The fate of nitrogen and phosphorus at the land-sea margin of the North Atlantic Ocean

Five large rivers that discharge on the western North Atlantic continental shelf carry about 45% of the nitrogen (N) and 70% of the phosphorus (P) that others estimate to be the total flux of these elements from the entire North Atlantic watershed, including North, Central and South America, Europe, and Northwest Africa. We estimate that 61 · 109 moles y^–1 of N and 20 · 10⁹ moles y^–1 of P from the large rivers are buried with sediments in their deltas, and that an equal amount of N and P from the large rivers is lost to the shelf through burial of river sediments that are deposited directly on the continental slope. The effective transport of active N and P from land to the shelf through the very large rivers is thus reduced to 292 · 10⁹ moles y^–1 of N and 13 · 10⁹ moles y^–1 of P.The remaining riverine fluxes from land must pass through estuaries. An analysis of annual total N and total P budgets for various estuaries around the North Atlantic revealed that the net fractional transport of these nutrients through estuaries to the continental shelf is inversely correlated with the log mean residence time of water in the system. This is consistent with numerous observations of nutrient retention and loss in temperate lakes. Denitrification is the major process responsible for removing N in most estuaries, and the fraction of total N input that is denitrified appears to be directly proportional to the log mean water residence time. In general, we estimate that estuarine processes retain and remove 30–65% of the total N and 10–55% of the total P that would otherwise pass into the coastal ocean. The resulting transport through estuaries to the shelf amounts to 172–335 · 10⁹ moles y^–1 of N and 11–19 · 10⁹ moles y^–1 of P. These values are similar to the effective contribution from the large rivers that discharge directly on the shelf.For the North Atlantic shelf as a whole, N fluxes from major rivers and estuaries exceed atmospheric deposition by a factor of 3.5–4.7, but this varies widely among regions of the shelf. For example, on the U.S. Atlantic shelf and on the northwest European shelf, atmospheric deposition of N may exceed estuarine exports. Denitrification in shelf sediments exceeds the combined N input from land and atmosphere by a factor of 1.4–2.2. This deficit must be met by a flux of N from the deeper ocean. Burial of organic matter fixed on the shelf removes only a small fraction of the total N and P input (2–12% of N from land and atmosphere; 1–17% of P), but it may be a significant loss for P in the North Sea and some other regions. The removal of N and P in fisheries landings is very small. The gross exchange of N and P between the shelf and the open ocean is much larger than inputs from land and, for the North Atlantic shelf as a whole, it may be much larger than the N and P removed through denitrification, burial, and fisheries. Overall, the North Atlantic continental shelf appears to remove some 700–950· 10⁹ moles of N each year from the deep ocean and to transport somewhere between 18 and 30 · 10⁹ moles of P to the open sea. If the N and P associated with riverine sediments deposited on the continental slope are included in the total balance, the net flux of N to the shelf is reduced by 60 · 10⁹ moles y^–1 and the P flux to the ocean is increased by 20 · 10⁹ moles y^–1. These conclusions are quite tentative, however, because of large uncertainties in our estimates of some important terms in the shelf mass balance.     

