Orbital-resolution Pliocene-Pleistocene simulations of CO2, sea level, global temperature and benthic d18O, and d11B-based proxy-CO2 data

During the past five million yrs, benthic d18O records indicate a large range of climates, from warmer than today during the Pliocene Warm Period to considerably colder during glacials. Antarctic ice cores have revealed Pleistocene glacial-interglacial CO2 variability of 60-100 ppm, while sea level fluctuations of typically 125 m are documented by proxy data. However, in the pre-ice core period, CO2 and sea level proxy data are scarce and there is disagreement between different proxies and different records of the same proxy. This hampers comprehensive understanding of the long-term relations between CO2, sea level and climate. Here, we drive a coupled climate-ice sheet model over the past five million years, inversely forced by a stacked benthic d18O record. We obtain continuous simulations of benthic d18O, sea level and CO2 that are mutually consistent. Our model shows CO2 concentrations of 300 to 470 ppm during the Early Pliocene. Furthermore, we simulate strong CO2 variability during the Pliocene and Early Pleistocene. These features are broadly supported by existing and new d11B-based proxy CO2 data, but less by alkenone-based records. The simulated concentrations and variations therein are larger than expected from global mean temperature changes. Our findings thus suggest a smaller Earth System Sensitivity than previously thought. This is explained by a more restricted role of land ice variability in the Pliocene. The largest uncertainty in our simulation arises from the mass balance formulation of East Antarctica, which governs the variability in sea level, but only modestly affects the modeled CO2 concentrations.

Adaptive variability to low-pH river discharges in Acartia tonsa and stress responses to high PCO2 conditions

Environmental transitions leading to spatial physical-chemical gradients are of ecological and evolutionary interest because they are able to induce variations in phenotypic plasticity. Thus, the adaptive variability to low-pH river discharges may drive divergent stress responses [ingestion rates (IR) and expression of stress-related genes such as Heat shock protein 70 (Hsp70) and Ferritin] in the neritic copepod Acartia tonsa facing changes in the marine chemistry associated to ocean acidification (OA). These responses were tested in copepod populations inhabiting two environments with contrasting carbonate system parameters (an estuarine versus coastal area) in the Southern Pacific Ocean, and assessing an in situ and 96-h experimental incubation under conditions of high pressure of CO2 (PCO2 1200 ppm). Adaptive variability was a determining factor in driving variability of copepods' responses. Thus, the food-rich but colder and corrosive estuary induced a traits trade-off expressed as depressed IR under in situ conditions. However, this experience allowed these copepods to tolerate further exposure to high PCO2 levels better, as their IRs were on average 43% higher than those of the coastal individuals. Indeed, expression of both the Hsp70 and Ferritin genes in coastal copepods was significantly higher after acclimation to high PCO2 conditions. Along with other recent evidence, our findings confirm that adaptation to local fluctuations in seawater pH seems to play a significant role in the response of planktonic populations to OA-associated conditions. Facing the environmental threat represented by the inter-play between multiple drivers of climate change, this biological feature should be examined in detail as a potential tool for risk mitigation policies in coastal management arrangements.

Permeable coral reef sediment dissolution driven by elevated pCO2 and pore water advection

Ocean acidification (OA) is expected to drive the transition of coral reef ecosystems from net calcium carbonate (CaCO3) precipitating to net dissolving within the next century. Although permeable sediments represent the largest reservoir of CaCO3 in coral reefs, the dissolution of shallow CaCO3 sands under future pCO2 levels has not been measured under natural conditions. In situ, advective chamber incubations under elevated pCO2 (~800 µatm) shifted the sediments from net precipitating to net dissolving. Pore water advection more than doubled dissolution rates (1.10 g CaCO3/m**2/day) when compared to diffusive conditions (0.42 g CaCO3/m**2 /day). Sediment dissolution could reduce net ecosystem calcification rates of the Heron Island lagoon by 8% within the next century, which is equivalent to a 25% reduction in the global average calcification rate of coral lagoons. The dissolution of CaCO3 sediments needs to be taken into account in order to address how OA will impact the net accretion of coral reefs under future predicted increases in CO2.

Responses of marine benthic microalgae to elevated CO2

Increasing anthropogenic CO2 emissions to the atmosphere are causing a rise in pCO2 concentrations in the ocean surface and lowering pH. To predict the effects of these changes, we need to improve our understanding of the responses of marine primary producers since these drive biogeochemical cycles and profoundly affect the structure and function of benthic habitats. The effects of increasing CO2 levels on the colonisation of artificial substrata by microalgal assemblages (periphyton) were examined across a CO2 gradient off the volcanic island of Vulcano (NE Sicily). We show that periphyton communities altered significantly as CO2 concentrations increased. CO2 enrichment caused significant increases in chlorophyll a concentrations and in diatom abundance although we did not detect any changes in cyanobacteria. SEM analysis revealed major shifts in diatom assemblage composition as CO2 levels increased. The responses of benthic microalgae to rising anthropogenic CO2 emissions are likely to have significant ecological ramifications for coastal systems.

