Climate change is altering global ocean phenology, the timing of annually occurring biological events. We examined the changing phenology of the phytoplankton accumulation season west of the Antarctic Peninsula to show that blooms are shifting later in the season over time in ice-associated waters. The timing of the start date and peak date of the phytoplankton accumulation season occurred later over time from 1997 to 2022 in the marginal ice zone and over the continental shelf. A divergence was seen between offshore waters and ice-associated waters, with offshore bloom timing becoming earlier, yet marginal ice zone and continental shelf bloom timing shifting later. Higher chlorophyll a (chl a) concentration in the fall season was seen in recent years, especially over the northern continental shelf. Minimal long-term trends in annual chl a occurred, likely due to the combination of later start dates in spring and higher chl a in fall. Increasing spring wind speed is the most likely mechanism for later spring start dates, leading to deeper wind mixing in a region experiencing sea ice loss. Later phytoplankton bloom timing over the marginal ice zone and continental shelf will have consequences for surface ocean carbon uptake, food web dynamics, and trophic cascades.

AbstractOrganic carbon (OC) sedimentation in marine sediments is the largest long‐term sink of atmospheric CO2 after silicate weathering. Understanding the mechanistic and quantitative aspects of OC delivery and preservation in marine sediments is critical for predicting the role of the oceans in modulating global climate. Yet, estimates of the global OC sedimentation in marginal settings span an order of magnitude, and the primary controls of OC preservation remain highly debated. Here, we provide the first global bottom‐up estimate of OC sedimentation along the margins using a synthesis of literature data. We quantify both terrestrial‐ and marine‐sourced OC fluxes and perform a statistical analysis to discern the key factors influencing their magnitude. We find that the margins host 23.2 ± 3.5 Tmol of OC sedimentation annually, with approximately 84% of marine origin. Accordingly, we calculate that only 2%–3% of OC exported from the euphotic zone escapes remineralization before sedimentation. Surprisingly, over half of all global OC sedimentation occurs below bottom waters with oxygen concentrations greater than 180 μM, while less than 4% occurs in settings with <50 μM oxygen. This challenges the prevailing paradigm that bottom‐water oxygen (BWO) is the primary control on OC preservation. Instead, our statistical analysis reveals that water depth is the most significant predictor of OC sedimentation, surpassing all other factors investigated, including BWO levels and sea‐surface chlorophyll concentrations. This finding suggests that the primary control on OC sedimentation is not production, but the ability of OC to resist remineralization during transit through the water column and while settling on the seafloor.

Climate change is leading to phenological shifts across a wide range of species globally. Polar oceans are hotspots of rapid climate change where sea ice dynamics structure ecosystems and organismal life cycles are attuned to ice seasonality. To anticipate climate change impacts on populations and ecosystem services, it is critical to understand ecosystem phenology to determine species activity patterns, optimal environmental windows for processes like reproduction, and the ramifications of ecological mismatches. Since 1991, the Palmer Antarctica Long-Term Ecological Research (LTER) program has monitored seasonal dynamics near Palmer Station. Here, we review the species that occupy this region as year-round residents, seasonal breeders, or periodic visitors. We show that sea ice retreat and increasing photoperiod in the spring trigger a sequence of events from mid-November to mid-February, including Adelie penguin clutch initiation, snow melt, calm conditions (low winds and warm air/sea temperature), phytoplankton blooms, shallow mixed layer depths, particulate organic carbon flux, peak humpback whale abundances, nutrient drawdown, and bacterial accumulation. Subsequently, from May to June, snow accumulates, zooplankton indicator species appear, and sea ice advances. The standard deviation in the timing of most events ranged from similar to 20 to 45 days, which was striking compared with Adelie penguin clutch initiation that varied < 1 week. In general, during late sea ice retreat years, events happened later (similar to 5 to > 30 days) than mean dates and the variability in timing was low (< 20%) compared with early ice retreat years. Statistical models showed the timing of some events were informative predictors (but not sole drivers) of other events. From an Adelie penguin perspective, earlier sea ice retreat and shifts in the timing of suitable conditions or prey characteristics could lead to mismatches, or asynchronies, that ultimately influence chick survival via their mass at fledging. However, more work is needed to understand how phenological shifts affect chick thermoregulatory costs and the abundance, availability, and energy content of key prey species, which support chick growth and survival. While we did not detect many long-term phenological trends, we expect that when sea ice trends become significant within our LTER time series, phenological trends and negative effects from ecological mismatches will follow.

