Bathymetric patterns of deep-sea gastropod species diversity in 10 basins of the Atlantic Ocean and Norwegian Sea

Bathymetric patterns of macrofaunal species diversity are best documented in the western North Atlantic where diversity is a unimodal function of depth, peaking in the mid-bathyal zone and being depressed in the upper slope and abyss. There are few inter-basin studies of diversity-depth trends that are controlled for taxonomy, sampling gear, and diversity measures. In this paper, we compare gastropod diversity gradients in the North American Basin of the Atlantic to estimates of diversity in 9 other regions: the Norwegian Sea, West European Basin, Guiana Basin, Gambia Basin, Equatorial Mid-Atlantic, Brazil Basin, Angola Basin, Cape Basin and Argentine Basin. All samples were collected with epibenthic sleds, and diversity calculated by the Sanders-Hurlbert normalized expected number of species. While sampling in other regions is generally less complete than in the western North Atlantic, results indicate that a unimodal pattern is not universal. Diversity can increase, decrease or show no relationship with depth. The level of diversity also varies among basins relative to the western North Atlantic, being depressed in the Norwegian Sea, at bathyal depths in the eastern North Atlantic, and below an oxygen minimum zone in the Cape Basin, and generally elevated at tropical latitudes and in abyssal regions where food supply is high. Associations between gastropod diversity and the ecology and geology of basins suggest that productivity, oxygen concentration, hydrographic disturbance and evolutionary-historical processes may be implicated in shaping bathymetric diversity gradients, but specific causes are difficult to discern. Much more intensive sampling, analyses of other major taxa, and more detailed ecological data are necessary to understand deep-sea biogeography at within- and between-basin spatial scales.     

Ecological classification of European transitional waters in the North-East Atlantic eco-region

A new methodology to classify European North-East Atlantic transitional waters into ecological types has been developed based on the most important hydrological and morphological features that are likely to determine the ecology of aquatic systems in transitional waters. Hydrological indicators help identifying if a transitional water area is dominated by fresh or sea water and/or by intertidal or subtidal areas, while morphological indicators allow an estimation of the complexity of the transitional water and the diversity of the habitats involved. Twelve transitional waters of the southern Bay of Biscay were classified using this methodology and the five hydro-morphological types obtained were validated with benthic macro-invertebrate data. Transitional waters with a complex morphology showed the highest values of species diversity, while those with a smaller tidal influence showed lower species diversity. The ‘Scrobicularia’ and ‘Abra’ assemblages, previously identified in the study area, were found to be related to different types of transitional waters. The ‘Abra’ assemblage only appeared in estuaries with a complex morphology and dominated by tidal influences, while the ‘Scrobicularia’ assemblage was detected in all the transitional waters except for a single coastal lagoon. This classification of transitional waters may therefore be useful to establish the biological reference conditions needed for European Directives.     

Atmospheric dust contribution to deep-sea particle fluxes in the subtropical Northeast Atlantic

The lithogenic flux of sediment trap material was analyzed from a three year time series (February 2002–March 2005) at 2000 m depth in the Northeast Atlantic (Kiel 276, 33°N, 22°W) with regards to the seasonal and interannual variability of flux intensity and mineralogy—by applying an automated particle SEM-EDX analysis (scanning electron microscope-energy dispersive X-ray analysis). The lithogenic flux shows strong interannual variations with highest lithogenic flux rates occurring during January–February and April–March coupled to the total particle flux. Mean lithogenic flux rates for the sample years are 7.1 (2002–2003), 5.1 (2003–2004) and 16.1 mg m^?2 d^?1 (2004–2005). Mineral assemblages from the three sample years reveal distinct major minerals related to specific source regions. Clay minerals dominate the lithogenic fraction within the years 2002 and 2004 with illite (2002–2003) and palygorskite (2003–2004) being the major clay minerals. During the year 2004–2005, quartz is the major lithogenic mineral accompanied by smectite. The mineral assemblages hint to a mixture of North African source areas with dominant sources in Mauritania and north western parts of NW Africa for the years 2002–2004 and central Sahara (Algeria–Mali) within the year 2004–2005.     

