Decadal variability in biogeochemical models: Comparison with a 50-year ocean colour dataset

Assessing the skill of biogeochemical models to hindcast past variability is challenging, yet vital in order to assess their ability to predict biogeochemical change. However, the validation of decadal variability is limited by the sparsity of consistent, long-term biological datasets. The Phytoplankton Colour Index (PCI) product from the Continuous Plankton Recorder survey, which has been sampling the North Atlantic since 1948, is an example of such a dataset. Converting the PCI to chlorophyll values using SeaWiFS data allows a direct comparison with model output. Here we validate decadal variability in chlorophyll from the GFDL TOPAZ model. The model demonstrates skill at reproducing interannual variability, but cannot simulate the regime shifts evident in the PCI data. Comparison of the model output, data and climate indices highlights under-represented processes that it may be necessary to include in future biogeochemical models in order to accurately simulate decadal variability in ocean ecosystems.

Drivers of euphausiid species abundance and numerical abundance in the Atlantic Ocean

Mid-ocean ridges are common features of the world’s oceans but there is a lack of understanding as to how their presence affects overlying pelagic biota. The Mid-Atlantic Ridge (MAR) is a dominant feature of the Atlantic Ocean. Here, we examined data on euphausiid distribution and abundance arising from several international research programmes and from the continuous plankton recorder. We used a generalized additive model (GAM) framework to explore spatial patterns of variability in euphausiid distribution on, and at either side of, the MAR from 60°N to 55°S in conjunction with variability in a suite of biological, physical and environmental parameters. Euphausiid species abundance peaked in mid-latitudes and was significantly higher on the ridge than in adjacent waters, but the ridge did not influence numerical abundance significantly. Sea surface temperature (SST) was the most important single factor influencing both euphausiid numerical abundance and species abundance. Increases in sea surface height variance, a proxy for mixing, increased the numerical abundance of euphausiids. GAM predictions of variability in species abundance as a function of SST and depth of the mixed layer were consistent with present theories, which suggest that pelagic niche availability is related to the thermal structure of the near surface water: more deeply-mixed water contained higher euphausiid biodiversity. In addition to exposing present distributional patterns, the GAM framework enables responses to potential future and past environmental variability including temperature change to be explored.

Growth and production of Pacific ocean perch (Sebastes alutus) in nursery habitats of the Gulf of Alaska

Nursery areas for juvenile fishes are often important for determining recruitment in marine populations by providing habitats that can maximize growth and thereby minimize mortality. Pacific ocean perch (POP, Sebastes alutus) have an extended juvenile period where they inhabit rocky nursery habitats. We examined POP nursery areas to link growth potential to recruitment. Juvenile POP were captured from nursery areas in 2004 and 2008, and estimated growth rates ranged from −0.19 to 0.60 g day−1 based on differences in size between June and August. Predicted growth rates from a bioenergetics model ranged from 0.05 to 0.49 g day−1 and were not significantly different than observed. Substrate preferences and the distribution of their preferred habitats were utilized to predict the extent of juvenile POP nursery habitat in the Gulf of Alaska. Based on densities of fish observed on underwater video transects and the spatial extent of nursery areas, we predicted 278 and 290 million juvenile POP were produced in 2004 and 2008. Growth potential for juvenile POP was reconstructed using the bioenergetics model, spring zooplankton bloom timing and duration and bottom water temperature for 1982–2008. When a single outlying recruitment year in 1986 was removed, growth potential experienced by juvenile POP in nursery areas was significantly correlated to the recruitment time-series from the stock assessment, explaining ∼30% of the variability. This research highlights the potential to predict recruitment using habitat-based methods and provides a potential mechanism for explaining some of the POP recruitment variability observed for this population.

Technical Note: Ensuring consistent, global measurements of short-lived halocarbon gases in the ocean and atmosphere

Very short-lived halocarbons are significant sources of reactive halogen in the marine boundary layer, and likely in the upper troposphere and lower stratosphere. Quantifying ambient concentrations in the surface ocean and atmosphere is essential for understanding the atmospheric impact of these trace gas fluxes. Despite the body of literature increasing substantially over recent years, calibration issues complicate the comparison of results and limit the utility of building larger-scale databases that would enable further development of the science (e.g. sea-air flux quantification, model validation, etc.). With this in mind, thirty-one scientists from both atmospheric and oceanic halocarbon communities in eight nations gathered in London in February 2008 to discuss the scientific issues and plan an international effort toward developing common calibration scales (http://tinyurl.com/c9cg58). Here, we discuss the outputs from this meeting, suggest the compounds that should be targeted initially, identify opportunities for beginning calibration and comparison efforts, and make recommendations for ways to improve the comparability of previous and future measurements.

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3-D coupled physical-biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate-change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom-up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.