Barium is widely used as a proxy for dissolved silicon and particulate organic carbon fluxes in seawater. However, these proxy applications are limited by insufficient knowledge of the dissolved distribution of Ba ([Ba]). For example, there is significant spatial variability in the barium-silicon relationship, and ocean chemistry may influence sedimentary Ba preservation. To help address these issues, we developed 4095 models for predicting [Ba] using Gaussian process regression machine learning. These models were trained to predict [Ba] from standard oceanographic observations using GEOTRACES data from the Arctic, Atlantic, Pacific, and Southern oceans. Trained models were then validated by comparing predictions against withheld [Ba] data from the Indian Ocean. We find that a model trained using depth, temperature, and salinity, as well as dissolved dioxygen, phosphate, nitrate, and silicate, can accurately predict [Ba] in the Indian Ocean with a mean absolute percentage deviation of 6.0 %. We use this model to simulate [Ba] on a global basis using these same seven predictors in the World Ocean Atlas. The resulting [Ba] distribution constrains the Ba budget of the ocean to 122(+/- 7) x 10(12) mol and reveals oceanographically consistent variability in the barium-silicon relationship. We then calculate the saturation state of seawater with respect to barite. This calculation reveals systematic spatial and vertical variations in marine barite saturation and shows that the ocean below 1000 m is at equilibrium with respect to barite. We describe a number of possible applications for our model outputs, ranging from use in mechanistic biogeochemical models to paleoproxy calibration. Our approach demonstrates the utility of machine learning in accurately simulating the distributions of tracers in the sea and provides a framework that could be extended to other trace elements. Our model, the data used in training and validation, and global outputs are available in Horner and Mete (2023, 10.26008/1912/bco-dmo.885506.2).

Marine heterotrophic Bacteria (or referred to as bacteria) play an important role in the ocean carbon cycle by utilizing, respiring, and remineralizing organic matter exported from the surface to deep ocean. Here, we investigate the responses of bacteria to climate change using a three-dimensional coupled ocean biogeochemical model with explicit bacterial dynamics as part of the Coupled Model Intercomparison Project Phase 6. First, we assess the credibility of the century-scale projections (2015-2099) of bacterial carbon stock and rates in the upper 100 m layer using skill scores and compilations of the measurements for the contemporary period (1988-2011). Second, we demonstrate that across different climate scenarios, the simulated bacterial biomass trends (2076-2099) are sensitive to the regional trends in temperature and organic carbon stocks. Bacterial carbon biomass declines by 5-10% globally, while it increases by 3-5% in the Southern Ocean where semi-labile dissolved organic carbon (DOC) stocks are relatively low and particle-attached bacteria dominate. While a full analysis of drivers underpinning the simulated changes in all bacterial stock and rates is not possible due to data constraints, we investigate the mechanisms of the changes in DOC uptake rates of free-living bacteria using the first-order Taylor decomposition. The results demonstrate that the increase in semi-labile DOC stocks drives the increase in DOC uptake rates in the Southern Ocean, while the increase in temperature drives the increase in DOC uptake rates in the northern high and low latitudes. Our study provides a systematic analysis of bacteria at global scale and a critical step toward a better understanding of how bacteria affect the functioning of the biological carbon pump and partitioning of organic carbon pools between surface and deep layers.

Heterotrophic marine bacteria utilize organic carbon for growth and biomass synthesis. Thus, their physiological variability is key to the balance between the production and consumption of organic matter and ultimately particle export in the ocean. Here we investigate a potential link between bacterial traits and ecosystem functions in the rapidly warming West Antarctic Peninsula (WAP) region based on a bacteria-oriented ecosystem model. Using a data assimilation scheme, we utilize the observations of bacterial groups with different physiological traits to constrain the group-specific bacterial ecosystem functions in the model. We then examine the association of the modeled bacterial and other key ecosystem functions with eight recurrent modes representative of different bacterial taxonomic traits. Both taxonomic and physiological traits reflect the variability in bacterial carbon demand, net primary production, and particle sinking flux. Numerical experiments under perturbed climate conditions demonstrate a potential shift from low nucleic acid bacteria to high nucleic acid bacteria-dominated communities in the coastal WAP. Our study suggests that bacterial diversity via different taxonomic and physiological traits can guide the modeling of the polar marine ecosystem functions under climate change.

