Regional and global impact of CO2 uptake in the Benguela Upwelling System through preformed nutrients.

Eastern Boundary Upwelling Systems (EBUS) are highly productive ecosystems. However, being poorly sampled and represented in global models, their role as atmospheric CO2 sources and sinks remains elusive. In this work, we present a compilation of shipboard measurements over the past two decades from the Benguela Upwelling System (BUS) in the southeast Atlantic Ocean. Here, the warming effect of upwelled waters increases CO2 partial pressure (pCO2) and outgassing in the entire system, but is exceeded in the south through biologically-mediated CO2 uptake through biologically unused, so-called preformed nutrients supplied from the Southern Ocean. Vice versa, inefficient nutrient utilization leads to preformed nutrient formation, increasing pCO2 and counteracting human-induced CO2 invasion in the Southern Ocean. However, preformed nutrient utilization in the BUS compensates with ~22-75 Tg C year-1 for 20-68% of estimated natural CO2 outgassing in the Southern Ocean's Atlantic sector (~ 110 Tg C year-1), implying the need to better resolve global change impacts on the BUS to understand the ocean's role as future sink for anthropogenic CO2.

Harmful algal blooms of the Benguela eastern boundary upwelling system.

The Benguela Upwelling System (BUS) is subject to a high incidence of HABs. Of the major shellfish poisoning syndromes associated with HABs, Paralytic and Diarrhetic Shellfish Poisoning (PSP and DSP) pose the greatest concern, but as documented herein there are several other HAB organisms that are also present. Blooms of Alexandrium catenella have been recognised as the typical cause of PSP since 1948. In addition to the risk posed to human health A. catenella has also been the cause of large shellfish and bird mortalities. An additional risk of PSP is provided by Alexandrium minutum first detected in Cape Town harbour in 2003. DSP was identified on the South African coast for the first time in 1991. Although several Dinophysis spp. known to cause DSP have been recognized as a component of the plankton of the region, it is accepted that DSP is usually attributed to D. acuminata or D. fortii. In the southern Benguela both Pseudo-nitzschia australis and Pseudo-nitzschia multiseries have been identified and shown to produce domoic acid. Multiple Pseudo-nitzschia spp. have been identified in the northern Benguela with the potentially toxigenic Pseudo-nitzschia pungens and P. australis dominant inshore. The yessotoxin (YTX) producing dinoflagellates Gonyaulax spinifera, Lingulodinium polyedrum and Protoceratium reticulatum are all known to form blooms and YTXs have been the cause of massive mortalities of farmed abalone. Prominent fish-killing blooms include Karlodinium veneficum in the northern Benguela and Karenia cristata in the southern Benguela. Shellfish farms in an embayment of the southern Benguela have suffered reduced growth rates due to the ecosystem disruptive blooms of Aureococcus anophagefferens. High biomass dinoflagellate blooms often attributed to Tripos and Prorocentrum spp. characterise the entire region and major mortalities of marine life are regularly attributed to their decay and the subsequent development of anoxic conditions.

Giant sulfur bacteria (Beggiatoaceae) from sediments underlying the Benguela upwelling system host diverse microbiomes.

Due to their lithotrophic metabolisms, morphological complexity and conspicuous appearance, members of the Beggiatoaceae have been extensively studied for more than 100 years. These bacteria are known to be primarily sulfur-oxidizing autotrophs that commonly occur in dense mats at redox interfaces. Their large size and the presence of a mucous sheath allows these cells to serve as sites of attachment for communities of other microorganisms. But little is known about their individual niche preferences and attached microbiomes, particularly in marine environments, due to a paucity of cultivars and their prevalence in habitats that are difficult to access and study. Therefore, in this study, we compare Beggiatoaceae strain composition, community composition, and geochemical profiles collected from sulfidic sediments at four marine stations off the coast of Namibia. To elucidate community members that were directly attached and enriched in both filamentous Beggiatoaceae, namely Ca. Marithioploca spp. and Ca. Maribeggiatoa spp., as well as non-filamentous Beggiatoaceae, Ca. Thiomargarita spp., the Beggiatoaceae were pooled by morphotype for community analysis. The Beggiatoaceae samples collected from a highly sulfidic site were enriched in strains of sulfur-oxidizing Campylobacterota, that may promote a more hospitable setting for the Beggiatoaceae, which are known to have a lower tolerance for high sulfide to oxygen ratios. We found just a few host-specific associations with the motile filamentous morphotypes. Conversely, we detected 123 host specific enrichments with non-motile chain forming Beggiatoaceae. Potential metabolisms of the enriched strains include fermentation of host sheath material, syntrophic exchange of H2 and acetate, inorganic sulfur metabolism, and nitrite oxidation. Surprisingly, we did not detect any enrichments of anaerobic ammonium oxidizing bacteria as previously suggested and postulate that less well-studied anaerobic ammonium oxidation pathways may be occurring instead.