Estimating denitrification in North Atlantic continental shelf sediments

A model of coupled nitrification/denitrification was developed for continental shelf sediments to estimate the spatial distribution of denitrification throughout shelf regions in the North Atlantic basin. Using data from a wide range of continental shelf regions, we found a linear relationship between denitrification and sediment oxygen uptake. This relationship was applied to specific continental shelf regions by combining it with a second regression relating sediment oxygen uptake to primary production in the overlying water. The combined equation was: denitrification (mmol N m^–2 d^–1)=0.019^* phytoplankton production (mmol C m^–2 d^–1). This relationship suggests that approximately 13% of the N incorporated into phytoplankton in shelf waters is eventually denitrified in the sediments via coupled nitrification/denitrification, assuming a C:N ratio of 6.625:1 for phytoplankton. The model calculated denitrification rates compare favorably with rates reported for several shelf regions in the North Atlantic.The model-predicted average denitrification rate for continental shelf sediments in the North Atlantic Basin is 0.69 mmol N m^{– 2} d^–1. Denitrification rates (per unit area) predicted by the model are highest for the continental shelf region in the western North Atlantic between Cape Hatteras and South Florida and lowest for Hudson Bay, the Baffin Island region, and Greenland. Within latitudinal belts, average denitrification rates were lowest in the high latitudes, intermediate in the tropics and highest in the mid-latitudes. Although denitrification rates per unit area are lowest in the high latitudes, the total N removal by denitrification (53 × 10¹⁰ mol N y^–1) is similar to that in the mid-latitudes (60 × 10¹⁰ mol N y^–1) due to the large area of continental shelf in the high latitudes. The Gulf of St. Lawrence/Grand Banks area and the North Sea are responsible for seventy-five percent of the denitrification in the high latitude region. N removal by denitrification in the western North Atlantic (96 × 10¹⁰ mol N y^–1) is two times greater than in the eastern North Atlantic (47 × 10¹⁰ mol N y^–1). This is primarily due to differences in the area of continental shelf in the two regions, as the average denitrification rate per unit area is similar in the western and eastern North Atlantic.We calculate that a total of 143 × 10¹⁰ mol N y^–1 is removed via coupled nitrification/denitrification on the North Atlantic continental shelf. This estimate is expected to underestimate total sediment denitrification because it does not include direct denitrification of nitrate from the overlying water. The rate of coupled nitrification/denitrification calculated is greater than the nitrogen inputs from atmospheric deposition and river sources combined, and suggests that onwelling of nutrient rich slope water is a major source of N for denitrification in shelf regions. For the two regions where N inputs to a shelf region from onwelling have been measured, onwelling appears to be able to balance the denitrification loss.     

Raiding the Landscape: Human Impact in the Scandinavian North Atlantic

Between ca. A.D. 800–1000, Scandinavian chiefly societies with a mixed maritime and agricultural economy expanded into the North Atlantic, colonizing Shetland, Orkney, Caithness, Hebrides, Faeroes, Iceland, Greenland, and Vinland. The settlers brought continental European economics and expectations to a widely varied set of island ecosystems. In many regions, rapid degradation of flora and soils took place associated with social and climate change. Recent research coordinated by the North Atlantic Bicultural Organization (NABO) highlights the extent of pre-modern impacts.     

North Atlantic marine communities through time

How and why ecological communities change their species membership over time and space is a central issue in ecology and evolution. Phylogeographic approaches based on animal mitochondrial DNA sequences have been important for revealing historical patterns of individual species and can provide qualitative comparisons among species. Exciting new methods, particularly implementing approximate Bayesian computation (ABC – Beaumont et al. 2002 ), now allow model‐based quantitative comparisons among species and permit the probabilistic exploration of alternative community‐level hypotheses (see review by Hickerson et al. 2010 ). In this issue of Molecular Ecology, Ilves et al. (2010) use an ABC approach to bring fresh insights into the well‐studied question of how North Atlantic coastal species contracted and expanded their ranges in response to late Pleistocene/Holocene climate fluctuations.     

Phylogeography and historical ecology of the North Atlantic intertidal

Abstract Recent glaciation covered the full extent of rocky intertidal habitat along the coasts of New England and the Canadian Maritimes. To test whether this glaciation in fact caused wholesale extinction of obligate rocky intertidal invertebrates, and thus required a recolonization from Europe, we compared American and European populations using allelic diversity and techniques adapted from coalescent theory. Mitochondrial DNA sequences were collected from amphi-Atlantic populations of three cold-temperate obligate rocky intertidal species (a barnacle, Semibalanus balanoides , and two gastropods, Nucella lapillus and Littorina obtusata ) and three cold-temperate habitat generalist species (a seastar, Asterias rubens ; a mussel, Mytilus edulis , and an isopod, Idotea balthica ). For many of these species we were able to estimate the lineage-specific mutation rate based on trans-Arctic divergences between Pacific and Atlantic taxa. These data indicate that some obligate rocky intertidal taxa have colonized New England from European populations. However, the patterns of persistence in North America indicate that other life-history traits, including mechanisms of dispersal, may be more important for surviving dramatic environmental and climatic change.     

