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Planktic foraminifera are key producers of pelagic carbonate, and their shell weight is suggested to have been influenced by the environment in which they calcify. However, there is debate about the use of size-normalised weight (SNW) as a proxy, as some authors invoke a carbonate system control on calcification (and by extension SNW as a pCO2 proxy), while others suggest that species optimum conditions, nutrient concentration, or temperature drive shell weight. To better understand this proxy, we investigate what drives SNW and whether discrepancies in the proposed control on weight are due to differing data collection methodologies and/or regionally different drivers. We integrate new and published SNW data with environmental hindcast data from the CMIP6 modelling suite. Using Bayesian regression modelling, we find that the environment alone does not explain the variability in SNW across species. Although physiology likely modulates the response to the environment, we find little evidence of a unifying driver at the ecogroup level. Instead, we identify species-specific responses associated with drivers including (but not limited to) the carbonate system, which are likely different between ocean basins. We hypothesise that this is partly influenced by cryptic species and regional phenotypic plasticity in changes to shell weight that are not well understood, such as the thickness of calcite deposited during some species' reproductive phases. Consequently, which species to use as a pCO2 proxy or whether multiple species should be used in parallel to reduce uncertainty should be carefully considered. We strongly encourage the regional testing and calibration of pCO2–SNW relationships.

The West Antarctic Peninsula (WAP) exhibits strong spatial and temporal oceanographic variability, resulting in highly heterogeneous biological productivity. Calcifying organisms that live in the waters off the WAP respond to temporal and spatial variations in ocean temperature and chemistry. These marine calcifiers are potentially threatened by regional climate change with waters already naturally close to carbonate undersaturation. Future projections of carbonate production in the Southern Ocean are challenging due to the lack of historical data collection and complex, decadal climate variability. Here we present a 6-year-long record of the shell fluxes, morphology and stable isotope variability of the polar planktic foraminifera Neogloboquadrina pachyderma (sensu stricto) from near Palmer Station, Antarctica. This species is fundamental to Southern Ocean planktic carbonate production as it is one of the very few planktic foraminifer species adapted to the marine polar environments. We use these new data to obtain insights into its ecology and to derive a robust assessment of the response of this polar species to environmental change. Morphology and stable isotope composition reveal the presence of different growth stages within this tightly defined species. Inter- and intra-annual variability of foraminiferal flux and size is evident and driven by a combination of environmental forcing parameters, most importantly food availability, temperature and sea ice duration and extent. Foraminiferal growth occurs throughout the austral year and is influenced by environmental change, a large portion of which is driven by the Southern Annular Mode and El Niño–Southern Oscillation. A distinct seasonal production is observed, with the highest shell fluxes during the warmest and most productive months of the year. The sensitivity of calcifying foraminifera to environmental variability in this region, from weeks to decades, has implications both for their response to future climatic change and for their use as palaeoclimate indicators. A longer ice-free season could increase carbonate production in this region at least while carbonate saturation is still high enough to allow for thick tests to grow.

Despite the important role of planktonic foraminifera in regulating the ocean carbonate production and their unrivalled value in reconstructing paleoenvironments, our knowledge on their ecology is limited. A variety of observational techniques such as plankton tows, sediment traps and experiments have contributed to our understanding of foraminifera ecology. But, fundamental questions around costs and benefits of calcification and the effect of nutrients, temperature and ecosystem structure on these organisms remain unanswered. To tackle these questions, we take a novel mechanistic approach to study planktonic foraminifera ecology based on trait theory. We develop a zero-dimensional (0-D) trait-based model to account for the biomass of prolocular (20 µm) and adult (160 µm) stages of non-spinose foraminifera species and investigate their potential interactions with phytoplankton and other zooplankton under different temperature and nutrient regimes. Building on the costs and benefits of calcification, we model two ecosystem structures to explore the effect of resource competition and temperature on planktonic foraminifera biomass. By constraining the model results with ocean biomass estimations of planktonic foraminifera, we estimate that the energetic cost of calcification could be about 10 %–50 % and 10 %–40 % for prolocular and adult stages respectively. Our result suggest that the shell provides protection against processes other than predation (e.g. pathogen invasion). We propose that the low standing biomass of planktonic foraminifera plays a key role in their survival from predation, along with their shell protection. Our model suggests a shift from temperature as a main regulator of foraminifera biomass in the early developmental stage to resource competition for adult biomass.

