A global seasonal surface ocean climatology of phytoplankton types based on CHEMTAX analysis of HPLC pigments

Much advancement has been made in recent years in field data assimilation, remote sensing and ecosystem modeling, yet our global view of phytoplankton biogeography beyond chlorophyll biomass is still a cursory taxonomic picture with vast areas of the open ocean requiring field validations. High performance liquid chromatography (HPLC) pigment data combined with inverse methods offer an advantage over many other phytoplankton quantification measures by way of providing an immediate perspective of the whole phytoplankton community in a sample as a function of chlorophyll biomass. Historically, such chemotaxonomic analysis has been conducted mainly at local spatial and temporal scales in the ocean. Here, we apply a widely tested inverse approach, CHEMTAX, to a global climatology of pigment observations from HPLC. This study marks the first systematic and objective global application of CHEMTAX, yielding a seasonal climatology comprised of ~1500 1°x1° global grid points of the major phytoplankton pigment types in the ocean characterizing cyanobacteria, haptophytes, chlorophytes, cryptophytes, dinoflagellates, and diatoms, with results validated against prior regional studies where possible. Key findings from this new global view of specific phytoplankton abundances from pigments are a) the large global proportion of marine haptophytes (comprising 32 ± 5% of total chlorophyll), whose biogeochemical functional roles are relatively unknown, and b) the contrasting spatial scales of complexity in global community structure that can be explained in part by regional oceanographic conditions. These publicly accessible results will guide future parameterizations of marine ecosystem models exploring the link between phytoplankton community structure and marine biogeochemical cycles.

Global distributions of pteropods (Gymnosomata, Thecosomata, Pseudothecosomata) abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

Pteropods are a group of holoplanktonic gastropods for which global biomass distribution patterns remain poorly resolved. The aim of this study was to collect and synthesize existing pteropod (Gymnosomata, Thecosomata and Pseudothecosomata) abundance and biomass data, in order to evaluate the global distribution of pteropod carbon biomass, with a particular emphasis on its seasonal, temporal and vertical patterns. We collected 25 902 data points from several online databases and a number of scientific articles. The biomass data has been gridded onto a 360 x 180° grid, with a vertical resolution of 33 WOA depth levels. Data has been converted to NetCDF format. Data were collected between 1951-2010, with sampling depths ranging from 0-1000 m. Pteropod biomass data was either extracted directly or derived through converting abundance to biomass with pteropod specific length to weight conversions. In the Northern Hemisphere (NH) the data were distributed evenly throughout the year, whereas sampling in the Southern Hemisphere was biased towards the austral summer months. 86% of all biomass values were located in the NH, most (42%) within the latitudinal band of 30-50° N. The range of global biomass values spanned over three orders of magnitude, with a mean and median biomass concentration of 8.2 mg C l-1 (SD = 61.4) and 0.25 mg C l-1, respectively for all data points, and with a mean of 9.1 mg C l-1 (SD = 64.8) and a median of 0.25 mg C l-1 for non-zero biomass values. The highest mean and median biomass concentrations were located in the NH between 40-50° S (mean biomass: 68.8 mg C l-1 (SD = 213.4) median biomass: 2.5 mg C l-1) while, in the SH, they were within the 70-80° S latitudinal band (mean: 10.5 mg C l-1 (SD = 38.8) and median: 0.2 mg C l-1). Biomass values were lowest in the equatorial regions. A broad range of biomass concentrations was observed at all depths, with the biomass peak located in the surface layer (0-25 m) and values generally decreasing with depth. However, biomass peaks were located at different depths in different ocean basins: 0-25 m depth in the N Atlantic, 50-100 m in the Pacific, 100-200 m in the Arctic, 200-500 m in the Brazilian region and >500 m in the Indo-Pacific region. Biomass in the NH was relatively invariant over the seasonal cycle, but more seasonally variable in the SH. The collected database provides a valuable tool for modellers for the study of ecosystem processes and global biogeochemical cycles.