Experiment: changing carbonate chemistry influence on coccoliths formed by Emiliania huxleyi

The coccolithophore Emiliania huxleyi is a marine phytoplankton species capable of forming small calcium carbonate scales (coccoliths) which cover the organic part of the cell. Calcification rates of E. huxleyi are known to be sensitive to changes in seawater carbonate chemistry. It has, however, not yet been clearly determined how these changes are reflected in size and weight of individual coccoliths and which specific parameter(s) of the carbonate system drive morphological modifications. Here, we compare data on coccolith size, weight, and malformation from a set of five experiments with a large diversity of carbonate chemistry conditions. This diversity allows distinguishing the influence of individual carbonate chemistry parameters such as carbon dioxide (CO2), bicarbonate (HCO3- ), carbonate ion (CO32-), and protons (H+) on the measured parameters. Measurements of fine-scale morphological structures reveal an increase of coccolith malformation with decreasing pH suggesting that H+ is the major factor causing malformations. Coccolith distal shield area varies from about 5 to 11 µm2. Changes in size seem to be mainly induced by varying [HCO3- ] and [H+] although influence of [CO32-] cannot be entirely ruled out. Changes in coccolith weight were proportional to changes in size. Increasing CaCO3 production rates are reflected in an increase in coccolith weight and an increase of the number of coccoliths formed per unit time. The combined investigation of morphological features and coccolith production rates presented in this study may help to interpret data derived from sediment cores, where coccolith morphology is used to reconstruct calcification rates in the water column.

Figure 5. Trace metal analysis of sediment core PS1506-1

An increase in whole ocean alkalinity during glacial periods could account, in part, for the drawdown of atmospheric CO2 into the ocean. Such an increase was inevitable due to the near elimination of shelf area for the burial of coral reef alkalinity. We present evidence, based on down-core measurements of benthic foraminiferal B/Ca and Mg/Ca from a core in the Weddell Sea, that the deep ocean carbonate ion concentration, [CO3 2-], was elevated by ~25 µmol/kg during each glacial period of the last 800 kyrs. The heterogeneity of the preservation histories in the different ocean basins reflects control of the carbonate chemistry of the deep glacial ocean in the Atlantic and Pacific by the changing ventilation and chemistry of Weddell Sea waters. These waters are more corrosive than interglacial northern sourced waters, but not as undersaturated as interglacial southern sourced waters. Our inferred increase in whole ocean alkalinity can be reconciled with reconstructions of glacial saturation horizon depth and the carbonate budget, if carbonate burial rates also increased above the saturation horizon as a result of enhanced pelagic calcification. The Weddell records display low [CO3 2-] during deglaciations and peak interglacial warmth, coincident with maxima in %CaCO3 in the Atlantic and Pacific Oceans. Should the burial rate of alkalinity in the more alkaline glacial deepwaters outstrip the rate of alkalinity supply, then pelagic carbonate production by the coccolithophores, at the end of the glacial maximum could drive a decrease in ocean [CO3 2-] and act to trigger the deglacial rise in pCO2.

Seawater carbonate chemistry and particulate organic particles during a semicontinuous batch culture experiment with Trichodesmium IMS101, 2007

Diazotrophic (N2-fixing) cyanobacteria provide the biological source of new nitrogen for large parts of the ocean. However, little is known about their sensitivity to global change. Here we show that the single most important nitrogen fixer in today's ocean, Trichodesmium, is strongly affected by changes in CO2 concentrations. Cell division rate doubled with rising CO2 (glacial to projected year 2100 levels) prompting lower carbon, nitrogen and phosphorus cellular contents, and reduced cell dimensions. N2 fixation rates per unit of phosphorus utilization as well as C:P and N:P ratios more than doubled at high CO2, with no change in C:N ratios. This could enhance the productivity of N-limited oligotrophic oceans, drive some of these areas into P limitation, and increase biological carbon sequestration in the ocean. The observed CO2 sensitivity of Trichodesmium could thereby provide a strong negative feedback to atmospheric CO2 increase.