Cold-water corals are important habitat formers in deep-water ecosystems and at high latitudes. Ocean acidification and the resulting change in aragonite saturation are expected to affect these habitats and impact coral growth. Counter to expectations, the deep water coral Lophelia pertusa has been found to be able to sustain growth even in undersaturated conditions. However, it is important to know whether such undersaturation modifies the skeleton and thus its ecosystem functioning. Here we used Synchrotron X-Ray Tomography and Raman spectroscopy to examine changes in skeleton morphology and fibre orientation. We combined the morphological assessment with boron isotope analysis to determine if changes in growth are related to changes in control of calcification pH. We compared the isotopic composition and structure formed in their natural environment to material grown in culture at lower pH conditions. Skeletal morphology is highly variable but shows no distinctive differences between natural and low pH conditions. Raman investigations found no difference in macromorphological skeletal arrangement of early mineralization zones and secondary thickening between the treatments. The δ11B analyses show that L. pertusa up-regulates the internal calcifying fluid pH (pHcf) during calcification compared to ambient seawater pH and maintains a similar elevated pHcf at increased pCO2 conditions. We suggest that as long as the energy is available to sustain the up-regulation, i.e. individuals are well fed, there is no detrimental effect to the skeletal morphology.

Coralline algae are important habitat formers found on all rocky shores. While the impact of future ocean acidification on the physiological performance of the species has been well studied, little research has focused on potential changes in structural integrity in response to climate change. A previous study using 2-D Finite Element Analysis (FEA) suggested increased vulnerability to fracture (by wave action or boring) in algae grown under high CO2 conditions. To assess how realistically 2-D simplified models represent structural performance, a series of increasingly biologically accurate 3-D FE models that represent different aspects of coralline algal growth were developed. Simplified geometric 3-D models of the genus Lithothamnion were compared to models created from computed tomography (CT) scan data of the same genus. The biologically accurate model and the simplified geometric model representing individual cells had similar average stresses and stress distributions, emphasising the importance of the cell walls in dissipating the stress throughout the structure. In contrast models without the accurate representation of the cell geometry resulted in larger stress and strain results. Our more complex 3-D model reiterated the potential of climate change to diminish the structural integrity of the organism. This suggests that under future environmental conditions the weakening of the coralline algal skeleton along with increased external pressures (wave and bioerosion) may negatively influence the ability for coralline algae to maintain a habitat able to sustain high levels of biodiversity.

Magnesium is an element critically involved in the carbon cycle, because weathering of Ca-Mg silicates removes atmospheric CO2 into rivers, and formation of Ca-Mg carbonates in the oceans removes carbon from the ocean-atmosphere system. Hence the Mg cycle holds the potential to provide valuable insights into Cenozoic climate-system history, and the shift during this time from a greenhouse to icehouse state. We present Mg isotope ratios for the past 40 Myr using planktic foraminifers as an archive. Modern foraminifera, which discriminate against elemental and isotopically heavy Mg during calcification, show no correlation between the Mg isotope composition (δ26Mg, relative to DSM-3) and temperature, Mg / Ca or other parameters such as carbonate saturation (ΔCO3). However, inter-species isotopic differences imply that only well-calibrated single species should be used for reconstruction of past seawater. Seawater δ26Mg inferred from the foraminiferal record decreased from ~0‰ at 15 Ma, to −0.83‰ at the present day, which coincides with increases in seawater lithium and oxygen isotope ratios. It strongly suggests that neither Mg concentrations nor isotope ratios are at steady state in modern oceans, given its ~10 Myr residence time. From these data, we have developed a dynamic box model to understand and constrain changes in Mg sources to the oceans (rivers) and Mg sinks (dolomitisation and hydrothermal alteration). Our estimates of seawater Mg concentrations through time are similar to those independently determined by pore waters and fluid inclusions. Modelling suggests that dolomite formation and the riverine Mg flux are the primary controls on the δ26Mg of seawater, while hydrothermal Mg removal and the δ26Mg of rivers are more minor controls. Using Mg riverine flux and isotope ratios inferred from the 87Sr / 86Sr record, the modelled Mg removal by dolomite formation shows minima in the Oligocene and at the present day (with decreasing trends from 15 Ma), both coinciding with rapid decreases in global temperatures.