Diversity of zooplankton assemblages at Station L4, Western English Channel, measured using next generation DNA sequencing

Background: Zooplankton play an important role in our oceans, in biogeochemical cycling and providing a food source for commercially important fish larvae. However, difficulties in correctly identifying zooplankton hinder our understanding of their roles in marine ecosystem functioning, and can prevent detection of long term changes in their community structure. The advent of massively parallel Next Generation Sequencing technology allows DNA sequence data to be recovered directly from whole community samples. Here we assess the ability of such sequencing to quantify the richness and diversity of a mixed zooplankton assemblage from a productive monitoring site in the Western English Channel. Methodology/Principle Findings: Plankton WP2 replicate net hauls (200 µm) were taken at the Western Channel Observatory long-term monitoring station L4 in September 2010 and January 2011. These samples were analysed by microscopy and metagenetic analysis of the 18S nuclear small subunit ribosomal RNA gene using the 454 pyrosequencing platform. Following quality control a total of 419,042 sequences were obtained for all samples. The sequences clustered in to 205 operational taxonomic units using a 97% similarity cut-off. Allocation of taxonomy by comparison with the National Centre for Biotechnology Information database identified 138 OTUs to species level, 11 to genus level and 1 to order, <2.5% of sequences were classified as unknowns. By comparison a skilled microscopic analyst was able to routinely enumerate only 75 taxonomic groups. Conclusions: The percentage of OTUs assigned to major eukaryotic taxonomic groups broadly aligns between the metagenetic and morphological analysis and are dominated by Copepoda. However, the metagenetics reveals a previously hidden taxonomic richness, especially for Copepoda and meroplankton such as Bivalvia, Gastropoda and Polychaeta. It also reveals rare species and parasites. We conclude that Next Generation Sequencing of 18S amplicons is a powerful tool for estimating diversity and species richness of zooplankton communities.

Global distributions of picophytoplankton abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The smallest marine phytoplankton, collectively termed picophytoplankton, have been routinely enumerated by flow cytometry since the late 1980s, during cruises throughout most of the world ocean. We compiled a database of 40,946 data points, with separate abundance entries for Prochlorococcus, Synechococcus and picoeukaryotes. We use average conversion factors for each of the three groups to convert the abundance data to carbon biomass. After gridding with 1° spacing, the database covers 2.4% of the ocean surface area, with the best data coverage in the North Atlantic, the South Pacific and North Indian basins. The average picophytoplankton biomass is 12 ± 22 µg C L-1 or 1.9 g C m-2. We estimate a total global picophytoplankton biomass, excluding N2-fixers, of 0.53 - 0.74 Pg C (17 - 39 % Prochlorococcus, 12 - 15 % Synechococcus and 49 - 69 % picoeukaryotes). Future efforts in this area of research should focus on reporting calibrated cell size, and collecting data in undersampled regions.

Global distributions of diatoms abundance, biovolume and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

This study is a first effort to compile the largest possible body of data available from different plankton databases as well as from individual published or unpublished datasets regarding diatom distribution in the world ocean. The data obtained originate from time series studies as well as spatial studies. This effort is supported by the Marine Ecosystem Data (MAREDAT) project, which aims at building consistent data sets for the main PFTs (Plankton Functional Types) in order to help validate biogeochemical ocean models by using converted C biomass from abundance data. Diatom abundance data were obtained from various research programs with the associated geolocation and date of collection, as well as with a taxonomic information ranging from group down to species. Minimum, maximum and average cell size information were mined from the literature for each taxonomic entry, and all abundance data were subsequently converted to biovolume and C biomass using the same methodology.

Altered neurotransmitter function in CO2-exposed stickleback (Gasterosteus aculeatus): a temperate model species for ocean acidification research

Studies on the consequences of ocean acidification for the marine ecosystem have revealed behavioural changes in coral reef fishes exposed to sustained near-future CO2 levels. The changes have been linked to altered function of GABAergic neurotransmitter systems, because the behavioural alterations can be reversed rapidly by treatment with the GABAA receptor antagonist gabazine. Characterization of the molecular mechanisms involved would be greatly aided if these can be examined in a well-characterized model organism with a sequenced genome. It was recently shown that CO2-induced behavioural alterations are not confined to tropical species, but also affect the three-spined stickleback, although an involvement of the GABAA receptor was not examined. Here, we show that loss of lateralization in the stickleback can be restored rapidly and completely by gabazine treatment. This points towards a worrying universality of disturbed GABAA function after high-CO2 exposure in fishes from tropical to temperate marine habitats. Importantly, the stickleback is a model species with a sequenced and annotated genome, which greatly facilitates future studies on underlying molecular mechanisms.