Tracing the upwelling process in the northern Benguela upwelling system (nBUS) by 129I.

New data on the presence of 129I in seawater in the Southern Hemisphere measured by Accelerator Mass Spectrometry (AMS) is presented. The samples were collected in 2014 along the Namibian coast during a cruise organised by the National Marine Information and Research Centre (NatMIRC), the national laboratories of the Ministry of Fisheries and Marine Resources (MFMR) in Namibia, and the IAEA Environment Laboratories (IAEA NAEL) in Monaco. The Benguela upwelling system is known as one of the most important marine upwelling regions in the world. Strong winds induce an offshore transport of surface seawater which is substituted by cool subsurface water inshore. As this water is nutrient-rich, which leads to high primary productivity, the Benguela upwelling system has a very important role as a fishing production area. The 129I concentrations in samples were between (0.66 ± 0.14) × 107 and (1.45 ± 0.30) × 107 atoms/kg. The highest 129I concentrations were found in the offshore surface samples. Deep-sea and inshore samples contained lower 129I concentrations, possibly as an effect of the upwelling process. A comparison with previously published studies suggests that the presence of 129I in the northern Benguela upwelling system (nBUS), is mainly due to the impact of nuclear weapons global fallout, without any evident impact of nuclear fuel reprocessing.

Presence of 236U and 237Np in a marine ecosystem: The northern Benguela Upwelling System, a case study.

The Benguela Upwelling System (BUS), off the south-western African coast, is one of the four major eastern boundary upwelling ecosystems in the oceans. However, this area has been overlooked in the field of environmental radioactivity. In this work, 236U and 237Np were collected off the coast of Namibia within the northern BUS. Surface seawater exhibited similar 236U and 237Np concentrations, ranging from 3.9·106 to 5.6·106 atoms kg-1 and from 4.6·106 to 8.5·106 atoms kg-1, respectively. The observed inventories in a water column from the continental margin, of (2.10 ± 0.11)·1012 atoms m-2 for 236U and (3.48 ± 0.13)·1012 atoms m-2 for 237Np, were in agreement with the global fallout (GF) source term in the Southern Hemisphere that was the main source of actinides to the region. A pattern was observed in the surface samples, with 237Np concentrations that decreased by 25-30% when moving from inshore to offshore stations, but such an effect could not be clearly discerned in the case of 236U within the data uncertainties. An explanation based on the larger particle reactivity of GF 237Np compared to GF 236U was proposed. Such an effect would have been important at the studied site due to the enhance presence of particles in the continental shelf triggered by the upwelling phenomenon. A value of 1.77 ± 0.20 was obtained for the 237Np/236U atom ratio for the GF source term in the marine environment.

Microbial communities associated with phosphogenic sediments and phosphoclast-associated DNA of the Benguela upwelling system.

The processes that lead to the precipitation of authigenic calcium phosphate minerals in certain marine pore waters remain poorly understood. Phosphogenesis occurs in sediments beneath some oceanic upwelling zones that harbor polyphosphate-accumulating bacteria. These bacteria are believed to concentrate phosphate in sediment pore waters, creating supersaturated conditions with respect to apatite precursors. However, the relationship between microbes and phosphorite formation is not fully resolved. To further study this association, we examined microbial community data generated from two sources: sediment cores recovered from the shelf of the Benguela upwelling region where phosphorites are currently forming, and DNA preserved within phosphoclasts recovered from a phosphorite deposit along the Benguela shelf. iTag and clone library sequencing of the 16S rRNA gene showed that many of our sediment-hosted communities shared large numbers of phylotypes with one another, and that the same metabolic guilds were represented at localities across the shelf. Sulfate-reducing bacteria and sulfur-oxidizing bacteria were particularly abundant in our datasets, as were phylotypes that are known to carry out nitrification and the anaerobic oxidation of ammonium. The DNA extracted from phosphoclasts contained the signature of a distinct microbial community from those observed in the modern sediments. While some aspects of the modern and phosphoclast communities were similar, we observed both an enrichment of certain common microbial classes found in the modern phosphogenic sediments and a relative depletion of others. The phosphoclast-associated DNA could represent a relict signature of one or more microbial assemblages that were present when the apatite or its precursors precipitated. While these taxa may or may not have contributed to the precipitation of the apatite that now hosts their genetic remains, several groups represented in the phosphoclast extract dataset have the genetic potential to metabolize polyphosphate, and perhaps modulate phosphate concentrations in pore waters where carbonate fluorapatite (or its precursors) are known to be precipitating.

