Redescription of the deep-sea octopod Benthoctopus normani (Massy 1907) and a description of a new species from the Northeast Atlantic

A long-synonymized species Benthoctopus normani (Massy 1907) (Cephalopoda: Octopodidae) is redescribed from material collected over 30 years by the National Oceanography Centre, Southampton and the National Museums of Scotland. It can be distinguished from other octopodid specimens found in deep waters of the Northeast Atlantic by its biserial suckers, lack of ink sac, and simple ligula, which lacks transverse ridges. Examination of the collections led to the identification of a new species of Benthoctopus from the Northeast Atlantic, which is described herein.

Threats and knowledge gaps for ecosystem services provided by kelp forests: a northeast Atlantic perspective.

Kelp forests along temperate and polar coastlines represent some of most diverse and productive habitats on the Earth. Here, we synthesize information from >60 years of research on the structure and functioning of kelp forest habitats in European waters, with particular emphasis on the coasts of UK and Ireland, which represents an important biogeographic transition zone that is subjected to multiple threats and stressors. We collated existing data on kelp distribution and abundance and reanalyzed these data to describe the structure of kelp forests along a spatial gradient spanning more than 10° of latitude. We then examined ecological goods and services provided by kelp forests, including elevated secondary production, nutrient cycling, energy capture and flow, coastal defense, direct applications, and biodiversity repositories, before discussing current and future threats posed to kelp forests and identifying key knowledge gaps. Recent evidence unequivocally demonstrates that the structure of kelp forests in the NE Atlantic is changing in response to climate- and non-climate-related stressors, which will have major implications for the structure and functioning of coastal ecosystems. However, kelp-dominated habitats along much of the NE Atlantic coastline have been chronically understudied over recent decades in comparison with other regions such as Australasia and North America. The paucity of field-based research currently impedes our ability to conserve and manage these important ecosystems. Targeted observational and experimental research conducted over large spatial and temporal scales is urgently needed to address these knowledge gaps.

Diatom carbon export enhanced by silicate upwelling in the North East Atlantic.

Diatom carbon export enhanced by silicate upwelling in the northeast Atlantic John T. Allen1,2, Louise Brown1,3, Richard Sanders1, C. Mark Moore1, Alexander Mustard1, Sophie Fielding1, Mike Lucas1, Michel Rixen4, Graham Savidge5, Stephanie Henson1 and Dan Mayor1 Top of pageDiatoms are unicellular or chain-forming phytoplankton that use silicon (Si) in cell wall construction. Their survival during periods of apparent nutrient exhaustion enhances carbon sequestration in frontal regions of the northern North Atlantic. These regions may therefore have a more important role in the 'biological pump' than they have previously been attributed1, but how this is achieved is unknown. Diatom growth depends on silicate availability, in addition to nitrate and phosphate2, 3, but northern Atlantic waters are richer in nitrate than silicate4. Following the spring stratification, diatoms are the first phytoplankton to bloom2, 5. Once silicate is exhausted, diatom blooms subside in a major export event6, 7. Here we show that, with nitrate still available for new production, the diatom bloom is prolonged where there is a periodic supply of new silicate: specifically, diatoms thrive by 'mining' deep-water silicate brought to the surface by an unstable ocean front. The mechanism we present here is not limited to silicate fertilization; similar mechanisms could support nitrate-, phosphate- or iron-limited frontal regions in oceans elsewhere.