Abstract. The West Antarctic Peninsula (WAP) is a rapidly warming region, with substantial ecological and biogeochemical responses to climate change and variability for the past decades, revealed by multi-decadal observations from the Palmer Antarctica Long-Term Ecological Research (LTER) program. The wealth of these long-term observations provides an important resource for ecosystem modelling, but there has been a lack of focus on the development of numerical models that simulate time-evolving plankton dynamics over the Austral growth season along the coastal WAP. Here we developed a one-dimensional, data assimilation planktonic ecosystem model (i.e., the WAP-1D-VAR model v1.0) equipped with a variational adjoint and model parameter optimization scheme. We first demonstrate the modified and newly added model schemes to the pre-existing food-web and biogeochemical components of the WAP-1D-VAR model, including diagnostic sea-ice forcing and trophic interactions specific to the WAP region. We then conducted model experiments by assimilating eleven different data types from an example Palmer LTER growth season (October 2002–March 2003) directly related to corresponding model state variables and intercompartmental flows. The iterative, data assimilation procedure reduced by 80 % the misfits between observations and model results, compared to before optimization, via an optimized set of 14 parameters out of total 72 free parameters. The optimized model results captured key WAP ecological features, such as blooms during seasonal sea-ice retreat, the lack of macronutrient limitation, and comparable values of the assimilated and non-assimilated model state variables and flows to other studies, as well as several important ecosystem metrics. One exception was slightly underestimated particle export flux, for which we discuss fully potential underlying reasons. The data assimilation scheme of the WAP-1D-VAR model enabled the available observational data to constrain previously poorly understood processes, including the partitioning of primary production by different phytoplankton groups, the optimal chlorophyll to carbon ratio of the WAP phytoplankton community, and the partitioning of dissolved organic carbon pools with different lability. The WAP-1D-VAR model was successfully employed to glue the snapshots from a range of the available data sets together to explain and understand the observed dynamics along the coastal WAP.

Figure S1: Agreement between observations and model output before and after optimization for 2010-11.Error as the difference and 5 1:1 agreement between optimized model outputs (blue dots) and model outputs before optimization (red dots) for each data type for 2010-11.

Abstract. Heterotrophic marine bacteria utilize organic carbon for growth and biomass synthesis. Thus, their variability is key to the balance between the production and consumption of organic matter and ultimately particle export in the ocean. Here we investigate a potential link between bacterial traits and ecosystem functions in a rapidly changing polar marine ecosystem based on a bacteria-oriented ecosystem model. Using a data-assimilation scheme we utilize the observations of bacterial groups with different physiological states to constrain the group-specific bacterial ecosystem functions. We also investigate the association of the modelled bacterial and other ecosystem functions with eight recurrent modes representative of different bacterial taxonomic traits. High nucleic acid (HNA) bacteria show relatively high cell-specific bacterial production, respiration, and utilization of the semi-labile dissolved organic carbon pool compared to low nucleic acid (LNA) bacteria. Both taxonomy and physiological states of the bacteria are strong predictors of bacterial carbon demand, net primary production, and particle export. Numerical experiments under perturbed climate conditions show overall increased bacterial activity and a potential shift from LNA- to HNA-dominated bacterial communities in a warming ocean. Microbial diversity via different taxonomic and physiological traits informs our ecosystem model, providing insights into key bacterial and ecosystem functions in a changing environment.

Analysis of plankton data sets from the Bermuda Atlantic Time‐series Study site (31°40′N, 64°10′W) demonstrates a long‐term decreasing trend of marine heterotrophic bacterial production (BP) in winter (November–February) over the past 24 years (1989–2012). We propose winter extratropical cyclones (storms) as a potential driver of the observed BP trend. Stormy winters were characterized by significantly enhanced phytoplankton biomass and productivity as well as organic carbon pools, as storms induce deeper convective mixing and increased nitrate fluxes into upper mixed‐layer. Counter to expectation, BP was lower by 24% due to storm‐mediated entrainment of cold water into upper mixed‐layer. The frequency of storm‐impacted months has increased over time, driven by a decline of the North Atlantic Oscillation (NAO) index. Our findings suggest that a NAO‐induced southward shift of winter storm tracks contributes to the decreasing BP trend, via temperature‐dependent decoupling of bacteria from phytoplankton, ultimately impacting carbon fluxes and cycling near Bermuda.