Global assessment of ocean carbon export by combining satellite observations and food-web models

The export of organic carbon from the surface ocean by sinking particles is an important, yet highly uncertain, component of the global carbon cycle. Here we introduce a mechanistic assessment of the global ocean carbon export using satellite observations, including determinations of net primary production and the slope of the particle size spectrum, to drive a food-web model that estimates the production of sinking zooplankton feces and algal aggregates comprising the sinking particle flux at the base of the euphotic zone. The synthesis of observations and models reveals fundamentally different and ecologically consistent regional-scale patterns in export and export efficiency not found in previous global carbon export assessments. The model reproduces regional-scale particle export field observations and predicts a climatological mean global carbon export from the euphotic zone of ~6 Pg C yr−1. Global export estimates show small variation (typically < 10%) to factor of 2 changes in model parameter values. The model is also robust to the choices of the satellite data products used and enables interannual changes to be quantified. The present synthesis of observations and models provides a path for quantifying the ocean's biological pump.

Quantifying ecosystem-level consequences of ocean acidification and warming by scaling up individual level responses of a predator snail and its trophic interactions

Understanding long‐term, ecosystem‐level impacts of climate change is challenging because experimental research frequently focuses on short‐term, individual‐level impacts in isolation. We address this shortcoming first through an interdisciplinary ensemble of novel experimental techniques to investigate the impacts of 14‐month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterize a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual‐level responses, while acidification had a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual‐level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large‐scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local‐environmental conditions and resource availability. Such changes in macro‐scale distributions cannot be predicted by investigating individual‐level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long‐term, multiscale responses to multiple stressors, in an ecosystem context.

Scaling up experimental ocean acidification and warming research: from individuals to the ecosystem

Understanding long-term, ecosystem-level impacts of climate change is challenging because experimental research frequently focuses on short-term, individual-level impacts in isolation. We address this shortcoming first through an inter-disciplinary ensemble of novel experimental techniques to investigate the impacts of 14-month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterise a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual level responses, while acidification has a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large-scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local environmental conditions and resource availability. Such changes in macro-scale distributions cannot be predicted by investigating individual level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long-term, multi-scale responses to multiple stressors, in an ecosystem context.

Air–sea fluxes of oxygenated volatile organic compounds across the Atlantic Ocean

We present air-sea fluxes of oxygenated volatile organics compounds (OVOCs) quantified by eddy covariance (EC) during the Atlantic Meridional Transect cruise in 2012. Measurements of acetone, acetaldehyde, and methanol in air as well as in water were made in several different oceanic provinces and over a wide range of wind speeds (1-18 m s(-1)). The ocean appears to be a net sink for acetone in the higher latitudes of the North Atlantic but a source in the subtropics. In the South Atlantic, seawater acetone was near saturation relative to the atmosphere, resulting in essentially zero net flux. For acetaldehyde, the two-layer model predicts a small oceanic emission, which was not well resolved by the EC method. Chemical enhancement of air-sea acetaldehyde exchange due to aqueous hydration appears to be minor. The deposition velocity of methanol correlates linearly with the transfer velocity of sensible heat, confirming predominant airside control. We examine the relationships between the OVOC concentrations in air as well as in water, and quantify the gross emission and deposition fluxes of these gases.

Challenges in integrative approaches to modelling the marine ecosystems of the North Atlantic: Physics to fish and coasts to ocean

It has long been recognised that there are strong interactions and feedbacks between climate, upper ocean biogeochemistry and marine food webs, and also that food web structure and phytoplankton community distribution are important determinants of variability in carbon production and export from the euphotic zone. Numerical models provide a vital tool to explore these interactions, given their capability to investigate multiple connected components of the system and the sensitivity to multiple drivers, including potential future conditions. A major driver for ecosystem model development is the demand for quantitative tools to support ecosystem-based management initiatives. The purpose of this paper is to review approaches to the modelling of marine ecosystems with a focus on the North Atlantic Ocean and its adjacent shelf seas, and to highlight the challenges they face and suggest ways forward. We consider the state of the art in simulating oceans and shelf sea physics, planktonic and higher trophic level ecosystems, and look towards building an integrative approach with these existing tools. We note how the different approaches have evolved historically and that many of the previous obstacles to harmonisation may no longer be present. We illustrate this with examples from the on-going and planned modelling effort in the Integrative Modelling Work Package of the EURO-BASIN programme.

Regional ocean-colour chlorophyll algorithms for the Red Sea

The Red Sea is a semi-enclosed tropical marine ecosystem that stretches from the Gulf of Suez and Gulf of Aqaba in the north, to the Gulf of Aden in the south. Despite its ecological and economic importance, its biological environment is relatively unexplored. Satellite ocean-colour estimates of chlorophyll concentration (an index of phytoplankton biomass) offer an observational platform to monitor the health of the Red Sea. However, little is known about the optical properties of the region. In this paper, we investigate the optical properties of the Red Sea in the context of satellite ocean-colour estimates of chlorophyll concentration. Making use of a new merged ocean-colour product, from the European Space Agency (ESA) Climate Change Initiative, and in situ data in the region, we test the performance of a series of ocean-colour chlorophyll algorithms. We find that standard algorithms systematically overestimate chlorophyll when compared with the in situ data. To investigate this bias we develop an ocean-colour model for the Red Sea, parameterised to data collected during the Tara Oceans expedition, that estimates remote-sensing reflectance as a function of chlorophyll concentration. We used the Red Sea model to tune the standard chlorophyll algorithms and the overestimation in chlorophyll originally observed was corrected. Results suggest that the overestimation was likely due to an excess of CDOM absorption per unit chlorophyll in the Red Sea when compared with average global conditions. However, we recognise that additional information is required to test the influence of other potential sources of the overestimation, such as aeolian dust, and we discuss uncertainties in the datasets used. We present a series of regional chlorophyll algorithms for the Red Sea, designed for a suite of ocean-colour sensors, that may be used for further testing.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.

