nm). UVR may act synergistically, antagonistically or independently with ocean acidification (high CO₂/low pH of seawater) to affect their calcification processes. We cultured the articulated coralline alga Corallina sessilis Yendo at 380 ppmv (low) and 1000 ppmv (high) CO₂ levels while exposing the alga to solar radiation treatments with or without UVR. The presence of UVR inhibited the growth, photosynthetic O₂ evolution and calcification rates by13%, 6% and 3% in the low and by 47%, 20% and 8% in the high CO₂ concentrations, respectively, reflecting a synergistic effect of CO₂ enrichment with UVR. UVR induced significant decline of pH in the CO₂‐enriched cultures. The contents of key photosynthetic pigments, chlorophyll a and phycobiliproteins decreased, while UV‐absorptivity increased under the high pCO₂/low pH condition. Nevertheless, UV‐induced inhibition of photosynthesis increased when the ratio of particulate inorganic carbon/particulate organic carbon decreased under the influence of CO₂‐acidified seawater, suggesting that the calcified layer played a UV‐protective role. Both UVA and UVB negatively impacted photosynthesis and calcification, but the inhibition caused by UVB was about 2.5–2.6 times that caused by UVA. The results imply that coralline algae suffer from more damage caused by UVB as they calcify less and less with progressing ocean acidification.     

Effects of elevated pCO2 on the metabolism of a temperate rhodolith Lithothamnion corallioides grown under different temperatures

Coralline algae are considered among the most sensitive species to near future ocean acidification. We tested the effects of elevated pCO₂ on the metabolism of the free‐living coralline alga Lithothamnion corallioides (“maerl”) and the interactions with changes in temperature. Specimens were collected in North Brittany (France) and grown for 3 months at pCO₂ of 380 (ambient pCO₂), 550, 750, and 1000 μatm (elevated pCO₂) and at successive temperatures of 10°C (ambient temperature in winter), 16°C (ambient temperature in summer), and 19°C (ambient temperature in summer +3°C). At each temperature, gross primary production, respiration (oxygen flux), and calcification (alkalinity flux) rates were assessed in the light and dark. Pigments were determined by HPLC. Chl a, carotene, and zeaxanthin were the three major pigments found in L. corallioides thalli. Elevated pCO₂ did not affect pigment content while temperature slightly decreased zeaxanthin and carotene content at 10°C. Gross production was not affected by temperature but was significantly affected by pCO₂ with an increase between 380 and 550 μatm. Light, dark, and diel (24 h) calcification rates strongly decreased with increasing pCO₂ regardless of the temperature. Although elevated pCO₂ only slightly affected gross production in L. corallioides, diel net calcification was reduced by up to 80% under the 1,000 μatm treatment. Our findings suggested that near future levels of CO₂ will have profound consequences for carbon and carbonate budgets in rhodolith beds and for the sustainability of these habitats.     

Consequences of elevated temperature and pCO2 on insect folivory at the ecosystem level: perspectives from the fossil record

Paleoecological studies document the net effects of atmospheric and climate change in a natural laboratory over timescales not accessible to laboratory or ecological studies. Insect feeding damage is visible on well‐preserved fossil leaves, and changes in leaf damage through time can be compared to environmental changes. We measured percent leaf area damaged on four fossil leaf assemblages from the Bighorn Basin, Wyoming, that range in age from 56.1 to 52.65 million years (Ma). We also include similar published data from three US sites 49.4 to ~45 Ma in our analyses. Regional climate was subtropical or warmer throughout this period, and the second oldest assemblage (56 Ma) was deposited during the Paleocene–Eocene Thermal Maximum (PETM), a geologically abrupt global warming event caused by massive release of carbon into the atmosphere. Total and leaf‐chewing damage are highest during the PETM, whether considering percent area damaged on the bulk flora, the average of individual host plants, or a single plant host that occurs at multiple sites. Another fossil assemblage in our study, the 52.65 Ma Fifteenmile Creek paleoflora, also lived during a period of globally high temperature and pCO₂, but does not have elevated herbivory. Comparison of these two sites, as well as regression analyses conducted on the entire dataset, demonstrates that, over long timescales, temperature and pCO₂ are uncorrelated with total insect consumption at the ecosystem level. Rather, the most important factor affecting herbivory is the relative abundance of plants with nitrogen‐fixing symbionts. Legumes dominate the PETM site; their prevalence would have decreased nitrogen limitation across the ecosystem, buffering generalist herbivore populations against decreased leaf nutritional quality that commonly occurs at high pCO₂. We hypothesize that nitrogen concentration regulates the opposing effects of elevated temperature and CO₂ on insect abundance and thereby total insect consumption, which has important implications for agricultural practices in today's world of steadily increasing pCO₂.     