Links between viruses and prokaryotes throughout the water column along a North Atlantic latitudinal transect

Viruses are an abundant, diverse and dynamic component of marine ecosystems and have a key role in the biogeochemical processes of the ocean by controlling prokaryotic and phytoplankton abundance and diversity. However, most of the studies on virus–prokaryote interactions in marine environments have been performed in nearshore waters. To assess potential variations in the relation between viruses and prokaryotes in different oceanographic provinces, we determined viral and prokaryotic abundance and production throughout the water column along a latitudinal transect in the North Atlantic. Depth-related trends in prokaryotic and viral abundance (both decreasing by one order of magnitude from epi- to abyssopelagic waters), and prokaryotic production (decreasing by three orders of magnitude) were observed along the latitudinal transect. The virus-to-prokaryote ratio (VPR) increased from ∼19 in epipelagic to ∼53 in the bathy- and abyssopelagic waters. Although the lytic viral production decreased significantly with depth, the lysogenic viral production did not vary with depth. In bathypelagic waters, pronounced differences in prokaryotic and viral abundance were found among different oceanic provinces with lower leucine incorporation rates and higher VPRs in the North Atlantic Gyre province than in the provinces further north and south. The percentage of lysogeny increased from subpolar regions toward the more oligotrophic lower latitudes. Based on the observed trends over this latitudinal transect, we conclude that the viral–host interactions significantly change among different oceanic provinces in response to changes in the biotic and abiotic variables.     

Causes and projections of abrupt climate-driven ecosystem shifts in the North Atlantic

Warming of the global climate is now unequivocal and its impact on Earth' functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.     

The North Atlantic Oscillation and the Arctic Oscillation favour harmful algal blooms in SW Europe

The ecological factors driving the occurrence of harmful algal blooms (HAB) have been traditionally analysed from a hydrological point of view, whereas the interplay between atmospheric oscillations and HAB has been scarcely explored. Here we address the possible link between atmospheric oscillations in SW Europe, using as proxies the North Atlantic Oscillation (NAO) and the Arctic Oscillation (AO) indices and the interannual variability in HAB. The yearly series (1973–2005) of mortality of water birds and fish in Doñana National Park (SW Spain) due to toxins associated with the cyanobacterium Microcystis aeruginosa, and the monthly incidence (January 1999 to December 2007) of the paralytic shellfish poisoning (PSP) toxin produced by the dinoflagellate Gymnodinium catenatum in the Rías Baixas (NW Spain), were selected as models of HAB in SW Europe. The incidences of both toxic algal events were fitted to a binary logistic regression as a function of the atmospheric oscillation indices (with different delays in accordance with each toxic event) and a favourability function was then computed. The favourability for the wildlife mortality in Doñana National Park was as a function of the AO averaged for the summer period (June–August) whereas the favourability for the incidence of PSP in the Rías Baixas was a function of the NAO recorded 1 month before the event. Since HAB have a relevant impact on ecosystems and human health, there is great interest in deciphering the ecological conditions favouring these events. Here we show that the atmospheric oscillations could be drivers of ecological processes linked to the occurrence of HAB in SW Europe. Moreover, the favourability functions relating NAO and AO indices with algal toxic events in SW Europe could be used as a powerful tool to predict toxic events.     

Water mass-specificity of bacterial communities in the North Atlantic revealed by massively parallel sequencing