Protecting the North-East Atlantic: enhancing synergies by institutional interplay

This article examines the interplay between three different international institutions on marine pollution in the North Sea and the wider North-East Atlantic: the International North Sea Conferences (INSC), the OSPAR-Convention and preceding conventions as well as the European Union (EU). It argues that interplay between these institutions have proved synergetic in two ways. First, the INSC-process has speeded up decision-making within OSPAR and the EU by means of leadership, change in membership and institutional arrangements. Second, OSPAR and the EU have subsequently facilitated domestic implementation of the North Sea Conference Declarations by means of higher authoritativeness and enforcement competence. These three institutions have fulfilled different functions all of which are needed to make international environmental cooperation effective.     

Wave climate variability in the North-East Atlantic Ocean over the last six decades

Ocean surface gravity waves play a major role in many engineering and environmental problems, both in the open ocean and in coastal zones. Therefore, it is essential to improve our knowledge on spatial and temporal variability of wave climate. This study aims at investigating this variability in the North-East Atlantic Ocean (25°W–0°W and 30°N–60° N), using a 57-year hindcast (1953–2009) obtained with a spectral wave model forced with reanalysis wind fields. The hindcast analysis reveals firstly strong seasonal fluctuations of wave climate, with winters characterized by large and long-period waves of mean direction spreading from south-west to north-west, and summers characterized by smaller and shorter-period waves originating from norther directions. From northern (55°N) to southern (35°N) latitudes, the significant wave height (Hs) decreases by roughly 40%, the mean wave direction (Mwd) rotates clockwise by about 25% while the peak period (Tp) only grows by 5%. These three parameters also exhibit a strong inter-annual variability, particularly when winter-means (from 1st of December to 1st of April) are considered. Linear trend analysis over the studied period shows spatially variable long-term trends, with a significant increase of Hs (up to 0.02 m yr^?1) and a counterclockwise shift of Mwd (up to ?0.1° yr^?1) at northern latitude, contrasting with a fairly constant trend for Hs and a clockwise shift of Mwd (up to +0.15° yr^?1) at southern latitudes. Long-term trends of Tp are less significant, with still a slight increase in the north-eastern part of the study area (up to +0.01 s yr^?1). Eventually, a comparison between the inter-annual variability of the winter-means of the three selected wave parameters and the North Atlantic Oscillation (NAO) reveals: (1) a strong positive correlation between Hs and the NAO index at northern latitudes (correlation coefficient up to R = 0.91) and a significant negative correlation at southern latitudes (up to R = ?0.6); (2) no significant correlation for Mwd north of 40°N and a clear positive correlation southward of 40°N (up to R = 0.8) and (3) a northward increasing correlation for Tp (up to R = 0.8). Long-term trends for Hs, Mwd and Tp are finally explained by a significant increase in the NAO index over the studied period.     

Influence of atmospheric inputs on the iron distribution in the subtropical North-East Atlantic Ocean

Aerosol (soluble and total) iron and water-column dissolved (DFe, < 0.2 μm) and total dissolvable (TDFe, unfiltered) iron concentrations were determined in the Canary Basin and along a transect towards the Strait of Gibraltar, in order to sample across the Saharan dust plume. Cumulative dust deposition fluxes estimated from direct aerosol sampling during our one-month cruise are representative of the estimated deposition fluxes based on near surface water dissolved aluminium concentrations measured on board. Iron inventories in near surface waters combined with flux estimates confirmed the relatively short residence time of DFe in waters influenced by the Saharan dust plume (6–14 months). Enhanced near surface water concentrations of DFe (5.90–6.99 nM) were observed at the Strait of Gibraltar mainly due to inputs from metal-rich rivers. In the Canary Basin and the transect towards Gibraltar, DFe concentrations (0.07–0.76 nM) were typical of concentrations observed in the surface North Atlantic Waters, with the highest concentrations associated with higher atmospheric inputs in the Canary Basin. Depth profiles showed that DFe and TDFe were influenced by atmospheric inputs in this area with an accumulation of aeolian Fe in the surface waters. The sub-surface minimum of both DFe and TDFe suggests that a simple partitioning between dissolved and particulate Fe is not obvious there and that export may occur for both phases. At depths of around 1000–1300 m, both regeneration and Meddies may explain the observed maximum. Our data suggest that, in deep waters, higher particle concentrations likely due to dust storms may increase the scavenging flux and thus decrease DFe concentrations in deep waters.     

