Bhavani E. Narayanaswamy

Global Patterns and Predictions of Seafloor Biomass Using Random Forests

A comprehensive seafloor biomass and abundance database has been constructed from 24 oceanographic institutions worldwide within the Census of Marine Life (CoML) field projects. The machine-learning algorithm, Random Forests, was employed to model and predict seafloor standing stocks from surface primary production, water-column integrated and export particulate organic matter (POM), seafloor relief, and bottom water properties. The predictive models explain 63% to 88% of stock variance among the major size groups. Individual and composite maps of predicted global seafloor biomass and abundance are generated for bacteria, meiofauna, macrofauna, and megafauna (invertebrates and fishes). Patterns of benthic standing stocks were positive functions of surface primary production and delivery of the particulate organic carbon (POC) flux to the seafloor. At a regional scale, the census maps illustrate that integrated biomass is highest at the poles, on continental margins associated with coastal upwelling and with broad zones associated with equatorial divergence. Lowest values are consistently encountered on the central abyssal plains of major ocean basins The shift of biomass dominance groups with depth is shown to be affected by the decrease in average body size rather than abundance, presumably due to decrease in quantity and quality of food supply. This biomass census and associated maps are vital components of mechanistic deep-sea food web models and global carbon cycling, and as such provide fundamental information that can be incorporated into evidence-based management.

Temporal variability in phytodetritus and megabenthic activity at the seabed in the deep Northeast Atlantic

We report a ten-year study of the abundance and activity of megabenthos on the Porcupine Abyssal Plain, northeast Atlantic, together with observations on the occurrence of phytodetritus at the deep-sea floor (4850 m). Using the Southampton Oceanography Centre time-lapse camera system, ‘Bathysnap’, we have recorded a radical change in the abundance and activity of megabenthos between the two periods of study (1991–1994 and 1997–2000). In 1991–1994, the larger megabenthos occurred at an abundance of c. 71.6/ha and were dominated by large holothurians. In addition, there were very substantial populations of smaller megabenthic ophiuroids (c. 4979/ha). Together, the total megabenthos are estimated to track over some 17 cm2/m2/d (exploiting 100% of the surface of the seabed in c. 2.5 years). In 1997–2000, the larger megabenthos increased to an abundance of c. 204/ha and were joined by exceptional numbers of a small holothurian species (Amperima rosea, 6457/ha) and ophiuroids (principally Ophiocten hastatum, 53,539/ha). The total megabenthos population was tracking at an estimnated rate of c. 247 cm2/m2/d (exploiting 100% of seabed in just 6 weeks). Coincident with these increases in the abundance and activity of the megabenthos, there were apparently no mass depositions of aggregated phytodetritus to the seabed in the summers of 1997–1999. Mass occurrences of phytodetritus had been noted during the summer months of the three years previously studied (1991, 1993 and 1994), with covering between 50 and 96% of the sediment surface. There is a statistically significant (p<0.02) negative correlation between maximum extent of this seabed cover of phytodetritus and seabed tracking by megabenthos. Additional studies [Lampitt et al., Progr. Ocean. 50 (2001)], indicate that there were no substantial changes in surface ocean primary productivity, in export flux, or in the composition of the flux that might otherwise account for the apparent absence of observable concentrations of phytodetritus during the summers of 1997–1999. We postulate that the marked increase in megabenthic tracking activity resulted in the removal (via consumption, disaggregation, burial etc.) of the bulk of the incoming phytodetrital flux during these years. A simple conceptual model, based on the apparent phytodetrital fluxes observed in 1991 and 1993, suggests that the megabenthos tracking rates estimated for 1997–1999 are sufficient to account for near-total removal of this flux. However, we are not able to estimate other processes removing phytodetritus (i.e. other elements of the benthos) that may also have increased between 1991–1994 and 1997–1999. Other independent studies [e.g. Ginger et al., Progr. Ocean. 50 (2001)] of flux constituents support the possibility that just a few species of megabenthos (e.g. A. rosea, and O. hastatum) could well have consumed a major proportion of the incoming flux and so substantially modified the composition of the organic matter available to other components of the benthos.