Bacterial assemblages from subsurface (100 m depth), meso- (200-1000 m depth) and bathy-pelagic (below 1000 m depth) zones at 10 stations along a North Atlantic Ocean transect from 60°N to 5°S were characterized using massively parallel pyrotag sequencing of the V6 region of the 16S rRNA gene (V6 pyrotags). In a dataset of more than 830,000 pyrotags, we identified 10,780 OTUs of which 52% were singletons. The singletons accounted for less than 2% of the OTU abundance, whereas the 100 and 1000 most abundant OTUs represented 80% and 96% respectively of all recovered OTUs. Non-metric Multi-Dimensional Scaling and Canonical Correspondence Analysis of all the OTUs excluding the singletons revealed a clear clustering of the bacterial communities according to the water masses. More than 80% of the 1000 most abundant OTUs corresponded to Proteobacteria of which 55% were Alphaproteobacteria, mostly composed of the SAR11 cluster. Gammaproteobacteria increased with depth and included a relatively large number of OTUs belonging to Alteromonadales and Oceanospirillales. The bathypelagic zone showed higher taxonomic evenness than the overlying waters, albeit bacterial diversity was remarkably variable. Both abundant and low-abundance OTUs were responsible for the distinct bacterial communities characterizing the major deep-water masses. Taken together, our results reveal that deep-water masses act as bio-oceanographic islands for bacterioplankton leading to water mass-specific bacterial communities in the deep waters of the Atlantic.     

Abyssal Megafaunal Necrophages: Latitudinal Differences in the Eastern North Atlantic Ocean

A pop-up time-lapse camera system (BATHYSNACK) was used to study megafaunal organisms attracted to bait at three abyssal localities in the eastern North Atlantic Ocean, a northern site on the Porcupine Abyssal Plain and two sub-tropical sites on the Madeira Abyssal Plain. Fish and decapod crustaceans appeared on 47% of photographs taken during deployments of up to seven days duration. The fishes Coryphaenoides (Nematonurus) armatus and Pachycara bulbiceps and the decapod Munidopsis spp. dominated appearances at the northern locality whereas over 95% of records at the two subtropical localities were of the decapod Plesiopenaeus armatus. Cyclical variations in necrophage abundance did not correlate with tidal cycles of near-bottom currents, although high P. armatus numbers were inversely related to current speed. Most of the frequently recorded entities showed contagious intraspecific distributions. In contrast, many of the interspecies relationships were non-random, with co-occurrences lower than expected, probably as a result of physical disturbance or potential predation. Population densities based on times of first arrival were calculated. Assemblages dominated by fish at higher latitudes and crustaceans at lower latitudes are discussed in terms of organic carbon supply to the deep sea and pelagic community structures.     

Review of the distribution of Mesoplodon species (order Cetacea, family Ziphiidae) in the North Atlantic

Four species of beaked whales of the genus Mesoplodon regularly occur in the North Atlantic. These are Sowerby's Beaked Whale Mesoplodon bidens, Blainville's Beaked Whale Mesoplodon densirostris, Gervais Beaked Whale Mesoplodon europaeus and True's Beaked Whale Mesoplodon mirus. These four species differ in their distribution within this area. M. bidens is the most northerly recorded species followed by M. mirus. M. densirostris and M. europaeus both generally occur further south and their distributions cross the equator. The distribution of Mesoplodon species may relate to variations in water temperature.     

Photoperiodic regulation of gametogenesis in a North Atlantic sea star,Asterias vulgaris

Summary

Individuals of the sea star Asterias vulgaris from New Hampshire were maintained under two contrasting regimes of seasonally changing photoperiod. In one regime the lights turned on and off in phase with local sunrise and sunset; in the other, the daily photoperiod was kept 6 months out of phase with ambient photo- period. When dissected after being maintained for 18 months, the gametogenic condition of the out-of-phase animals was found to be 6 months out of phase with that of the in-phase animals. This experiment demonstrates that photoperiodic regulation of reproduction is important even in marine animals that normally experience marked seasonal changes in sea temperature.     

Regional differences in bacterial flora in harbour porpoises from the North Atlantic: environmental effects?