Bottlenose dolphin Tursiops truncatus group dynamics,site fidelity,residency and movement patterns in the Madeira Archipelago (North-East Atlantic)

The bottlenose dolphin Tursiops truncatus is one of the most frequently sighted cetacean species in the Madeira Archipelago (North-East Atlantic); however, little is known about its population ecology in these waters. Photo-identification undertaken during systematic, non-systematic and opportunistic surveys conducted between 2001 and 2012 was used to assess group dynamics, site fidelity, residency and movement patterns of bottlenose dolphins in the archipelago. Three different patterns of residency were assigned, based on individual sighting histories. Group size ranged between 2 and 90 individuals (median 12), with no significant monthly difference in group size. All resident dolphins formed a core, complex network with migrant and transient dolphins. In contrast, satellite clusters were formed exclusively by transient individuals. The lagged identification rate (LIR) model indicated that a dolphin remained in the area for an average of 90 days, whereas the average time an individual spent outside the study area was approximately 313 days. The Madeira Archipelago appears to be only a section of a much larger home range for this population, with the dolphins exhibiting three different patterns of occurrence that associate with each other. The information gained about this poorly studied population contributes to the overall knowledge of bottlenose dolphins in insular oceanic habitats.     

Bacterial abundance and activity in deep-sea sediments from the eastern North Atlantic

Results are presented from four cruises to the Porcupine Abyssal Plain (PAP site) that took place during the BENGAL project from September 1996 to March 1998, and two cruises to the PAP and an oligotrophic site (EUMELI) that took place during the DEEPSEAS project between September 1993 and March 1994. Bacterial abundances in sediment and sediment contact water were measured by epifluorescence microscopy. Bacterial activity was determined by ³H-thymidine incorporation as a measure of DNA synthesis, and by ³H-leucine incorporation as a measure of protein synthesis. Activities were measured under atmospheric and in situ pressures and temperatures. Bacterial activity was usually higher in samples incubated at in situ pressure than those incubated at atmospheric pressure indicating that a barophilic community was dominant. Inter-cruise comparisons of abundance and activity during the BENGAL project showed no firm evidence of there being a seasonal response in the benthic microbial community to any episodic phytodetritus event. This was probably because of inter-annual variations in the quality and quantity of phytodetritus deposition at the PAP site, the rapid remineralization of fresh organic material by the microbial communities and the timing of cruises to the study area. ³H-thymidine and ³H-leucine incorporation in sediments was higher during the BENGAL period than the DEEPSEAS programme. A methodological change in the ³H-thymidine incorporation technique for sediments may explain the differences in DNA synthesis observed between the two projects, whereas the lower levels of protein synthesis observed during the DEEPSEAS programme probably reflected both inter-annual variations in activity at the PAP site and the lower productivity that prevailed at surface at the EUMELI oligotrophic site. Rates of ³H-thymidine incorporation in sediment contact water were similar during both projects.     

Nitrogen cycling in deep-sea sediments of the Porcupine Abyssal Plain, NE Atlantic

