Sediment DNA metabarcoding and morphology provide complementary insight into macrofauna and meiobenthos response to environmental gradients in an Arctic glacial fjord.

Arctic fjords ecosystems are highly dynamic, with organisms exposed to various natural stressors along with productivity clines driven by advection of water masses from shelves. The benthic response to these environmental clines has been extensively studied using traditional, morphology-based approaches mostly focusing on macroinvertebrates. In this study we analyse the effects of glacially mediated disturbance on the biodiversity of benthic macrofauna and meiobenthos (meiofauna and Foraminifera) in a Svalbard fjord by comparing morphology and eDNA metabarcoding. Three genetic markers targeting metazoans (COI), meiofauna (18S V1V2) and Foraminifera (18S 37f) were analyzed. Univariate measures of alpha diversity and multivariate compositional dissimilarities were calculated and tested for similarities in response to environmental gradients using correlation analysis. Our study showed different taxonomic composition of morphological and molecular datasets for both macrofauna and meiobenthos. Some taxonomic groups while abundant in metabarcoding data were almost absent in morphology-based inventory and vice versa. In general, species richness and diversity measures in macrofauna morphological data were higher than in metabarcoding, and similar for the meiofauna. Both methodological approaches showed different patterns of response to the glacially mediated disturbance for the macrofauna and the meiobenthos. Macrofauna showed an evident distinction in taxonomic composition and a dramatic cline in alpha diversity indices between the outer and inner parts of fjord, while the meiobenthos showed a gradual change and more subtle responses to environmental changes along the fjord axis. The two methods can be seen as complementing rather than replacing each other. Morphological approach provides more accurate inventory of larger size species and more reliable quantitative data, while metabarcoding allows identification of inconspicuous taxa that are overlooked in morphology-based studies. As different taxa may show different sensitivities to environmental changes, both methods shall be used to monitor marine biodiversity in Arctic ecosystems and its response to dramatically changing environmental conditions.

Shell mineralogy and chemistry - Arctic bivalves in a global context.

This study provided new data on shell mineralogy in 23 Arctic bivalve species. The majority of examined species had purely aragonitic shells. Furthermore, we measured concentrations of Al, Ba, Ca, Fe, K, Mg, Mn, Na, P, S, Sr and Zn in 542 shells representing 25 Arctic bivalve species. Species-related differences in concentrations of specific elements were significant and occurred regardless of locations and water depths. This observation implies the dominance of biological processes regulating elemental uptake into the skeleton over factors related to the variability of abiotic environmental conditions. Analysis of the present study and literature data revealed that the highest concentrations of metals were observed in bivalves collected in the temperate zone, with intermediate levels in the tropics and the lowest levels in polar regions. This trend was ascribed mainly to the presence of higher anthropogenic pressure at temperate latitudes being a potential source of human-mediated metal pollution.

Will shrinking body size and increasing species diversity of crustaceans follow the warming of the Arctic littoral?

Over thirty species of littoral marine Gammaridea occur along the coasts of the North Atlantic. From one to several species can coexist in a single region. There is an evident, inverse relationship between egg incubation time and temperature (from 14 to >120 days) and consequent trends in the size of the animals on reaching maturity (from 5 mm in warmer waters to 30 mm in the coldest ones) and in lifespan (from <6 months to >5 years). Littoral gammarids are a good example of the shrinking size effect of increasing temperatures and size-related species diversity. In large species, the annual cohorts of the population (3-5 annual size groups) functionally replace the adults of smaller species. The ongoing warming of the European Arctic seas may extend the distribution limits of boreal species so that more Gammarus species may appear on northern coasts hitherto occupied by just one or at most two species.

Latitudinal consistency of biomass size spectra - benthic resilience despite environmental, taxonomic and functional trait variability.

Global warming is expected to cause reductions in organism body size, a fundamental biological unit important in determining biological processes. Possible effects of increasing temperature on biomass size spectra in coastal benthic communities were investigated. We hypothesized higher proportions of smaller size classes in warmer conditions. Soft bottom infauna samples were collected in six Norwegian and Svalbard fjords, spanning wide latitudinal (60-81°N) and bottom water temperature gradients (from -2 to 8 °C). Investigated fjords differed in terms of environmental settings (e.g., pigments or organic carbon in sediments). The slopes of normalised biomass size spectra (NBSS) did not differ among the fjords, while the benthic biomass and NBSS intercepts varied and were related to chlorophyll a and δ13C in sediments. The size spectra based on both abundance and biomass remained consistent, regardless of the strong variability in macrofauna taxonomic and functional trait composition. Variable relationships between temperature and body size were noted for particular taxa. Our results indicate that while benthic biomass depends on the nutritional quality of organic matter, its partitioning among size classes is consistent and independent of environmental and biological variability. The observed size structure remains a persistent feature of studied communities and may be resilient to major climatic changes.

Bryozoan zooid size variation across a bathymetric gradient: a case study from the Icelandic shelf and continental slope.

Body size is one of the most important biological characters, as it defines many aspects of organismal functioning at the individual and community level. As body size controls many ecological aspects of species, it is often used as a proxy for the status of the ecosystem. So far no consistent mechanism driving size shift has been proposed. In this study, we investigated bathymetric variability in zooid's size and shape in aquatic colonial animals, Bryozoa. Although the response of bryozoan zooid size to temperature or food concentration has been experimentally proven, the effects of natural environmental variability on marine bryozoan populations has been much less explored. The presented investigation is aimed to assess the bathymetric patterns and environmental drivers of bryozoan zooid size on continental shelf and slope of southern Iceland. 196 colonies of 11 species representing different colonial forms and taxonomic groups were selected for zooid characteristics measurements. A pattern of depth-related increase in zooid size was documented for Bicellarina alderi, Chartella barleei and Sarsiflustra abyssicola, no statistically significant effects were detected for the other eight species. Two species Bicellarina alderi and Caberea ellisii had significantly longer zooids in deeper water, shape of the remaining species did not change along the bathymetric gradient. Intercolonial coefficient of variation in zooid size did not change across the depth gradient. Temperature differences along studied depth could be responsible for the observed pattern.

