Leanne K. Armand

Effect of natural iron fertilization on carbon sequestration in the Southern Ocean

The availability of iron limits primary productivity and the associated uptake of carbon over large areas of the ocean. Iron thus plays an important role in the carbon cycle, and changes in its supply to the surface ocean may have had a significant effect on atmospheric carbon dioxide concentrations over glacial–interglacial cycles. To date, the role of iron in carbon cycling has largely been assessed using short-term iron-addition experiments. It is difficult, however, to reliably assess the magnitude of carbon export to the ocean interior using such methods, and the short observational periods preclude extrapolation of the results to longer timescales. Here we report observations of a phytoplankton bloom induced by natural iron fertilization—an approach that offers the opportunity to overcome some of the limitations of short-term experiments. We found that a large phytoplankton bloom over the Kerguelen plateau in the Southern Ocean was sustained by the supply of iron and major nutrients to surface waters from iron-rich deep water below. The efficiency of fertilization, defined as the ratio of the carbon export to the amount of iron supplied, was at least ten times higher than previous estimates from short-term blooms induced by iron-addition experiments. This result sheds new light on the effect of long-term fertilization by iron and macronutrients on carbon sequestration, suggesting that changes in iron supply from below—as invoked in some palaeoclimatic and future climate change scenarios—may have a more significant effect on atmospheric carbon dioxide concentrations than previously thought.

Late Quaternary sea ice history in the Indian sector of the Southern Ocean as recorded by diatom assemblages

Abstract A Modern Analog Technique (MAT5201/31) has been applied to fossil diatom assemblages to provide down-core estimates of February sea-surface temperatures (SSTs) and of sea ice duration over the past 220 000 years at 56°40′S, 160°14′E. At the core location, sea ice progression lagged the SST drop by ∼1 ka at interglacial–glacial transitions, and sea ice retreat was almost synchronous to the SST increase at glacial–interglacial terminations. Sea ice increased continuously during glacial periods to reach its maximum extent at the end of glacial times, although SSTs were almost constant during glacials. This indicates that SSTs are the major parameter determining the advance and retreat of sea ice at transitions, but that the sea ice advance during glacial conditions may be related to positive feedbacks of the ice on albedo, air temperature and meridional wind stress. The strong correlation (r=0.75) between sea ice duration at the core location and the Vostok CO2 record argues for a control of Antarctic sea ice extent on atmospheric CO2 concentration via the modification of the ocean-to-atmosphere gas balance.

Continental shelf drift deposit indicates non-steady state Antarctic bottom water production in the Holocene

A late Quaternary, current-lain sediment drift deposit over 30 m in thickness has been discovered on the continental shelf of East Antarctica in an 850 m deep glacial trough off George Vth Land. Radiocarbon dating indicates that a period of rapid deposition on the drift (averaging 290 cm/kyr) occurred in the mid-Holocene, between about 3000 and 5000 yr before present. Slower rates of around 10 cm/kyr, during the past 0-3000 yr and from 5000 to about 13000 yr BP, coincides with deposition of bioturbated, ice-rafted debris (IRD) rich, sandy mud under an energetic bottom current regime. In contrast, the mid-Holocene (3000-5000 yr BP) sediments are fine-grained, laminated to cross-laminated with minimal IRD content, and are contemporaneous with a period of warmer marine conditions with less sea ice production. This pattern suggests that bottom currents were weaker than present day in the mid-Holocene, and that the rate of dense bottom water production was reduced at that time. This scenario is consistent with the hypothesis of non-steady state rates of Antarctic bottom water production through the Holocene as recently proposed by Broecker and his colleagues.

DIATOM SPECIES OF THE GENUS RHIZOSOLENIA FROM SOUTHERN OCEAN SEDIMENTS: DISTRIBUTION AND TAXONOMIC NOTES

The classification of diatom taxa in the genus Rhizosolenia Brightwell in Southern Ocean waters and sediments remains unclear. Our review demonstrates that considerable intertaxa confusion has occurred and continues to occur as a result of misapplied taxonomy. In general, valves of Rhizosolenia taxa within total diatom sediment assemblages are a minor, though common component, with only a few taxa sufficiently abundant for statistical analysis. We have endeavoured to clarify through literature and data set assessment the currently accepted taxon concepts in terms of otaria morphology, this being one of the most useful features available for the discrimination of fossil rhizosolenioids in Southern Ocean sediments. Otaria morphology allows separation of incomplete valves of preserved Rhizosolenia specimens to be determined from descriptions related to the complete cell. The identification of R. styliformis Brightwell, R. antennata f. semispina Sundström, and R. polydactyla Castracane f. polydactyla are addressed here in detail. The presence of Rhizosolenia species A, a taxon with similarities to the otaria-lacking, original descriptions of R. hebetata f. semispina (Hensen) Gran, is described and illustrated. Taxonomic notes, light microscopic representation and distributional data for several other Antarctic Rhizosolenia taxa are provided. The documentation of down-core occurrences of fossil rhizosolenioids in published material is poor. Our core material indicates relative abundance increases for Rhizosolenia species A and R. antennata f. semispina in glacial time sequences, which may constitute a valuable biostratigraphic tool for palaeoenvironmental reconstructions. Comments on winter resting stages are provided.

