Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Environmental influence on phytoplankton communities in the northern Benguela ecosystem

An investigation of surface phytoplankton communities was undertaken on the shelf of the northern Benguela upwelling ecosystem during austral autumn (May) and spring (September), along latitudinal transects at 20° S and 23° S, from 2 to 70 nautical miles offshore, as well as on a zigzag grid located between these transects. Microscopic identification of the phytoplankton and CHEMTAX analysis of pigment biomarkers were used to characterise the community composition. During May 2014, warmer, more-saline water with a shallower upper mixed layer corresponding to periods of less-intense offshore Ekman transport was encountered on the shelf. Satellite imagery indicated high phytoplankton biomass extending for a considerable distance from the coast, and CHEMTAX indicated diatoms as dominant at most of the stations (52–92%), although dinoflagellates were dominant at some inshore localities (57–74%). Species of Chaetoceros, Bacteriastrum and Cylindrotheca were the most abundant, with abundance of the Pseudo-nitzschia ‘seriata-group’ being particularly high at a number of stations. In September 2014, more-intense wind-forcing resulted in a deeper upper mixed layer and stronger upwelling of colder, less-saline water. Elevated phytoplankton biomass was confined close to the coast, where diatoms accounted for most of the population (54–87%), whereas small flagellates, such as prasinophytes, haptophytes and cryptophytes, as well as the cyanobacterium Synechococcus, dominated the communities (58–90%) farther from the coast. It is hypothesised that stronger upwelling and deeper vertical mixing in September of that year were not conducive for widespread diatom growth, and that small flagellates populated the water column by being entrained from offshore onto the shelf in the upwelled water that moved in towards the coast.     

Inferring upwelling rates in the equatorial Atlantic using Be measurements in the upper ocean

Ocean upwelling rates are difficult to measure because of the relatively small velocities involved, and therefore are typically inferred from indirect methods such as heat budget estimates or tracer observations. Here we present the first results using a novel technique, based on the isotope ⁷Be, to infer rates of upwelling along the equator. Beryllium-7 (half-life=53.3 d) is a cosmic-ray produced radioactive nuclide that is deposited by rainfall upon the ocean surface and subsequently enriched and homogenized within the mixed layer. Previous investigations have utilized the penetration of characteristically high mixed layer concentrations into the upper thermocline to trace ocean ventilation and subduction over seasonal timescales. Here, the tracer is used in a reverse sense; that is, the ⁷Be concentration in the usually ⁷Be-rich surface mixed layer will be diluted from penetration of ⁷Be “dead” water upwelled from below. This dilution provides a means to infer upwelling rates. Furthermore, with knowledge of upwelling rates, ⁷Be profiles can be used to constrain vertical diffusivity within the upper thermocline. These ideas were tested with samples collected during the Tropical Atlantic Climate Experiment (TACE) cruise (May 22-June 27, 2009). The observations indicated a nearly linear relationship between ⁷Be inventory and mixed layer temperature, as with increased upwelling, lower mixed layer temperatures correspond to greater ⁷Be dilution from depth. With this data, upwelling rates were estimated at a number of stations near the equator between 0°E and 30°W within and adjacent to the equatorial cold tongue. The derived upwelling rates ranged from 0 to 2.2 m/d, with maximum values found between the equator and 2°S. The corresponding K_z values derived for the upper thermocline were in the range 1-4×10⁻⁴ m²/s.     

Dissolved Iodate and Total Iodine During an Extreme Hypoxic Event in the Southern Benguela System

Dissolved iodate and total iodine were studied in St Helena Bay, South Africa, during a period of acute hypoxia, following upwelling off Cape Columbine. Despite the generally high concentrations of chlorophyll α (10–30 mg m⁻³) total iodine concentration was essentially constant in the main part of the Bay, and similar to that found elsewhere in the oceans. Occasional, lower concentrations of total iodine (0·28 to 0·42 μM) were found with exceptionally high chlorophyll α concentrations (500 mg m⁻³) in shallow waters. In contrast, iodate was found to be reduced to iodide at both the surface and the bottom of the Bay. The implications of these changes are discussed, given that the surface waters reflect sustained eutrophication while the bottom waters are hypoxic as a result of the organic-rich sediment from the waters above.     

Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela

A regime shift is considered to be a sudden shift in structure and functioning of a marine ecosystem, affecting several living components and resulting in an alternate state. According to this definition, regime shifts differ from species replacement or alternation of species at similar trophic levels, whereby the ecosystem is not necessarily significantly altered in terms of its structure and function; only its species composition changes. This paper provides an overview of regime shifts, species replacements and alternations that have been observed in the northern and southern Benguela ecosystems over the past few decades. Bottom-up control, initiating and sustaining regime shifts or species replacements via environmental forcing, is documented for both the southern and the northern Benguela ecosystems. Fishing (a case of top-down control) appears to have played an important role in regime shift processes in the Namibian ecosystem. Very low biomass levels of exploited fish stocks associated with less efficient energy transfer in the northern Benguela are indicative of a regime shift. Very high biomass levels have been reached in the southern Benguela in the 2000s. However the alternation between sardine and anchovy that has been observed in the southern Benguela over the last two decades appears not to have had major effects on the overall functioning of the ecosystem. The consequences of regime shifts for exploitation are highlighted, suggesting that fisheries managers should move towards a more effective ecosystem approach to fisheries.     

The reduction of plankton biomass induced by mesoscale stirring: A modeling study in the Benguela upwelling

Recent studies, both based on remote sensed data and coupled models, showed a reduction of biological productivity due to vigorous horizontal stirring in upwelling areas. In order to better understand this phenomenon, we consider a system of oceanic flow from the Benguela area coupled with a simple biogeochemical model of Nutrient-Phyto-Zooplankton (NPZ) type. For the flow three different surface velocity fields are considered: one derived from satellite altimetry data, and the other two from a regional numerical model at two different spatial resolutions. We compute horizontal particle dispersion in terms of Lyapunov exponents, and analyzed their correlations with phytoplankton concentrations. Our modeling approach confirms that in the south Benguela there is a reduction of biological activity when stirring is increased. Two-dimensional offshore advection and latitudinal difference in primary production, also mediated by the flow, seem to be the dominant processes involved. We estimate that mesoscale processes are responsible for 30–50% of the offshore fluxes of biological tracers. In the northern area, other factors not taken into account in our simulation are influencing the ecosystem. We suggest explanations for these results in the context of studies performed in other eastern boundary upwelling areas.     

Accumulation of diarrhetic shellfish poisoning toxins in the oyster Crassostrea gigas and the mussel Choromytilus meridionalis in the southern Benguela ecosystem

Diarrhetic shellfish poisoning (DSP) poses a significant threat to the safe consumption of shellfish in the southern Benguela ecosystem. The accumulation of DSP toxins was investigated in two cultivated bivalve species, the Pacific oyster Crassostrea gigas and the mussel Choromytilus meridio-nalis, suspended from a mooring located off Lambert's Bay on the west coast of South Africa. The dinoflagellate Dinophysis acuminata, a known source of polyether toxins associated with DSP, was common through most of the study period. The toxin composition of the dinoflagellate was dominated by okadaic acid (OA) (91%), with lesser quantities of the dinophysistoxin DTX-1 (6.5%) and pecteno-toxin PTX-2 (2.4%), and traces of PTX-2sa and PTX-11. The mean cell toxin quota of D. acuminata was 7.8 pg OA cell^–1. The toxin profile in shellfish was characterised by a notably higher relative content of DTX-1. The study showed the average concentration of DSP toxins in the mussels to exceed that in the oysters by approximately 20-fold. The results indicate a need to establish species-specific sampling frequencies in shellfish safety monitoring programmes.     

A generalized coordinate ocean model and a comparison of the bottom boundary layer dynamics in terrain-following and in z-level grids