The uptake of silica during the spring bloom in the North East Atlantic Ocean

A full understanding of the biogeochemical cycling of silica in the North Atlantic is hampered by a lack of estimates of silica uptake by phytoplankton. We applied the ${}^{32}\text{Si}$ radiotracer incubation technique to determine silica uptake rates at 10 sites during the UK-(Natural Environment Research Council) Faroes-Iceland-Scotland hydrographic and environmental survey (FISHES) cruise in the Northeast Atlantic, May 2001. Column silica uptake rates ranged between 6 and 166 mmol Si $\text{m}^{-2}\ \text{d}^{-1}$; this data set was integrated with concurrent hydrographic, chemical, and primary productivity data to explain these changes in silica uptake in terms of the progress of the spring bloom. In order to interpret data covering a relatively large spatial and temporal scale, we used mean photic zone silica concentration as a proxy time-series measure of diatom bloom progression. Both absolute and specific silica uptake rates were highest at dissolved silica concentrations >2 mmol $\text{L}^{-1}$. Si and C uptake were vertically decoupled at those stations where surface silica was strongly depleted. Absolute primary productivity was not strongly correlated with dissolved silica concentrations, owing to either exhaustion of silica at diatom-dominated stations or to dominance of the community by other phytoplankton. Silica uptake as a function of increased substrate concentration was linear up to 25 $\mu \text{mol}\ \text{L}^{-1}$; we consider some possible reasons for the nonhyperbolic response.

Uptake of carbon, nitrogen and phosphorus by phytoplankton along the 20 degrees W meridian in the NE Atlantic between 57.5 degrees N and 37 degrees N

Phytoplankton biomass and rate of production were measured along a transect from 57.54 degreesN to 37.01 degreesN in the northeast Atlantic during July 1996 and at a series of stations over a 7-day period at 37 degreesN 20 degreesW. Surface nutrient concentrations ranged from 4 mu mol l(-1) NO3-, and 0.35 mu mol l(-1) PO43- at 57.54 degreesN to <10 nmol l(-1) NO3- and similar to 10 nmol l(-1) PO43- at 37.01 degreesN. The greatest phytoplankton biomass and production were measured in the vicinity of a frontal system at 50 degreesN, and there was a general decline in total phytoplankton biomass and production to the south of the transect. Production was measured in three size fractions. At the station with the highest chlorophyll concentrations (50.34 degreesN), phytoplankton cells larger than 5 mum dominated the assemblage, accounting for 72% of the chlorophyll concentration (22.9 mg m(-2)) and 51% of primary production (54.1 mmol Cm-2 d(-1)), but picophytoplankton production was also high (43%). At 57 degreesN, carbon fixation by the > 5 mum fraction accounted for 75% of the daily production of 60.75 mmol Cm-2 d(-1). At 37 degreesN, picophytoplankton was the dominant group, accounting for similar to 58% (10 mg m(-2)) of chlorophyll and similar to 64% (46 mmol Cm-2 d(-1)), of primary production. Nitrate, ammonium and phosphate uptake rates also were determined. Although high nitrate uptake rates were measured in the surface water at similar to 50 degreesN, the greatest uptake rates of both depth-integrated nitrate and ammonium were at the south of the transect. At 37 degreesN, a deep euphotic zone was present and light penetrated through the nitracline; total nitrate uptake was enhanced because of assimilation at the base of the euphotic zone. As a consequence, high values of depth-integrated f-ratio were measured in the oligotrophic waters at the south of the transect. Phosphate was predominantly incorporated into the picoplankton fraction, which included heterotrophic and autotrophic components, at all stations and a significant proportion of phosphate uptake occurred in the dark. The C:N:P assimilation ratios were variable throughout the region; phosphate uptake was generally greater than would be expected if nutrient assimilation were in proportion to the Redfield ratio. (C) 2001 Elsevier Science Ltd. All rights reserved.

High-resolution genetic analysis reveals extensive gene flow within the jellyfish Pelagia noctiluca (Scyphozoa) in the North Atlantic and Mediterranean Sea

Despite the importance of gelatinous zooplankton as components of marine ecosystems, both ecologically and socio-economically, relatively little is known about population persistence or connectivity in jellyfish. In the present study, we employed a combination of nuclear microsatellite markers and sequence data from the mitochondrial cytochrome oxidase I (COI) gene to determine levels and patterns of population genetic structuring in the holoplanktonic jellyfish Pelagia noctiluca across the northeast Atlantic Ocean and Mediterranean Sea. Our results indicate a high degree of connectivity in P. noctiluca, with little evidence of geographical structuring of genetic variation. A small but significant differentiation of Atlantic Ocean and Mediterranean stocks was detected based on the microsatellite data, but no evidence of differentiation was observed with the mtDNA, probably due to the higher power of the microsatellites to detect low levels of genetic structuring. Two clearly distinct groups of genotypes were observed within the mtDNA COI, which probably diverged in the early Pleistocene, but with no evidence of geographical structuring. Palaeodistribution modelling of P. noctiluca at the Last Glacial Maximum (LGM; ca. 21 KYA) indicated large areas of suitable habitat south of the species’ current-day distribution, with little reduction in area. The congruent evidence for minimal genetic differentiation from the nuclear microsatellites and the mtDNA, coupled with the results of the palaeodistribution modelling, supports the idea of long-term population stability and connectivity, thus providing key insights into the population dynamics and demography of this important species