The Relationship between Phytoplankton Distribution and Water Column Characteristics in North West European Shelf Sea Waters

Phytoplankton underpin the marine food web in shelf seas, with some species having properties that are harmful to human health and coastal aquaculture. Pressures such as climate change and anthropogenic nutrient input are hypothesized to influence phytoplankton community composition and distribution. Yet the primary environmental drivers in shelf seas are poorly understood. To begin to address this in North Western European waters, the phytoplankton community composition was assessed in light of measured physical and chemical drivers during the “Ellett Line” cruise of autumn 2001 across the Scottish Continental shelf and into adjacent open Atlantic waters. Spatial variability existed in both phytoplankton and environmental conditions, with clear differences not only between on and off shelf stations but also between different on shelf locations. Temperature/salinity plots demonstrated different water masses existed in the region. In turn, principal component analysis (PCA), of the measured environmental conditions (temperature, salinity, water density and inorganic nutrient concentrations) clearly discriminated between shelf and oceanic stations on the basis of DIN∶DSi ratio that was correlated with both salinity and temperature. Discrimination between shelf stations was also related to this ratio, but also the concentration of DIN and DSi. The phytoplankton community was diatom dominated, with multidimensional scaling (MDS) demonstrating spatial variability in its composition. Redundancy analysis (RDA) was used to investigate the link between environment and the phytoplankton community. This demonstrated a significant relationship between community composition and water mass as indexed by salinity (whole community), and both salinity and DIN∶DSi (diatoms alone). Diatoms of the Pseudo-nitzschia seriata group occurred at densities potentially harmful to shellfish aquaculture, with the potential for toxicity being elevated by the likelihood of DSi limitation of growth at most stations and depths.

Biodiversity Trends along the Western European Margin

The seas along the western European margin encompass a vast geographical area comprising numerous different habitats, and are home to more than 10,000 metazoan species. Although research in this extensive region has been undertaken since the early 1800s, many new species are being described and distributional patterns identified. Recent studies incorporating the most extensive data series ever used in such European studies have failed to find any relationship between latitude and infaunal shelf biodiversity. Along the European shelf, species richness generally increases to a depth of 200 m and then decreases from 300–500 m. In the deep Northeast Atlantic, a unimodal curve illustrates how macrofaunal species diversity changes with depth whilst the megafauna appear to have a bimodal distribution. Regional studies are equivocal in that poleward increases in species diversity have been observed in some studies or taxa, but not in others. In the North Sea, arguably the best studied system in European waters, there appears to be a distinct increase in diversity with increasing latitude. Since this trend is confounded by similar latitudinal gradients in depth and trawling intensity, there is no clear explanation for the biodiversity pattern. Climatic shifts in diversity patterns and species ranges have recently been observed. Here we report previously unpublished data on changes in species richness that have been observed along the Norwegian coast over the past two decades, with the most northerly region seeing more than a 15% increase in the number of species being discovered there. This review synthesizes published and new biodiversity data across multiple spatial and temporal scales, and from the coast to the deep-sea, to provide an overview of what is known along the western European margin. Threats to the biodiversity of the region are highlighted, as well as identifying where there are still gaps in our knowledge.

Synthesis of Knowledge on Marine Biodiversity in European Seas: From Census to Sustainable Management

The recently completed European Census of Marine Life, conducted within the framework of the global Census of Marine Life programme (2000–2010), markedly enhanced our understanding of marine biodiversity in European Seas, its importance within ecological systems, and the implications for human use. Here we undertake a synthesis of present knowledge of biodiversity in European Seas and identify remaining challenges that prevent sustainable management of marine biodiversity in one of the most exploited continents of the globe. Our analysis demonstrates that changes in faunal standing stock with depth depends on the size of the fauna, with macrofaunal abundance only declining with increasing water depth below 1000 m, whilst there was no obvious decrease in meiofauna with increasing depth. Species richness was highly variable for both deep water macro- and meio- fauna along latitudinal and longitudinal gradients. Nematode biodiversity decreased from the Atlantic into the Mediterranean whilst latitudinal related biodiversity patterns were similar for both faunal groups investigated, suggesting that the same environmental drivers were influencing the fauna. While climate change and habitat degradation are the most frequently implicated stressors affecting biodiversity throughout European Seas, quantitative understanding, both at individual and cumulative/synergistic level, of their influences are often lacking. Full identification and quantification of species, in even a single marine habitat, remains a distant goal, as we lack integrated data-sets to quantify these. While the importance of safeguarding marine biodiversity is recognised by policy makers, the lack of advanced understanding of species diversity and of a full survey of any single habitat raises huge challenges in quantifying change, and facilitating/prioritising habitat/ecosystem protection. Our study highlights a pressing requirement for more complete biodiversity surveys to be undertaken within contrasting habitats, together with investigations in biodiversity-ecosystem functioning links and identification of separate and synergistic/cumulative human-induced impacts on biodiversity.

