(Table S1) Uranium and Thorium isotopes and U/Th ages of IODP Exp310 cores from offshore Tahiti

The timing of sea-level change provides important constraints on the mechanisms driving Earth's climate between glacial and interglacial states. Fossil corals constrain the timing of past sea level by their suitability for dating and their growth position close to sea level. The coral-derived age for the last deglaciation is consistent with climate change forced by Northern Hemisphere summer insolation (NHI), but the timing of the penultimate deglaciation is more controversial. We found, by means of uranium/thorium dating of fossil corals, that sea level during the penultimate deglaciation had risen to ~85 meters below the present sea level by 137,000 years ago, and that it fluctuated on a millennial time scale during deglaciation. This indicates that the penultimate deglaciation occurred earlier with respect to NHI than the last deglacial, beginning when NHI was at a minimum.

Meiofaunal abundance, biomass, taxon richness and indices of nematode diversity of samples from the Santa Maria di Leuca Bank (southern Adriatic margin)

Deep-water coral ecosystems are hot spots of biodiversity and provide habitats and refuges for several deep-sea species. However, their role in shaping the biodiversity of the surrounding open slopes is still poorly known. We investigated how meiofaunal biodiversity varies with and is related to the occurrence of deep-water living scleractinian corals and coral rubble in two deep-sea areas (the Rockall Bank, northeastern Atlantic) and the Santa Maria di Leuca (central Mediterranean). In both areas, replicated sampling on alive and dead coral areas and from the adjacent slope sediments without corals (at the same and increasing depths) allowed us to demonstrate that sediments surrounding the living corals and coral rubble were characterised by higher meiofaunal biodiversity (as number of higher taxa, and nematode species richness) than the slope sediments. Despite the soft sediments surrounding the living coral having a higher nutritional value than those not associated with corals, with the opposite seen for coral rubble, the presence of both alive and dead corals had a significant effect on nematode assemblages. Our data suggest that, due particularly to the effects on habitat heterogeneity/complexity, both living coral and coral rubble promoted higher biodiversity levels than in surrounding slope sediments. We conclude that the protection of deep-water corals can be crucial to preserve the biodiversity of surrounding open slopes, and that the protection of dead corals, a so-far almost neglected habitat in terms of biological conservation, can further contribute to the maintenance of a high deep-sea biodiversity along continental margins.

Effect of ocean warming and acidification on the early life stages of subtropical Acropora spicifera

This study investigated the impacts of acidified seawater (pCO2 900 µatm) and elevated water temperature (+3 °C) on the early life history stages of Acropora spicifera from the subtropical Houtman Abrolhos Islands (28°S) in Western Australia. Settlement rates were unaffected by high temperature (27 °C, 250 µatm), high pCO2 (24 °C, 900 µatm), or a combination of both high temperature and high pCO2 treatments (27 °C, 900 µatm). There were also no significant differences in rates of post-settlement survival after 4 weeks of exposure between any of the treatments, with survival ranging from 60 to 70 % regardless of treatment. Similarly, calcification, as determined by the skeletal weight of recruits, was unaffected by an increase in water temperature under both ambient and high pCO2 conditions. In contrast, high pCO2 significantly reduced early skeletal development, with mean skeletal weight in the high pCO2 and combined treatments reduced by 60 and 48 %, respectively, compared to control weights. Elevated temperature appeared to have a partially mitigative effect on calcification under high pCO2; however, this effect was not significant. Our results show that rates of settlement, post-settlement survival, and calcification in subtropical corals are relatively resilient to increases in temperature. This is in marked contrast to the sensitivity to temperature reported for the majority of tropical larvae and recruits in the literature. The subtropical corals in this study appear able to withstand an increase in temperature of 3 °C above ambient, indicating that they may have a wider thermal tolerance range and may not be adversely affected by initial increases in water temperature from subtropical 24 to 27 °C. However, the reduction in skeletal weight with high pCO2 indicates that early skeletal formation will be highly vulnerable to the changes in ocean pCO2 expected to occur over the twenty-first century, with implications for their longer-term growth and resilience.