Rates of transformation, recycling and burial of nitrogen and their temporal and spatial variability were investigated in deep-sea sediments of the Porcupine Abyssal Plain (PAP), NE Atlantic during eight cruises from 1996 to 2000. Benthic fluxes of ammonium (NH₄) and nitrate (NO₃) were measured in situ using a benthic lander. Fluxes of dissolved organic nitrogen (DON) and denitrification rates were calculated from pore water profiles of DON and NO₃, respectively. Burial of nitrogen was calculated from down core profiles of nitrogen in the solid phase together with ¹⁴C-based sediment accumulation rates and dry bulk density. Average NH₄ and NO₃-effluxes were 7.4 ± 19 μmol m⁻² d⁻¹ (n = 7) and 52 ± 30 μmol m⁻² d⁻¹ (n = 14), respectively, during the period 1996–2000. During the same period, the DON-flux was 11 ± 5.6 μmol m⁻² d⁻¹ (n = 5) and the denitrification rate was 5.1 ± 3.0 μmol m⁻² d⁻¹ (n = 22). Temporal and spatial variations were only found in the benthic NO₃ fluxes. The average burial rate was 4.6 ± 0.9 μmol m⁻² d⁻¹. On average over the sampling period, the recycling efficiency of the PON input to the sediment was 94% and the burial efficiency hence 6%. The DON flux constituted 14% of the nitrogen recycled, and it was of similar magnitude as the sum of burial and denitrification. By assuming the PAP is representative of all deep-sea areas, rates of denitrification, burial and DON efflux were extrapolated to the total area of the deep-sea floor (>2000 m) and integrated values of denitrification and burial of 8 ± 5 and 7 ± 1 Tg N year⁻¹, respectively, were obtained. This value of total deep-sea sediment denitrification corresponds to 3–12% of the global ocean benthic denitrification. Burial in deep-sea sediments makes up at least 25% of the global ocean nitrogen burial. The integrated DON flux from the deep-sea floor is comparable in magnitude to a reported global riverine input of DON suggesting that deep-sea sediments constitute an important source of DON to the world ocean.     

Neogene sedimentation patterns in north-central peninsular Florida

Sea level fluctuations and shoreline migrations are responsible for commonly oriented ridges and hills of north-central Florida. Lake Wales Ridge is the principal geomorphic feature of the region, with younger ridges, including Trail Ridge, occurring to the north and east.Sand is the dominant sediment size of the deposits forming these ridges, but gravels and clay beds also occur. Cross-laminations and burrowing are prominent in certain horizons. Generalized sections resemble marine regressive patterns and a marine barrier environment. The deposits reflect a prograding shoreline and the subaerial evolution of the Florida peninsula.     

First insights into macro- and meiofaunal colonisation patterns on paired wood/slate substrata at Atlantic deep-sea hydrothermal vents

In 2006, paired wood and slate panels, each equipped with a temperature probe, were deployed on three different localities on and around the Eiffel Tower edifice (Lucky Strike vent field, Mid-Atlantic Ridge) within close proximity of visible hydrothermal activity. Recovery of these panels took place in 2008. For this two-year deployment period, the composition of colonising organisms (both macro-and meiofauna) was assessed, along with image analyses of the deployment sites in 2006 and 2008. Very few significant differences in colonisation between organic (wood) and inorganic (slate) panels were revealed. Rather, the locality of deployment and the local environmental conditions and hydrothermal activity were found to influence taxonomic composition. Variability in microhabitat conditions and biological interactions were hypothesised to interact jointly in shaping new faunal communities on the colonisation substrata.     

Cenozoic alongslope processes and sedimentation on the NW European Atlantic margin

Based on studies of sediment accumulations deposited from-and erode by-alongslope flowing ocean currents on the European continental margin from Porcupine (Ireland) to Lofoten (Norway), the evolution of the Cenozoic paleocirculation was reconstructed as part of the STRATAGEM project. There is evidence of ocean current-controlled erosion and deposition in the Rockall Trough, in the Faeroe-Shetland Channel and on the Vøring Plateau since the late Eocene, although the circulation pattern remains ambiguous. The late Palaeogene flow in the Rockall Trough was almost probably driven by southerly-derived Tethyan Outflow Water. The extent and strength of any northerly-derived flow is uncertain. From the early Neogene (early-mid-Miocene), there was a massive regional expansion of contourite drift development both in the North Atlantic and in the Norwegian-Greenland Sea. This was most probably related to the development of the Faroe Conduit, the opening of the Fram Strait and the general subsidence of the Greenland-Scotland Ridge. These may have combined to cause a considerable acceleration in the exchange and overflow of deep waters between the Arctic and Atlantic Oceans. An early late Neogene (late early Pliocene) regional erosional event has been ascribed to a vigorous pulse of bottom-current activity, most probably the result of a global reorganisation of ocean currents associated with the closure of the Central American Seaway. During the late Neogene, contourites and sediment drifts developed in deep-water basins, between units of glacigenic sediments as well as infill of several paleo-slide scars. These sediments were derived from areas of bottom-current erosion as well as from the development of Plio-Pleistocene prograding sediment wedges, incorporating the extensive sediment supply derived from shelf-wide ice sheets. Presently a profound winnowing prevails along the shelf and upper slope due to the inflowing currents of Atlantic water. Depocentres of sediments derived from the winnowing are located (locally) in lower slope embayments and in slide scars.     