Morphology and toxicity of Pseudo-nitzschia species in the northern Benguela Upwelling System.

The Benguela upwelling system, considered the world's most productive marine ecosystem, has a long record of potentially toxic diatoms belonging to the genus Pseudo-nitzschia. Species of Pseudo-nitzschia were reported as early as 1936 from the northern Benguela upwelling system (nBUS). For the current study, long-term phytoplankton monitoring data (2004-2011) for the Namibian coast were analysed to examine inshore and offshore temporal distribution of Pseudo-nitzschia species, their diversity and ultrastructure. The potentially toxigenic P. pungens and P. australis were the dominant inshore species, whereas offshore Pseudo-nitzschia showed a higher diversity that also included potentially toxic species. During a warming event, a community shift from P. pungens and P. australis dominance to P. fraudulenta and P. multiseries was documented in the central nBUS. A case study of a toxic event (August 2004) revealed that P. australis and P. pungens were present at multiple inshore and offshore stations, coincident with fish (pilchard) and bird mortalities reported from the central part of Namibia. Toxin analyses (LC-MS/MS) of samples collected from June to August 2004 revealed the presence of particulate domoic acid (DA) in seawater at multiple stations (maximum ∼180 ng DA/L) in the >0.45 µm size-fraction, as well as detectable DA (0.12 µg DA/g) in the gut of one of two pilchard samples tested. These findings indicate that DA may have been associated with the fish and bird mortalities reported from this event in the nBUS. However, the co-occurrence of very high biomass phytoplankton blooms suggests that other explanations may be possible.

Morphological, molecular and toxigenic characteristics of Namibian Pseudo-nitzschia species - including Pseudo-nitzschia bucculenta sp. nov.

A field study was undertaken to investigate the occurrence and toxin production of species in the diatom genus Pseudo-nitzschia in Namibian waters, in the extremely productive Benguela upwelling system. From surveys conducted on the R/V Mirabilis and the R/V !Anichab, 52 strains were morphologically determined to species level, supported by nuclear ITS rDNA data. Seven species were identified; P. australis, P. decipiens, P. dolorosa, P. fraudulenta, P. plurisecta, P. pungens var. cingulata, and the new species P. bucculenta F. Gai, C. K. Hedemand, N. Lundholm & Ø. Moestrup sp. nov. Molecular and morphological diversity of the Namibian Pseudo-nitzschia species is discussed. Most importantly, P. bucculenta is both morphologically and phylogenetically most similar to P. dolorosa differing mainly in valve width and densities of striae, poroids and band striae as well as by four hemi-compensatory base changes in the ITS2. Morphological and molecular differences among the strains of P. decipiens suggest a temperate and a warm water subdivision. The geographical and toxigenic characteristics of the identified Pseudo-nitzschia species are described and compared to previous studies. Initial tests of toxin production in all seven species revealed production of domoic acid (DA) in two species: one strain of P. australis (0.074 pg DA cell-1) and two strains of P. plurisecta (0.338 pg DA cell-1 and 0.385 pg DA cell-1).

Spatio-temporal variability of copepod abundance along the 20 °S monitoring transect in the Northern Benguela upwelling system from 2005 to 2011.

Long-term data sets are essential to understand climate-induced variability in marine ecosystems. This study provides the first comprehensive analysis of longer-term temporal and spatial variations in zooplankton abundance and copepod community structure in the northern Benguela upwelling system from 2005 to 2011. Samples were collected from the upper 200 m along a transect at 20 °S perpendicular to the coast of Namibia to 70 nm offshore. Based on seasonal and interannual trends in surface temperature and salinity, three distinct time periods were discernible with stronger upwelling in spring and extensive warm-water intrusions in late summer, thus, high temperature amplitudes, in the years 2005/06 and 2010/11, and less intensive upwelling followed by weaker warm-water intrusions from 2008/09 to 2009/10. Zooplankton abundance reflected these changes with higher numbers in 2005/06 and 2010/11. In contrast, zooplankton density was lower in 2008/09 and 2009/10, when temperature gradients from spring to late summer were less pronounced. Spatially, copepod abundance tended to be highest between 30 and 60 nautical miles off the coast, coinciding with the shelf break and continental slope. The dominant larger calanoid copepods were Calanoides carinatus, Metridia lucens and Nannocalanus minor. On all three scales studied, i.e. spatially from the coast to offshore waters as well as temporally, both seasonally and interannually, maximum zooplankton abundance was not coupled to the coldest temperature regime, and hence strongest upwelling intensity. Pronounced temperature amplitudes, and therefore strong gradients within a year, were apparently important and resulted in higher zooplankton abundance.