Environmental selection on transcriptome-derived SNPs in a high gene flow marine fish, the Atlantic herring (Clupea harengus)

High gene flow is considered the norm for most marine organisms and is expected to limit their ability to adapt to local environments. Few studies have directly compared the patterns of differentiation at neutral and selected gene loci in marine organisms. We analysed a transcriptome-derived panel of 281 SNPs in Atlantic herring (Clupea harengus), a highly migratory small pelagic fish, for elucidating neutral and selected genetic variation among populations and to identify candidate genes for environmental adaptation. We analysed 607 individuals from 18 spawning locations in the northeast Atlantic, including two temperature clines (5-12 °C) and two salinity clines (5-35‰). By combining genome scan and landscape genetic analyses, four genetically distinct groups of herring were identified: Baltic Sea, Baltic-North Sea transition area, North Sea/British Isles and North Atlantic; notably, samples exhibited divergent clustering patterns for neutral and selected loci. We found statistically strong evidence for divergent selection at 16 outlier loci on a global scale, and significant correlations with temperature and salinity at nine loci. On regional scales, we identified two outlier loci with parallel patterns across temperature clines and five loci associated with temperature in the North Sea/North Atlantic. Likewise, we found seven replicated outliers, of which five were significantly associated with low salinity across both salinity clines. Our results reveal a complex pattern of varying spatial genetic variation among outlier loci, likely reflecting adaptations to local environments. In addition to disclosing the fine scale of local adaptation in a highly vagile species, our data emphasize the need to preserve functionally important biodiversity.

Gene-associated markers can assign origin in a weakly structured fish, Atlantic herring

Regulations on the exploitation of populations of commercially important fish species and the ensuing consumer interest in sustainable products have increased the need to accurately identify the population of origin of fish and fish products. Although genomics-based tools have proven highly useful, there are relatively few examples in marine fish displaying accurate origin assignment. We synthesize data for 156 single-nucleotide polymorphisms typed in 1039 herring, Clupea harengus L., spanning the Northeast Atlantic to develop a tool that allows assignment of individual herring to their regional origin. We show the method's suitability to address specific biological questions, as well as management applications. We analyse temporally replicated collections from two areas, the Skagerrak (n = 81, 84, 66) and the western Baltic (n = 52, 52). Both areas harbour heavily fished mixed-origin stocks, complicating management issues. We report novel genetic evidence that herring from the Baltic Sea contribute to catches in the North Sea, and find support that western Baltic feeding aggregations mainly constitute herring from the western Baltic with contributions from the Eastern Baltic. Our study describes a general approach and outlines a database allowing individual assignment and traceability of herring across a large part of its East Atlantic distribution.

Fronts, jumps and secondary introductions suggested as different invasion patterns in marine species, with an increase in spread rates over time

Not all introduced (invasive) species in a region will spread from a single point of introduction. Long-distance dispersal or further introductions can obscure the pattern of spread, but the regional importance of such processes is difficult to gauge. These difficulties are further compounded when information on the multiple scale process of invasive species range expansion is reduced to one-dimensional estimates of spread (e. g. km yr(-1)). We therefore compared the results of two different metrics of range expansion: maximum linear rate of spread and accumulation of occupied grid squares (50 x 50 km) over time. An analysis of records for 54 species of introduced marine macrophytes in the Mediterranean and northeast Atlantic revealed cases where the invasion process was probably missed (e. g. Atlantic Bonnemaisonia hamifera) and suggested cases of secondary introductions or erratic jump dispersal (Dasysiphonia sp. and Womersleyella setacea). A majority of species analysed showed evidence for an accumulation of invaded sites without a clear invasion front. Estimates of spread rate are increasing for more recent introductions. The increase is greater than can be accounted for by temporally varying search effort and implies a historical increase in vector efficiency and/or a decreased resistance of native communities to invasion.