Geotechnical characteristics of deep-sea sediments from the North Atlantic and North Pacific oceans

Geotechnical properties from a series of deep-sea sites in the North Atlantic and North Pacific oceans are examined to evaluate overall trends and to compare with similar fine-grained soils found on land. The study areas encompass a range of sedimentary environments dominated by combinations of turbidite and pelagic deposits. Carbonate content in excess of 20% is seen to result in a decrease in liquid limit and compressibility. Vertical profiles of geotechnical properties in the North Pacific show broader changes in down-core geotechnical properties compared to the North Atlantic and reflect the effects of long-term climatic changes and seafloor spreading. Sediments in the North Atlantic indicate significant differences depending on location, which is attributed to variability in turbidite deposition, water depth, distance from sediment sources, and the effects of bottom currents. Compared to equivalent fine-grained soils on land, deep-sea sediments are generally softer, more compressible and have higher friction angles at comparable Atterberg limits. Deeper and older sediments in the North Pacific are characterized by unusually large plastic limits, which are attributed to the presence of volcanic fractions. Empirical relationships for compression index and friction angle are discussed for sediments from both oceans.     

Bathymetric patterns of megafaunal assemblages from the arctic deep-sea observatory HAUSGARTEN

Five photographic transects, covering some 830 m² of seafloor in total, were analyzed to characterize the megabenthic community along a bathymetric gradient covering water depths from 1200 to 5500 m in the eastern Fram Strait. Megafaunal densities ranged between 11 and 38 ind. m⁻². The highest densities were found at 1650 m and the lowest densities occurred at 3000 m depth. The number of taxa and morphotypes ranged between 4 at 5500 m and 27 at 1650 m water depth. Ophiocten gracilis, a small white unidentified amphipod, Kolga hyalina, and Bathycrinus carpenteri were the dominant and characteristic species on the slope and continental rise. Elpidia heckeri dominated in the Molloy Hole, the deepest depression known in the Arctic Ocean. Megafaunal zonation patterns appeared to be mainly controlled by food availability, as indicated by phytodetrital matter measured at the seafloor, and by benthic biomass in the sediments, as indicated by sediment-bound particulate proteins and phospholipids. By contrast, physical factors, including water depth and seabed properties such as sediment porosity and hard substrata (e.g., dropstones), appear to play a secondary role in determining megabenthic zonation patterns along the bathymetric HAUSGARTEN gradient.     

Species richness in Atlantic deep-sea fishes assessed in terms of the mid-domain effect and Rapoport's rule

A decrease in species richness with increasing latitude has been documented for a broad range of taxonomic groups. A number of hypotheses relating to biological, environmental, and historical factors have been proposed to explain this phenomenon, and the mid-domain effect (MDE) has been proposed in the form of a null model. This model considers only the geometry of spatial gradients and species’ range extents, excluding any assumptions of environmental, biological or historical causes, and predicts that species richness will peak in the centre of a domain in which species occur when their ranges are randomly distributed. This model has been applied to observed latitudinal, elevational and depth gradients as a test to quantify the extent to which non-random processes influence species richness patterns in comparison to those based on geographical boundary constraints alone. We apply the MDE model to empirical datasets for the ranges of the bottom-living fish species occurring in the Faroe-Iceland Ridge, Denmark Strait, Southern New England and Northern Gulf of Mexico regions of the North Atlantic Ocean. The observed patterns show a decline in richness with depth, and do not match the richness patterns produced by the null model. Therefore it can be said that non-random processes have resulted in the observed patterns. Applied to bathymetric ranges, Rapoport's rule predicts that richness decreases and range size increases with depth and latitude. The rule explained decreasing fish species richness with depth and between latitudes, but did not appear to explain increasing range size with depth.     

