Dissolution dominating calcification process in polar pteropods close to the point of aragonite undersaturation

Thecosome pteropods are abundant upper-ocean zooplankton that build aragonite shells. Ocean acidification results in the lowering of aragonite saturation levels in the surface layers, and several incubation studies have shown that rates of calcification in these organisms decrease as a result. This study provides a weight-specific net calcification rate function for thecosome pteropods that includes both rates of dissolution and calcification over a range of plausible future aragonite saturation states (Omega_Ar). We measured gross dissolution in the pteropod Limacina helicina antarctica in the Scotia Sea (Southern Ocean) by incubating living specimens across a range of aragonite saturation states for a maximum of 14 days. Specimens started dissolving almost immediately upon exposure to undersaturated conditions (Omega_Ar,0.8), losing 1.4% of shell mass per day. The observed rate of gross dissolution was different from that predicted by rate law kinetics of aragonite dissolution, in being higher at Var levels slightly above 1 and lower at Omega_Ar levels of between 1 and 0.8. This indicates that shell mass is affected by even transitional levels of saturation, but there is, nevertheless, some partial means of protection for shells when in undersaturated conditions. A function for gross dissolution against Var derived from the present observations was compared to a function for gross calcification derived by a different study, and showed that dissolution became the dominating process even at Omega_Ar levels close to 1, with net shell growth ceasing at an Omega_Ar of 1.03. Gross dissolution increasingly dominated net change in shell mass as saturation levels decreased below 1. As well as influencing their viability, such dissolution of pteropod shells in the surface layers will result in slower sinking velocities and decreased carbon and carbonate fluxes to the deep ocean

Shelled pteropods in peril: Assessing vulnerability in a high CO2 ocean

The impact of anthropogenic ocean acidification (OA) on marine ecosystems is a vital concern facing marine scientists and managers of ocean resources. Euthecosomatous pteropods (holoplanktonic gastropods) represent an excellent sentinel for indicating exposure to anthropogenic OA because of the sensitivity of their aragonite shells to the OA conditions less favorable for calcification. However, an integration of observations, experiments and modelling efforts is needed to make accurate predictions of how these organisms will respond to future changes to their environment. Our understanding of the underlying organismal biology and life history is far from complete and must be improved if we are to comprehend fully the responses of these organisms to the multitude of stressors in their environment beyond OA. This review considers the present state of research and understanding of euthecosomatous pteropod biology and ecology of these organisms and considers promising new laboratory methods, advances in instrumentation (such as molecular, trace elements, stable isotopes, palaeobiology alongside autonomous sampling platforms, CT scanning and high-quality video recording) and novel field-based approaches (i.e. studies of upwelling and CO2 vent regions) that may allow us to improve our predictive capacity of their vulnerability and/or resilience. In addition to playing a critical ecological and biogeochemical role, pteropods can offer a significant value as an early-indicator of anthropogenic OA. This role as a sentinel species should be developed further to consolidate their potential use within marine environmental management policy making

Interpretation and design of ocean acidification experiments in upwelling systems in the context of carbonate chemistry co-variation with temperature and oxygen

Coastal upwelling regimes are some of the most productive ecosystems in the ocean but are also among the most vulnerable to ocean acidification (OA) due to naturally high background concentrations of CO₂. Yet our ability to predict how these ecosystems will respond to additional CO₂ resulting from anthropogenic emissions is poor. To help address this uncertainty, researchers perform manipulative experiments where biological responses are evaluated across different CO₂ partial pressure (pCO₂) levels. In upwelling systems, however, contemporary carbonate chemistry variability remains only partly characterized and patterns of co-variation with other biologically important variables such as temperature and oxygen are just beginning to be explored in the context of OA experimental design. If co-variation among variables is prevalent, researchers risk performing OA experiments with control conditions that are not experienced by the focal species, potentially diminishing the ecological relevance of the experiment. Here, we synthesized a large carbonate chemistry dataset that consists of carbonate chemistry, temperature, and oxygen measurements from multiple moorings and ship-based sampling campaigns from the California Current Ecosystem (CCE), and includes fjord and tidal estuaries and open coastal waters. We evaluated patterns of pCO₂ variability and highlight important co-variation between pCO₂, temperature, and oxygen. We subsequently compared environmental pCO₂–temperature measurements with conditions maintained in OA experiments that used organisms from the CCE. By drawing such comparisons, researchers can gain insight into the ecological relevance of previously published OA experiments, but also identify species or life history stages that may already be influenced by contemporary carbonate chemistry conditions. We illustrate the implications co-variation among environmental variables can have for the interpretation of OA experimental results and suggest an approach for designing experiments with pCO₂ levels that better reflect OA hypotheses while simultaneously recognizing natural co-variation with other biologically relevant variables.Keywords: hypoxia, multistressor experiment, California Current, pH, climate chang

Systematic Review and Meta-Analysis Toward Synthesis of Thresholds of Ocean Acidification Impacts on Calcifying Pteropods and Interactions With Warming