Ecology of bathyal polychaete fauna at an Arctic–Atlantic boundary (Faroe–Shetland Channel, North-east Atlantic)

By reference to a series of 15 sampling stations spanning the West Shetland Slope (150–1000 m; Faroe–Shetland Channel, North-east Atlantic) we examined the potential environmental controls on the standing stock, diversity and composition of the polychaete fauna. In contrast to the majority of studied bathyal environments, the Faroe–Shetland Channel has a highly complex and dynamic hydrographic regime, particularly notable for extreme thermal variability at mid-slope depths (i.e. 7°C range at ca. 500 m). Contrary to general expectation, polychaete biomass increased (rather than decreased) with depth. Species diversity exhibited a parabolic pattern with depth, maximum diversity occurring at depths of 350–550 m, rather shallower than observed in other bathyal studies, and possibly linked with a maximum in habitat temperature range. Multivariate analyses of faunal composition suggested a separation of the sampling stations into a shallower and a deeper group, with temperature exerting a major control on polychaete species distributions. The decline in diversity below 600 m (i.e. the descending limb of the parabolic relationship) may be a result of historically limited immigration/recolonization of the thermally isolated Arctic deep-water basins that feed the cold-water flow through the Faroe–Shetland Channel. The bathymetric distribution of polychaetes and other benthos in this region appears to be intimately linked with the thermal regime, having a long-term impact (geological timescales) on the deep-water species pool and leading to local enhancement of diversity where cold- and warm-water masses meet and mix.

Deep-sea litter: A comparison of seamounts, banks and a ridge in the Atlantic and Indian Oceans reveals both environmental and anthropogenic factors impact accumulation and composition

Marine litter is a global challenge that has recently received policymakers’ attention, with new environmental targets in addition to changes to old legislation. There are no global estimates of benthic litter because of the scarcity of data and only patchy survey coverage. However, estimates of baseline abundance and composition of litter are vital in order to implement litter reduction policies and adequate monitoring schemes. Two large-scale surveys of submarine geomorphological features in the Indian and Atlantic Oceans reveal that litter was found at all locations, despite their remoteness. Litter abundance was patchy, but both surveyed oceans had sites of high litter density. There was a significant difference in the type of litter found in the two oceans, with the Indian Ocean sites being dominated by fishing gear, whereas the Atlantic Ocean sites displayed a greater mix of general refuse. This study suggests that seabed litter is ubiquitous on raised benthic features, such as seamounts. It also concludes that the pattern of accumulation and composition of the litter is determined by a complex range of factors both environmental and anthropogenic. We suggest that the tracing of fishing effort and gear type would be an important step to elucidate hotspots of litter abundance on seamounts, ridges and banks.

Optimisation of enzymatic digestion and validation of specimen preservation methods for the analysis of ingested microplastics

Microplastics are considered to be a widespread environmental contaminant. Due to their small size microplastics have the potential to be ingested by a range of aquatic organisms which mistake them for a food source and can suffer adverse impacts as a result. Development of standardised methods is imperative to provide reliable and meaningful data when analysing microplastic ingestion by marine fauna. A range of proteolytic digestive enzymes (trypsin, papain and collagenase) were tested to establish optimum digestion efficacy of biological samples and assess the effects of enzymes on microplastics; additionally the applicability of freezing and formaldehyde followed by ethanol as specimen preservation techniques for microplastic research was investigated. Of the enzymes investigated, trypsin yielded the greatest digestive efficacy based on weight reduction (88% ± 2.52 S.D.) at the lowest concentration (0.3125%) with no observed impacts on microplastics. Enumeration of microplastics from wild collected Mytilus edulis revealed mean numbers of 1.05 ± 0.66 S.D. (minimum) to 4.44 ± 3.03 S.D. (maximum) microplastic particles per g wet weight mussel tissue depending on location. There was no significant difference based on preservation method on the quantification of ingested microplastics and no detrimental impacts were observed on the microplastics directly. Enzymatic digestion using trypsin therefore provides a suitable, time and cost effective method to extract microplastics from M. edulis. Furthermore the preservation methods did not have detrimental effects on microplastics, serving to highlight the suitability of biological samples preserved either way for future inquiries into ingested microplastics.