Sensitivity studies with a new generalized coordinate ocean model are performed in order to compare the behavior of bottom boundary layers (BBLs) when terrain-following (sigma or combined sigma and z-level) or z-level vertical grids are used, but most other numerical aspects remain unchanged. The model uses a second-order turbulence closure scheme that provides surface and BBL mixing and results in a quite realistic climatology and deep water masses after 100 year simulations with a coarse resolution (1° × 1°) basin-scale terrain-following grid. However, with the same turbulence scheme but using a z-level grid, the model was unable to produce dense water masses in the deep ocean. The latter is a known problem for coarse resolution z-level models, unless they include highly empirical BBL schemes.A set of dense water overflow experiments with high-resolution grids (10 and 2.5 km) are used to investigate the influence of model parameters such as horizontal diffusivity, vertical mixing, horizontal resolution, and vertical resolution on the simulation of bottom layers for the different coordinate systems. Increasing horizontal diffusivity causes a thinner BBL and a bottom plume that extends further downslope in a sigma grid, but causes a thicker BBL and limited downslope plume extension in a z-level grid. A major difference in the behavior of the BBL in the two grids is due to the larger vertical mixing generated by the turbulence scheme over the step-like topography in the z-level grid, compared to a smaller vertical mixing and a more stably stratified BBL in the sigma grid. Therefore, the dense plume is able to maintain its water mass better and penetrates farther downslope in the sigma grid than in the z-level grid. Increasing horizontal and vertical resolution in the z-level grid converges the results toward those obtained by a much coarser resolution sigma coordinate grid, but some differences remain due to the basic differences in the mixing process in the BBL.     

Physical and biological features across an upwelling front in the southern Benguela

The upwelling front of the Cape Columbine upwelling centre was intensively studied, physically and biologically, along a repeated transect during December 1984 following a quiescent phase in the upwelling cycle. Three distinct zones were evident, an inshore zone influenced by upwelling, an offshore warm oligotrophic zone and a transitional frontal zone separating the two. Salinity proved to be a useful indicator of recent water movements. There was evidence of intrusions and mixing of water types within the frontal zone, possibly accounting for the elevated phytoplankton biomass recorded there. Floral and faunal changes occurred between the frontal and offshore zones, corresponding to the thermal front. The predominant flow was alongshore, with strong equatorward jet currents, making the interpretation of cross-shelf gradients difficult in this dynamic area. Aspects of the distributions of organisms and their productivity across the upwelling front are described with respect to the hydrographic parameters and associated flow-field.     

Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.     

Thermal Stratification in Saldanha Bay (South Africa) and Subtidal, Density-driven Exchange with the Coastal Waters of the Benguela Upwelling System

A study was conducted to understand the mechanisms driving observed subtidal variability in the stratification of Saldanha Bay, located in the southern Benguela system. It was found that the 6–8 day period variability in bay stratification was caused by the inflow and outflow of cold upwelled water driven by changing baroclinic pressure gradients between the coastal and bay domains. The direction and magnitude of the pressure gradients were governed by coastal upwelling activity and a lag in the response of the bay to changes in density structure in the coastal ocean. When the pressure gradients were bayward and cold water was being driven into the bay the cycle was termed to be in an ‘ active phase ’ and the reverse was termed the ‘ relaxation phase ’. The upwelling-favourable equatorward wind stress impacted the bay stratification in two ways: on the regional scale, wind drives upwelling and governs the inflow–outflow of cold upwelled bottom water, which strengthens stratification; conversely, on the local bay scale, wind drives vertical mixing, which weakens stratification. A four-phase model is used to describe the observed variability in stratification in the bay. The associated density-driven exchange flows are capable of flushing the bay in 6–8 days, about one-third of the time for tidal exchange alone (c. 25 days). These inflows of cold bottom water are ecologically critical as they supply nutrients to the bay and thus impose a control on new production within the bay environment. Further ecological implications of this bay–ocean exchange include export of phytoplankton new production to the coast, limitation of the risk of harmful algal blooms (HABs) and the division of the system into two distinct ecosystems (bay and lagoon).     

The cold-core eddy and strong upwelling off the coast of New South Wales in early 2007