As a result of anthropogenic pCO2 increases, future oceans are growing warmer and lower in pH and oxygen, conditions that are likely to impact planktic communities. Past intervals of elevated and changing pCO2 and temperatures can offer a glimpse into the response of marine calcifying plankton to changes in surface oceans under conditions similar to those projected for the future. Here we present new records of planktic foraminiferal and coccolith calcification (weight and size) from Deep Sea Drilling Project Site 607 (mid-North Atlantic) and Ocean Drilling Program Site 999 (Caribbean Sea) from the Pliocene, the last time that pCO2 was similar to today, and extending through a global cooling event into the intensification of Northern Hemisphere glaciation (3.3 to 2.6 million years ago). Test weights of both surface-dwelling Foraminifera Globigerina bulloides and thermocline-dwelling Foraminifera Globorotalia puncticulata vary with a potential link to regional temperature variation in the North Atlantic, whereas in the tropics Globigerinoides ruber test weight remains stable. In contrast, reticulofenestrid coccoliths show a narrowing size range and a decline in the largest lith diameters over this interval. Our results suggest no major changes in plankton calcite production during the high pCO2 Pliocene or during the transition into an icehouse world.

The observed variation in the calcification responses of coccolithophores to changes in carbonate chemistry paints a highly incoherent picture, particularly for the most commonly cultured "species", Emiliania huxleyi. The disparity between magnitude and potentially even sign of the calcification change under simulated end-of-century ocean surface chemical changes (higher pCO2, lower pH and carbonate saturation), raises challenges to quantifying future carbon cycle impacts and feedbacks because it introduces significant uncertainty in parameterizations used for global models. Here we compile the results of coccolithophore carbonate chemistry manipulation experiments and review how ocean carbon cycle models have attempted to bridge the gap from experiments to global impacts. Although we can rule out methodological differences in how carbonate chemistry is altered as introducing an experimental bias, the absence of a consistent calcification response implies that model parameterizations based on small and differing subsets of experimental observations will lead to varying estimates for the global carbon cycle impacts of ocean acidification. We highlight two pertinent observations that might help: (1) the degree of coccolith calcification varies substantially, both between species and within species across different genotypes, and (2) the calcification response across mesocosm and shipboard incubations has so-far been found to be relatively consistent. By analogy to descriptions of plankton growth rate vs. temperature, such as the "Eppley curve", which seek to encapsulate the net community response via progressive assemblage change rather than the response of any single species, we posit that progressive future ocean acidification may drive a transition in dominance from more to less heavily calcified coccolithophores. Assemblage shift may be more important to integrated community calcification response than species-specific response, highlighting the importance of whole community manipulation experiments to models in the absence of a complete physiological understanding of the underlying calcification process. However, on a century time-scale, regardless of the parameterization adopted, the atmospheric pCO2 impact of ocean acidification is minor compared to other global carbon cycle feedbacks.

The impacts of climate change on marine organisms have been increasingly documented in laboratory and experimental studies. However, the use of different taxonomic groupings and the assessment of a range of processes make identifying overall trends challenging. Meta-analysis has been used to determine general trends, but coarse taxonomic granularity may mask phylogenetically specific responses. Bivalve molluscs are a data-rich clade of ecologically and economically important calcifying marine taxa that allow for the assessment of species-specific vulnerability across developmental stages. Drawing on the large body of available literature, we conduct a meta-analysis of 203 unique experimental set-ups in order to examine how bivalve growth responds to increased water temperature, acidity, deoxygenation, and changes in salinity in 10 climate change stressor combinations. This is the most complete examination of bivalve responses to date and shows that anthropogenic climate change will disproportionally affect particular families, suggesting taxonomic differentiation in climate change response. Specifically, Mytilidae, Ostreidae, and Pectinidae (67 % of experiments) respond with negative effect sizes for all individual stressors, whereas responses in Pinnidae, Tellinidae, and Veneridae are more complex. Our analysis shows that earlier studies reporting negative impacts on bivalves are driven by only three or four well-studied, commercially important families. Despite the taxonomic differentiation, almost all drivers and their combinations have significant negative effects on growth. The synergistic impacts of deoxygenation, acidification, and temperature result in the largest negative effect size. Infaunal taxa, including Tellinidae and Veneridae, appear more resistant to warming and oxygen reduction than epifaunal or motile taxa, but this difference between the two taxa is also based on a small number of data points. The current focus of experimental set-ups on commercially important taxa and families within a small geographic range creates gaps in the understanding of global impacts on these economically important foundation organisms.