Glacial cold-water coral: ages, isotope concentrations and ratios

A set of 40 Uranium-series datings obtained on the reef-forming scleractinian cold-water corals Lophelia pertusa and Madrepora oculata revealed that during the past 400 kyr their occurrence in the Gulf of Cádiz (GoC) was almost exclusively restricted to glacial periods. This result strengthens the outcomes of former studies that coral growth in the temperate NE Atlantic encompassing the French, Iberian and Moroccan margins dominated during glacial periods, whereas in the higher latitudes (Irish and Norwegian margins) extended coral growth prevailed during interglacial periods. Thus it appears that the biogeographical limits for sustained cold-water coral growth along the NE Atlantic margin are strongly related to climate change. By focussing on the last glacial-interglacial cycle, this study shows that palaeo-productivity was increased during the last glacial. This was likely driven by the fertilisation effect of an increased input of aeolian dust and locally intensified upwelling. After the Younger Dryas cold event, the input of aeolian dust and productivity significantly decreased concurrent with an increase in water temperatures in the GoC. This primarily resulted in reduced food availability and caused a widespread demise of the formerly thriving coral ecosystems. Moreover, these climate induced changes most likely caused a latitudinal shift of areas withoptimum coral growth conditions towards the northern NE Atlantic where more suitable environmental conditions established with the onset of the Holocene.

Colony-specific calcification and mortality under ocean acidification in the branching coral Montipora digitata

Ocean acidification (OA) threatens calcifying marine organisms including reef-building corals. In this study, we examined the OA responses of individual colonies of the branching scleractinian coral Montipora digitata. We exposed nubbins of unique colonies (n = 15) to ambient or elevated pCO2 under natural light and temperature regimes for 110 days. Although elevated pCO2 exposure on average reduced calcification, individual colonies showed unique responses ranging from declines in positive calcification to negative calcification (decalcification) to no change. Similarly, mortality was greater on average in elevated pCO2, but also showed colony-specific patterns. High variation in colony responses suggests the possibility that ongoing OA may lead to natural selection of OA-tolerant colonies within a coral population.

(Table 5, page 206) The uranium contents of some manganese nodules

The uranium concentrations in marine calcareous material of a biological origin varied between 0.0X and 0.X p.p.m. with the exception of corals which had concentrations of several p.p.m. The aragonitic oolites and aragonite precipitated from sea-water had values similar to those of the corals. A geochronology based on the growth of ionium (thorium-230) from uranium is applicable not only to corals, as previous investigators have pointed out, but also to oolites. Several examples of "oolite ages" are given. The uranium content of ferromanganese minerals from pelagic deposits is of the order of from 4 to 5 p.p.m.

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)

Seismic reflection data, calcium/iron ratios, stable istopes, and magnetic susceptibility of sediment cores from the Pen Duick Escarpment, Gulf of Cadiz

Here we present a case study of three cold-water coral mounds in a juvenile growth stage on top of the Pen Duick Escarpment in the Gulf of Cadiz; Alpha, Beta and Gamma mounds. Although cold-water corals are a common feature on the adjacent cliffs, mud volcanoes and open slope, no actual living cold-water coral has been observed. This multidisciplinary and integrated study comprises geophysical, sedimentological and (bio)geochemical data and aims to present a holistic view on the interaction of both environmental and geological drivers in cold-water coral mound development in the Gulf of Cadiz. Coring data evidences (past or present) methane seepage near the Pen Duick Escarpment. Several sources and pathways are proposed, among which a stratigraphic migration through uplifted Miocene series underneath the escarpment. The dominant morphology of the escarpment has influenced the local hydrodynamics within the course of the Pliocene, as documented by the emplacement of a sediment drift. Predominantly during post-Middle Pleistocene glacial episodes, favourable conditions were present for mound growth. An additional advantage for mound formation near the top of Pen Duick Escarpment is presented by seepage-related carbonate crusts which might have offered a suitable substrate for coral settling. The spatially and temporally variable character and burial stage of the observed open reef frameworks, formed by cold-water coral rubble, provides a possible model for the transition from cold-water coral reef patches towards juvenile mound. These rubble "graveyards" not only act as sediment trap but also as micro-habitat for a wide range of organisms. The presence of a fluctuating Sulphate-Methane Transition Zone has an important effect on early diagenetic processes, affecting both geochemical and physical characteristics, transforming the buried reef into a solid mound. Nevertheless, the responsible seepage fluxes seem to be locally variable. As such, the origin and evolution of the cold-water coral mounds on top of the Pen Duick Escarpment is, probably more than any other NE Atlantic cold-water coral mound province, located on the crossroads of environmental (hydrodynamic) and geological (seepage) pathways.