Environmental stability affects phenotypic evolution in a globally distributed marine picoplankton

Marine phytoplankton can evolve rapidly when confronted with aspects of climate change because of their large population sizes and fast generation times. Despite this, the importance of environment fluctuations, a key feature of climate change, has received little attention-selection experiments with marine phytoplankton are usually carried out in stable environments and use single or few representatives of a species, genus or functional group. Here we investigate whether and by how much environmental fluctuations contribute to changes in ecologically important phytoplankton traits such as C:N ratios and cell size, and test the variability of changes in these traits within the globally distributed species Ostreococcus. We have evolved 16 physiologically distinct lineages of Ostreococcus at stable high CO2 (1031±87?µatm CO2, SH) and fluctuating high CO2 (1012±244?µatm CO2, FH) for 400 generations. We find that although both fluctuation and high CO2 drive evolution, FH-evolved lineages are smaller, have reduced C:N ratios and respond more strongly to further increases in CO2 than do SH-evolved lineages. This indicates that environmental fluctuations are an important factor to consider when predicting how the characteristics of future phytoplankton populations will have an impact on biogeochemical cycles and higher trophic levels in marine food webs.

Investigations on the Chilean kelp crab Taliepus dentatus

Physiological responses of larval stages can differ from those of the adults, affecting key ecological processes. Therefore, developing a mechanistic understanding of larval responses to environmental conditions is essential vis-à-vis climate change. We studied the thermal tolerance windows, defined by lower and upper pejus (Tp) and critical temperatures (Tc), of zoea I, II, and megalopa stages of the Chilean kelp crab Taliepus dentatus. Tp limits determine the temperature range where aerobic scope is maximal and functioning of the organism is unrestrained and were estimated from direct observations of larval activity. Tc limits define the transition from aerobic to anaerobic metabolism, and were estimated from the relationship between standard metabolic rate and temperature. Zoea I showed the broadest, Zoea II an intermediate, and megalopae the narrowest tolerance window (Tp). Optimum performance in megalopae was limited to Tp between 11 and 15°C, while their Tc ranged between 7 and 19°C. Although Tc may be seldom encountered by larvae, the narrower Tp temperatures can frequently expose larvae to unfavorable conditions that can drastically constrain their performance. Temperatures beyond the Tp range of megalopae have been observed in most spring and summer months in central Chile, and can have important consequences for larval swimming performance and impair their ability to avoid predators or settle successfully. Besides the well-documented effects of temperature on development time, variability in field temperatures beyond Tp can affect performance of particular larval stages, which could drive large-scale variability in recruitment and population dynamics of T. dentatus and possibly other invertebrate species.

R-codes, and coral abundance, water temperature and transect characteristics of nearshore reefs in Belize

Coral reefs are increasingly threatened by global and local anthropogenic stressors, such as rising seawater temperature and nutrient enrichment. These two stressors vary widely across the reef face and parsing out their influence on coral communities at reef system scales has been particularly challenging. Here, we investigate the influence of temperature and nutrients on coral community traits and life history strategies on lagoonal reefs across the Belize Mesoamerican Barrier Reef System (MBRS). A novel metric was developed using ultra-high-resolution sea surface temperatures (SST) to classify reefs as enduring low (lowTP), moderate (modTP), or extreme (extTP) temperature parameters over 10 years (2003 to 2012). Chlorophyll-a (chl a) records obtained for the same interval were employed as a proxy for bulk nutrients and these records were complemented with in situ measurements to "sea truth" nutrient content across the three reef types. Chl a concentrations were highest at extTP sites, medial at modTP sites and lowest at lowTP sites. Coral species richness, abundance, diversity, density, and percent cover were lower at extTP sites compared to lowTP and modTP sites, but these reef community traits did not differ between lowTP and modTP sites. Coral life history strategy analyses showed that extTP sites were dominated by hardy stress-tolerant and fast-growing weedy coral species, while lowTP and modTP sites consisted of competitive, generalist, weedy, and stress-tolerant coral species. These results suggest that differences in coral community traits and life history strategies between extTP and lowTP/modTP sites were driven primarily by temperature differences with differences in nutrients across site types playing a lesser role. Dominance of weedy and stress-tolerant genera at extTP sites suggests that corals utilizing these two life history strategies may be better suited to cope with warmer oceans and thus may warrant further protective status during this climate change interval. Data associated with this project are archived here, including: -SST data -Satellite Chl a data -Nutrient measurements -Raw coral community survey data For questions contact Justin Baumann (j.baumann3 gmail.com)