Ever since marine heterotrophic bacteria were understood to mediate carbon fluxes to upper trophic levels via the microbial loop, quantifying the coupling between bacteria and phytoplankton has been one of the main priorities for marine microbiologists. The complex nature of phytoplankton-bacterial interactions may lead to nonlinear and regime-dependent coupling, for which conventional statistical approaches such as cross-correlation provide an incomplete picture of the dynamics. Here, we employed a nonlinear method for detecting causality, called convergent cross mapping (CCM), to examine a causal linkage between phytoplankton (primary production) and bacteria (bacterial production) at the Bermuda Atlantic Time-series Study (BATS) site. First, we verified the robustness of our CCM models with synthetic time series output from the Fasham-Ducklow-McKelvie ecosystem model. Initially, we hypothesized a strong bi-directional causal link between phytoplankton and bacteria due to the importance of the microbial loop at BATS. However, the results from our CCM models highlight that phytoplankton-bacterial causal coupling depends on seasonally distinct mixing regimes. While there was no evidence for bi-directional causality between phytoplankton and bacteria during the mixing period, moderate unidirectional causality of bacteria on phytoplankton was detected during the stratification period. Our study reveals causal associations between the 2 major microbial loop processes, for which better quantification is needed to improve our understanding of carbon cycling and export via microbial food webs.

The West Antarctic Peninsula (WAP) is a climatically sensitive region where periods of strong warming have caused significant changes in the marine ecosystem and food-web processes. Tight coupling between phytoplankton and higher trophic levels implies that the coastal WAP is a bottom-up controlled system, where changes in phytoplankton dynamics may largely impact other food-web components. Here, we analysed the inter-decadal time series of year-round chlorophyll-a (Chl) collected from three stations along the coastal WAP: Carlini Station at Potter Cove (PC) on King George Island, Palmer Station on Anvers Island and Rothera Station on Adelaide Island. There were trends towards increased phytoplankton biomass at Carlini Station (PC) and Palmer Station, while phytoplankton biomass declined significantly at Rothera Station over the studied period. The impacts of two relevant climate modes to the WAP, the El Niño-Southern Oscillation and the Southern Annular Mode, on winter and spring phytoplankton biomass appear to be different among the three sampling stations, suggesting an important role of local-scale forcing than large-scale forcing on phytoplankton dynamics at each station. The inter-annual variability of seasonal bloom progression derived from considering all three stations together captured ecologically meaningful, seasonally co-occurring bloom patterns which were primarily constrained by water-column stability strength. Our findings highlight a coupled link between phytoplankton and physical and climate dynamics along the coastal WAP, which may improve our understanding of overall WAP food-web responses to climate change and variability.This article is part of the theme issue 'The marine system of the West Antarctic Peninsula: status and strategy for progress in a region of rapid change'.

The West Antarctic Peninsula (WAP) is a climatically-sensitive region where periods of strong warming have caused significant changes in marine ecosystem and food web processes. Tight coupling between phytoplankton and higher trophic levels implies that the coastal WAP is a bottom-up controlled system, where changes in phytoplankton dynamics may largely impact other food web components. Here, we analyzed the interdecadal time series of year-round chlorophyll-a (Chl) collected from three stations along the coastal WAP, Carlini Station at Potter Cove (PC) on King George Island, Palmer Station on Anvers Island, and Rothera Station on Adelaide Island. There were trends toward increased phytoplankton biomass at Carlini Station (PC) and Palmer Station, while phytoplankton biomass declined significantly at Rothera Station over the studied period. The impacts of two relevant climate modes to the WAP, the El Nino-Southern Oscillation and the Southern Annular Mode, on winter and spring phytoplankton biomass appear to be different among three sampling stations, suggesting a possibly more important role of local-scale forcing than large-scale forcing on phytoplankton dynamics at each station. The interannual variability of seasonal bloom progression derived from considering all three stations together captured ecologically meaningful, seasonally co-occurring bloom patterns which were primarily constrained by water-column stability strength. Our findings highlight a coupled link between phytoplankton and physical and climate dynamics along the coastal WAP, understanding of which is crucial in predicting overall WAP food web responses to climate change and variability.