Intraspecific facilitation by allelochemical mediated grazing protection within a toxigenic dinoflagellate population, link to supplementary material

Dinoflagellates are a major cause of harmful algal blooms, with consequences for coastal marine ecosystem functioning and services. Alexandrium tamarense is one of the most abundant and widespread toxigenic species in the temperate northern and southern hemisphere, and produces paralytic shellfish poisoning toxins as well as lytic allelochemical substances. These bioactive compounds may support the success of A. tamarense and its ability to form blooms. Here we investigate the impact of grazing on monoclonal and mixed set-ups of highly (Alex2) and moderately (Alex4) allelochemically active A. tamarense strains and on a non-allelochemically active conspecific (Alex5) by the heterotrophic dinoflagellate Polykrikos kofoidii. While Alex4 and particularly Alex5 were strongly grazed by P. kofoidii when offered alone, both strains grew well in the mixed assemblages (Alex4+Alex5 and Alex2+Alex5). Hence, the allelochemical active strains facilitated growth of the non-active strain by protecting the population as a whole against grazing. Based on our results, we argue that facilitation among clonal lineages within a species may partly explain the high genotypic and phenotypic diversity of Alexandrium populations. Populations of Alexandrium may comprise multiple cooperative traits that act in concert with intraspecific facilitation, and hence promote the success of this notorious harmful algal bloom species.

Anemone abundance and productivity at North Vulcano Island in May 2011

Increased seawater pCO2, and in turn 'ocean acidification' (OA), is predicted to profoundly impact marine ecosystem diversity and function this century. Much research has already focussed on calcifying reef-forming corals (Class: Anthozoa) that appear particularly susceptible to OA via reduced net calcification. However, here we show that OA-like conditions can simultaneously enhance the ecological success of non-calcifying anthozoans, which not only play key ecological and biogeochemical roles in present day benthic ecosystems but also represent a model organism should calcifying anthozoans exist as less calcified (soft-bodied) forms in future oceans. Increased growth (abundance and size) of the sea anemone (Anemonia viridis) population was observed along a natural CO2 gradient at Vulcano, Italy. Both gross photosynthesis (PG) and respiration (R) increased with pCO2 indicating that the increased growth was, at least in part, fuelled by bottom up (CO2 stimulation) of metabolism. The increase of PG outweighed that of R and the genetic identity of the symbiotic microalgae (Symbiodinium spp.) remained unchanged (type A19) suggesting proximity to the vent site relieved CO2 limitation of the anemones' symbiotic microalgal population. Our observations of enhanced productivity with pCO2, which are consistent with previous reports for some calcifying corals, convey an increase in fitness that may enable non-calcifying anthozoans to thrive in future environments, i.e. higher seawater pCO2. Understanding how CO2-enhanced productivity of non- (and less-) calcifying anthozoans applies more widely to tropical ecosystems is a priority where such organisms can dominate benthic ecosystems, in particular following localized anthropogenic stress.

Biometrics, fatty acid composition and stable isotopic ratios of five seabird species from Kongsfjorden, Svalbard

Marine birds are important predators in the marine ecosystem, and dietary studies can give useful information about their feeding ecology, food webs and oceanographic variability. The aim of this study was to increase our understanding of the diet and trophic level of the seabirds breeding in Kongsfjorden, Svalbard. We have used fatty acids and stable isotopes, both of which integrate diet information over space and time, to determine trophic relationships in marine food webs. Fatty acid compositions of muscle from Little auk (Alle alle), Brünnich's guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) were determined and compared with their prey species. Canonical analysis (CA) showed that fatty acid composition differed among the five seabird species. Little auk, Black-legged kittiwake and Northern fulmar had high levels of the Calanus markers 20:1n9 and 22:1, indicating that these seabirds are a part of the Calanus food chain. Brünnich's guillemot differed from the other species with much lower levels of 20:1n9 and 22:1. Brünnich's guillemot is a pursuit diver feeding on fish and amphipods deeper in the water column, below 30 m. Glaucous gull also differed from the other seabird species, with a larger variation in the fatty acid composition indicating a more diverse diet. Trophic level analysis placed Little auk at the lowest trophic level, Brünnich's guillemot and Black-legged kittiwake at intermediate levels and Glaucous gull and Northern fulmar at the highest trophic level.

Global distributions of microzooplankton abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

Microzooplankton database. Originally published in: Buitenhuis, Erik, Richard Rivkin, Sévrine Sailley, Corinne Le Quéré (2010) Biogeochemical fluxes through microzooplankton. Global Biogeochemical Cycles Vol. 24, GB4015, doi:10.1029/2009GB003601 This new version has had some mistakes corrected.

(Table 1) Movement parameters of nine adult female snowy owls (Bubo scandiacus) tracked during the winter period 07/08 and 08/09 around Hudson Strait, Canada

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea-ice between December and April. Analysis of high-resolution satellite images of sea-ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea-ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea-ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.