Ocean acidification and warming scenarios increase microbioerosion of coral skeletons

Biological mediation of carbonate dissolution represents a fundamental component of the destructive forces acting on coral reef ecosystems. Whereas ocean acidification can increase dissolution of carbonate substrates, the combined impact of ocean acidification and warming on the microbioerosion of coral skeletons remains unknown. Here, we exposed skeletons of the reef‐building corals, Porites cylindrica and Isopora cuneata, to present‐day (Control: 400 μatm – 24 °C) and future pCO₂–temperature scenarios projected for the end of the century (Medium: +230 μatm – +2 °C; High: +610 μatm – +4 °C). Skeletons were also subjected to permanent darkness with initial sodium hypochlorite incubation, and natural light without sodium hypochlorite incubation to isolate the environmental effect of acidic seawater (i.e., Ω_aragonite <1) from the biological effect of photosynthetic microborers. Our results indicated that skeletal dissolution is predominantly driven by photosynthetic microborers, as samples held in the dark did not decalcify. In contrast, dissolution of skeletons exposed to light increased under elevated pCO₂–temperature scenarios, with P. cylindrica experiencing higher dissolution rates per month (89%) than I. cuneata (46%) in the high treatment relative to control. The effects of future pCO₂–temperature scenarios on the structure of endolithic communities were only identified in P. cylindrica and were mostly associated with a higher abundance of the green algae Ostreobium spp. Enhanced skeletal dissolution was also associated with increased endolithic biomass and respiration under elevated pCO₂–temperature scenarios. Our results suggest that future projections of ocean acidification and warming will lead to increased rates of microbioerosion. However, the magnitude of bioerosion responses may depend on the structural properties of coral skeletons, with a range of implications for reef carbonate losses under warmer and more acidic oceans.     

CO2 supersaturation along the aquatic conduit in Swedish watersheds as constrained by terrestrial respiration,aquatic respiration and weathering

We tested the hypothesis that CO₂ supersaturation along the aquatic conduit over Sweden can be explained by processes other than aquatic respiration. A first generalized‐additive model (GAM) analysis evaluating the relationships between single water chemistry variables and pCO₂ in lakes and streams revealed that water chemistry variables typical for groundwater input, e.g., dissolved silicate (DSi) and Mg²⁺ had explanatory power similar to total organic carbon (TOC). Further GAM analyses on various lake size classes and stream orders corroborated the slightly higher explanatory power for DSi in lakes and Mg²⁺ for streams compared with TOC. Both DSi and TOC explained 22–46% of the pCO₂ variability in various lake classes (0.01–>100 km²) and Mg²⁺ and TOC explained 11–41% of the pCO₂ variability in the various stream orders. This suggests that aquatic pCO₂ has a strong groundwater signature. Terrestrial respiration is a significant source of the observed supersaturation and we may assume that both terrestrial respiration and aquatic respiration contributed equally to pCO₂ efflux. pCO₂ and TOC concentrations decreased with lake size suggesting that the longer water residence time allow greater equilibration of CO₂ with the atmosphere and in‐lake mineralization of TOC. For streams, we observed a decreasing trend in pCO₂ with stream orders between 3 and 6. We calculated the total CO₂ efflux from all Swedish lakes and streams to be 2.58 Tg C yr^?1. Our analyses also demonstrated that 0.70 Tg C yr^?1 are exported to the ocean by Swedish watersheds as HCO₃^? and CO₃^2? of which about 0.56 Tg C yr^?1 is also a residual from terrestrial respiration and constitute a long‐term sink for atmospheric CO₂. Taking all dissolved inorganic carbon (DIC) fluxes along the aquatic conduit into account will lower the estimated net ecosystem C exchange (NEE) by 2.02 Tg C yr^?1, which corresponds to 10% of the NEE in Sweden.     