(Table 1) Details on the Uranium/Thorium age determinations carried out on core GeoB11569-2

Cold-water corals are widely distributed along the Atlantic continental margin with varying growth patterns in relation to their specific environment. Here, we investigate the long-term development of cold-water corals that once thrived on a low-latitude (17°40'N) cold-water coral mound in the Banda Mound Province off Mauritania during the last glacial-interglacial cycle. U/Th dates obtained from 20 specimens of the cold-water coral Lophelia pertusa, revealed three distinct periods of coral growth during the last glacial at 65 to 57 kyr BP, 45 to 32 kyr BP and 14 kyr BP, thus comprising the cool periods of Marine Isotopic Stages (MIS) 2-4. These coral growth periods occur during periods of increased productivity in the region, emphasizing that productivity seems to be the major steering factor for coral growth off Mauritania, which is one of the major upwelling regions in the world. This pattern differs from the well studied coral mounds off Ireland, where the current regime predominantly influences the prosperity of the cold-water corals. Moreover, coral growth off Ireland takes place during rather warm interglacial and interstadial periods, whereas off Mauritania coral growth is restricted to glacial and stadial periods. However, the on-mound sedimentation patterns off Mauritania largely resemble the observations reported from the Irish mounds. The bulk of the preserved sediments derives from periods of coral growth, whereas during periods without corals hardly any net sedimentation or mound growth took place.

Seawater carbonate chemistry, calcification and dissolution response, and skeletal mineralogy of benthic orhanisms during experiments, 2011

Increasing atmospheric pCO2 reduces the saturation state of seawater with respect to the aragonite, high-Mg calcite (Mg/Ca > 0.04), and low-Mg calcite (Mg/Ca < 0.04) minerals from which marine calcifiers build their shells and skeletons. Notably, these polymorphs of CaCO3 have different solubilities in seawater: aragonite is more soluble than pure calcite, and the solubility of calcite increases with its Mg-content. Although much recent progress has been made investigating the effects of CO2-induced ocean acidification on rates of biological calcification, considerable uncertainties remain regarding impacts on shell/skeletal polymorph mineralogy. To investigate this subject, eighteen species of marine calcifiers were reared for 60-days in seawater bubbled with air-CO2 mixtures of 409 ± 6, 606 ± 7, 903 ± 12, and 2856 ± 54 ppm pCO2, yielding aragonite saturation states of 2.5 ± 0.4, 2.0 ± 0.4, 1.5 ± 0.3, and 0.7 ± 0.2. Calcite/aragonite ratios within bimineralic calcifiers increased with increasing pCO2, but were invariant within monomineralic calcifiers. Calcite Mg/Ca ratios (Mg/CaC) also varied with atmospheric pCO2 for two of the five high-Mg-calcite-producing organisms, but not for the low-Mg-calcite-producing organisms. These results suggest that shell/skeletal mineralogy within some-but not all-marine calcifiers will change as atmospheric pCO2 continues rising as a result of fossil fuel combustion and deforestation. Paleoceanographic reconstructions of seawater Mg/Ca, temperature, and salinity from the Mg/CaC of well-preserved calcitic marine fossils may also be improved by accounting for the effects of paleo-atmospheric pCO2 on skeletal Mg-fractionation.

Coral energy reserves and calcification in a high-CO2 world at two temperatures

Rising atmospheric CO2 concentrations threaten coral reefs globally by causing ocean acidification (OA) and warming. Yet, the combined effects of elevated pCO2 and temperature on coral physiology and resilience remain poorly understood. While coral calcification and energy reserves are important health indicators, no studies to date have measured energy reserve pools (i.e., lipid, protein, and carbohydrate) together with calcification under OA conditions under different temperature scenarios. Four coral species, Acropora millepora, Montipora monasteriata, Pocillopora damicornis, Turbinaria reniformis, were reared under a total of six conditions for 3.5 weeks, representing three pCO2 levels (382, 607, 741 µatm), and two temperature regimes (26.5, 29.0°C) within each pCO2 level. After one month under experimental conditions, only A. millepora decreased calcification (-53%) in response to seawater pCO2 expected by the end of this century, whereas the other three species maintained calcification rates even when both pCO2 and temperature were elevated. Coral energy reserves showed mixed responses to elevated pCO2 and temperature, and were either unaffected or displayed nonlinear responses with both the lowest and highest concentrations often observed at the mid-pCO2 level of 607 µatm. Biweekly feeding may have helped corals maintain calcification rates and energy reserves under these conditions. Temperature often modulated the response of many aspects of coral physiology to OA, and both mitigated and worsened pCO2 effects. This demonstrates for the first time that coral energy reserves are generally not metabolized to sustain calcification under OA, which has important implications for coral health and bleaching resilience in a high-CO2 world. Overall, these findings suggest that some corals could be more resistant to simultaneously warming and acidifying oceans than previously expected.