(Table S1) Age determination of sediment cores SK99_17 and MD76-131

Productivity in the Arabian Sea is one of the highest in the world. It is controlled by seasonally reversing monsoonal wind-driven upwelling of nutrient-rich deeper waters which fuel phytoplankton growth. The detailed history of upwelling-induced productivity in the eastern Arabian Sea is unknown. Here we present paleoproductivity records from a composite sediment core at the millennial scale during the last 80 kyr B.P. These records are based on relative abundance counts of planktonic foraminifera and organic carbon contents, which are shown to mainly vary in concert. The eastern Arabian Sea upwelling-induced productivity was higher in the glacial period than in the Holocene, but it fell repeatedly on millennial timescales. These productivity declines occurred during cold events in the North Atlantic region, with the most pronounced changes prevailing during the Heinrich events. Hence, seasonal monsoon winds that drive upwelling-induced productivity in the east were weak when the North Atlantic was cold. These weak winds resulted in stratification of the water column, comparable to today's Arabian Sea stratification in the intermonsoonal period. Combining the new eastern with published western Arabian Sea results shows that the entire biological factory was severely diminished during the North Atlantic Heinrich events, and the seasonal productivity change in the Arabian Sea monsoon system was reduced with year-round low productivity.

Seawater carbonate chemistry, tissue-skeleton interface and calcification in reef corals in a laboratory experiment

Insight into the response of reef corals and other major marine calcifiers to ocean acidification is limited by a lack of knowledge about how seawater pH and carbonate chemistry impact the physiological processes that drive biomineralization. Ocean acidification is proposed to reduce calcification rates in corals by causing declines in internal pH at the calcifying tissue-skeleton interface where biomineralization takes place. Here, we performed an in vivo study on how partial-pressure CO(2)-driven seawater acidification impacts intracellular pH in coral calcifying cells and extracellular pH in the fluid at the tissue-skeleton interface [subcalicoblastic medium (SCM)] in the coral Stylophora pistillata. We also measured calcification in corals grown under the same conditions of seawater acidification by measuring lateral growth of colonies and growth of aragonite crystals under the calcifying tissue. Our findings confirm that seawater acidification decreases pH of the SCM, but this decrease is gradual relative to the surrounding seawater, leading to an increasing pH gradient between the SCM and seawater. Reductions in calcification rate, both at the level of crystals and whole colonies, were only observed in our lowest pH treatment when pH was significantly depressed in the calcifying cells in addition to the SCM. Overall, our findings suggest that reef corals may mitigate the effects of seawater acidification by regulating pH in the SCM, but they also highlight the role of calcifying cell pH homeostasis in determining the response of reef corals to changes in external seawater pH and carbonate chemistry.

Sea surface temperature and primary productivity reconstruction of sediment core SO130-275KL from the northeastern Arabian Sea during the late Holocene

Variability in the oceanic environment of the Arabian Sea region is strongly influenced by the seasonal monsoon cycle of alternating wind directions. Prominent and well studied is the summer monsoon, but much less is known about late Holocene changes in winter monsoon strength with winds from the northeast that drive convective mixing and high surface ocean productivity in the northeastern Arabian Sea. To establish a high-resolution record of winter monsoon variability for the late Holocene, we analyzed alkenone-derived sea surface temperature (SST) variations and proxies of primary productivity (organic carbon and d15N) in a well-laminated sediment core from the Pakistan continental margin. Weak winter monsoon intensities off Pakistan are indicated from 400 B.C. to 250 A.D. by reduced productivity and relatively high SST. At about 250 A.D., the intensity of the winter monsoon increased off Pakistan as indicated by a trend to lower SST. We infer that monsoon conditions were relatively unstable from ~500 to 1300 A.D., because primary production and SST were highly variable. Declining SST and elevated biological production from 1400 to 1900 A.D. suggest invigorated convective winter mixing by strengthening winter monsoon circulation, most likely a regional expression of colder climate conditions during the Little Ice Age on the Northern Hemisphere. The comparison of winter monsoon intensity with records of summer monsoon intensity suggests that an inverse relationship between summer and winter monsoon strength exists in the Asian monsoon system during the late Holocene, effected by shifts in the Intertropical Convergence Zone.

A 5 million year reconstruction of sea level, temperature, and sea water d18O

Marine sediment records from the Oligocene and Miocene reveal clear 400,000-year (400-kyr) climate cycles related to variations in orbital eccentricity. These cycles are also observed in the Plio-Pleistocene records of the global carbon cycle. However they are absent in the Late Pleistocene ice-age record over the past 1.5 million years. Here, we present a simulation of global ice volume over the past 5 million years with a coupled system of four 3-D ice-sheet models. Our simulation shows that the 400-kyr long eccentricity cycles of Antarctica vary coherently with d13C records during the Pleistocene suggesting that they drive the long-term carbon cycle changes throughout the past 35 million years. The 400-kyr response of Antarctica is eventually suppressed by the dominant 100-kyr glacial cycles of the large ice sheets in the Northern Hemisphere (NH).