We investigated the dynamics of heterotrophic bacteria in the coastal western Antarctic Peninsula (WAP), using decadal (2002-2014) time series of two bacterial variables, bacterial production (BP) via 3H-leucine incorporation rates and bacterial biomass (BB) via bacterial abundance, collected at Palmer Antarctica Long Term Ecological Research (LTER) Station B (64.8°S, 64.1°W) over a full austral growing season (October-March). Strong seasonal and interannual variability in the degree of bacterial coupling with phytoplankton processes were observed with varying lags. On average, BP was only 4% of primary production (PP), consistent with low BP:PP ratios observed in polar waters. BP was more strongly correlated with chlorophyll (Chl), than with PP, implying that bacteria feed on DOC produced from a variety of trophic levels (e.g. zooplankton sloppy feeding and excretion) as well as directly on phytoplankton-derived DOC. The degree of bottom-up control on bacterial abundance was moderate and relatively consistent across entire growing seasons, suggesting that bacteria in the coastal WAP are under consistent DOC limitation. Temperature also influenced BP rates, though its effect was weaker than DOC. We established generalized linear models (GLMs) for monthly composites of BP and BB via stepwise regression to explore a set of physical and biogeochemical predictors. Physically, high BP and large BB were shaped by a stratified water-column, similar to forcing mechanisms favoring phytoplankton blooms, but high sea surface temperature (SST) also significantly promoted bacterial processes. High BP and large BB were influenced by high PP and bulk DOC concentrations. Based on these findings, we suggest an increasingly important role of marine heterotrophic bacteria in the coastal WAP food-web as climate change introduces a more favorable environmental setting for promoting BP, with increased DOC from retreating glaciers, a more stabilized upper water-column from ice-melt, and a baseline shift of water temperature due to more frequent delivery of warming Upper Circumpolar Deep Water (UCDW) onto the WAP shelf.

Offshore geologic storage of carbon dioxide (CO2), known as offshore carbon capture and sequestration (CCS), has been under active investigation as a safe, effective mitigation option for reducing CO2 levels from anthropogenic fossil fuel burning and climate change. Along with increasing trends in implementation plans and related logistics on offshore CCS, thorough risk assessment (i.e. environmental impact monitoring) needs to be conducted to evaluate potential risks, such as CO2 gas leakage at injection sites. Gas leaks from offshore CCS may affect the physiology of marine organisms and disrupt certain ecosystem functions, thereby posing an environmental risk. Here, we synthesize current knowledge on environmental impact monitoring of offshore CCS with an emphasis on biological aspects and provide suggestions for better practice. Based on our critical review of preexisting literatures, this paper: 1) discusses key variables sensitive to or indicative of gas leakage by summarizing physico-chemical and ecological variables measured from previous monitoring cruises on offshore CCS; 2) lists ecosystem and organism responses to a similar environmental condition to CO2 leakage and associated impacts, such as ocean acidification and hypercapnia, to predict how they serve as responsive indicators of short- and long-term gas exposure, and 3) discusses the designs of the artificial gas release experiments in fields and the best model simulation to produce realistic leakage scenarios in marine ecosystems. Based on our analysis, we suggest that proper incorporation of biological aspects will provide successful and robust long-term monitoring strategies with earlier detection of gas leakage, thus reducing the risks associated with offshore CCS.