Latitudinal and temporal distributions of diatom populations in the pelagic waters of the Subantarctic and Polar Frontal zones of the Southern Ocean and their role in the biological pump

Abstract. The Subantarctic and Polar Frontal zones (SAZ and PFZ) represent a large portion of the total area of the Southern Ocean and serve as a strong sink for atmospheric CO2. These regions are central to hypotheses linking particle fluxes and climate change, yet multi-year records of modern flux and the organisms that control it are, for obvious reasons, rare. In this study, we examine two sediment trap records of the flux of diatoms and bulk components collected by two bottom-tethered sediment traps deployed at mesopelagic depths (~ 1 km) in the SAZ (2-year record; July 1999–October 2001) and in the PFZ (6-year record; September 1997–February 1998, July 1999–August 2000, November 2002–October 2004 and December 2005–October 2007) along the 140° E meridian. These traps provide a direct measure of transfer below winter mixed layer depths, i.e. at depths where effective sequestration from the atmosphere occurs, in contrast to study of processes in the surface ocean. Total mass fluxes were about twofold higher in the PFZ (24 ± 13 g m−2 yr−1) than in the SAZ (14 ± 2 g m−2 yr−1). Bulk chemical composition of the particle fluxes mirrored the composition of the distinct plankton communities of the surface layer, being dominated by carbonate in the SAZ and by biogenic silica in the PFZ. Particulate organic carbon (POC) export was similar for the annual average at both sites (1.0 ± 0.1 and 0.8 ± 0.4 g m−2 yr−1 for the PFZ and SAZ, respectively), indicating that the particles in the SAZ were relatively POC rich. Seasonality in the particle export was more pronounced in the PFZ. Peak fluxes occurred during summer in the PFZ and during spring in the SAZ. The strong summer pulses in the PFZ are responsible for a large fraction of the variability in carbon sequestration from the atmosphere in this region. The latitudinal variation of the total diatom flux was found to be in line with the biogenic silica export with an annual flux of 31 ± 5.5 × 108 valves m−2 yr−1 at the PFZ compared to 0.5 ± 0.4 × 108 m−2 yr−1 at the SAZ. Fragilariopsis kerguelensis dominated the annual diatom export at both sites (43 % at the SAZ and 59 % in the PFZ). POC fluxes displayed a strong positive correlation with the relative contribution of a group of weakly silicified and bloom-forming species in the PFZ. Several lines of evidence suggests that the development of these species during the growth season facilitates the formation of aggregates and carbon export. Our results confirm previous work suggesting that F. kerguelensis plays a major role in the decoupling of the carbon and silicon cycles in the high-nutrient low-chlorophyll waters of the Southern Ocean.

Composition of diatom communities and their contribution to plankton biomass in the naturally iron-fertilized region of Kerguelen in the Southern Ocean

In the naturally iron-fertilized surface waters of the northern Kerguelen Plateau region, the early spring diatom community composition and contribution to plankton carbon biomass were investigated and compared with the high nutrient, low chlorophyll (HNLC) surrounding waters. The large iron-induced blooms were dominated by small diatom species belonging to the genera Chaetoceros (Hyalochaete) and Thalassiosira, which rapidly responded to the onset of favorable light-conditions in the meander of the Polar Front. In comparison, the iron-limited HNLC area was typically characterized by autotrophic nanoeukaryote-dominated communities and by larger and more heavily silicified diatom species (e.g. Fragilariopsis spp.). Our results support the hypothesis that diatoms are valuable vectors of carbon export to depth in naturally iron-fertilized systems of the Southern Ocean. Furthermore, our results corroborate observations of the exported diatom assemblage from a sediment trap deployed in the iron-fertilized area, whereby the dominant Chaetoceros (Hyalochaete) cells were less efficiently exported than the less abundant, yet heavily silicified, cells of Thalassionema nitzschioides and Fragilariopsis kerguelensis Our observations emphasize the strong influence of species-specific diatom cell properties combined with trophic interactions on matter export efficiency, and illustrate the tight link between the specific composition of phytoplankton communities and the biogeochemical properties characterizing the study area.