The effect of ocean acidification and temperature on the fertilization and embryonic development of the Sydney rock oyster Saccostrea glomerata (Gould 1850)

This study investigated the synergistic effects of ocean acidification (caused by elevations in the partial pressure of carbon dioxide pCO₂) and temperature on the fertilization and embryonic development of the economically and ecologically important Sydney rock oyster, Saccostrea glomerata (Gould 1850). As pCO₂ increased, fertilization significantly decreased. The temperature of 26 °C was the optimum temperature for fertilization, as temperature increased and decreased from this optimum, fertilization decreased. There was also an effect of pCO₂ and temperature on embryonic development. Generally as pCO₂ increased, the percentage and size of D‐veligers decreased and the percentage of D‐veligers that were abnormal increased. The optimum temperature was 26 °C and embryonic development decreased at temperatures that were above and below this temperature. Abnormality of D‐veligers was greatest at 1000 ppm and 18 and 30 °C (≥90%) and least at 375 ppm and 26 °C (≤4%). Finally prolonged exposure of elevated pCO₂ and temperature across early developmental stages led to fewer D‐veligers, more abnormality and smaller sizes in elevated CO₂ environments and may lead to lethal effects at suboptimal temperatures. Embryos that were exposed to the pCO₂ and temperature treatments for fertilization and embryonic development had fewer D‐veligers, greater percentage of abnormality and reduced size than embryos that were exposed to the treatments for embryonic development only. Further at the elevated temperature of 30 °C and 750–1000 ppm, there was no embryonic development. The results of this study suggest that predicted changes in ocean acidification and temperature over the next century may have severe implications for the distribution and abundance of S. glomerata as well as possible implications for the reproduction and development of other marine invertebrates.     

Stable isotope fractionation of strontium in coccolithophore calcite: Influence of temperature and carbonate chemistry

Marine calcifying eukaryotic phytoplankton (coccolithophores) is a major contributor to the pelagic production of CaCO₃ and plays an important role in the biogeochemical cycles of C, Ca and other divalent cations present in the crystal structure of calcite. The geochemical signature of coccolithophore calcite is used as palaeoproxy to reconstruct past environmental conditions and to understand the underlying physiological mechanisms (vital effects) and precipitation kinetics. Here, we present the stable Sr isotope fractionation between seawater and calcite (Δ^88/86Sr) of laboratory cultured coccolithophores in individual dependence of temperature and seawater carbonate chemistry. Coccolithophores were cultured within a temperature and a pCO₂ range from 10 to 25°C and from 175 to 1,240 μatm, respectively. Both environmental drivers induced a significant linear increase in coccolith stable Sr isotope fractionation. The temperature correlation at constant pCO₂ for Emiliania huxleyi and Coccolithus braarudii is expressed as Δ^88/86Sr = ?7.611 × 10^?3 T + 0.0061. The relation of Δ^88/86Sr to pCO₂ was tested in Emiliania huxleyi at 10 and 20°C and resulted in Δ^88/86Sr = ?5.394 × 10^?5 pCO₂ – 0.0920 and Δ^88/86Sr = ?5.742 × 10^?5 pCO₂ – 0.1351, respectively. No consistent relationship was found between coccolith Δ^88/86Sr and cellular physiology impeding a direct application of fossil coccolith Δ^88/86Sr as coccolithophore productivity proxy. An overall significant correlation was detected between the elemental distribution coefficient (D_Sr) and Δ^88/86Sr similar to inorganic calcite with a physiologically induced offset. Our observations indicate (i) that temperature and pCO₂ induce specific effects on coccolith Δ^88/86Sr values and (ii) that strontium elemental ratios and stable isotope fractionation are mainly controlled by precipitation kinetics when embedded into the crystal lattice and subject to vital effects during the transmembrane transport from seawater to the site of calcification. These results provide an important step to develop a coccolith Δ^88/86Sr palaeoproxy complementing the existing toolbox of palaeoceanography.     

Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest

Variation in soil temperature can account for most of the seasonal and diel variation in soil CO₂ efflux, but the temperature effect is not always consistent, and other factors such as soil water content are known to influence soil respiration. The objectives of this research were to study the spatial and temporal variation in soil respiration in a temperate forested landscape and to evaluate temperature and soil water functions as predictors of soil respiration. Soil CO₂ fluxes were measured with chambers throughout an annual cycle in six study areas at the Harvard Forest in central Massachusetts that include soil drainage classes from well drained to very poorly drained. The mean annual estimate of soil CO₂ efflux was 7.2 Mg ha^–1, but ranged from 5.3 in the swamp site to 8.5 in a well-drained site, indicating that landscape heterogeneity is related to soil drainage class. An exponential function relating CO₂ fluxes to soil temperature accounted for 80% of the seasonal variation in fluxes across all sites (Q₁₀ = 3.9), but the Q₁₀ ranged from 3.4 to 5.6 for the individual study sites. A significant drought in 1995 caused rapid declines in soil respiration rates in August and September in five of the six sites (a swamp site was the exception). This decline in CO₂ fluxes correlated exponentially with decreasing soil matric potential, indicating a mechanistic effect of drought stress. At moderate to high water contents, however, soil water content was negatively correlated with soil temperature, which precluded distinguishing between the effects of these two confounded factors on CO₂ flux. Occurrence of high Q₁₀ values and variation in Q₁₀ values among sites may be related to: (i) confounding effects of high soil water content; (ii) seasonal and diel patterns in root respiration and turnover of fine roots that are linked to above ground phenology and metabolism; and (iii) variation in the depth where CO₂ is produced. The Q₁₀ function can yield reasonably good predictions of annual fluxes of CO₂, but it is a simplification that masks responses of root and microbial processes to variation in temperature and water content throughout the soil.     

SHORT‐ VERSUS LONG‐TERM RESPONSES TO CHANGING CO2 IN A COASTAL DINOFLAGELLATE BLOOM: IMPLICATIONS FOR INTERSPECIFIC COMPETITIVE INTERACTIONS AND COMMUNITY STRUCTURE

Increasing pCO₂ (partial pressure of CO₂) in an “acidified” ocean will affect phytoplankton community structure, but manipulation experiments with assemblages briefly acclimated to simulated future conditions may not accurately predict the long‐term evolutionary shifts that could affect inter‐specific competitive success. We assessed community structure changes in a natural mixed dinoflagellate bloom incubated at three pCO₂ levels (230, 433, and 765 ppm) in a short‐term experiment (2 weeks). The four dominant species were then isolated from each treatment into clonal cultures, and maintained at all three pCO₂ levels for approximately 1 year. Periodically (4, 8, and 12 months), these pCO₂‐conditioned clones were recombined into artificial communities, and allowed to compete at their conditioning pCO₂ level or at higher and lower levels. The dominant species in these artificial communities of CO₂‐conditioned clones differed from those in the original short‐term experiment, but individual species relative abundance trends across pCO₂ treatments were often similar. Specific growth rates showed no strong evidence for fitness increases attributable to conditioning pCO₂ level. Although pCO₂ significantly structured our experimental communities, conditioning time and biotic interactions like mixotrophy also had major roles in determining competitive outcomes. New methods of carrying out extended mixed species experiments are needed to accurately predict future long‐term phytoplankton community responses to changing pCO₂.