Seawater carbonate chemistry, larval settlement and growth rate during experiments with coral Porites astreoides, 2008

In response to the increases in pCO2 projected in the 21st century, adult coral growth and calcification are expected to decrease significantly. However, no published studies have investigated the effect of elevated pCO2 on earlier life history stages of corals. Porites astreoides larvae were collected from reefs in Key Largo, Florida, USA, settled and reared in controlled saturation state seawater. Three saturation states were obtained, using 1 M HCl additions, corresponding to present (380 ppm) and projected pCO2 scenarios for the years 2065 (560 ppm) and 2100 (720 ppm). The effect of saturation state on settlement and post-settlement growth was evaluated. Saturation state had no significant effect on percent settlement; however, skeletal extension rate was positively correlated with saturation state, with ~50% and 78% reductions in growth at the mid and high pCO2 treatments compared to controls, respectively.

Seawater carbonate chemistry and processes during experiments with coral Acropora eurystoma, 2006

The rise in atmospheric CO2 has caused significant decrease in sea surface pH and carbonate ion (CO3-2) concentration. This decrease has a negative effect on calcification in hermatypic corals and other calcifying organisms. We report the results of three laboratory experiments designed specifically to separate the effects of the different carbonate chemistry parameters (pH, CO3-2, CO2 [aq], total alkalinity [AT], and total inorganic carbon [CT]) on the calcification, photosynthesis, and respiration of the hermatypic coral Acropora eurystoma. The carbonate system was varied to change pH (7.9-8.5), without changing CT; CT was changed keeping the pH constant, and CT was changed keeping the pCO2 constant. In all of these experiments, calcification (both light and dark) was positively correlated with CO3-2 concentration, suggesting that the corals are not sensitive to pH or CT but to the CO3-2 concentration. A decrease of ~30% in the CO3-2 concentration (which is equivalent to a decrease of about 0.2 pH units in seawater) caused a calcification decrease of about 50%. These results suggest that calcification in today's ocean (pCO2 = 370 ppm) is lower by ~20% compared with preindustrial time (pCO2 = 280 ppm). An additional decrease of ~35% is expected if atmospheric CO2 concentration doubles (pCO2 = 560 ppm). In all of these experiments, photosynthesis and respiration did not show any significant response to changes in the carbonate chemistry of seawater. Based on this observation, we propose a mechanism by which the photosynthesis of symbionts is enhanced by coral calcification at high pH when CO2(aq) is low. Overall it seems that photosynthesis and calcification support each other mainly through internal pH regulation, which provides CO3-2 ions for calcification and CO2(aq) for photosynthesis.

Short-term metabolic and growth responses of the cold-water coral lophelia pertusa to ocean acidification

Cold-water corals are amongst the most three-dimensionally complex deep-sea habitats known and are associated with high local biodiversity. Despite their importance as ecosystem engineers, little is known about how these organisms will respond to projected ocean acidification. Since preindustrial times, average ocean pH has already decreased from 8.2 to ~ 8.1. Predicted CO2 emissions will decrease this by up to another 0.3 pH units by the end of the century. This decrease in pH may have a wide range of impacts upon marine life, and in particular upon calcifiers such as cold-water corals. Lophelia pertusa is the most widespread cold-water coral (CWC) species, frequently found in the North Atlantic. Data here relate to a short term data set (21 days) on metabolism and net calcification rates of freshly collected L. pertusa from Mingulay Reef Complex, Scotland. These data from freshly collected L. pertusa from the Mingulay Reef Complex will help define the impact of ocean acidification upon the growth, physiology and structural integrity of this key reef framework forming species.

Physiological and ecological variables measured at the high and low pCO2 reef sections

Experiments have shown that ocean acidification due to rising atmospheric carbon dioxide concentrations has deleterious effects on the performance of many marine organisms. However, few empirical or modelling studies have addressed the long-term consequences of ocean acidification for marine ecosystems. Here we show that as pH declines from 8.1 to 7.8 (the change expected if atmospheric carbon dioxide concentrations increase from 390 to 750 ppm, consistent with some scenarios for the end of this century) some organisms benefit, but many more lose out. We investigated coral reefs, seagrasses and sediments that are acclimatized to low pH at three cool and shallow volcanic carbon dioxide seeps in Papua New Guinea. At reduced pH, we observed reductions in coral diversity, recruitment and abundances of structurally complex framework builders, and shifts in competitive interactions between taxa. However, coral cover remained constant between pH 8.1 and ~7.8, because massive Porites corals established dominance over structural corals, despite low rates of calcification. Reef development ceased below pH 7.7. Our empirical data from this unique field setting confirm model predictions that ocean acidification, together with temperature stress, will probably lead to severely reduced diversity, structural complexity and resilience of Indo-Pacific coral reefs within this century.