High-resolution view of the spring bloom initiation and net community production in the Subantarctic Southern Ocean using glider data

In the Southern Ocean, there is increasing evidence that seasonal to subseasonal temporal scales, and meso- to submesoscales play an important role in understanding the sensitivity of ocean primary productivity to climate change. This drives the need for a high-resolution approach to re- solving biogeochemical processes. In this study, 5.5 months of continuous, high-resolution (3 h, 2 km horizontal resolution) glider data from spring to summer in the Atlantic Subantarctic Zone is used to investigate: (i) the mechanisms that drive bloom initiation and high growth rates in the region and (ii) the seasonal evolution of water column production and respiration. Bloom initiation dates were analysed in the context of upper ocean boundary layer physics highlighting sensitivities of different bloom detection methods to different environmental processes. Model results show that in early spring (September to mid-November) increased rates of net community production (NCP) are strongly affected by meso- to submesoscale features. In late spring/early summer (late-November to mid-December) seasonal shoaling of the mixed layer drives a more spatially homogenous bloom with maximum rates of NCP and chlorophyll biomass. A comparison of biomass accumulation rates with a study in the North Atlantic highlights the sensitivity of phytoplankton growth to fine-scale dynamics and emphasizes the need to sample the ocean at high resolution to accurately resolve phytoplankton phenology and improve our ability to estimate the sensitivity of the biological carbon pump to climate change.

Temporal changes in total and size-fractioned chlorophyll-a in surface waters of three provinces in the Atlantic Ocean (September to November) between 2003 and 2010

Phytoplankton total chlorophyll concentration (TCHLa) and phytoplankton size structure are two important ecological indicators in biological oceanography. Using high performance liquid chromatography (HPLC) pigment data, collected from surface waters along the Atlantic Meridional Transect (AMT), we examine temporal changes in TCHLa and phytoplankton size class (PSC: micro-, nano- and pico-phytoplankton) between 2003 and 2010 (September to November cruises only), in three ecological provinces of the Atlantic Ocean. The HPLC data indicate no significant change in TCHLa in northern and equatorial provinces, and an increase in the southern province. These changes were not significantly different to changes in TCHLa derived using satellite ocean-colour data over the same study period. Despite no change in AMT TCHLa in northern and equatorial provinces, significant differences in PSC were observed, related to changes in key diagnostic pigments (fucoxanthin, peridinin, 19′-hexanoyloxyfucoxanthin and zeaxanthin), with an increase in small cells (nano- and pico-phytoplankton) and a decrease in larger cells (micro-phytoplankton). When fitting a three-component model of phytoplankton size structure — designed to quantify the relationship between PSC and TCHLa to each AMT cruise, model parameters varied over the study period. Changes in the relationship between PSC and TCHLa have wide implications in ecology and marine biogeochemistry, and provide key information for the development and use of empirical ocean-colour algorithms. Results illustrate the importance of maintaining a time-series of in-situ observations in remote regions of the ocean, such as that acquired in the AMT programme.

Ecological controls on biogeochemical fluxes in the western Antarctic Peninsula studied with an inverse foodweb model

Sea ice in the western Antarctic Peninsula (WAP) region is both highly variable and rapidly changing. In the Palmer Station region, the ice season duration has decreased by 92 d since 1978. The sea-ice changes affect ocean stratification and freshwater balance and in turn impact every component of the polar marine ecosystem. Long-term observations from the WAP nearshore and offshore regions show a pattern of chlorophyll (Chl) variability with three to five years of negative Chl anomalies interrupted by one or two years of positive anomalies (high and low Chl regimes). Both field observations and results from an inverse food-web model show that these high and low Chl regimes differed significantly from each other, with high primary productivity and net community production (NCP) and other rates associated with the high Chl years and low rates with low Chl years. Gross primary production rates (GPP) averaged 30 mmolC.m-2.d-1 in the low Chl years and 100 mmolC.m-2.d-1 in the high Chl years. Both large and small phytoplankton were more abundant and more productive in high Chl years than in low Chl years. Similarly, krill were more important as grazers in high Chl years, but did not differ from microzooplankton in high or low Chl years. Microzooplankton did not differ between high and low Chl years. Net community production differed significantly between high and low Chl years, but mobilized a similar proportion of GPP in both high and low Chl years. The composition of the NCP was uniform in high and low Chl years. These results mphasize the importance of microbial components in the WAP plankton system and suggest that food webs dominated by small phytoplankton can have pathways that funnel production into NCP, and likely, export.