Interpreting the vulnerability of pelagic calcifiers to ocean acidification (OA) is enhanced by an understanding of their critical thresholds and how these thresholds are modified by other climate change stressors (e.g., warming). To address this need, we undertook a three-part data synthesis for pteropods, one of the calcifying zooplankton group. We conducted the first meta-analysis and threshold analysis of literature characterizing pteropod responses to OA and warming by synthetizing dataset comprising of 2,097 datapoints. Meta-analysis revealed the extent to which responses among studies conducted on differing life stages and disparate geographies could be integrated into a common analysis. The results demonstrated reduced calcification, growth, development, and survival to OA with increased magnitude of sensitivity in the early life stages, under prolonged duration, and with the concurrent exposure of OA and warming, but not species-specific sensitivity. Second, breakpoint analyses identified OA thresholds for several endpoints: dissolution (mild and severe), calcification, egg development, shell growth, and survival. Finally, consensus by a panel of pteropod experts was used to verify thresholds and assign confidence scores for five endpoints with a sufficient signal: noise ratio to develop life-stage specific, duration-dependent thresholds. The range of aragonite saturation state from 1.5–0.9 provides a risk range from early warning to lethal impacts, thus providing a rigorous basis for vulnerability assessments to guide climate change management responses, including an evaluation of the efficacy of local pollution management. In addition, meta-analyses with OA, and warming shows increased vulnerability in two pteropod processes, i.e., shell dissolution and survival, and thus pointing toward increased threshold sensitivity under combined stressor effect

Biogeography and genetic diversity of the atlantid heteropods.

The atlantid heteropods are regularly encountered, but rarely studied marine planktonic gastropods. Relying on a small (<14 mm), delicate aragonite shell and living in the upper ocean means that, in common with pteropods, atlantids are likely to be affected by imminent ocean changes. Variable shell morphology and widespread distributions indicate that the family is more diverse than the 23 currently known species. Uncovering this diversity is fundamental to determining the distribution of atlantids and to understanding their environmental tolerances. Here we present phylogenetic analyses of all described species of the family Atlantidae using 437 new and 52 previously published cytochrome c oxidase subunit 1 mitochondrial DNA (mtCO1) sequences. Specimens and published sequences were gathered from 32 Atlantic Ocean stations, 14 Indian Ocean stations and 21 Pacific Ocean stations between 35°N and 43°S. DNA barcoding and Automatic Barcode Gap Discovery (ABGD) proved to be valuable tools for the identification of described atlantid species, and also revealed ten additional distinct clades, suggesting that the diversity within this family has been underestimated. Only two of these clades displayed obvious morphological characteristics, demonstrating that much of the newly discovered diversity is hidden from morphology-based identification techniques. Investigation of six large atlantid collections demonstrated that 61% of previously described (morpho) species have a circumglobal distribution. Of the remaining 39%, two species were restricted to the Atlantic Ocean, five occurred in the Indian and Pacific oceans, one species was only found in the northeast Pacific Ocean, and one occurred only in the Southern Subtropical Convergence Zone. Molecular analysis showed that seven of the species with wide distributions were comprised of two or more clades that occupied distinct oceanographic regions. These distributions may suggest narrower environmental tolerances than the described morphospecies. Results provide an updated biogeography and mtCO1 reference dataset of the Atlantidae that may be used to identify atlantid species and provide a first step in understanding their evolutionary history and accurate distribution, encouraging the inclusion of this family in future plankton research

Global Synthesis of the Status and Trends of Ocean Acidification Impacts on Shelled Pteropods

The accumulation of anthropogenic CO₂ in the ocean has major ecological, socioeconomic, and biogeochemical impacts, with repercussions for the ocean as a critical carbon sink. Ocean acidification (OA) disproportionately affects marine calcifiers, among which pelagic zooplanktonic pteropods play a significant role in carbonate export. The pteropod, due to the susceptibility of its aragonite shell to rapid dissolution, is one of most vulnerable groups and a key indicator for OA regional monitoring, but its regional sensitivities have not yet been extrapolated over global scales. To delineate spatial and temporal changes in pteropod shell dissolution, global OA status and the OA rate of change were evaluated, based on gridded climatologies of observations and using a Regional Ocean Modeling System (ROMS) biogeochemical/ecosystem model. Pteropods, which dominate in the polar and subpolar regions, are characterized by low aragonite saturation state and low buffering capacity, where extended pteropod subsurface dissolution is projected. We show that pteropods are most susceptible to OA in the polar regions, subpolar North Pacific, and eastern boundary upwelling system regions, particularly the California and Humboldt Current Systems. Rates of acidification and corresponding increases in pteropod shell dissolution are projected to be the fastest in the North and South Equatorial Currents. These are the regions with the greatest impacts on pteropods and biogeochemical implications.ISSN:1042-8275ISSN:2377-617