Polyopes lancifolius (Halymeniales, Rhodophyta), a new component of the Japanese marine flora introduced to Europe

Polyopes lancifolius (Harvey) S. Kawaguchi & H.W. Wang has been recorded for the first time in Europe, during the summer or 2008. A small population was discovered in the Gulf of Morbihan (northeast Atlantic, France). This is the first observation of P. lancifolius outside its native range. Vegetative and reproductive morphological features are compared with previous descriptions. rbcL sequences show no divergence from Japanese populations. Imports of Pacific oysters Crassostrea gigas (Thunberg 1793) are likely to be responsible for its accidental introduction into the Gulf of Morbihan, either directly from northwest Pacific regions or indirectly (secondary dispersal) by transfers from another European oyster farming site. The history of previous algal introductions from Japan suggests that if it becomes successfully established at Morbihan, the species is likely to spread to other European coastal areas.

Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment

Leatherback turtles (Dermochelys coriacea) are obligate predators of gelatinous zooplankton. However, the spatial relationship between predator and prey remains poorly understood beyond sporadic and localized reports. To examine how jellyfish (Phylum Cnidaria: Orders Semaeostomeae and Rhizostomeae) might drive the broad-scale distribution of this wide ranging species, we employed aerial surveys to map jellyfish throughout a temperate coastal shelf area bordering the northeast Atlantic. Previously unknown, consistent aggregations of Rhizostoma octopus extending over tens of square kilometers were identified in distinct coastal

Why the size structure of marine communities can require decades to recover from fishing

A dynamic food-web model of more than 1000 species was used to quantify the recovery trajectory of marine community size-structure under different hypothetical fishing regimes, using the Northeast Atlantic as an example. Size-structure was summarised by four indicators: the Large Fish Indicator (LFI), the Large Species Indicator (LSI), the biomass-weighted mean maximum length of fish species (EMBED Equation.3) and the biomass-weighted mean maturation length of fish species (EMBED Equation.3). Time-series of these indicators recorded recovery following release from fishing with various size-selectivities, intensities and durations. In model simulations, fishing-induced trophic cascades were observed to distort fish community size-structure, but these did not have a large influence on recovery level or duration as measured by the four indicators. However, simulations showed that local extinctions of large fish species increased in number with both fishing intensity and duration, and could strongly limit the recovery level. Recovery of fish community size-structure to near equilibrium frequently took multiple decades in simulations; these long transient periods suggest that management interventions for size-structure recovery may require much longer than previously thought. Our results demonstrate the need for community-level modelling to set realistic targets for management of community size-structure.

Identifying marine pelagic ecosystem management objectives and indicators

International policy frameworks such as the Common Fisheries Policy and the European Marine Strategy Framework Directive define high-level strategic goals for marine ecosystems. Strategic goals are addressed via general and operational management objectives. To add credibility and legitimacy to the development of objectives, for this study stakeholders explored intermediate level ecological, economic and social management objectives for Northeast Atlantic pelagic ecosystems. Stakeholder workshops were undertaken with participants being free to identify objectives based on their own insights and needs. Overall 26 objectives were proposed, with 58% agreement in proposed objectives between two workshops. Based on published evidence for pressure-state links, examples of operational objectives and suitable indicators for each of the 26 objectives were then selected. It is argued that given the strong species-specific links of pelagic species with the environment and the large geographic scale of their life cycles, which contrast to demersal systems, pelagic indicators are needed at the level of species (or stocks) independent of legislative region. Pelagic community indicators may be set at regional scale in some cases. In the evidence-based approach used in this study, the selection of species or region specific operational objectives and indicators was based on demonstrated pressure-state links. Hence observed changes in indicators can reliably inform on appropriate management measures