Biogeochemical cycling of dissolved zinc along the GEOTRACES South Atlantic transect GA10 at 40°S

The biogeochemical cycle of zinc (Zn) in the South Atlantic, at 40°S, was investigated as part of the UK GEOTRACES program. To date there is little understanding of the supply of Zn, an essential requirement for phytoplankton growth, to this highly productive region. Vertical Zn profiles displayed nutrient-like distributions with distinct gradients associated with the watermasses present. Surface Zn concentrations are among the lowest reported for theworld’s oceans (<50 pM). A strong Zn-Si linear relationshipwas observed (Zn (nM)= 0.065 Si (μM), r2=0.97, n = 460). Our results suggest that the use of a global Zn-Si relationship would lead to an underestimation of dissolved Zn in deeper waters of the South Atlantic. By utilizing Si* and a new tracer Zn* our data indicate that the preferential removal of Zn in the Southern Ocean prevented a direct return path for dissolved Zn to the surface waters of the South Atlantic at 40°S and potentially the thermocline waters of the South Atlantic subtropical gyre. The importance of Zn for phytoplankton growth was evaluated using the Zn-soluble reactive phosphorus (SRP) relationship. We hypothesize that the low Zn concentrations in the South Atlantic may select for phytoplankton cells with a lower Zn requirement. In addition, a much deeper kink at ~ 500m in the Zn:SRP ratio was observed compared to other oceanic regions.

Seasonal ITCZ migration dynamically controls the location of the (sub)tropical Atlantic biogeochemical divide

Inorganic nitrogen depletion restricts productivity in much of the low-latitude oceans, generating a selective advantage for diazotrophic organisms capable of fixing atmospheric dinitrogen (N2). However, the abundance and activity of diazotrophs can in turn be controlled by the availability of other potentially limiting nutrients, including phosphorus (P) and iron (Fe). Here we present high-resolution data (∼0.3°) for dissolved iron, aluminum, and inorganic phosphorus that confirm the existence of a sharp north–south biogeochemical boundary in the surface nutrient concentrations of the (sub)tropical Atlantic Ocean. Combining satellite-based precipitation data with results from a previous study, we here demonstrate that wet deposition in the region of the intertropical convergence zone acts as the major dissolved iron source to surface waters. Moreover, corresponding observations of N2 fixation and the distribution of diazotrophic Trichodesmium spp. indicate that movement in the region of elevated dissolved iron as a result of the seasonal migration of the intertropical convergence zone drives a shift in the latitudinal distribution of diazotrophy and corresponding dissolved inorganic phosphorus depletion. These conclusions are consistent with the results of an idealized numerical model of the system. The boundary between the distinct biogeochemical systems of the (sub)tropical Atlantic thus appears to be defined by the diazotrophic response to spatial–temporal variability in external Fe inputs. Consequently, in addition to demonstrating a unique seasonal cycle forced by atmospheric nutrient inputs, we suggest that the underlying biogeochemical mechanisms would likely characterize the response of oligotrophic systems to altered environmental forcing over longer timescales.

Enhancement of primary production in the North Atlantic outside of the spring bloom, identified by remote sensing of ocean colour and temperature

The heterogeneity in phytoplankton production in the North Atlantic after the spring bloom is poorly understood. We analysed merged microwave and infrared satellite sea surface temperature (SST) data and ocean colour phytoplankton size class biomass, primary production (PP) and new production (ExP) derived from SeaWiFS data, to assess the spatial and temporal frequency of surface thermal fronts and areas of enhanced PP and ExP. Strong and persistent surface thermal fronts occurred at the Reykjanes Ridge (RR) and sub-polar front (SPF), which sustain high PP and ExP and, outside of the spring bloom, account for 9% and 15% of the total production in the North Atlantic. When normalised by area, PP at the SPF is four times higher than the RR. Analysis of 13 years of satellite ocean colour data from SeaWiFS, and compared with MODIS-Aqua and MERIS, showed that there was no increase in Chla from 1998 to 2002, which then decreased in all areas from 2002 to 2007 and was most pronounced in the RR. These time series also illustrated that the SPF exhibited the highest PP and the lowest variation in Chla over the ocean colour record. This implies that the SPF provides a high and consistent supply of carbon to the benthos irrespective of fluctuations in the North Atlantic Oscillation.