History of heavy metal accumulation in the Svalbard area: Distribution, origin and transport pathways.

In this study temporal changes of Pb, Zn, Cd and Cu concentrations were studied in 19 dated sediment cores collected from Svalbard fjords and the Barents Sea. The main aim was to study spatial and historical variations in heavy metal concentrations, deposition rates and sources in the context of different metal transport pathways. Metal concentrations ranged from 5.7 to 45.8 mg kg-1 for Pb, from 13.4 to 54.5 mg kg-1 for Cu, from 0.01 to 0.90 mg kg-1 for Cd and from 55.6 to 130.4 mg kg-1 for Zn. Some fjords were unpolluted by heavy metals while in others a clear signal of metal enrichment was found (outer Kongsfjorden, Hornsund, Adventfjorden). Large-scale processes such as atmospheric and oceanic transport were found to be important drivers of heavy metal contaminant distribution. The significance of global drivers varied among the fjords, due to coupling with local processes. Outer fjord parts, the most impacted by oceanic transport, were characterized by the excess 206Pb/207Pb values of ∼1.17, while the inner basins were characterized by the excess 206Pb/207Pb of ∼1.14 suggesting possible different importance of Pb sources (marine currents and atmospheric transport).

Unexpected Levels of Biological Activity during the Polar Night Offer New Perspectives on a Warming Arctic.

The current understanding of Arctic ecosystems is deeply rooted in the classical view of a bottom-up controlled system with strong physical forcing and seasonality in primary-production regimes. Consequently, the Arctic polar night is commonly disregarded as a time of year when biological activities are reduced to a minimum due to a reduced food supply. Here, based upon a multidisciplinary ecosystem-scale study from the polar night at 79°N, we present an entirely different view. Instead of an ecosystem that has entered a resting state, we document a system with high activity levels and biological interactions across most trophic levels. In some habitats, biological diversity and presence of juvenile stages were elevated in winter months compared to the more productive and sunlit periods. Ultimately, our results suggest a different perspective regarding ecosystem function that will be of importance for future environmental management and decision making, especially at a time when Arctic regions are experiencing accelerated environmental change [1].

Seagrass vegetation and meiofauna enhance the bacterial abundance in the Baltic Sea sediments (Puck Bay).

This study presents the first report on bacterial communities in the sediments of eelgrass (Zostera marina) meadows in the shallow southern Baltic Sea (Puck Bay). Total bacterial cell numbers (TBNs) and bacteria biomass (BBM) assessed with the use of epifluorescence microscope and Norland's formula were compared between bare and vegetated sediments at two localities and in two sampling summer months. Significantly higher TBNs and BBM (PERMANOVA tests, P < 0.05) were recorded at bottom covered by the seagrass meadows in both localities and in both sampling months. The relationships between bacteria characteristics and environmental factors (grain size, organic matter, photopigments in sediments), meiofauna and macrofauna densities, as well as macrophyte vegetation characteristics (shoot density, phytobenthos biomass) were tested using PERMANOVA distance-based linear model (DISTLM) procedures and showed that the main factors explaining bacteria characteristics are bottom type (vegetated vs. unvegetated) and meiofauna density. These two factors explained together 48.3% of variability in TBN and 40.5% in BBM, and their impacts did not overlap (as indicated by DISTLM sequential tests) demonstrating the different natures of these relationships. The effects of seagrass were most probably related to the increase of organic matter and providing habitat while higher numbers of meiofauna organisms may have stimulated the bacterial growth by increased grazing.

Evidence of season-dependency in vegetation effects on macrofauna in temperate seagrass meadows (Baltic Sea).

Seagrasses and associated macrophytes are important components of coastal systems as ecosystem engineers, habitat formers, and providers of food and shelter for other organisms. The positive impacts of seagrass vegetation on zoobenthic abundance and diversity (as compared to bare sands) are well documented, but only in surveys performed in summer, which is the season of maximum canopy development. Here we present the results of the first study of the relationship between the seasonal variability of seagrass vegetation and persistence and magnitude of contrasts in faunal communities between vegetated and bare sediments. The composition, abundance, biomass, and diversity of macrozoobenthos in both habitats were compared five times throughout the year in temperate eelgrass meadows in the southern Baltic Sea. Significant positive effects of macrophyte cover on invertebrate density and biomass were recorded only in June, July, and October when the seagrass canopy was relatively well developed. The effects of vegetation cover on faunal species richness, diversity, and composition persisted throughout the year, but the magnitude of these effects varied seasonally and followed changes in macrophyte biomass. The strongest effects were observed in July and coincided with maximums in seagrass biomass and the diversity and biomass of other macrophytes. These observations indicate that in temperate, clearly seasonal systems the assessment of macrophyte impact cannot be based solely on observations performed in just one season, especially when that season is the one in which macrophyte growth is at its maximum. The widely held belief that macrophyte cover strongly influences benthic fauna in marine coastal habitats, which is based on summer surveys, should be revisited and complemented with information obtained in other seasons.

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.