During the Austral summer of 2006-07 a series of extreme oceanic events occurred in the Tasman Sea. Following a series of strong wind-driven upwelling events, an intense cold-core eddy developed off Sydney, Australia. A data-assimilating, eddy-resolving ocean model is used to create a three-dimensional time-varying reanalysis of these events. The reanalysis indicates that the cold anomalies associated with the upwellings were in excess of −5 °C near the coast, where sea level decreased by as much as 0.2 m. The reanalysed three-dimensional structure of the cold-core eddy shows the eddy "leaning" to the west-north-west, in towards the continental shelf. The diameter of the eddy is about 100 km and the sea-level anomaly at the eddy centre peaks at around −1 m, with an associated sub-surface temperature anomaly in excess of −8 °C at 200 m depth, corresponding to an upward isotherm excursion of 600 m. The circulation around the cyclonic eddy is ageostrophic, with upwelling in the southern sector of the eddy (where flow is onshore and climbing the continental slope) and downwelling in the northern sector (where flow is descending off the slope). Three-dimensional trajectories of water parcels around the eddy involve 50-100-m vertical excursions. Based on the reanalysed circulation and composite satellite images of Chlorophyll-a, we hypothesise that the circulation around the eddy led to significant nutrient enrichment in the euphotic zone around the perimeter of the eddy.     

Spatio‐temporal variability of zooplankton biomass and environmental control in the Northern Benguela Upwelling System: field investigations and model simulation

Spatial and temporal distribution patterns of zooplankton are highly variable in the Northern Benguela Upwelling System. We studied the distribution of zooplankton (size class ≥ 0.33 mm) and used field data from four cruises that took place between March 2008 and February 2011, as well as simulation results of a regional ecosystem model. Remotely sensed sea surface temperatures (SST) and surface chlorophyll concentrations were analysed to investigate environmental influences on zooplankton biomass. The Intense Benguela Upwelling Index showed a distinct seasonal signal throughout the years and the highest upwelling peaks in August/September. Even though surface chlorophyll concentrations were very variable throughout the year, the highest concentrations were always detected in September, following the upwelling of nutrient‐rich water. In field catches, zooplankton biomass concentration in the upper 200 m was highest above the outer shelf and shelf‐break in December 2010 and February 2011, i.e. 6 months after the upwelling peaks. In contrast, zooplankton biomass simulated by the model in the surface water was highest in September. In March/April, biomass maxima were typically measured in the field at intermediate water depths, but the vertical distribution was also affected by extensive oxygen minimum zones. The ecosystem model reproduced this vertical pattern. Although general trends were similar, simulation data of zooplankton standing stocks overestimated the field data by a factor of 3. In upwelling systems, food webs are generally considered to be short and dominated by large cells. However, our field data indicate more small‐sized zooplankton organisms above the shelf than offshore.     

Contributions of shortwave radiation to the formation of temperature inversions in the Bay of Bengal and eastern equatorial Indian Ocean: A modeling approach

Variations in incoming shortwave radiation influence the net surface heat flux, contributing to the formation of a temperature inversion. The effects of shortwave radiation on the temperature inversions in the Bay of Bengal and eastern equatorial Indian Ocean have never been investigated. Thus, a high-resolution (horizontal resolution of 0.07°×0.07° with 50 vertical layers) Regional Ocean Modeling System (ROMS) model is utilized to quantify the contributions of shortwave radiation to the temperature inversions in the study domain. Analyses of the mixed layer heat and salt budgets are performed, and different model simulations are compared. The model results suggest that a 30% change in shortwave radiation can change approximately 3% of the temperature inversion area in the Bay of Bengal. Low shortwave radiation reduces the net surface heat flux and cools the mixed layer substantially; it also reduces the evaporation rate, causing less evaporative water vapor losses from the ocean than the typical situation, and ultimately enhances haline stratification. Thus, the rudimentary outcome of this research is that a decrease in shortwave radiation produces more temperature inversion in the study region, which is primarily driven by the net surface cooling and supported by the intensive haline stratification. Moreover, low shortwave radiation eventually intensifies the temperature inversion layer by thickening the barrier layer. This study could be an important reference for predicting how the Indian Ocean climate will respond to future changes in shortwave radiation.     

北太平洋西边界流研究综述

综述了近20年来国内外学者在研究北太平洋西边界流的平均结构及NEC分叉动力机制、NM K流系平均输运的分配及变化、NM K流系季节及年际变化规律及其与EN SO之间的关系、NM K流系在热带和亚热带水交换中的作用以及水团的平均分布特征等方面所取得的主要成果。通过分析,发现东亚季风、R ossby波和K e lv in波等是影响北太平洋西边界流的主要因素;而缺乏长期直接的海流观测资料是深入研究北太平洋西边界流遇到的最大障碍。