Aims:  Microbiological findings in harbour porpoises from different regions of the North Atlantic were compared. Results in animals from the North and Baltic Seas were evaluated over a period of 18 years for changes in the microbiological flora.
Methods and Results:  Microbiological investigations were performed on 1429 organ samples from the lung, liver, kidney, spleen, intestine, and mesenteric lymph nodes from harbour porpoises of the German North and Baltic Seas, Greenlandic, Icelandic and Norwegian waters. A large variety of bacteria, including potentially pathogenic bacteria like Brucella sp., Clostridium perfringens, Escherichia coli, Erysipelothrix rhusiopathiae , β-haemolytic streptococci and Staphylococcus aureus were isolated . Those bacteria were associated with bronchopneumonia, gastroenteritis, hepatitis, pyelonephritis, myocarditis and septicemia.
Conclusions:  Organs from animals originating from Greenlandic and Icelandic waters showed clearly less bacterial growth and fewer associated pathological lesions compared to animals from the German North and Baltic Seas and Norwegian waters.
Significance and Impact of the Study:  Differences in bacterial findings and associated lesions between harbour porpoises from the German North and Baltic Seas and animals from Greenlandic, Norwegian and Icelandic waters may result from higher stress due to anthropogenic activities such as chemical pollutants in the North and Baltic Seas.     

Reassessing the impact of North Atlantic Oscillation on the sub-Saharan vegetation productivity

The Northern Atlantic Oscillation (NAO) has been shown to have a significant impact on the terrestrial ecosystem in the Sahelian region of Africa during the 1980s, and it has been strongly suggested that NAO may be a reliable predictor for the response of the Sahelian ecosystem to global climate variability. Using data from an extended period, we provide a reassessment for the impact of NAO on the Sahelian climate and ecosystem, and show that there is no consistent relationship between NAO and the ecosystem over Sahel. Statistical analysis on the NAO, vegetation, and precipitation data indicates that NAO influences the Sahelian vegetation productivity exclusively through its impact on precipitation. However, the relationship between the NAO index and Sahelian precipitation varies substantially with time. The correlation coefficient fluctuates between positive and negative values, and does not pass the 5% significance test during most of the twentieth century. The NAO system, although documented to govern the ecosystem dynamics over many other regions, does not have a consistent impact on the ecosystem over the Sahel. Therefore, the NAO index cannot produce a useful prediction on the ecosystem variability and changes in this region. This study provides an example that correlations based on short climate and ecological records (less than 20 years in this case) can be spurious and potentially misleading.     

Iron limits primary productivity during spring bloom development in the central North Atlantic

We present in situ biophysical measurements and bioassay experiments that demonstrate iron limitation of primary productivity during the spring bloom in the central North Atlantic. Mass balance calculations indicate that nitrate drawdown is iron (Fe)-limited and that aeolian Fe supply to this region cannot support maximal phytoplankton growth during the bloom. Using a simple simulation model, we show that relief of Fe limitation during the spring bloom can increase nitrate drawdown and, hence, new primary production, by 70%. We conclude that the episodic nature of iron supplied by dust deposition is an important factor controlling the dynamics of the spring bloom. From this, we hypothesize that variability in the timing and magnitude of the spring bloom in response to aeolian Fe supply will affect carbon drawdown and food web dynamics in the central North Atlantic.     

Reconstruction of pollutant lead invasion into the tropical North Atlantic during the twentieth century

In this study we reconstruct pollutant lead transport and input into the Caribbean basin in relation to the meteorological changes and past industrial activities. These reconstructions are based on the geochemical data obtained from massive corals collected in 1998 in Mona Island. Lead concentration profile mimics the general atmospheric lead trend obtained from the previous investigations on ice cores, carbonate skeletons and in sediments from the North Atlantic. This similarity corroborates the significance of anthropogenic impact on all reservoirs of the North Atlantic as associated with the proliferation of the automobile and use of leaded gasoline during the twentieth century. Our high-resolution record reveals a 2–4 years shift between the maximum pollutant lead input to Mona Island and leaded gasoline consumption peaks in North America, suggesting a possible longer secondary oceanic transport of US lead within the Subtropical North Atlantic Gyre system. A striking relationship is found between lead isotopic imprints and the winter North Atlantic oscillation (NAO) index for two specific periods (1914–1929 and 1965–1997). These periods are characterized by distinct isotopic signatures from the main pollutant lead emitter to the North Atlantic troposphere, North America and Western Europe. This involves a noteworthy influence of the NAO on pollutant input and distribution in the North Atlantic that strengthens the usefulness of lead isotopes as tracers of paleo-atmospheric circulation and changes in marine circulation pathways.     