Estimating the economic loss of recent North Atlantic fisheries management

It is accepted that world’s fisheries are not generally exploited at their biological or their economic optimum. Most fisheries assessments focus on the biological capacity of fish stocks to respond to harvesting and few have attempted to estimate the economic efficiency at which ecosystems are exploited. The latter is important as fisheries contribute considerably to the economic development of many coastal communities. Here we estimate the overall potential economic rent for the fishing industry in the North Atlantic to be B€ 12.85, compared to current estimated profits of B€ 0.63. The difference between the potential and the net profits obtained from North Atlantic fisheries is therefore B€ 12.22. In order to increase the profits of North Atlantic fisheries to a maximum, total fish biomass would have to be rebuilt to 108 Mt (2.4 times more than present) by reducing current total fishing effort by 53%. Stochastic simulations were undertaken to estimate the uncertainty associated with the aggregate bioeconomic model that we use and we estimate the economic loss NA fisheries in a range of 2.5 and 32 billion of euro. We provide economic justification for maintaining or restoring fish stocks to above their MSY biomass levels. Our conclusions are consistent with similar global scale studies.

Organ damage in Atlantic herring larvae as a result of ocean acidification

The dissolution of anthropogenically emitted excess carbon dioxide lowers the pH of the world's ocean water. The larvae of mass spawning marine fishes may be particularly vulnerable to such ocean acidification (OA), yet the generality of earlier results is unclear. Here we show the detrimental effects of OA on the development of a commercially important fish species, the Atlantic herring (Clupea harengus). Larvae were reared at three levels of CO2: today (0.0385 kPa), end of next century (0.183 kPa), and a coastal upwelling scenario (0.426 kPa), under near-natural conditions in large outdoor tanks. Exposure to elevated CO2 levels resulted in stunted growth and development, decreased condition, and severe tissue damage in many organs, with the degree of damage increasing with CO2 concentration. This complements earlier studies of OA on Atlantic cod larvae that revealed similar organ damage but at increased growth rates and no effect on condition.

Thermal front variability along the North Atlantic Current observed using satellite microwave and infrared data

Thermal fronts detected using multiple satellite sensors have been integrated to provide new information on the spatial and seasonal distribution of oceanic fronts in the North Atlantic. The branching of the North Atlantic Current (NAC) as it encounters the Mid-Atlantic Ridge (MAR) is reflected in surface thermal fronts, which preferentially occur at the Charlie Gibbs Fracture Zone (CGFZ) and several smaller fracture zones. North of the CGFZ there are few thermal fronts, contrasting with the region to the south, where there are frequent surface thermal fronts that are persistent seasonally and interannually. The alignment of the fronts confirms that the shallower Reykjanes Ridge north of the CGFZ is more of a barrier to water movements than the ridge to the south. Comparison of front distributions with satellite altimetry data indicates that the MAR influence on deep ocean currents is also frequently exhibited in surface temperature. The improved spatial and temporal resolution of the front analysis has revealed consistent seasonality in the branching patterns. These results contribute to our understanding of the variability of the NAC, and the techniques for visualising oceanic fronts can be applied in other regions to reveal details of surface currents that cannot be resolved using satellite altimetry or in situ measurements.

Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. “Gut flux” for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.

The future of the northeast Atlantic benthic flora in a high CO2 world

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO2 emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf-forming seaweeds.

Environmental variability and biodiversity of megabenthos on the Hebrides Terrace Seamount (Northeast Atlantic)

We present the first remotely operated vehicle investigation of megabenthic communities (1004-1695 m water depth) on the Hebrides Terrace Seamount (Northeast Atlantic). Conductivity-temperature-depth casts showed rapid light attenuation below the summit and an oceanographic regime on the flanks consistent with an internal tide, and high short-term variability in water temperature, salinity, light attenuation, aragonite and oxygen down to 1500 m deep. Minor changes in species composition (3-14%) were explained by changes in depth, substratum and oceanographic stability, whereas environmental variability explained substantially more variation in species richness (40-56%). Two peaks in species richness occurred, the first at 1300-1400 m where cooler Wyville Thomson Overflow Water (WTOW) mixes with subtropical gyre waters and the second at 1500-1600 m where WTOW mixes with subpolar mode waters. Our results suggest that internal tides, substrate heterogeneity and oceanographic interfaces may enhance biological diversity on this and adjacent seamounts in the Rockall Trough.

Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar)

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater (parr') stage to the migratory stage where they descend streams and enter salt water (smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 degrees C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.

Family-level AMBI is valid for use in the north-eastern Atlantic but not for assessing the health of microtidal Australian estuaries

Analysis of benthic macroinvertebrate samples at a higher taxonomic level than species, e.g. family, potentially provides a more cost-effective protocol for environmental impact assessments and monitoring as it requires less time, funds and taxonomic expertise. Using the AMBI database, species ecological group scores are shown to be coherent within families. Faunal data from a wide range of environmental impact scenarios in the north-eastern Atlantic demonstrate that AMBI, calculated from mean values for families, exhibits a strong linear relationship with species-level AMBI, the correlation improving by using square-root transformed rather than untransformed abundances. In many regions of the world, however, the sensitivity of benthic macroinvertebrates to environmental perturbations is unknown, precluding the use of AMBI for environmental assessments. Yet the families are essentially the same as in the AMBI database. The utility of family-level AMBI is tested using data for four south-western Australian estuaries previously subjected to environmental quality assessments, but where only 17 species of the 144 taxa are included in the AMBI database. Although family-level AMBI scores reflect differences in environmental quality spatially and temporally within an estuary, they do not follow variations in environmental quality among estuaries. Indeed, south-western Australia estuaries are numerically dominated by families with high AMBI scores, probably due to the detrimental effects of natural accumulations of organic material in estuaries with long residence times. As taxonomic distinctness follows trends in environmental quality among estuaries, as well as temporally and spatially within a system, it provides an appropriate substitute for assessing the 'heath' of microtidal estuaries.

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.

Comparison of new and primary production models using SeaWiFS data in contrasting hydrographic zones of the northern North Atlantic.

The accuracy of two satellite models of marine primary (PP) and new production (NP) were assessed against 14C and 15N uptake measurements taken during six research cruises in the northern North Atlantic. The wavelength resolving model (WRM) was more accurate than the Vertical General Production Model (VGPM) for computation of both PP and NP. Mean monthly satellite maps of PP and NP for both models were generated from 1997 to 2010 using SeaWiFS data for the Irminger basin and North Atlantic. Intra- and inter-annual variability of the two models was compared in six hydrographic zones. Both models exhibited similar spatio-temporal patterns: PP and NP increased from April to June and decreased by August. Higher values were associated with the East Greenland Current (EGC), Iceland Basin (ICB) and the Reykjanes Ridge (RKR) and lower values occurred in the Central Irminger Current (CIC), North Irminger Current (NIC) and Southern Irminger Current (SIC). The annual PP and NP over the SeaWiFS record was 258 and 82 gC m-2 yr-1 respectively for the VGPM and 190 and 41 gC m-2 yr-1 for the WRM. Average annual cumulative sum in the anomalies of NP for the VGPM were positively correlated with the North Atlantic Oscillation (NAO) in the EGC, CIC and SIC and negatively correlated with the multivariate ENSO index (MEI) in the ICB. By contrast, cumulative sum of the anomalies of NP for the WRM were significantly correlated with NAO only in the EGC and CIC. NP from both VGPM and WRM exhibited significant negative correlations with Arctic Oscillation (AO) in all hydrographic zones. The differences in estimates of PP and NP in these hydrographic zones arise principally from the parameterisation of the euphotic depth and the SST dependence of photo-physiological term in the VGPM, which has a greater sensitivity to variations in temperature than the WRM. In waters of 0 to 5C PP using the VGPM was 43% higher than WRM, from 5 to 10C the VGPM was 29% higher and from 10 to 15C the VGPM was 27% higher.

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.

Basking sharks and oceanographic fronts: quantifying associations in the north‐east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.