The diatom genus Pseudo-nitzschia (Bacillariophyceae) in New South Wales, Australia: Morphotaxonomy, molecular phylogeny, toxicity, and distribution

Species belonging to the potentially harmful diatom genus Pseudo‐nitzschia, isolated from 16 localities (31 sampling events) in the coastal waters of south‐eastern Australia, were examined. Clonal isolates were characterized by (i) light and transmission electron microscopy; (ii) phylogenies, based on sequencing of nuclear‐encoded ribosomal deoxyribonucleic acid (rDNA) regions and, (iii) domoic acid (DA) production as measured by liquid chromatography–mass spectrometry (LC‐MS/MS). Ten taxa were unequivocally confirmed as Pseudo‐nitzschia americana, P. arenysensis, P. calliantha, P. cuspidata, P. fraudulenta, P. hasleana, P. micropora, P. multiseries, P. multistriata, and P. pungens. An updated taxonomic key for south‐eastern Australian Pseudo‐nitzschia is presented. The occurrence of two toxigenic species, P. multistriata (maximum concentration 11 pg DA per cell) and P. cuspidata (25.4 pg DA per cell), was documented for the first time in Australia. The Australian strains of P. multiseries, a consistent producer of DA in strains throughout the world, were nontoxic. Data from 5,888 water samples, collected from 31 oyster‐growing estuaries (2,000 km coastline) from 2005 to 2009, revealed 310 regulatory exceedances for “Total Pseudo‐nitzschia,” resulting in six toxic episodes. Further examination of high‐risk estuaries revealed that the “P. seriata group” had highest cell densities in the austral summer, autumn, or spring (species dependent), and lowest cell densities in the austral winter, while the “P. delicatissima group” had highest in winter and spring.

First reports of Pseudo-nitzschia micropora and P. hasleana (Bacillariaceae) from the Southern Hemisphere: Morphological, molecular and toxicological characterization

Pseudo‐nitzschia H. Peragallo is a marine diatom genus found worldwide in polar, temperate, subtropical and tropical waters. It includes toxigenic representatives that produce domoic acid (DA), a neurotoxin responsible for Amnesic Shellfish Poisoning. In this study we characterized two species of Pseudo‐nitzschia collected from Port Stephens and the Hawkesbury River (south eastern Australia) previously unreported from Australian waters. Clonal isolates were sub‐sampled for (i) light and transmission electron microscopy; (ii) DNA sequencing, based on the nuclear‐encoded partial large subunit ribosomal RNA gene and internal transcribed spacer (ITS)‐ITS1, 5.8S and ITS2 rDNA regions and, (iii) DA production as measured by liquid chromatography‐mass spectrometry. Morphological and molecular data unambiguously revealed the species to be Pseudo‐nitzschia micropora Priisholm, Moestrup & Lundholm (Port Stephens) and Pseudo‐nitzschia hasleana Lundholm (Hawkesbury River). This is the first report of the occurrence of these species from the Southern Hemisphere and the first report of P. micropora in warm‐temperate waters. Cultures of P. micropora, tested for DA production for the first time, proved to be non‐toxic. Similarly, no detectable toxin concentrations were observed for P. hasleana. Species resolution and knowledge on the toxicity of local Pseudo‐nitzschia species has important implications for harmful algal bloom monitoring and management.

Abundance and richness of key Antarctic seafloor fauna correlates with modelled food availability

Most seafloor communities at depths below the photosynthesis zone rely on food that sinks through the water column. However, the nature and strength of this pelagic–benthic coupling and its influence on the structure and diversity of seafloor communities is unclear, especially around Antarctica where ecological data are sparse. Here we show that the strength of pelagic–benthic coupling along the East Antarctic shelf depends on both physical processes and the types of benthic organisms considered. In an approach based on modelling food availability, we combine remotely sensed sea-surface chlorophyll-a, a regional ocean model and diatom abundances from sediment grabs with particle tracking and show that fluctuating seabed currents are crucial in the redistribution of surface productivity at the seafloor. The estimated availability of suspended food near the seafloor correlates strongly with the abundance of benthic suspension feeders, while the deposition of food particles correlates with decreasing suspension feeder richness and more abundant deposit feeders. The modelling framework, which can be modified for other regions, has broad applications in conservation and management, as it enables spatial predictions of key components of seafloor biodiversity over vast regions around Antarctica.Combining data on sea-surface chlorophyll-a with a regional ocean model and diatom abundance from sediment grabs, the authors determine the strength of pelagic–benthic coupling across the George V region in East Antarctica.

First observations of living sea-ice diatom agglomeration to tintinnid loricae in East Antarctica

Tintinnid ciliates are an important link in marine food webs as they feed on phytoplankton and bacteria while providing nutrients to higher trophic levels. Tintinnids are known to agglutinate mineral particles or dead biogenic material such as diatom frustules to their shell-like housing (lorica), however, reasons for this agglutination remain questioned. We report on our observation of agglomeration of the living diatoms Fragilariopsis curta , F. cylindrus, F. pseudonana and F. rhombica to loricae of the Antarctic tintinnid ciliates Laackmanniella naviculaefera and Codonellopsis gaussi. These unusual associations between living diatoms and tintinnids were exclusively observed south of 63.59°S. We discuss the significance of our new finding and generate hypotheses to be tested by future research. It remains unclear where these living diatom–tintinnid associations are initially formed (in or near sea ice or also further north when abundances of L. naviculaefera, C. gaussi , F. curta , F. cylindrus , F. pseudonana and F. rhombica happen to be relatively high); who the beneficiary is in this association; what the exact benefits are; and how they might influence the Southern Ocean carbon cycle. Nevertheless, our observation provides a key step forward towards illuminating the largely unknown ecology of two Southern Ocean-endemic tintinnid species.