We analyzed 20 years (1993–2013) of observations of dissolved inorganic macronutrients (nitrate, N; phosphate, P; and silicate, Si) and chlorophyll a (Chl) at Palmer Station, Antarctica (64.8°S, 64.1°W) to elucidate how large‐scale climate and local physical forcing affect the interannual variability in the seasonal phytoplankton bloom and associated drawdown of nutrients. The leading modes of nutrients (N, P, and Si empirical orthogonal functions 1, EOF1) represent overall negative anomalies throughout growing seasons, showing a mixed signal of variability in the initial levels and drawdown thereafter (low‐frequency dynamics). The second most common seasonal patterns of nitrate and phosphate (N and P EOF2) capture prolonged drawdown events during December–March, which are correlated to Chl EOF1. Si EOF2 captures a drawdown event during November–December, which is correlated to Chl EOF2. These different drawdown patterns are shaped by different sets of physical and climate forcing mechanisms. N and P drawdown events during December–March are influenced by the winter and spring Southern Annular Mode (SAM) phase, where nutrient utilization is enhanced in a stabilized upper water column as a consequence of SAM‐driven winter sea ice and spring wind dynamics. Si drawdown during November–December is influenced by early sea ice retreat, where ice breakup may induce abrupt water column stratification and a subsequent diatom bloom or release of diatom cells from within the sea ice. Our findings underscore that seasonal nutrient dynamics in the coastal WAP are coupled to large‐scale climate forcing and related physics, understanding of which may enable improved projections of biogeochemical responses to climate change.

We investigated the effects of pH on movement behaviors of the harmful algal bloom causing raphidophyte Heterosigma akashiwo. Motility parameters from >8000 swimming tracks of individual cells were quantified using 3D digital video analysis over a 6-h period in 3 pH treatments reflecting marine carbonate chemistry during the pre-industrial era, currently, and the year 2100. Movement behaviors were investigated in two different acclimation-to-target-pH conditions: instantaneous exposure and acclimation of cells for at least 11 generations. There was no negative impairment of cell motility when exposed to elevated PCO2 (i.e., low pH) conditions but there were significant behavioral responses. Irrespective of acclimation condition, lower pH significantly increased downward velocity and frequency of downward swimming cells (p < 0.001). Rapid exposure to lower pH resulted in 9% faster downward vertical velocity and up to 19% more cells swimming downwards (p < 0.001). Compared to pH-shock experiments, pre-acclimation of cells to target pH resulted in similar to 30% faster swimming speed and up to 46% faster downward velocities (all p < 0.001). The effect of year 2100 PCO2 levels on population diffusivity in pre-acclimated cultures was >2-fold greater than in pH-shock treatments (2.2 x 10(5) mu m(2) s(-1) vs. 8.4 x 10(4) mu m(2) s(-1)). Predictions from an advection-diffusion model, suggest that as PCO2 increased the fraction of the population aggregated at the surface declined, and moved deeper in the water column. Enhanced downward swimming of H. akashiwo at low pH suggests that these behavioral responses to elevated PCO2 could reduce the likelihood of dense surface slick formation of H. akashiwo through reductions in light exposure or growth independent surface aggregations. We hypothesize that the HAB alga's response to higher PCO2 may exploit the signaling function of high PCO2 as indicative of net heterotrophy in the system, thus indicative of high predation rates or depletion of nutrients. (c) 2013 Elsevier B.V. All rights reserved.

Here we determined (1) how accurately the semi‐automated Coulter Counter (CC) measured the abundance, volume, and growth rates of mono‐specific plankton cultures compared to light microscopy and (2) whether CC‐derived particle size‐abundance spectra of whole water samples provided a near‐real time assessment of plankton community composition. A significant, positive relationship was found between CC particle and microscope cell abundance and volume estimates for 16 phylogenetically diverse plankton species. However, abundance estimates of the 2 methods differed significantly from a 1:1 relationship, with CC counts on average 15% greater than microscopy counts. Growth rate estimates showed excellent agreement for both methods for all species and were consistent over time. No significant differences in abundance, growth rate, or size estimates were observed due to light levels (Scrippsiella trochoidea) or strain variation (Heterosigma akashiwo). CC‐based, ship‐board characterization of whole plankton communities, from Georges Bank, captured changes in size‐abundance spectra and thus provided a coarse picture of in‐situ community composition. In the laboratory, the CC appears particularly well‐suited to measuring relative changes in cell abundance, such as for growth and grazing rate measurements of mono‐specific laboratory cultures, or when predator and prey species are well separated by size differences. Although microscopy remains essential for plankton analyses, these results suggest that the CC can increase sample size, improving accuracy, and precision of studies requiring frequent sampling or high degrees of replication. Deployment of these automated techniques in the field may aid in quantifying spatial and temporal variability in plankton community structure by forthcoming ocean‐observing systems.