The role of the North Atlantic Drift in the millennial timescale glacial climate fluctuations

The Greenland ice core records show that the overall cold climate of the last glacial period was repeatedly interrupted by short, rapid warmings. The events, known as Dansgaard-Oeschger (D-O) events, are strongly imprinted in the North Atlantic marine records suggesting that they were linked to North Atlantic circulation changes. However, the causes of the D-O events are poorly understood and they represent one of the most intriguing puzzles of the last glacial period. In order to investigate a possible mechanism we have studied variations in the distribution of benthic and planktonic foraminifera, oxygen isotopes and ice rafted debris during the last glacial period in eight cores from the North Atlantic and the Nordic Seas taken at mid depth from 853 to 1760 m. The parameters indicate, in agreement with previous studies, that the circulation system during the interstadials resembled the present system. Atlantic surface water flowed north into the Nordic Seas, where most of it sank through convection and, as cold deep water, flowed back into the North Atlantic. During the stadials, a halocline was established in the Nordic Seas and in the northernmost part of the North Atlantic and the outflow from the Nordic Seas stopped. However, below the cold, light surface layer, the relatively warm water of the North Atlantic Drift continued to flow across the North Atlantic and into the Nordic Seas, here warming up the deep-water masses. We suggest that the warm water gradually made the water column unstable. The pivotal abrupt climate warmings were caused by sudden upwelling of the deep, warm water masses causing overturning of the entire water column and onset of convection.     

New records of Echiura and Sipuncula in the North Atlantic Ocean, with the description of a new species of Jacobia

Data on Echiura and Sipuncula collected by the MAR-ECO expedition on the RV G.O. Sars to the northern Mid-Atlantic Ridge are presented. Eight specimens of echiuran and 13 specimens of sipunculan worms were obtained from eight stations. Two species of Echiura and two species of Sipuncula were identified. One echiuran is a new species of Jacobia.     

The biogeographical division of the Arctic and North Atlantic by the mysid (Crustacea: Mysidacea) fauna

The biogeographical analysis of Arctic and North Atlantic waters north of 30°N is based upon the distribution of 150 Mysidacea (Crustacea) species. The reasons for biogeographical divisions conducted independently by faunae of pelagic and benthopelagic mysids are adduced. The original schemes of the Arctic and North Atlantic division are proposed. Using the fauna of pelagic mysids, one biogeographical realm, one province and one transitional zone are designated in cold and temperate waters. Using the fauna of benthopelagic mysids, one biogeographical realm and four provinces are designated in those waters.     

'SSW to NNE'– North Atlantic Oscillation affects the progress of seasons across Europe

Three European plant phenological network datasets were analysed for latitudinal and longitudinal gradients of nine phenological ‘seasons’ spanning the entire year. The networks were: (1) the historical first European Phenological Network (1882–1941) by Hoffmann & Ihne, (2) the network of the International Phenological Gardens in Europe (1959–1998), founded by Schnelle & Volkert in 1957 and based on cloned plants, and (3) a dataset (1951–1998) that was recently collated during the EU Fifth Framework project POSITIVE, which included network data of seven Central and Eastern European countries. Our study is most likely the first, for over a century, to analyse average onset and year‐to‐year variability of the progress of seasons across a continent. For early, mid, and late spring seasons we found a marked progress of the seasonal onset from SW to NE throughout Europe, more precisely from WSW to ENE in early spring, then from SW to NE and finally from SSW to NNE in late spring, as exhibited by the relationship between latitudinal and longitudinal gradients. The movement of summer was more south to north directed, as the longitudinal gradient (west–east component) strongly declined or was even of opposite sign. Autumn, as shown by leaf colouring dates, arrived from NE to SW. Possible reasons for the differences among the three datasets are discussed. The annual variability of latitudinal and longitudinal gradients of the seasons across Europe was closely related to the North Atlantic Oscillation (NAO) index; in years with high NAO in both winter and spring, the west–east component of progress was more pronounced; in summer and autumn, the pattern of the seasons may be more uniform.