Intra-annual variability of extremely rapid sedimentation onto Gardar Drift in the northern North Atlantic

North Atlantic sediment drifts are valuable archives for paleoceanographic reconstructions spanning various timescales. However, the short-term dynamics of such systems are poorly known, and this impinges on our ability to quantitatively reconstruct past change. Here we describe a high-resolution 319-day time-series of hydrodynamics and near-bottom (4 m) particulate matter flux variability at a 2600 m deep site with an extremely high sediment accumulation rate on the southern Gardar Drift in the North Atlantic. We compare our findings with the actual deposits at the site. The total annual particle flux amounted to ～360 g m^?2 yr^?1, varied from ～0.15 to >5.0 g m^?2 day^?1 and displayed strong seasonal compositional changes, with the highest proportion of fresh biogenic matter arriving after the spring bloom in June and July. Flux variability also depended on the changing input of lithogenic matter that had been (re)suspended for a longer time (decades). Active focussing of material from both sources is required to account for the composition and the magnitude of the total flux, which exceed observations elsewhere by an order of magnitude. The enhanced focussing or increased delivery appeared to be positively related to current velocity. The intercepted annual particle flux accounted for only 60% of the sediment accumulation rate of 600±20 g m^?2 yr^?1 (0.20±0.07 cm yr^?1), indicating higher intra- and inter-annual variability of both the biogenic and lithogenic fluxes and/or advection of additional sediment closer to the seafloor (i.e. <4 m). This temporal variability in the composition and amount of material deposited highlights intra-annual changes in the flux of lithogenic material, but also underscores the importance of (reworked) sediment focussing and seasonality of the biogenic flux. All should be taken into account in the interpretation of the paleorecord from such depositional settings.     

Biogeochemical patterns in the Atlantic Inflow through the Strait of Gibraltar

The effects of tidal forcing on the biogeochemical patterns of surface water masses flowing through the Strait of Gibraltar are studied by monitoring the Atlantic Inflow (AI) during both spring and neap tides. Three main phenomena are defined depending on the strength of the outflowing phase predicted over the Camarinal Sill: non-wave events (a very frequent phenomenon during the whole year); type I Internal wave events (a very energetic event, occurring during spring tides); and type II Internal wave events (less intense, occurring during neap tides).During neap tides, a non-wave event comprising oligotrophic open-ocean water from the Gulf of Cádiz is the most frequent and clearly dominant flow through the Strait. In this tidal condition, the inflow of North Atlantic Central Water (NACW) provides the main nutrient input to the surface layer of the Alboran Sea, supplying almost 70% of total annual nitrate transport to the Mediterranean basin. A low percentage of active and large phytoplankton cells and low average concentrations of chlorophyll (0.3–0.4 mg m⁻³) are found in this tidal phase. Around 50% of total annual phytoplankton biomass transport into the Mediterranean Sea through the Strait presents these oligotrophic characteristics.In contrast, during spring tides, patches of water with high chlorophyll levels (0.7–1 mg m⁻³) arrive intermittently, and these are recorded concurrently with the passage of internal waves coming from the Camarinal Sill (type I internal wave events). When large internal waves are arrested over the Camarinal Sill this implies strong interfacial mixing and the probable concurrent injection of coastal waters into the main channel of the Strait. These processes result in a mixed water column in the AI and can account for around 30% of total annual nitrate transport into the Mediterranean basin. Associated with type I internal wave events there is a regular inflow of large and active phytoplankton cells, transported in waters with relatively high nutrient concentrations, which constitutes a significant supply of planktonic resources to the pelagic ecosystem of the Alboran Sea (almost 30% of total annual phytoplankton biomass transport).