Feeding Preferences of Abyssal Macrofauna Inferred from In Situ Pulse Chase Experiments

Climatic fluctuations may significantly alter the taxonomic and biochemical composition of phytoplankton blooms and subsequently phytodetritus, the food source for the majority of deep-sea communities. To examine the response of abyssal benthic communities to different food resources we simulated a food sedimentation event containing diatoms and coccolithophorids at Station M in the NE Pacific. In one set of experiments we measured incorporation of diatomC and coccoN into the macrofauna using isotopically enriched 13C-diatoms and 15N-coccolithophores. In a second experiment we measured incorporation of C and N from dual-labelled (13C and 15N) diatoms. The second experiment was repeated 2 months later to assess the effect of seasonality. The simulated food pulses represented additions of 650 – 800 mg C m−2 and 120 mg N m−2 to the seafloor. In all cases rapid incorporation of tracer was observed within 4 days, with between 20% and 52% of the macrofauna displaying evidence of enrichment. However, incorporation levels of both diatomC and coccoN were low (<0.05% and 0.005% of the added C and N). Incorporation of labelled diatoms was similar during both June and September suggesting that the community was not food limited during either period. We found no evidence for selective ingestion of the different food types in the metazoan fauna suggesting that macrofauna do not have strong preferences for diatom vs. coccolithophore dominated phytodetrital pulses. C∶N ratios from both experiments suggest that the metazoan macrofauna community appear to have higher C demands and/or assimilation efficiencies compared to N. Concomitantly, the foraminifera preferentially selected for diatomN over coccoN, and we suggest that this may be related to foraminiferal requirements for intracellular nitrate. These experiments provide evidence that abyssal faunal feeding strategies are in part driven by an organisms internal stoichiometric budgets and biochemical requirements.

Benthic Assemblages of the Anton Dohrn Seamount (NE Atlantic): Defining Deep-Sea Biotopes to Support Habitat Mapping and Management Efforts with a Focus on Vulnerable Marine Ecosystems

In 2009 the NW and SE flanks of Anton Dohrn Seamount were surveyed using multibeam echosounder and video ground-truthing to characterise megabenthic biological assemblages (biotopes) and assess those which clearly adhere to the definition of Vulnerable Marine Ecosystems, for use in habitat mapping. A combination of multivariate analysis of still imagery and video ground-truthing defined 13 comprehensive descriptions of biotopes that function as mapping units in an applied context. The data reveals that the NW and SE sides of Anton Dohrn Seamount (ADS) are topographically complex and harbour diverse biological assemblages, some of which agree with current definitions of ‘listed’ habitats of conservation concern. Ten of these biotopes could easily be considered Vulnerable Marine Ecosystems; three coral gardens, four cold-water coral reefs, two xenophyophore communities and one sponge dominated community, with remaining biotopes requiring more detailed assessment. Coral gardens were only found on positive geomorphic features, namely parasitic cones and radial ridges, found both sides of the seamount over a depth of 1311–1740 m. Two cold-water coral reefs (equivalent to summit reef) were mapped on the NW side of the seamount; Lophelia pertusa reef associated with the cliff top mounds at a depth of 747–791 m and Solenosmilia variabilis reef on a radial ridge at a depth of 1318-1351 m. Xenophyophore communities were mapped from both sides of the seamount at a depth of 1099–1770 m and were either associated with geomorphic features or were in close proximity (< 100 m) to them. The sponge dominated community was found on the steep escarpment either side of the seamount over at a depth of 854-1345 m. Multivariate diversity revealed the xenophyophore biotopes to be the least diverse, and a hard substratum biotope characterised by serpulids and the sessile holothurian, Psolus squamatus, as the most diverse.